Osmotic adjustment (OA) is a major physiological mechanism associated with maintenance of cell turgor in response to dehydration stress. The objectives of this study were to examine changes in capacity for OA in relation to plant tolerance to drought stress for two cool-season turfgrass species, creeping bentgrass (Agrostis stolonifera L.) and velvet bentgrass (A. canina L.), and to determine major solutes contributing to OA in these grass species. Plants of `L-93' creeping bentgrass and `Greenwich' velvet bentgrass were grown in a growth chamber in polyvinyl chloride (PVC) tubes (5 cm diameter, 40 cm high) filled with a 1:3 (v/v) sterilized mixture of sand and sandy loam soil. The experiment consisted of two soil moisture treatments: 1) well-watered control, irrigated three times per week to maintain soil moisture near pot capacity; and 2) drought stress, irrigation completely withheld. Velvet bentgrass exhibited higher drought tolerance compared to creeping bentgrass, as manifested by higher visual turfgrass quality (TQ) and leaf relative water content (RWC) under drought stress. Both creeping bentgrass and velvet bentgrass exhibited OA in response to drought stress; however, velvet bentgrass exhibited 50% to 60% higher magnitude of OA, which could be related to the maintenance of higher leaf RWC and TQ for greater drought duration compared to creeping bentgrass. OA for both creeping bentgrass and velvet bentgrass was associated with accumulation of water soluble carbohydrates during the early period of drought and increases in proline content following prolonged period of drought; however, inorganic ion content (Ca2+ and K+) did not considerably change under drought stress and did not seem to contribute to OA in these species.
Efficient carbon distribution and utilization may enhance drought survival and recovery ability for perennial grasses. The objectives of this study were to examine changes in carbon partitioning and carbohydrate accumulation patterns in shoots and roots of colonial bentgrass (Agrostis capillaris L.), creeping bentgrass (A. stolonifera L.), and velvet bentgrass (A. canina L.) in response to drought and re-watering following drought, and to determine whether species variation in drought tolerance and recuperative potential is related to differences in the patterns of carbon partitioning and accumulation. The experiment consisted of three treatments: 1) well-watered control; 2) drought, irrigation completely withheld for 18 days; and 3) drought recovery, a group of drought-stressed plants were re-watered at the end of the drought treatment (18 days). Drought tolerance and recuperative ability of three species was evaluated by measuring turf quality and leaf relative water content. These parameters indicated that velvet bentgrass was most drought tolerant while colonial bentgrass had highest recuperative ability among the three species. Plants were labeled with 14CO2 to determine carbon partitioning to shoots and roots. Carbohydrate accumulation was assessed by total nonstructural carbohydrate (TNC) content. The proportion of newly photosynthesized 14C partitioned to roots increased at 12 days of drought compared to the pre-stress level, to a greater extent for velvet bentgrass (45%) than for colonial bentgrass (35%) and creeping bentgrass (30%). In general, the proportion of 14C was highest in roots, intermediate in stems, and lowest in leaves at 12 days of drought treatment for all three bentgrass species. As drought duration and severity increased (18 days), 14C partitioning increased more in leaves and stems relative to that in roots for all three species. Stem TNC content was significantly greater for drought-stressed plants of colonial bentgrass and velvet bentgrass compared to their respective well-watered control plants, whereas no differences in stem TNC content were observed between drought-stressed and well-watered creeping bentgrass. Our results suggest that increased carbon partitioning to roots during initial drought stress represented an adaptive response of bentgrass species to short-term drought stress, and increased carbon partitioning and carbohydrate accumulation in stems during prolonged period of drought stress could be beneficial for rapid recovery of turf growth and water status upon re-watering.
Previous investigations identified velvet bentgrass (Agrostis canina L.) as having higher drought resistance among bentgrass species. This study was designed to determine whether species variation in drought resistance for colonial bentgrass (A. capillaris L.), creeping bentgrass (A. stolonifera L.), and velvet bentgrass was associated with differences in antioxidant enzyme levels in response to drought. Plants of ‘Tiger II’ colonial bentgrass, ‘L-93’ creeping bentgrass, and ‘Greenwich’ velvet bentgrass were maintained in a growth chamber under two watering treatments: 1) well-watered control and 2) irrigation completely withheld for 28 d (drought stress). Prolonged drought stress caused oxidative damage in all three bentgrass species as exhibited by a general decline in antioxidant enzyme activities and an increase in lipid peroxidation. Compared among the three species, velvet bentgrass maintained antioxidant enzyme activities for a greater duration of drought treatment compared with both colonial bentgrass and creeping bentgrass. Higher antioxidant enzyme capacity for velvet bentgrass was associated with less lipid peroxidation and higher turf quality, leaf relative water content, and photochemical efficiency for a greater duration of stress compared with colonial bentgrass and creeping bentgrass. These results suggest that bentgrass resistance to drought stress could be associated with higher oxidative scavenging ability, especially for velvet bentgrass.
Abscisic acid (ABA) and cytokinins are two groups of plant hormones that play important roles in regulating plant responses to decreases in soil water availability. The primary objective for this study was to determine whether species variability in drought survival and recovery for colonial bentgrass (Agrostis capillaris L.), creeping bentgrass (A. stolonifera L.), and velvet bentgrass (A. canina L.) were related to changes in ABA and cytokinin content. Plants of ‘Tiger II’ colonial bentgrass, ‘L-93’ creeping bentgrass, and ‘Greenwich’ velvet bentgrass were subjected to two soil moisture treatments: 1) well-watered controls, irrigated three times per week; and 2) drought, irrigation completely withheld for 16 days. For recovery, previously drought-stressed plants were rewatered and irrigated three times per week to evaluate the recovery potential for each species. Drought stress resulted in significant declines in turf quality (TQ), shoot extension rates, canopy net photosynthetic rate (Pn), daily evapotranspiration rate (ET), and cytokinin content, and significant increases in ABA content for all three bentgrass species. Velvet bentgrass exhibited less severe drought injury, as exhibited by higher TQ, Pn, and daily ET rate compared with colonial bentgrass and creeping bentgrass. Velvet bentgrass also had significantly less ABA accumulation, which could allow for continued gas exchange and sustained plant survival during drought stress compared with colonial bentgrass and creeping bentgrass. Upon rewatering after drought stress, colonial bentgrass exhibited more rapid recovery in turfgrass growth and water use compared with creeping bentgrass and velvet bentgrass. The higher recuperative ability of colonial bentgrass could be associated with its more rapid decline in ABA content and increases in cytokinin content compared with creeping bentgrass and velvet bentgrass.
Reestablishment of damaged golf greens and fairways planted to creeping bentgrass (Agrostis stolonifera), colonial bentgrass (A. capillaris), and velvet bentgrass (A. canina) is a common practice following winter injuries. Identifying bentgrass species (Agrostis sp.) and cultivars with the potential to establish under low soil temperatures would be beneficial to achieving more mature stands earlier in the spring. Twelve bentgrass cultivars, including seven cultivars of creeping bentgrass (007, 13-M, Declaration, L-93, Memorial, Penncross, and T-1), two colonial bentgrass cultivars (Capri and Tiger II), and three velvet bentgrass cultivars (Greenwich, SR-7200, and Villa), along with ‘Barbeta’ perennial ryegrass (Lolium perenne) were evaluated for grass cover in the field during early spring. Bentgrass species and cultivars were seeded in the field at the same seed count per unit area. Soil temperatures were monitored in unseeded check plots from initial planting date on 8 Apr. to termination on 29 May 2013. Soil temperatures increased linearly during the 52-day experimental period from 4.7 to 23.5 °C. All species and cultivars emerged at ≈10 °C soil temperature. Bentgrass species and cultivars varied only 2 to 3 days in their initial seedling emergence, while days varied among bentgrasses from 5.5 days (to 10% cover) to 8.6 days (to 90% cover). All velvet bentgrass cultivars required higher soil temperatures (13.6 °C) and more time (26 days) following initial seedling emergence to establish to 90% cover in the early spring. Creeping bentgrass cultivars 007, 13-M, and Memorial, along with colonial bentgrass cultivars Capri and Tiger II, were statistically equal to ‘Barbeta’ perennial ryegrass in their capacity after seedling emergence to achieve faster cover at lower soil temperatures. Heavier (larger) bentgrass seed was associated with faster cover during the early stages of establishment, but seed size was uncorrelated with establishment during later stages from 50% to 90% cover.
Reseeding of creeping bentgrass (Agrostis stolonifera L.) under unfavorable temperature (≈10 °C) is a common practice on golf putting greens and fairways. Seed priming to enhance germination and early emergence increases seeding success. Seed priming comparing abscisic acid (ABA), gibberellic acid (GA), glycinebetaine (GB), hydrogen peroxide (H2O2), and polyethylene glycol (PEG) has not been investigated in turfgrass. Our objective was to compare these chemical primers at three concentrations with water- and unprimed-seed at two competing germinating temperatures (10 vs. 25 °C). Two seed lots of ‘T-1’ creeping bentgrass were compared. Curve fitting of daily germination was used to compute days to 50% germination (D50) and maximum germination percentage (Gmax). Cold (10 °C) significantly inhibited emergence (higher D50) more than Gmax. The effects of primers and their rates varied with the seed lot and temperature. Enhancement of seed germination measured as early emergence (lower D50) and/or higher Gmax were only detected at 10 °C. Osmotic primers (GB and PEG) were most effective in promoting germination relative to unprimed seed followed by hormone primers (ABA and GA) with redox primers (H2O2) least effective. Glycinebetaine primed seed was the only primer effective at all concentrations, with the 100 mм concentration the only concentration to enhance germination by increasing both Gmax and early emergence (lower D50) compared with unprimed seed.
Cold acclimation improves freezing tolerance in various plants, including perennial grass species. The objectives of this study were to determine protein changes in crowns of velvet bentgrass (Agrostis canina) during cold acclimation in association with freezing tolerance. Treatments consisted of: 1) nonacclimated (NA) plants maintained at 18/12 °C (day/night); 2) plants acclimated at a constant 2 °C for 4 weeks with a 10-hour photoperiod [A4 (cold acclimation)]; and 3) plants acclimated at a constant 2 °C for 4 weeks with additional subzero acclimation (SZA) at a constant –2 °C for 2 weeks (A4 + SZA2). Exposing plants to A4 significantly increased freezing tolerance, but additional SZA had no further beneficial effects on freezing tolerance, as demonstrated by the lethal temperature for 50% of the test population (LT50). Thirteen protein spots with increased abundance (up-regulated) or with decreased abundance (down-regulated) during cold acclimation were identified for biological functions. Proteins up-regulated after cold acclimation (A4 or A4 + SZA2) included methionine synthase, serine hydroxymethyltransferase, aconitase, UDP-D-glucuronate decarboxylase, and putative glycine-rich protein. Cold acclimation-responsive proteins involved in amino acid metabolism, energy production, stress defense, and secondary metabolism could contribute to the improved freezing tolerance induced by cold acclimation in velvet bentgrass.
Carbohydrate metabolism is important for plant adaptation to drought stress. The objective of this study was to examine major forms of carbohydrates associated with superior drought tolerance and post-drought recovery in kentucky bluegrass (Poa pratensis) by comparing responses of different forms of carbohydrates with drought stress and re-watering in two cultivars contrasting in drought tolerance. Plants of drought-tolerant ‘Midnight’ and drought-sensitive ‘Brilliant’ were maintained well watered or subjected to drought stress for 10 days by withholding irrigation, and drought-stressed plants were re-watered for 3 days. Physiological analysis (turf quality, relative water content, and electrolyte leakage) confirmed the genetic variability of the two cultivars in drought tolerance. The two cultivars exhibited differential responses to drought stress and re-watering for the content of water-soluble sugars (sucrose, fructose, and glucose) and storage carbohydrates (starch and fructan), and ‘Midnight’ maintained higher sucrose content at 10 days of drought stress and more fructan at 3 days of re-watering. The greater accumulation of sucrose in ‘Midnight’ under drought stress corresponded with higher activities of two sucrose-synthesizing enzymes (sucrose phosphate synthase and sucrose synthase) but was not related to the sucrose-degrading enzyme activity (acid invertase). These results suggested that increased sucrose accumulation resulting from the maintenance of active sucrose synthesis could be associated with superior turf performance during drought stress, whereas increased fructan accumulation could contribute to rapid re-growth and post-drought recovery on re-watering in kentucky bluegrass.