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TURFGRASS SCIENCE

Relative NaCl Tolerance of Kentucky Bluegrass, Texas Bluegrass, and Their Hybrids

^a Dep. of Hortic. and Landscape Architecture, Colorado State Univ., Fort Collins, CO 80523-1173
^b Texas A&M Univ. Res. and Ext. Center, 17360 Coit Rd., Dallas, TX 85252-6599

* Corresponding author (mploense{at}earthlink.net)

	ABSTRACT

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 
Government regulations to force water conservation have escalated the use of secondary water high in soluble salts for turfgrass irrigation in the arid and semiarid western USA, thus increasing the need for more salt-tolerant turfgrasses. This study was initiated to determine the variability in salt tolerance within and among two Poa species and their hybrids. Two experiments were conducted during 2000 in the greenhouse at Fort Collins, CO, in solution culture to examine the effects of NaCl on leaf firing and shoot and root growth reduction of nine Kentucky bluegrass (KBG; Poa pratensis L.) cultivars representing three ecotypes, three Texas bluegrass (TBG; P. arachnifera Torr.) accessions, and five of their hybrids (P. pratensis x P. arachnifera). In Exp. I, conducted during late winter through spring 2000, overall salt tolerance based on leaf firing and electrical conductivity (EC) of 50% shoot growth reduction (EC_shoot 50) placed seven KBG cultivars in the most tolerant group. In Exp. II, conducted during summer though early fall 2000, overall salt tolerance rankings placed 4 KBG and 3 TBG cultivars in the most tolerant group. On the basis of percentage leaf firing and the salinity levels that caused 25 and 50% shoot growth reduction, compact (low, compact growth habit) and aggressive (aggressive, lateral growth habit) KBG ecotypes showed more salt tolerance than common ecotypes in both Exp. I and II. A broad range of variability in leaf firing and shoot and root growth reduction in response to salinity was found to exist within and among these Poa species and their hybrids, indicating that improvement in the salt tolerance of bluegrasses may be possible. Additionally, differences in salt tolerance of KBG and TBG between Exp. I and Exp. II suggested that environmental conditions could affect bluegrass salt tolerance expression.

Abbreviations: EC, electrical conductivity • HBG, hybrid bluegrass • KBG, Kentucky bluegrass • TBG, Texas bluegrass

	INTRODUCTION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 
DEMANDS ON FRESH WATER RESOURCES in the western USA have resulted in codified limitations on the permissible area devoted to turfgrass (Clark Co., NV, 1992; City of Aurora, CO, 1998) as well as the quantity of potable water relegated to its irrigation (Hull and Pearson, 1999; City of Albuquerque, NM, 2000). As a result, secondary or effluent water, often containing high concentrations of NaCl, the only municipal water supply expected to increase in this region in the future (Oad and DiSpigno, 1996), is increasingly being used as an irrigation source for turfgrass whenever available (Gill and Rainville, 1994; Marcum et al., 1998; San Diego County, CA, 2000). Freshwater conservation mandates in rapidly growing metropolitan areas of the arid and semiarid western USA, where saline soils are common, has increased the need for more salt-tolerant turfgrasses.

Kentucky bluegrass, considered to be salt-sensitive with an average threshold EC of 3 dS m^-1 (Carrow and Duncan, 1998), is the most widely used cool-season turfgrass species in the more temperate regions of the western USA (Christians, 1998). Texas bluegrass is native to the southern Great Plains, and persists under extended periods of high temperature (Gould, 1975). ‘Reveille’ hybrid bluegrass (HBG), a hybrid between KBG and TBG and recently released as a heat-resistant hybrid bluegrass for the southwestern USA (Read et al., 1999), may increase the area of bluegrass planted under saline conditions.

Several studies have been conducted to assess the effect of salinity on KBG (Grueb et al., 1983; Horst and Taylor, 1983; Torello and Spokas, 1983; Torello and Symington, 1984; Butler et al., 1985; Qian et al., 2001). However, screening of the considerable amount of KBG plant material released during the past 15 yr for salt tolerance has received little attention. Although KBG is generally ranked as a salt-sensitive turfgrass, variability in salt tolerance has been shown to exist among cultivars. Horst and Taylor (1983), examining germination and initial growth in saline solution culture, reported significant differences in salt tolerance during germination and initial growth between 44 KBG cultivars. Variability in salt tolerance at the species level has likewise been demonstrated in bluegrass. Grueb et al. (1983) found Rough stalk bluegrass (P. trivialis L.) to be more salt-tolerant than a group of six KBG cultivars, within which group there was significant variability in visual appearance under salt stress. Demonstrated inter- and intra- specific variability in salt tolerance among the Poa spp. suggests that genetic improvement in the salt tolerance of bluegrasses is possible. More current information regarding the salt tolerance of KBG is needed. Nothing is known of the salt tolerance of TBG or HBG.

Therefore, the objective of this study was to examine the growth and turf quality responses to salinity within and among KBG cultivars, TBG accessions, and their hybrids.

	MATERIALS AND METHODS

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 
Plant Material
Five KBG cultivars were chosen for this study: ‘Huntsville’, ‘Dellwood Fine’, and ‘H-86-386’ because they served as the pollen source for the HBG examined in this study, and ‘Kenblue’ and ‘Bensuns A-34’ because of their contrasting growth characteristics. Dellwood Fine, Huntsville, and Kenblue represented three common KBG ecotypes (erect growth habit, narrow leaf blade) and Bensuns A-34 and H86-386 represented two aggressive ecotypes (aggressive lateral growth habit, high shoot density) as described by Murphy et al. (1997). Additionally, four unidentified Kentucky bluegrass plants discovered persisting on a highly sodic site at College Station, TX, which exhibit the appearance of compact ecotypes (low, compact growth habit) were collected and included in this study. Three TBG accessions, 20-11, 10-30, and 10-24, which served as the female parents of the released hybrid Reveille, and four HBG experimental lines (TXKY 96-260-6, TXKY 96-260-7, TXKY 96-260-22, and TXKY 94-8-8) were examined.

Plant Culture and Treatment Procedures
From 30 Dec. 1999 to 30 May 2000 (Exp. I) and from 1 June to 30 Oct. 2000 (Exp. II), plant material was screened for salinity tolerance in a greenhouse at the W.D. Holly Plant Environmental Research Center at Colorado State University, Fort Collins, CO, in a solution culture system (Qian et al., 2000). New sets of plants not previously subjected to salinity treatment were used at the initiation of each experiment. Greenhouse air temperatures ranged from 20.9 to 28.2°C in Exp. I and from 20.8 to 36.1°C in Exp. II. Because no supplemental light source was used during either experiment, mean photosynthetically active radiation in the greenhouse increased from 33.8 to 78.1 W m^-2 in Exp. I (December to May) and decreased from 35.7 to 80.6 W m^-2 in Exp. II (June to October) (Table 1) . Sod pieces of each grass were planted into 7-cm diam. by 4-cm-deep plastic cups filled with a 1-cm layer of Isolite (Sundine Enterprises Inc., Arvada, CO). The cup bottom was removed and covered with nylon screen to hold the Isolite and allow roots to grow through. Seventeen cups consisting of one entry per cup were placed into holes of a 1.5-cm-thick wooden lid, with the lid suspended over a 38-L tank. A total of 15 tanks were used with each accommodating 17 cups. The tanks contained 36 L of constantly aerified full strength Hoagland solution, which was replaced weekly. This volume allowed the bottom of each cup to be submersed 2 cm into the solution.

Data Collection and Analyses
Data were collected on shoot and root growth and leaf firing percentage. After reaching the designated treatment level, shoots were clipped weekly to a 2.5-cm height and discarded. Beginning 4 wk after salinity treatment when grasses had fully exhibited response to the salt treatments, clippings were collected weekly for 6 wk. Clippings were immediately dried for 48 h at 70°C and weighed. Six of these harvests were combined to determine shoot growth. At the conclusion of each experiment, roots were harvested and dried for 48 h at 70°C, then weighed for dry mass. Leaf firing percentage was determined by visually estimating the total percentage of chlorotic leaf area at the conclusion of each experiment.

Analysis of variance indicated significant difference in all measured parameters between Exp. I and Exp. II (Table 2) . Therefore, data on shoot and root growth and leaf firing percentage are presented separately. Linear and quadratic regression analysis was conducted to determine relationships between shoot and root growth vs. salinity level. By regression analysis, threshold EC (salinity level at which growth reduction began relative to control) and the slope of growth reduction were determined for each entry. The regression slopes were then used to derive salinity levels that caused 25 and 50% shoot and root growth reductions (EC_shoot 25, EC_shoot 50, EC_root 25, and EC_root 50) (Maas and Hoffman, 1977). Resulting data were then subjected to ANOVA tests, and cultivar means were separated by Fisher's LSD (SAS Institute, 1990). Pearson's correlation analysis between percentage leaf firing and threshold EC, EC_shoot 25, and EC_shoot 50 was conducted using the CORR procedure of SAS (SAS Institute, 1990). Group response comparisons were made using the TTEST procedure assuming both equal and unequal variance (SAS Institute, 1990). Using two factors, leaf firing percentage at 5 dS m^-1 and EC_shoot 50 as source data, cluster analysis was performed on all 17 entries using the nonhierarchical FASTCLUS procedure to place entries into groups not defined a priori (SAS Institute, 1990). Electrical conductivity of 50% shoot growth reduction values was transformed (max EC_shoot 50 - xEC_shoot 50) so that a lower score indicated greater tolerance by both factors. Thus, both axis values increased with increasing distance from the point of origin.

	RESULTS

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

Shoot Growth Reduction
Experiment I
Differences were found for threshold EC, EC_shoot 25, and EC_shoot 50 (Table 3). Threshold EC ranged from 1.0 to 3.0 dS m^-1. The highest threshold EC of 3.0 was observed in TBG 20-11, Kenblue, H-86-386 KBG, and TXKY 94-8-8 HBG. Salinity levels that caused 25% shoot growth reduction ranged from 2.0 dS m^-1 in Reveille HBG to 7.8 dS m^-1 in CS ST 4 KBG. Salinity levels that caused 50% shoot growth reduction ranged from 4.0 dS m^-1 in Reveille HBG to 12.6 dS m^-1 in CS ST 4 KBG. Variability in shoot growth reduction both within and among Poa species and their hybrids increased from EC_sh4o_ot 25 to EC_shoot 50, indicating the influence of slope on shoot growth reduction in response to increasing salinity. No differences were found in threshold EC, EC_shoot 25, and EC_shoot 50 between Poa species, their hybrids, or among KBG ecotypes as groups (Table 4).

Experiment II
Again, significant variability in shoot growth reduction parameters was found within and among Poa species and their hybrids (Table 3). Threshold EC ranged from 1.0 to 5.0 dS m^-1, with the highest threshold EC of 5.0 observed in Bensuns A-34 KBG, H-86-386 KBG, and TXKY 94-8-8 HBG. Values of EC 25 ranged from 3.1 dS m^-1 in TXKY 96-260-7 HBG to 10.0 dS m^-1 in Bensuns A-34 KBG. Both HBG and KBG significantly increased their mean EC_shoot 25 from Exp. I to Exp. II. Among KBG ecotypes, aggressive types exhibited higher EC_shoot 25 than both common and compact types (Table 4).

Salinity levels that caused 50% shoot growth reduction values ranged from 6.2 dS m^-1 in TXKY 96-260-7 HBG to 15.0 in Bensuns A-34 KBG. Although the most and least tolerant entries were the same from EC_shoot 25 to EC_shoot 50, greater difference in EC_shoot 50 was found between KBG and HBG than EC_shoot 25, indicating influence of slope of reduction on decreased shoot growth. Both species and their hybrids increased their mean EC_shoot 50 in Exp. II compared with Exp. I (Table 4). Among KBG ecotypes, aggressive types exhibited higher EC_shoot 50 than common types.

Root Growth Reduction
Experiment I
Relationships between root growth and level of salinity were determined for each entry. Variability existed within and among Poa species and their hybrids for each measured or derived root growth reduction parameter (threshold EC, EC that caused 25 and 50% root growth reduction) (Table 3). Threshold EC ranged from 1.0 to 7.3 dS m^-1. The highest threshold EC of 7.3 was observed in CS ST 1 KBG. Salinity levels that caused 25% root growth reduction ranged from as low as 2.2 to 2.5 dS m^-1 in Reveille HBG, TXKY 96-260-6 HBG, TXKY 96-260-7 HBG, Dellwood Fine KBG, and TBG to as high as 13.3 dS m^-1 in CS ST 1 KBG. Electrical conductivity of 50% root growth reduction ranged from 4.4 dS m^-1 in Reveille HBG to 15.8 in CS ST 1 KBG. No difference in EC_root 25 was found between Poa species and their hybrids. No difference in any root growth reduction parameter was exhibited among KBG ecotypes.

Experiment II
Significant variability in all measured and derived root growth reduction parameters were found within and among Poa species and their hybrids (Table 3). Threshold EC ranged from 1.0 to 5.0 dS NaCl m^-1, with the highest threshold EC of 5.0 observed in TBG 10-24 and TXKY 94-8-8 HBG. Values of EC_root 25 ranged from a low of 1.9 dS m^-1 in TXKY 96-260-7 HBG to 8.5 dS m^-1 in TBG 10-24. No difference was found among KBG ecotypes for EC_root 25.

Although salinity levels that caused EC_root 50 ranged from 3.9 dS m^-1 in TXKY 96-260-7 HBG to 12.1 dS m^-1 in TBG 10-24, no difference in EC_root 50 was found between Poa species, their hybrids, or among KBG ecotypes. Even though the most and least tolerant entries ranked the same from EC_root 25 to 50, variability within and among Poa species and their hybrids was greater at EC_root 50 than at EC_root 25.

Kentucky bluegrasses exhibited a significantly lower (7.3 to 3.4 dS m^-1) EC_root 25 in Exp. II than in Exp. I because of poorer performance of the compact and aggressive ecotypes in Exp. II. The same occurred for EC_root 50; however, at this level it was primarily attributable to the poorer performance of only compact ecotypes.

Salt Tolerance Ranking
To assess the overall salt tolerance of all entries, cluster analysis was performed based on relative leaf firing and EC_shoot 50 (Fig. 1) . These factors were chosen because turfgrass appearance and growth are two important factors influencing overall turf performance, and the highest correlation between relative leaf firing and shoot growth was found between EC_shoot 50 scores (Table 5) .

View larger version (13K): [in this window] [in a new window]   Fig. 1. Cluster analysis of Exp. I and Exp. II based on relative leaf firing at 5 dS m-1 and salinity level resulting in 50% shoot growth reduction (ECshoot 50) for three Texas bluegrasses (TXBG), nine Kentucky bluegrasses (KBG), and five of their hybrids (HBG). A = TXKY 96-260-7 HBG, B = TXBG 10-30, C = TXBG 10-24, D = ‘Dellwood Fine’ KBG, E = TXBG 20-11, F = ‘Huntsville’ KBG, G = H-86-386 KBG, H = 96-260-6 HBG, I = CS ST 1 KBG, J = ‘Kenblue’ KBG, K = CS ST 4 KBG, L = CS ST 2 KBG, M = TXKY 94-8-8 HBG, N = ‘Bensuns A-34’ KBG, O = TXKY 96-260-22 HBG, P = ‘Reveille’ HBG, Q = CS ST 3 KBG.

Experiment II
In Exp. II, the most salt-tolerant group (Cluster 1) was composed of four KBG cultivars (Bensuns A-34, CS ST 1, CS ST 3, and CS ST 4), including three compact and one aggressive ecotype, and all three TBG accessions examined (10-24, 10-30, and 20-11) (Fig. 1). Again, the intermediate and least tolerant groups (Clusters 2 and 3, respectively) included KBG, TBG, and HBG.

	DISCUSSION

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 
Results indicate that significant variability in leaf firing and shoot and root growth responses to salinity exist within and among Poa species and their hybrids. The range of salinity tolerance among the nine KBG cultivars in relative leaf firing response at 5 dS m^-1 in both Exp. I (1.7–58.3%) and Exp. II (5.0–93.3%) was considerably broad. Among the KBG examined in this study, greatest salt tolerance was seen in the aggressive and compact ecotypes. Qian et al. (2001) also reported that ‘Limousine’ KBG, an aggressive ecotype, was more salt-tolerant than ‘Kenble’, a common ecotype KBG. Nevertheless, KBG is still a salt-sensitive turfgrass species when compared with other species such as tall fescue (Festuca arundinacea Schreb.) and creeping bentgrass {Agrostis palustris Huds. [= A. stolonifera var. palustris (Huds.) Farw.]} (Carrow and Duncan, 1998). Selecting relatively salt-tolerant KBG cultivars is beneficial for sites where salinity level is only marginal.

Differences between Exp. I and Exp. II suggested that environmental conditions could substantially affect bluegrass salt tolerance (Table 4). In a previous study (Qian and Suplick, 2001), involving the interactive effect of temperature and salinity on KBG seed germination, the effect of salinity became more pronounced as temperature deviated from optimum growth range. Optimum temperature range for KBG shoot growth is 16 to 24°C, and 10 to 18°C for root growth (Beard, 1973). Optimum shoot and root growth temperature ranges have not been determined for TBG or HBG.

Texas bluegrass is distributed throughout several vegetative regions of Texas, where mean maximum summer temperatures range from 34 to 37°C (Gould, 1975; Northeast Regional Climate Center, 2000). This region of origin suggests that TBG may have a warmer optimum temperature range than KBG. Current and ongoing research (Suplick and Qian, 2000) suggests that HBG may have a broader temperature adaptation than those of KBG and TBG.

This speculation may be supported by the differences in performance from Exp. I to Exp. II of the three Poa groups examined in this study in many measured and derived parameters (Table 4). Experiment I was conducted during winter and spring. Experiment II was conducted throughout summer into fall when daily warm temperatures were higher, and their duration prolonged, possibly creating an environment more favorable to TBG but less favorable to KBG growth. This change in environment may explain several observations: (i) There was an 81% decrease in TBG mean leaf firing at 5 dS m^-1 from Exp. I to Exp. II; (ii) KBG root threshold EC, EC_root 25, and EC_root 50 were significantly decreased from Exp. I to Exp. II, whereas no change was seen in TBG or HBG; and (iii) there was an improvement in overall salt tolerance for all three TBG accessions, demotion of several KBG cultivars, and relatively steady ranking of HBG from Exp. I to Exp. II.

This research supports that of Horst and Taylor (1983) and Grueb et al. (1983), in that significant variability in salt tolerance exists within and among Poa species, indicating that improvement in the salt tolerance of bluegrasses may be possible. Additionally, our findings, with respect to the effect of sub- and supraoptimum temperature regimes on the expression of that tolerance, are in agreement with previous research (Qian and Suplick, 2001), confirming the importance of temperature in evaluating the salt tolerance of Poa species.

	NOTES

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 
Funding was provided by Gardner Turfgrass, Inc. and the Colorado Agric. Exp. Stn. (project 780). Manuscript submitted by the senior author in partial fulfillment of the requirements for the Ph.D. degree from Colorado State Univ.

Received for publication September 20, 2001.

	REFERENCES

TOP NOTES ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS DISCUSSION REFERENCES

 

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