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Crop Science 41:451-456 (2001)
© 2001 Crop Science Society of America

TURFGRASS SCIENCE

Spring Dead Spot Resistance and Quality of Seeded Bermudagrasses under Different Mowing Heights

D.L. Martina, G.E. Bella, J.H. Bairdb, C.M. Taliaferroc, N.A. Tisseratd, R.M. Kuzmice, D.D. Dobsona and J.A. Andersona

a Dep. of Horticulture & Landscape Architecture, Oklahoma State Univ. (OSU), 360 AG Hall, Stillwater, OK 74078-6027
b Dep. of Plant & Soil Sci., Michigan State Univ., East Lansing, MI 48824
c Dep. of Plant & Soil Sci., OSU, OK 74078
d Dep. of Plant Pathology, Kansas State Univ., Manhattan, KS 66506
e Dep. of Forestry, OSU, Stillwater, OK 74078

Corresponding author (hortdm1{at}okstate.edu)


   ABSTRACT
TOP
 NOTES
 ABSTRACT
INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
Bermudagrass (Cynodon spp. L.C. Rich.) cultivars resistant to spring dead (SDS) caused by Ophiosphaerella herpotricha (Fr.) Walker and winter-kill are important for residential and commercial use in transition zone climates. Field research was conducted on ‘Mirage’, ‘OKS 91-11’, and ‘Jackpot’ seeded bermudagrasses [Cynodon dactylon (L.) Pers. var. dactylon] at Stillwater, OK, between 1996 and 1999 to determine their resistance to SDS. Plots were inoculated in 1996 with O. herpotricha isolate KS 188 and mowed to a height of either 1.3 cm or 3.8 cm. The SDS necrotic patch area of all cultivars increased each year and patch area was greater in turf at the higher mowing height. OKS 91-11 had a smaller SDS necrotic patch area than the other cultivars. Mirage had necrotic patch area equal to Jackpot at 1.3 cm and less necrotic area than Jackpot at 3.8 cm. OKS 91-11 had greater shoot survival density in necrotic patches than Jackpot or Mirage in 2 of 3 yr at 1.3 cm. Cultivars did not differ in shoot survival density at 3.8 cm. Higher overall quality was provided by OKS 91-11, when compared with Mirage and Jackpot in 3 of 4 and 2 of 4 yr, respectively. Jackpot had better quality than Mirage in 2 of 4 yr. Early spring green-up was negatively correlated with SDS patch size and positively correlated with shoot survival in SDS patches. Winter-kill was positively correlated with SDS patch size and negatively correlated with shoot survival in SDS patches.

Abbreviations: KS 188, Kansas State University Ophiosphaerella herpotricha isolate 188 • SAS, Statistical Analysis Systems • NP, necrotic patch • SDS, spring dead spot


   INTRODUCTION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
SPRING DEAD SPOT is one of the most serious diseases of bermudagrass (C. dactylon and C. dactylon x C. transvaalensis Burtt-Davy) in the transition zone between the cool temperate and subtropical climatic zones. Spring dead spot is caused by at least three different ectotrophic, root-rotting fungi, including Ophiosphaerella herpotricha, O. korrae Walker and Smith (synonym Leptosphaeria korrae Walker and Smith), and O. narmari Walker and Smith (synonym L. narmari Walker and Smith) (Crahay et al., 1988; Endo et al., 1985; Smith, 1971; Tisserat et al., 1989; and Walker and Smith, 1972; Wetzel et al., 1999). Ophiosphaerella herpotricha is the most common causal agent of SDS in the southern great plains region of Kansas, Oklahoma, and Texas (Tisserat et al., 1989).

A large number of turf-type seeded bermudagrasses entered the market during the 1990s. Knowledge of the aesthetic value and functional performance, including SDS and winter-kill resistance, of these grasses is essential to determine regional adaptation. Mirage, OKS 91-11, and Jackpot (Samudio and Brede, 1997) ranked highest among seeded bermudagrasses for overall turf quality in a National Turfgrass Evaluation Program (NTEP) study at 22 sites (NTEP, 1997). The visual quality rating is generally accepted as the best indicator of overall adaptation of a turfgrass cultivar to a particular site. Visual quality ratings integrate density, texture, uniformity, color, growth habit, and smoothness (Turgeon, 1991). Baird et al. (1997) reported on the SDS resistance of 25 seeded bermudagrasses and 11 vegetatively propagated bermudagrasses in Kansas and Oklahoma using a lab and field inoculation technique. Under field conditions, Mirage was found to have improved O. herpotricha (isolate KS 188) resistance over Jackpot. Additionally, Baird et al. (1997) found that Mirage had significantly less winter-kill than Jackpot at Stillwater, OK. The cultivar OKS 91-11 was not included in trials by Baird et al. (1997) and has not been evaluated for SDS resistance.

Cultural management practices are thought to influence bermudagrass response to SDS. Pair et al. (1986) suggested that the disease is generally associated with cold winters and management factors that reduce hardiness, delay autumn dormancy, or promote early spring growth. Low mowing height, thatch accumulation, and soil compaction are generally associated with increased SDS development (Lucas, 1980), but these factors have not been studied in detail. Dernoeden et al. (1991) reported that acidification of soil by NH4-N sources reduced the severity of SDS (O. korrae) on ‘Tufcote’ bermudagrass (C. dactylon var. dactylon) over time. Tisserat and Fry (1997) found a combination of core aeration plus vertical mowing, performed twice each year, was moderately effective in reducing SDS (O. herpotricha) on ‘Midlawn’ bermudagrass (C. dactylon x C. transvaalensis). Vertical mowing or aeration alone, however, did not significantly reduce disease severity. Yearly removal of infected sod followed by regrowth of bermudagrass from rhizomes also was effective in reducing SDS severity (Tisserat and Fry, 1997). Sod removal and regrowth, however, is not practical on most turfgrass sites. The effect of mowing height on SDS severity has not been studied although Couch (1995), Lucas (1980) and Vincelli and Williams (1998) suggested that increasing the mowing height may reduce damage from the disease.

Improved, cold hardy, turf-type seeded bermudagrasses need to be screened for resistance to SDS, and management practices that reduce disease severity require further investigation. The objectives of this research were to: (i) evaluate the field SDS response of Mirage, Jackpot, and OKS 91-11 inoculated with O. herpotricha; (ii) determine the influence of mowing height on severity of SDS; and (iii) evaluate the visual quality response of these three cultivars under field conditions.


   MATERIALS AND METHODS
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
Plant Materials
Jackpot, Mirage, and OKS 91-11 bermudagrasses were established from seed at the Oklahoma State University. Turfgrass Research Center at Stillwater, OK, on 5 July 1995. Soil was an Easpur silty clay loam (a fine-loamy, mixed, thermic Fluventic Haplustoll). Soil pH ranged from 6.9 to 7.0 during the study. Mean phosphorus (P) and potassium (K) soil levels were 92 and 375 kg ha-1, respectively. The seeding rate was 49 kg ha-1 of pure live seed. The cultivars were planted in a randomized complete block with three replications. Upon establishment, main plots were split into two, 3.8- by 4.5-m subplots and two mowing height treatments of 1.3 and 3.8 cm were administered 2 to 3 times wk-1 during the growing season with a reel mower. The 1.3-cm mowing height is typically used on golf course fairways, while the 3.8-cm mowing height is used for high maintenance roughs and home lawns in the southern great plains region of the USA. The turf received 49 kg N ha-1 mo-1 from May through September each year in the form of urea.

Spring Dead Spot
The cultivars were inoculated in September 1996 with O. herpotricha isolate KS 188 by the method described by Baird et al. (1997) and Tisserat et al. (1989). The isolate was originally obtained from bermudagrass turf at the test site. Six soil-turf plugs were removed to a depth of 10 cm from each cultivar x mowing height subplot with a standard 10-cm diam (84-cm2 area) golf cup cutter. Ten grams of infested oat (Avena sativa L.) were placed into three of the holes with the remaining holes receiving 10 g of sterile oat. The plugs then were replaced and the turf was irrigated to prevent desiccation. The inoculation points were mapped so that plots could be monitored for disease development.

Disease evaluations were made 1 May 1997, 21 April 1998, and 13 April 1999 when symptoms were prominent, by the method described by Baird et al. (1997). The diameter of each necrotic patch (NP) was measured in two directions, and the average radius was used to calculate disease area by the equation for the area of a circle. Values reported for disease area or NP represent the average of the three inoculated areas taken per replication for a total of nine area measurements per cultivar.

The number of live shoots within each NP was determined at the same time as the NP area measurements were made. In addition, the number of live shoots were counted within a 46-cm2 plug, taken from a disease-free area in each cultivar x mowing height subplot. Percentage shoot survival within the NP was calculated by dividing the number of live shoots per unit area within the NP by the number of live shoots per unit area in unaffected turf and multiplying by 100. This transformation would account for inherent differences in shoot density that might be present among the cultivars and mowing heights.

Cultivar Quality Assessment
Green-up ratings were visually assessed each spring by a 1-to-9 scale, where 1 = dead or dormant turf and 9 = completely green, healthy turf. Green-up ratings were taken on 28 March 1997, 24 April 1998, and 24 March 1999. Late-season color retention ratings were visually determined on 19 Nov. 1996, 20 Oct. 1997, and 8 Dec. 1998, by a 1-to-9 scale, where 1 = completely brown turf and 9 = completely green turf. A winter-kill rating was made 16 April 1999 by visually estimating by a 0-to-100% scale, where 0 = no winter-kill and 100 = complete winter-kill, the percent area of above ground shoots that had been killed. Visual quality ratings were obtained monthly from June to September 1996 and from May to September in 1997 to 1999. Quality was rated visually on a 1-to-9 scale, where 1 = very poor quality and 9 = very high quality.

Statistical Analysis
Analyses of variance were performed on the dependent variables, NP area, shoot survival density in NP, percent shoot survival, shoot density in nonaffected turf, spring green-up, color retention, percent winter-kill, and visual quality ratings by Statistical Analysis System (SAS) software (SAS Institute Inc., Cary, NC). The range in percent shoot survival was only 0 to 12% and less than 40% of the scale, so arcsin transformation was not performed on these data (Little and Hills, 1978). The arcsin transformation was performed on percent winter-kill ratings after converting them to a decimal fraction. For all data other than quality and winter-kill ratings, the experiment was analyzed as a split-split plot split in time with cultivars as main plots, cultivar x mowing heights as subplots, and years within cultivar x mowing heights as sub-sub plots. The cultivar x rep and mowing height x rep within cultivar error terms initially were subjected to the F test by the year x rep within cultivar x mowing height error term. These error terms were not significantly different at P <= 0.05, and all error terms were pooled into a residual error term to test all effects in the model (Carmer et al., 1969). When treatment effects were found significant with the F test, means were separated by the least significant difference (LSD) test at P <= 0.05.

Because of an unequal number of rating dates in 1996 compared with other years, quality data were analyzed by year and the same split-split plot split in time analysis was conducted on these data where cultivars were main plots, cultivar x mowing heights were subplots and rating month within cultivar x mowing heights were sub-sub plots. The cultivar x rep and mowing height x rep within cultivar x mowing height error terms were significant for 1996, but not in 1997, 1998, and 1999, so a single pooled residual error term was used in each of the latter three years for the F test.

Winter-kill data were analyzed as a split-split plot with cultivars as main plots, and cultivar x mowing heights as subplots. A pooled residual error term was used when the cultivar x rep and mowing height x rep within cultivar terms were not statistically significant. Winter-kill means were separated as previously discussed, and the nontransformed means are presented for ease of interpretation.


   RESULTS AND DISCUSSION
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
CONCLUSIONS
 REFERENCES
 
Spring Dead Spot Severity
Spring dead spot appeared at all inoculation sites in all cultivars during the 3 yr following inoculation. No symptoms were present in the uninoculated control sites where sterile oat were used. Significant differences in SDS NP size were present among years (Table 1). Disease severity, as evidenced by NP size, became greater in each cultivar in each successive year of the study (Table 2). The cultivar OKS 91-11 exhibited smaller NP than Jackpot and Mirage in all 3 yr and 2 of 3 yr, respectively (Table 2). In general, NP area was numerically equal to or slightly less in Mirage than Jackpot, but the difference was not significant. Baird et al. (1997) reported less severe SDS in Mirage than Jackpot under simulated golf course fairway conditions using the same O. herpotricha isolate. Differences in methodology between our work and that of Baird et al. (1997) included age of the stands and seed lots used.


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  		Table 1. Mean squares from the analysis of variance conducted on spring dead spot resistance indices, shoot density, green-up ratings, and color retention collected from three bermudagrass cultivars

 

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  		Table 2. Influence of cultivar and mowing height on bermudagrass response to spring dead spot

 
Mowing height influenced SDS severity, and NP were generally larger in turf maintained at the higher mowing height (Tables 1 and 2). In Mirage, however, the mean NP area was slightly smaller in the plots maintained to a height of 3.8 cm. We are unaware of any previous research concerning the effects of mowing height on SDS severity, although the disease is most often associated with low mowing (Couch, 1995; Lucas, 1980; Vincelli and Williams, 1998). The reason for the difference in NP size as a function of mowing height in our study is not known. The environment as well as the thatch depth in the soil-thatch interface of turf maintained at various mowing heights deserves further study to determine their influence on SDS severity.

The overall percent shoot survival within NP caused by O. herpotricha was substantially greater in the bermudagrasses during 1999 than in 1997 or 1998. A significantly larger percentage of shoots survived within NP in OKS 91-11 than in Jackpot or Mirage during 1998 and 1999, with no significant differences present among cultivars in 1997. No statistical differences in percent shoot survival were observed between Mirage and Jackpot during any year. Baird et al. (1997) reported Mirage as having greater shoot survival within NP than Jackpot under simulated golf course fairway conditions nearly identical to those used in our study.

No differences were observed in shoot survival density among cultivars at the higher mowing height in all 3 yr of this study (data not shown). No differences in shoot survival density were observed among the cultivars at the lower mowing height in 1997. However, differences were present in 1998 and 1999 when OKS 91-11 had greater shoot survival density (1.9 and 19.0 dm-2) than Jackpot (0.1 and 1.8 dm-2) or Mirage (0.1 and 0.4 dm-2). Overall, shoot survival density was greater in 1997 (3.2 dm-2) and 1999 (4.7 dm-2) than in 1998 (0.4 dm-2).

Shoot Density
Shoot density in noninfected areas was greater for OKS 91-11 (189 dm-2) than for Mirage (123 dm-2) (data not shown), while the density of Jackpot (151 dm-2) did not differ from OKS 91-11 or Mirage. Shoot density of all cultivars decreased during the study, with mean shoot densities of 193, 183, and 88 dm-2 being observed in 1997, 1998, and 1999, respectively. The shoot density in unaffected areas was greatest in turf maintained to a height of 1.3 cm (182 dm-2), when compared with 3.8-cm (127-dm-2) mowing height.

Spring Green-Up
Differences in the earliness of green-up among cultivars were observed during 1998 and 1999 (data not shown). In 1998, OKS 91-11 greened-up earlier (5.5 rating) than Mirage (4.3 rating) or Jackpot (4.2 rating). During 1999, OKS 91-11 provided earlier green-up (2.3 rating) than Mirage (1.3 rating), while the mean green-up rating of Jackpot (1.8 rating) did not differ from that of OKS 91-11 or Mirage. Turf maintained at the lower mowing height exhibited earlier green-up (3.5 rating) than was observed under higher mowing (2.5 rating). This effect may be due to warmer soil temperatures under the shorter cut turf due to less solar radiation reflection and less insulation by the tan, dead turf canopy. The earlier green-up at a lower mowing height is consistent with the findings in other bermudagrass evaluation trials conducted at our site (NTEP, 1997).

Late-Season Color Retention
Overall, OKS 91-11 had better late-season color retention (5.6 rating) than Mirage (4.8 rating) or Jackpot (4.8 rating) (data not shown). Bermudagrass maintained at the lower mowing height (6.2 rating) also remained greener longer into the autumn than that at the higher mowing height (3.9 rating).

Visual Quality
The cultivars did not differ in quality ratings during 1996 at either mowing height (Table 3). Very few differences in quality were present among cultivars on individual dates in 1997 (data not shown). Differences did occur in May and June 1997 at the lower mowing height, and OKS 91-11 generally had better quality (6.7 and 7.3 rating) than Mirage (5.3 and 6.0 rating) or Jackpot (5.7 and 5.3 rating). The quality of Mirage, however, did not differ from the other two cultivars. The only date in 1997 when differences in quality occurred at the higher mowing height was in July, when OKS 91-11 provided better quality (7.3 rating) than Jackpot (6.6 rating) or Mirage (6.3 rating). The overall 1997 seasonal mean quality of OKS 91-11 (6.7 rating) exceeded that of Jackpot (6.4 rating) and Mirage (6.2 rating). In 1998, few differences in quality again were present on individual rating dates. In June 1998 at the lower mowing height, quality of OKS 91-11 (6.7 rating) exceeded that of Mirage (6.0 rating), while quality of Jackpot (6.7 rating) did not differ from either cultivar. At the higher mowing height, OKS 91-11 (7.0 rating) and Jackpot (7.0 rating) had better quality than Mirage (6.0 rating) in July, while OKS 91-11 provided better quality (7.0 rating) than Jackpot (6.0 rating) and Mirage (6.0 rating) in September of 1998. The overall seasonal mean quality in 1998 for OKS 91-11 (6.6 rating) and Jackpot (6.4 rating) did not differ, while both had better quality than Mirage (6.1 rating). In 1999, OKS 91-11 had better quality (6.7 rating) than Jackpot (6.0 rating), and Jackpot had better quality than Mirage (5.8 rating). On individual rating dates, the quality of Mirage and Jackpot did not significantly differ at either mowing height. At the low mowing height, OKS 91-11 provided better quality than Mirage and Jackpot on four of five and three of five rating dates, respectively in 1999. With higher mowing, OKS 91-11 provided better quality than Mirage or Jackpot on two of five and one of five rating dates, respectively.


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  		Table 3. Mean squares from the analysis of variance conducted on visual quality ratings and winter-kill ratings

 
Winter-Kill
A large amount of winter-kill occurred in bermudagrass throughout Oklahoma during the 1998-1999 winter (D.L. Martin, unpublished data). This provided the opportunity to examine winter hardiness of the bermudagrasses (Table 4). No statistical differences in winter-kill among cultivars occurred at the higher mowing height. However, at the lower mowing height, OKS 91-11 had substantially less winter-kill than either Mirage or Jackpot. The latter two cultivars did not differ in winter-kill levels. Overall, the cultivars maintained at the lower mowing height did not suffer as much winter-kill as turf managed at the higher mowing height.


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  		Table 4. Winter-kill ratings collected from three common bermudagrasses on 16 April 1999

 
Correlations among Disease Resistance, Green-Up, Color Retention, and Winter-Kill
Correlation analyses were performed to relate the indices of SDS resistance with green-up, late-season color retention, and winter-kill ratings. Of the correlations examined, winter-kill ratings in 1999 most closely correlated with total shoot survival in NP (negative), shoot survival density in NP (negative), percent shoot survival in NP (negative), and NP area (positive) (Table 5). Significant correlations also were found between green-up ratings and total shoot survival in NP (positive), and shoot survival density in NP (positive) and NP area (negative) in 1998 and 1999. Only in 1998 was there a significant and positive correlation between green-up and percent shoot survival in NP. These correlations suggest that shoot survival in NP is greatest for cultivars having early spring green-up, and that cultivars having early spring green-up tend to have smaller NP. Improved shoot survival within NP is generally presumed to improve the recovery rate of a bermudagrass once optimum growing conditions prevail; however, research needs to be conducted to confirm our findings. Additionally, our correlation analyses suggested that cultivars with improved resistance to winter-kill tend to have smaller SDS NP, and increased shoot survival in NP. The only other significant correlation was between total shoot survival in NP in spring 1999 and late-season color retention (positive) during 1998. Overall, there appeared to be no significant relationships among late-season color retention and indices of SDS resistance in the three seeded bermudagrasses. While turfgrass management practices that delay autumn dormancy may increase severity of SDS (Pair et al., 1986), breeding and selection for cultivars with improved late-season color retention need not inextricably link the desirable late-season color retention characteristic with decreased resistance to SDS caused by O. herpotricha.


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  		Table 5. Correlation coefficients{dagger} among parameters measured on three bermudagrasses evaluated for resistance to spring dead spot (SDS)

 

   CONCLUSIONS
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
REFERENCES
 
The field inoculation technique using oat infested with a known virulent strain of O. herpotricha was very effective in screening cultivars for resistance to SDS. Spring dead spot severity increased each year in the 3-yr time period of this study. The cultivar OKS 91-11 was more resistant to O. herpotricha than Mirage or Jackpot using NP size and shoot survival density as well as percent shoot survival in NP as indices of SDS resistance. This work confirmed earlier findings by Baird et al. (1997) that Mirage has slightly more resistance to O. herpotricha, when compared with Jackpot. We did not confirm the differences in percent shoot survival in NP between Jackpot and Mirage as reported by Baird et al. (1997). Additional work is needed to determine the response of these bermudagrasses to the other known causal agents of SDS. Severity of SDS as measured by NP size was greater in all cultivars mowed to a height of 3.8 cm, when compared with the 1.3-cm mowing height. The mechanism for increased NP size with increasing mowing height needs to be studied to better improve cultural recommendations used in managing SDS.

The cultivars OKS 91-11 and Jackpot appear to have greater shoot density than Mirage in noninfected areas. As expected, shoot density of all cultivars was greater at a 1.3-cm mowing height, when compared with a 3.8-cm mowing height. Expressing shoot survival in NP as a percentage of shoot density in nonaffected areas accounted for inherent differences in density among cultivars and between mowing heights. This transformation allowed for clear comparisons of treatment effects on SDS severity.

The cultivar OKS 91-11 had improved winter hardiness when compared with Jackpot and Mirage at a 1.3-cm mowing height, with no substantial differences observed among cultivars maintained at the 3.8-cm mowing height. Tissue cold hardiness testing of these cultivars would be beneficial to validate our field observations.

The cultivar OKS 91-11 provided slightly better quality than Mirage and Jackpot in three of four and two of four growing seasons, respectively. Jackpot provided slightly better quality than Mirage during two of four growing seasons. The quality of these grasses may vary on the basis of different management regimes and local climatic factors. On the basis of the limited amount of work performed on OKS 91-11 in this study and elsewhere (NTEP, 1997), this cultivar shows promise for use on well maintained golf course fairways, roughs, and home lawns where bermudagrass is adapted, establishment by seed is necessary, and where improved winter hardiness and SDS resistance are valued.


   ACKNOWLEDGMENTS
 
This research was supported by grants from the United States Golf Association and the Oklahoma and Kansas Turfgrass Research Foundations. The authors would also like to thank the Oklahoma and Kansas Agricultural Experiment Stations for their support of this work. Appreciation is expressed to Keith Reed for his maintenance of the research site and to Stephanie Larimer for her assistance in preparation of this manuscript.


   NOTES
TOP
 NOTES
ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 
This study was funded by the United States Golf Assoc., Far Hills, NJ 07931 under grant AG-98-RS-026. Additional funding was received from the Oklahoma Turfgrass Research Foundation, Broken Arrow, OK 74012. Approved for publication by the Director, Oklahoma Agricultural Experiment Station. Research conducted under Oklahoma Agric. Exp. Stn. Proj. OKLO 2222. Kansas State Agric. Exp. Stn. Paper No. 00-85-J.

Received for publication March 27, 2000.


   REFERENCES
TOP
 NOTES
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 CONCLUSIONS
 REFERENCES
 




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