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Zhaolong Wang, Bingru Huang and Qingzhang Xu

Abscisic acid (ABA) is an important hormone regulating plant response to drought stress. The objective of this study was to investigate effects of exogenous ABA application on turf performance and physiological activities of kentucky bluegrass (Poa pratensis L.) in response to drought stress. Plants of two kentucky bluegrass cultivars, `Brilliant' (drought susceptible) and `Midnight' (drought tolerant), were treated with ABA (100 μm) or water by foliar application and then grown under drought stress (no irrigation) or well-watered (irrigation on alternate days) conditions in a growth chamber. The two cultivars responded similarly to ABA application under both watering regimes. Foliar application of ABA had no effects on turf quality or physiological parameters under well-watered conditions. ABA application, however, helped maintain higher turf quality and delayed the quality decline during drought stress, compared to the untreated control. ABA-treated plants exposed to drought stress had higher cell membrane stability, as indicated by less electrolyte leakage of leaves, and higher photochemical efficiency, expressed as Fv/Fm, compared to untreated plants. Leaf water potential was not significantly affected, whereas leaf turgor pressure increased with ABA application after 9 and 12 d of drought. Osmotic adjustment increased with ABA application, and was sustained for a longer period of drought in `Midnight' than in `Brilliant'. The results suggested that exogenous ABA application improved turf performance during drought in both drought-sensitive and tolerant cultivars of kentucky bluegrass. This positive effect of ABA could be related to increased osmotic adjustment, cell turgor maintenance, and reduced damage to cell membranes and the photosynthetic system.

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Qingzhang Xu, Bingru Huang and Zhaolong Wang

High air and soil temperatures are major factors limiting growth of cool-season grasses. A previous study by the authors reported that a soil temperature reduction of only 3 °C when air temperature was maintained at 35 °C significantly improved shoot and root growth of creeping bentgrass [Agrostis stolonifera L. var. palustris (Huds.) Farw. (syn. A. palustris Huds.)]. This study was designed to investigate the responses of photosynthetic activities of creeping bentgrass to lowered root-zone temperatures from the supraoptimal level when shoots were exposed to high air temperature. Two cultivars of creeping bentgrass, `L-93' and `Penncross', were exposed to the following air/root-zone temperature regimes in growth chambers and water baths: 1) optimal air and soil temperatures (20/20 °C, control); 2) lowering soil temperature by 3, 6, and 11 °C from 35 °C at high air temperatures (35/32, 35/29, and 35/24 °C); and 3) high air and soil temperatures (35/35 °C). Soil temperature was reduced from 35 °C by circulating cool water (18 °C) in water baths at variable flow rates. Both cultivars had similar responses to high or low root-zone temperatures with high air temperature. High air and root-zone temperatures caused significant reductions in canopy photosynthetic rate (Pcanopy), single-leaf photosynthetic rate (Pleaf), leaf chlorophyll content, photochemical efficiency (Fv/Fm), and ribulose-1,5-bisphosphate carboxylase/oxygenase (Rubisco) activity, beginning on day 1 of high air and soil temperature stress for Pcanopy and Pleaf, and day 7 for chlorophyll content, Fv/Fm, and Rubisco activity. The 3 °C reduction in root-zone temperature at high air temperature had no effect on those photosynthetic parameters, except chlorophyll content. Reducing root-zone temperature by 6 °C or 11 °C while maintaining air temperature at 35 °C significantly improved Pcanopy, Poleaf, leaf chlorophyll content, Fv/Fm, and Rubisco activity. Single leaf photosynthetic rate at 35/24 °C was not different from the control level, but Pcanopy at 35/24 °C was lower than the control level. A reduction in root-zone temperature enhanced canopy and single-leaf photosynthetic capacity even though shoots were exposed to supraoptimal air temperature, which could contribute to improved turfgrass growth.

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Hongmei Du, Zhaolong Wang, Wenjuan Yu and Bingru Huang

The accumulation of different types of metabolites may reflect variations in plant adaptation to different severities or durations of drought stress. The objectives of this project are to examine changes in metabolomic profiles and determine predominant metabolites in response to short-term (6 days) and long-term (18 days) drought stress with gas chromatography–mass spectrometry analysis in a C4 perennial grass species. Plants of hybrid bermudagrass (Cynodon dactylon × C. transvaalensis cv. Tifdwarf) were unirrigated for 18 days to induce drought stress in growth chambers. Physiological responses to drought stress were evaluated by visual rating of grass quality, relative water content, photochemical efficiency, and electrolyte leakage (EL). All parameters decreased significantly at 6 and 18 days of drought stress, except EL, which increased with the duration of drought stress. Under short-term drought stress (6 days), the content did not change significantly for most metabolites, except methionine, serine, γ-aminobutyric acid (GABA), isoleucine, and mannose. Most metabolites showed higher accumulation under long-term drought stress compared with that under the well-watered conditions, including three organic acids (malic acid, galacturonic acid, and succinic acid), 10 amino acids (proline, asparagine, phenylalanine, methionine, serine, 5-hydroxynorvaline, GABA, glycine, theorine, valine), seven sugars (sucrose, glucose, galactose, fructose, mannose, maltose, xylose), one nitrogen compound (ethanolamine), and two-sugar alcohol (myo-inositol). The accumulation of those metabolites, especially malic acid, proline, and sucrose, could be associated with drought adaptation of C4 hybrid bermudagrass to long-term or severe drought stress.

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Jingjin Yu, Hongmei Du, Ming Xu and Bingru Huang

Heat is a major factor limiting growth of C3 grass species. Elevated CO2 may mitigate the adverse effects of heat stress or enhance heat tolerance. The objective of this study was to determine metabolic changes associated with improvement of heat tolerance by elevated atmospheric CO2 concentration in tall fescue (Festuca arundinacea). Plants (cv. Rembrandt) were exposed to ambient day/night temperature (25/20 °C) or heat stress (35/30 °C) and ambient CO2 concentration (400 ± 10 μmol·mol−1) or double ambient CO2 concentration (800 ± 10 μmol·mol−1) in growth chambers. Turf quality (TQ), shoot growth rate, and leaf electrolyte leakage results demonstrated that heat stress at ambient CO2 concentration inhibits turf growth and reduces cell membrane stability, whereas heat-stressed plants under elevated CO2 concentration exhibit improved TQ, shoot growth rate, and membrane stability. Plants exposed to heat stress under elevated CO2 exhibited a significantly greater amount of several organic acids (shikimic acid, malonic acid, threonic acid, glyceric acid, galactaric acid, and citric acid), amino acids (serine, valine, and 5-oxoproline), and carbohydrates (sucrose and maltose) compared with heat-stressed plants at ambient CO2. The increased production or maintenance of metabolites with important biological functions such as those involved in photosynthesis, respiration, and protein metabolism could play a role in elevated CO2 mitigation of heat stress damage. Therefore, elevated CO2 conditions may contribute to improved heat stress tolerance as exhibited by better TQ and shoot growth of heat-stressed plants. Practices to harness the power of CO2 may be incorporated into turfgrass management for plant adaptation to increasing temperatures, particularly during summer months.

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Jinpeng Xing, Yan Xu, Jiang Tian, Thomas Gianfagna and Bingru Huang

Cytokinins have been associated with delaying or suppressing leaf senescence in plants. The objectives of this study were to determine whether the expression of the ipt gene that encodes adenine isopentenyltransferase would delay leaf senescence induced by shade or heat stress in a perennial grass species. Creeping bentgrass (Agrostis stolonifera cv. Penncross) was transformed with ipt isolated from agrobacterium (Agrobacterium tumefaciens) using two gene constructs (SAG12-ipt and HSP18-ipt) designed to activate cytokinin synthesis during shade or heat stress. Whole plants of nine SAG12-ipt transgenic lines and the nontransgenic control plants were incubated in darkness at 20 °C for 20 days. Chlorophyll content of all transgenic lines and the control line decreased after dark treatment, but the decline was less pronounced in transgenic lines. All transgenic lines had higher isopentenyladenine (iP/iPA) content than the control line after 20 days of treatment. In six of the transgenic lines, iP/iPA content remained the same or higher after dark treatment. Whole plants of nine HSP18-ipt transgenic lines and the control plants were incubated at 35 °C for 7 days. Chlorophyll and iP/iPA content declined in the control plants, but the nine transgenic lines had a significantly higher concentration of iP/iPA and were able to maintain chlorophyll content at the prestress level. Our results suggest that expression of SAG12-ipt or HSP18-ipt in creeping bentgrass resulted in increases in cytokinin production, which may have led to the delay and suppression of leaf senescence induced by shade or heat stress.

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Zhimin Yang, Jingjin Yu, Emily Merewitz and Bingru Huang

Abscisic acid (ABA) and glycine betaine (GB) may regulate plant responses to drought or salinity stress. The objectives of this controlled-environment study were to determine whether foliar application of ABA or GB improves turf quality under drought or salinity and whether improved stress responses were associated changes in antioxidant metabolism in two C3 turfgrass species, creeping bentgrass (Agrostis stolonifera) and kentucky bluegrass (Poa pratensis). Physiological parameters evaluated included turf quality, leaf relative water content, membrane electrolyte leakage (EL), membrane lipid peroxidation [expressed as malondialdehyde (MDA) content], and activity of superoxide dismutase (SOD), peroxidase (POD), and ascorbate peroxidase (APX). Abscisic acid and GB were both effective in mitigating physiological damage resulting from drought or salinity for both grass species, but effects were more pronounced on kentucky bluegrass. The most notable effects of ABA or GB application were the suppression of EL and MDA accumulation and an increase in APX, POD, and SOD activities after prolonged periods of drought (21 days) or salinity stress (35 days). These results suggest foliar application of ABA or GB may alleviate physiological damage by drought or salinity stress in turfgrass and the maintenance of membrane stability and active antioxidant metabolism could contribute to the positive effects in the stress mitigation effects.

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Yan Zhang, Cuiyue Liang, Yan Xu, Thomas Gianfagna and Bingru Huang

The objective of the study was to determine whether the expression of a cytokinin (CK) biosynthesis gene encoding adenine isopentenyl transferase (ipt) would delay or suppress leaf senescence induced by nitrogen (N) or phosphorus (P) deficiency in a C3 grass species, creeping bentgrass (Agrostis stolonifera). The ipt gene was ligated to a senescence-associated promoter, SAG12, and was transferred into creeping bentgrass using an agrobacterium (Agrobacterium tumefaciens)-mediated transformation technique. Plants from an SAG12-ipt transgenic line (S41) and a null transformant (NT) control line were grown in nutrient solutions with all essential elements or without N (−N) or P (−P) for 21 days. Significant declines in leaf photochemical efficiency (Fv/Fm) and chlorophyll content of mature leaves were detected in NT and SAG12-ipt plants exposed to N or P deficiency. Compared to the NT control line, SAG12-ipt plants had higher levels of Fv/Fm, chlorophyll, and CK contents in leaves, and these differences between the NT control and SAG12-ipt line became more pronounced with treatment duration. The ipt expression was detected in the −P-treated and the −N-treated plants after 21 days, although the level of expression decreased under N or P deficiency. Under −P treatment, root acid phosphatase activity was greater in SAG12-ipt line than in the NT control line. No significant differences in nitrate reductase activity were detected in leaves or roots between the SAG12-ipt and the NT control lines. Our results demonstrated that SAG12-ipt expression suppressed leaf senescence induced by N or P deficiency in a perennial grass species. The suppressing effects on leaf senescence under P deficiency may be related to CK regulation of more efficient use of P in roots of the SAG12-ipt plants.

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Qi Chai, Fang Jin, Emily Merewitz and Bingru Huang

The objective of this study was to determine physiological traits for drought survival and post-drought recovery upon re-watering in two C3 perennial grass species, kentucky bluegrass [KBG (Poa pratensis)] and perennial ryegrass [PRG (Lolium perenne)]. Plants were maintained well watered or exposed to drought stress by withholding irrigation and were then re-watered in a growth chamber. KBG had significantly higher grass quality and leaf photochemical efficiency, and lower electrolyte leakage than PRG during 20 days of drought. After 7 days of re-watering, drought-damaged leaves were rehydrated to the control level in KBG, but could not fully recover in PRG. KBG produced a greater number of new roots, while PRG had more rapid elongation of new roots after 16 days of re-watering. Superior drought tolerance in KBG was associated with osmotic adjustment, higher cell wall elasticity, and lower relative water content at zero turgor. Osmotic adjustment, cell wall elasticity, and cell membrane stability could play important roles in leaf desiccation tolerance and drought survival in perennial grass species. In addition, post-drought recovery of leaf hydration level and physiological activity could be associated with the accumulation of carbohydrates in leaves and rhizomes during drought stress and new root production after re-watering.

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Yali Song, Patrick Burgess, Hairong Han and Bingru Huang

Turfgrass growth and physiological activities are sensitive to temperatures and are affected by mowing height. Increasing temperatures associated with global climate change may limit photosynthetic capacity of established turfgrass stands. The objective of this study was to determine the effects of mowing height on carbon exchange of a turfgrass system and consequential effects on turfgrass growth in response to temperature variations across the growing season in kentucky bluegrass (Poa pratensis cv. Baron) stands. Mature (8 years old) turfgrass was mowed at 7.6 cm [high mowing height (HM)] or 3.8 cm [low mowing height (LM)] during 2012 and 2013. Both LM and HM plots displayed significant decline in turf quality (TQ), shoot biomass, and canopy photosynthetic rate (Pn) with increasing air temperature above 23–24 °C in both years and the decline was more pronounced for LM plots. Turf plots were carbon emitters when total respiration rate of shoots, roots, and soil (Rtotal) exceeded canopy Pn under high temperatures during July–September but maintained net carbon gain during cooler seasons (May and June) due to greater Pn to Rtotal ratio (Pn:Rtotal). Lowering mowing height accelerated carbon loss by reducing canopy Pn, particularly under high temperatures. Our results suggested that whether mature turfgrass stands fix or emit carbon is heavily dependent on interaction between seasonal temperatures and mowing height gauging whole-stand photosynthetic capacity. Furthermore, increasing mowing height during summer months may offset the deleterious effects of high temperature by maintaining positive carbon balance within the turfgrass system.

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Lili Zhuang, Mengxian Liu, Xiuyun Yuan, Zhimin Yang and Bingru Huang

Aquaporin (AQP) proteins serve important roles in regulating water movement across cellular membranes and affect plant responses to drought stress. The objective of this study was to characterize and examine functions of an AQP gene FaPIP2;1, isolated from a drought-tolerant perennial grass species tall fescue (Festuca arundinacea), for involvement in leaf dehydration status during water stress by overexpressing the gene in arabidopsis (Arabidopsis thaliana). FaPIP2;1 had characteristic transmembrane domains and Asn–Pro–Ala motifs and was similar to PIP2;1 in rice (Oryza sativa) and maize (Zea mays). Quantitative real-time reverse transcriptase polymerase chain reaction analysis showed that FaPIP2;1 was upregulated during moderate water stress (hydroponic culture, osmotic potential (ΨS) at −0.47 and −0.78 MPa) and the transcript level decreased as ΨS further decreased. Transgenic arabidopsis plants overexpressing FaPIP2;1 showed greater number of leaves per plant and improved survival rate compared with the wild type (WT) during drought stress. Transgenic plants also maintained higher leaf relative water content (RWC), chlorophyll content (Chl), net photosynthetic rate (Pn), and lower leaf electrolyte leakage (EL) than the WT. However, there was no difference in root length between the transgenic and WT plants following drought stress. The results demonstrated that overexpressing FaPIP2;1 could improve plant tolerance to drought stress by enhancing leaf water status, Chl, and photosynthetic rate, as well as maintaining improved cellular membrane stability relative to the WT plants. FaPIP2;1 may be used as a candidate gene for genetic modification of perennial grasses to develop new drought-tolerant germplasm and cultivars.