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Crop Science 41:457-465 (2001)
© 2001 Crop Science Society of America

TURFGRASS SCIENCE

Trinexapac-Ethyl and Iron Effects on Supina and Kentucky Bluegrasses Under Low Irradiance

J.C. Stiera and J.N. Rogers, IIIb

a Dep. of Horticulture, Univ. of Wisconsin, Madison, WI 53706-1590
b III, Dep. of Crop and Soil Sciences, Michigan State Univ., East Lansing, MI 48824-1325

Corresponding author (jstier{at}facstaff.wisc.edu)


    ABSTRACT
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Turf use in covered stadiums and other environments with reduced irradiance is limited due to lack of suitable turf species and management practices. This study compared the tolerance of supina bluegrass (Poa supina Schrad.) and Kentucky bluegrass (P. pratensis L.) with reduced irradiance of approximately 1 to 5 mol m-2 d-1. Treatments included trinexapac-ethyl {[4-(cyclopropyl-{alpha}-hydroxy-methylene)-3,5-dioxo-cyclohexane-carboxylic acid ethyl ester]} (TE), foliar iron, and simulated athletic traffic inside a covered stadium simulator facility. Analysis of variance showed supina bluegrass was more responsive to TE than Kentucky bluegrass. Trinexapac-ethyl reduced supina bluegrass clipping yields approximately 60%; Kentucky bluegrass yields were reduced by 20% or less. In non-trafficked turf, TE increased supina bluegrass tillers by 50% and leaves by 33% but did not change tillering and leaf number of Kentucky bluegrass. Without traffic, TE-treated supina bluegrass provided an acceptable turf at 10 to 15% solar irradiance for at least 4 to 6 mo, while Kentucky bluegrass and untreated supina bluegrass became unacceptable within 2 to 4 mo. Under traffic, TE-treated supina bluegrass provided acceptable turf for up to 5 wk, while Kentucky bluegrass did not provide acceptable turf for more than 2 to 4 wk. Trinexapac-ethyl enhanced supina bluegrass color and increased chlorophyll levels of both species. Foliar applications of iron had negligible effects on all of the parameters evaluated. Supina bluegrass is a useful turf for reduced irradiance situations but requires more than 5 mol m-2 d-1 to sustain traffic for periods longer than 5 wk.

Abbreviations: CSSF, Covered Stadium Simulator Facility • GA, gibberellic acid • LAI, leaf area index • PAR, photosynthetically active radiation • PGR, plant growth regulator • TE, trinexapac-ethyl


    INTRODUCTION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
COMMONLY USED COOL-SEASON TURFGRASSES are thought to have evolved near the margins of forests in Eurasia where light would not have been limited (Beard, 1973). Consequently, most commonly used cool-season turfgrass species have relatively poor shade tolerance. Kentucky bluegrass (Poa pratensis L.) is the most commonly used cool-season turfgrass but its growth can be severely limited in the shade due to insufficient light and enhanced disease susceptibility (Beard, 1973; Vargas and Beard, 1981). Fine fescues (e.g., Festuca rubra L., F. rubra var. commutata Gaud.) and rough bluegrass have better shade tolerance than Kentucky bluegrass but have poor traffic tolerance (Beard, 1973). A relatively shade- and traffic-tolerant cool-season turfgrass species is desirable for athletic fields subjected to reduced irradiance, including covered stadia.

Supina bluegrass is a cool-season stoloniferous turfgrass capable of forming a dense turf at low mowing heights suitable for athletic and golf course turf (Berner, 1980; Pietsch, 1989). The stolons are significantly more robust and have shorter internodes compared with rough bluegrass. Supina bluegrass is endemic to moist, shaded sites, often subject to high traffic (e.g., human and cattle paths) in the sub-alpine regions of the European Alps (Pietsch, 1989). In Germany, supina bluegrass often encroaches and fills in high-wear areas on sports fields (Köck and Walch, 1977). Berner (1980) demonstrated good wear tolerance of supina bluegrass due in part to a rapid recuperative rate.

The ability to persist in moist, shaded, high-traffic environments makes supina bluegrass a suitable candidate for use as a turf for shaded golf course or athletic field situations (e.g., partially or wholly covered stadia). Limitations for the use of supina bluegrass include light green leaf color and undefined management regimes (Berner, 1980). Leaf color is an adjustable parameter that could increase the acceptability of supina bluegrass if a darker color can be easily obtained.

Flurprimidol {{alpha}-(1-methylethyl)-{alpha}-[4-(trifluoromethoxy)phenyl]-5-pyrimidine-methanol}, a plant growth regulator that suppresses gibberellic acid (GA) biosynthesis, can significantly enhance turf color and quality in reduced irradiance (<30% full sunlight) (Stier et al., 1999). Trinexapac-ethyl (TE), the most recently labeled GA inhibitor for turf, reduced clipping yields and enhanced creeping bentgrass quality under normal field conditions (Calhoun and Branham, 1995). Its effects on cool-season turf in reduced irradiance have not been reported.

Turf color decreases as chlorophyll content is reduced under extremely low irradiances (Beard, 1973). Foliar applications of iron have been used to darken turf color (Glinski et al., 1992) and to negate the transient phytotoxicity that can result from plant growth regulator (PGR) application under normal irradiance (Carrow and Johnson, 1990). Lee et al. (1996) showed above-normal levels of iron in nutrient culture enhanced granal development in chloroplasts with a commensurate increase of chlorophyll b levels in Kentucky bluegrass, a situation characteristic of shade-adapted plants (Nobel, 1991). The effect of foliar iron applications, and the potential interaction with PGRs, on turf color and chlorophyll in reduced irradiance is unknown.

Our research objectives were to compare the tolerance of supina bluegrass and Kentucky bluegrass under reduced irradiance, with and without traffic, and to determine the effects of multiple applications of TE and iron on the growth and quality of supina bluegrass and Kentucky bluegrass grown under reduced irradiance.


    MATERIALS AND METHODS
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Experimental Environment
The research was conducted inside the air-supported Covered Stadium Simulator Facility (CSSF) at the Hancock Turfgrass Research Center, East Lansing, MI. Three forced-air furnaces (3.15 J each) heated the CSSF during the winter months. In the spring and summer, when outdoor ambient temperatures were above 17°C, the temperature in the CSSF was maintained at outdoor ambient temperatures using an automated system of ventilation fans and curtains located on the southern and northern ends of the CSSF, respectively. The CSSF was covered in October 1994 with an experimental fiberglass fabric, Sheerfill IV (Chemical Fabrics, Buffalo, NY). Quality of the irradiance transmitted through the Sheerfill IV was equivalent to that transmitted by Sheerfill II (Stier et al., 1999).

Daily totals of photosynthetically active radiation (PAR) in the CSSF were determined, based on the percentage of PAR transmitted through the fabric onto the turf canopy inside the CSSF. A spectroradiometer (model LI-1800, Li-Cor, Lincoln, NE) was used to scan irradiance weekly on each plot inside the CSSF within 1 h of the solar zenith using a 300- to 850-nm bandwith at 2-nm intervals. Two scans were collected outside the CSSF with the spectroradiometer immediately before, midway through, and immediately after collecting data inside the CSSF. Data from inside the CSSF were divided by data collected outside to determine the percent transmission. Average daily PAR values were determined by collecting solar radiation data outside the CSSF daily with a pyranometer (model Li-Cor PY 14226, Li-Cor, Lincoln, NE) located approximately 15 m north of the CSSF. Radiometric units from the pyranometer were converted to quantum units using the following equation, which was based on conversion units from Thimijan and Heins (1983):

(1)

The average percentage of light transmitted into the CSSF was used to determine the daily PAR inside the CSSF, based on the data recorded outside with the pyranometer.

Temperature and relative humidity were recorded daily with a sling psychrometer. Temperature inside the CSSF averaged 16.6°C ± 1.5°C with a range of 12 to 24°C. Relative humidity averaged 46% ± 12% with a range of 28 to 63%.

Plot Construction and Maintenance
Portable plots were established as previously described (Stier et al., 1999). The project was conducted twice from December 1994 through June 1996. A first set of plots was sodded 28 Sept. 1994 and a second set was sodded 28 Aug. 1995. Starter fertilizer was applied to the root zone mix surface immediately before sodding (Table 1). Additional fertilizer was applied twice in 1994 and once in 1995 during establishment. The turf was mowed at 3-cm height once to twice weekly, depending on growth rate, and irrigation was applied as necessary to prevent visible drought stress.


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Table 1. Root zone mixture conditions and fertility applications during establishment.{dagger}

 
Plots were moved into the CSSF for testing from 12 Dec. 1994 through 12 Apr. 1995 and from 8 Dec. 1995 through 11 June 1996. A reel mower was used to maintain turf height at 3 cm. The turf was mowed once to twice weekly to prevent removal of more than one-third of the leaf tissue. Plots were fertilized at 30-d intervals with 24, 2, and 20 kg ha-1 N, P, and K, respectively (18-3-18). Approximately 1.25 cm of water was applied immediately following fertilizer application. Additional irrigation was applied as necessary to prevent drought stress (approximately 1.25 cm at 7- to 14-d intervals). Industrial fans were occasionally used for 24- to 72-h periods to dry the turf surface following irrigation to discourage fungal pathogen activity. Iprodione [3-(3,5-dichlorophenyl)-N-(1-methylethyl)-2,4-dioxo-1-imidazolidinecarboximide] was applied to all plots on 23 Dec. 1994 (3 kg ha-1), 6 Mar. 1995 (6 kg ha-1), and 14 Apr. 1995 (6 kg ha-1) to control Microdochium patch [Microdochium nivale (Fr.) Samuels and I.C. Hallett]. Fungicides were not applied in 1995–1996; instead, fans were used to dry the turf surface when mycelia were noticed.

Treatments and Experimental Design
A completely randomized design with three replications was used, since there was no apparent spatial or environmental variability that required blocking. Data were analyzed as a 2 x 2 x 2 factorial using MSTAT analysis of variance procedures (MSTAT, 1988). The factors were turfgrass species, TE, and foliar iron. When appropriate, means for interactions were separated using Fisher's Protected LSD ({alpha} = 0.05).

Supina bluegrass cv. Supranova and Kentucky bluegrass cv. Victa/Abbey (50:50 v/v) were used both years. Washed sod was used for establishment except in 1994, when supina bluegrass, produced on a plastic surface in composted bark media, was used. Trinexapac-ethyl at 0.19 kg ha-1 in 896 L of H2O ha-1 was applied on 3 Oct. 1994 and 9 Oct. 1995, using a CO2-powered backpack sprayer equipped with 8002 flat fan nozzles. Additional TE (0.08 kg ha-1 in 896 L of H2O ha-1) was applied on 21 Dec. 1994, 20 Jan. 1995, 18 Feb. 1995, and 16 Mar. 1995 for the first year's testing and on 31 Jan. 1996, 15 Mar. 1996, and 26 Apr. 1996 for the second year's testing. Iron (1.14 kg of Fe ha-1 as FeSO4·7H2O) was applied to foliage using Ferromec AC (PBI Gordon, Kansas City, MO) on 10 Jan. 1995, 14 Feb. 1995, 17 Mar. 1995, 28 Feb. 1996, and 13 May 1996.

Two experiments were designed to determine treatment effects in both non-trafficked (Exp. I) and trafficked (Exp. II) conditions. Traffic was applied by having a person (approximately 70–75 kg) jog 50 passes at 7-d intervals while wearing molded soccer cleats. Traffic was applied 28 Dec. 1994 through 16 Mar. 1995 (600 passes) and 26 Jan. 1995 through 26 Apr. 1996 (700 passes).

Data Collection
Clippings were collected from a 41- by 117-cm strip through the center of each trafficked and non-trafficked plot, dried in a forced-air oven at 60°C for 48 h, then weighed. Clippings were collected from the remainder of the plot area and discarded. Turf color and quality were evaluated visually on a one-to-nine scale at approximately 14- to 21-d intervals. A rating of one represented 100% necrotic turf/bare soil; a rating of nine represented dark green or ideal turf. A rating of five was considered the minimum acceptable value. Turf rooting and strength were evaluated periodically using an Eijkelkamp shear vane apparatus (Eijkelkamp, Giesbeek, the Netherlands). The torque required to tear the turf with the shear vane was recorded as an average of two measurements per plot (Rogers and Waddington, 1990). On 24 March 1995 and 30 May 1996, plant densities were determined by counting the number of plants in eight random squares (32.7 cm2 each) of a 0.4-m2 quadrat (Skogley and Sawyer, 1992). On 12 Apr. 1995 and 31 May 1996, samples of 10 randomly selected plants were collected from each plot for biomass assessments. Average leaf number shoot-1, average shoot number plant-1, and average oven-dry weight plant-1 (specific weight) were determined for each sample.

Leaf samples from 10 randomly selected plants were collected from each non-trafficked plot for chlorophyll analysis on 4 Apr. 1995 and 29 May 1996 (trafficked plots were not sampled because adequate plant material was often not available). A 10-mm segment from the youngest fully expanded leaf of each plant was excised, starting 5 mm above the leaf collar. Chlorophyll was extracted by placing 10 leaf segments in 3 mL of N,N-dimethlyformamide and incubating them in the dark at 4°C for 48 h (Moran and Porath, 1980). A double-beam spectrophotometer was used to determine absorbances, and the extinction coefficients described by Inskeep and Bloom (1985) were used to calculate levels of chlorophyll a, b, and total chlorophyll.


    RESULTS AND DISCUSSION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS AND DISCUSSION
 REFERENCES
 
Environmental Data
Photosynthetically active radiation inside the CSSF tripled between December (0.9–1.5 mol m-2 d-1) and April (2.8–4.5 mol m-2 d-1) of each year due to increased solar radiation between the winter solstice and the vernal equinox (Table 2). Extensive solar bleaching of the Sheerfill IV fabric covering the CSSF occurred during spring and summer 1995, increasing solar transmission from 10.5% during the first season to more than 15% during the second season. Irradiances of 5 mol m-2 d-1 and greater inside the CSSF were not reached until after the vernal equinox of the second year.


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Table 2. Mean values of photosynthetically active radiation (PAR){dagger} outside and inside the Covered Stadium Simulator Facility (CSSF) at the Hancock Turfgrass Research Center, East Lansing, MI

 
These data are crucial because the bleaching requirement of the fiberglass fabric for maximal solar transmission could limit the use of natural turfgrass in stadiums covered with fiberglass fabric following the first year of construction. When GA inhibitors were not used, a minimum of 5 to 9 mol m-2 d-1 were required to maintain acceptable Kentucky bluegrass turf for periods of 2 mo or longer when subjected to light traffic (Stier et al., 1999). Even though 15% solar transmission is considered low irradiance (McBee and Holt, 1966), the 5% increase between years improved turf performance, particularly of the supina bluegrass. The ultra-low PAR of the winter months may restrict the long-term use of natural grass for covered stadiums.

Turf Quality
Experiment I: Turf Not Subjected to Traffic
The main effects of both species and TE were significant on most rating dates, but their individual importance was overridden by interaction between the two factors (Table 3). The species x TE interaction significantly affected turf quality on 5 of 7 dates in 1994–1995 and on 9 of 10 dates in 1995–1996 (Table 3). The species effect was inconsistent between years due to differences in disease susceptibility and reaction to TE. Although TE affected the turf quality of both species, supina bluegrass was more responsive than Kentucky bluegrass. Supina bluegrass treated with TE was the only species treatment combination to provide acceptable turf quality for the duration of the test periods (16 wk in 1994–1995 and 27 wk in 1995–1996) (Fig. 1) . Unlike previous reports of TE-induced phytotoxicity on turf subjected to stress (Quian and Engelke, 1999), no phytotoxicity was observed due to TE applications. This was probably due to the relatively low rates of TE.


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Table 3. Significance of treatment effects on quality, color, yield, and shear resistance of non-trafficked turfgrass in reduced-irradiance conditions, East Lansing, MI

 


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Fig. 1. Interaction of trinexapac-ethyl (TE) and species on turf quality in reduced-irradiance conditions during 1994–1995 (a) and 1995–1996 (b), East Lansing, MI. Quality was rated visually on a one-to-nine scale: 1 = 100% dead turf/bare soil; 9 = dense, uniform turf; 5 was the minimum value for acceptable athletic field turf. LSD values are for comparing within or between species or TE levels

 
Turf quality of untreated supina bluegrass was usually not better than untreated Kentucky bluegrass during the first year (Fig. 1a). During the second year, powdery mildew (Erysiphe graminis DC.) caused a significant thinning of the Kentucky bluegrass. Supina bluegrass appeared resistant to the disease. Microdochium patch was more prevalent on Supina bluegrass than Kentucky bluegrass (data not shown), but the disease was controlled with fungicide and by using fans to dry the turf.

These data indicate that untreated supina bluegrass is not necessarily more tolerant of reduced irradiance than Kentucky bluegrass at <=5 mol m-2 d-1. A companion study in the CSSF during 1995–1996 using supplemental irradiance (5–9 mol m-2 d-1) showed untreated supina bluegrass was superior to untreated Kentucky bluegrass (unpublished data, 1996). Previous reports of supina bluegrass' superior shade tolerance did not indicate the amount of irradiation (Berner, 1980; Pietsch, 1989).

The sensitivity of supina bluegrass to TE had both negative and positive impacts. For example the turf, which was semi-dormant when brought into the CSSF in both years, recovered within 2 wk except in year two when greenup of the TE-treated supina bluegrass took 4 wk. Daytime temperatures, which dropped to near 0°C, and the occurrence of snow cover within 1 mo after the first application of TE in the second season (during October) seemed to delay the metabolism of TE in the supina bluegrass. The delayed greenup in the second year is important because turf brought into a covered stadium for repair of damaged areas will need to be able to quickly recover from winter dormancy. Thus PGR applications made during the autumn should be applied well in advance of potential winter dormancy so as not to inhibit recovery. Additional work is needed to determine metabolic rates of TE during hardening and de-hardening periods.

Experiment II: Turf Subjected to Traffic
As in the non-trafficked turf, the interaction between species and TE had significant impact on most rating dates with TE-treated supina bluegrass providing the best turf on 7 of 13 dates during both years of the study. On a practical basis, during both years, approximately 5 wk after initiating traffic (200 jogging passes), turf quality was reduced to unacceptable levels for all treatments (Fig. 2) . Supina bluegrass treated with TE provided higher quality turf for the first 4 wk after traffic was started in 1994. Similar differences were not statistically significant during the second year, perhaps because traffic was initiated 7 wk (instead of 2 wk) after installation in the CSSF to allow TE-treated supina bluegrass to recover from dormancy. Supina bluegrass treated with TE recovered more quickly than Kentucky bluegrass or untreated supina within 3 wk after traffic ceased in 1996 (Fig. 2b). Although turf quality did not reach acceptable levels 8 wk after traffic ceased, under greater irradiances supina bluegrass treated with TE could be expected to recover to acceptable levels faster than untreated supina bluegrass or Kentucky bluegrass.



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Fig. 2. Interaction of trinexapac-ethyl (TE) and species on quality of turf subjected to traffic in reduced-irradiance conditions during 1994–1995 (a) and 1995–1996 (b), East Lansing, MI. Quality was rated visually on a one-to-nine scale: 1 = 100% dead turf/bare soil; 9 = dense, uniform turf; 5 was the minimum value for acceptable athletic field turf. LSD values are for comparing within or between species or TE levels. Plots were installed inside the Covered Stadium Simulator Facility (CSSF) on 12 Dec. 1994 and 8 Dec. 1995. Traffic was applied by a person wearing soccer cleats who jogged across the turf 50 passes weekly from 28 Dec. 1994 (2 wk after installation) to 16 March 1995 (600 passes) and 26 Jan. 1996 (7 wk after installation) to 26 April 1996 (700 passes)

 
Turf Color
Kentucky bluegrass had significantly darker green color compared with supina bluegrass (avg. rating of 6.4 vs. 5.3, respectively, on a one-to-nine scale) on 12 of 18 rating dates between 1994 and 1996 (data not shown). Trinexapac-ethyl darkened the color of both species on most rating dates (Tables 3 and 4), but interactions between TE and species showed the color of supina bluegrass was enhanced to a significantly greater degree than Kentucky bluegrass. Averaged over both years, TE increased the green color rating of non-trafficked supina bluegrass from 3.4 ± 0.9 to 6.3 ± 0.4 and of trafficked supina bluegrass from 3.0 ± 1.3 to 5.0 ± 1.6. In Kentucky bluegrass, TE increased the green color rating of non-trafficked turf from 5.2 ± 1.2 to 6.7 ± 0.8. In trafficked Kentucky bluegrass, the green color rating averaged 4.7 ± 1.1 for untreated turf and 5.6 ± 1.8 for TE-treated turf.


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Table 4. Significance of treatment effects on quality, color, yield, and shear resistance of turfgrass subjected to traffic in reduced-irradiance conditions, East Lansing, MI

 
Color enhancement from TE and other GA inhibitors may result from the concentration of chloroplasts due to decreased leaf elongation. Flurprimidol, another GA inhibitor, was previously reported to enhance Kentucky bluegrass color under similarly reduced irradiance (Stier et al., 1999). The degree of color adjustment can depend on rates of the GA inhibitor (Stier et al., 1999), application frequency (Quian and Engelke, 1999), and nitrogen fertility (unpublished data, 1996).

Chlorophyll Content
Trinexapac-ethyl increased the amount of chlorophyll on a leaf-area basis from 26.7 to 36.7 µg cm-2 (Table 5). Kentucky bluegrass averaged 35.9 µg of total chlorophyll cm-2 leaf area vs. 27.4 µg cm-2 in supina bluegrass. Unlike the results seen for color, there was no species x TE interaction on chlorophyll content.


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Table 5. Species and trinexapac-ethyl effects on chlorophyll (Chl) content of turfgrasses in reduced-irradiance conditions, East Lansing, MI

 
The ratio of chlorophyll a:b was 3.1 for both species. Trinexapac-ethyl apparently decreased the ratio from 3.2 to 3.0 in 1995 but not in 1996 (Table 5). A ratio of approximately 3.1 is typical of sun-adapted plants (Nobel, 1991). In shade-adapted leaves, the increased stacking of grana reduces the ratio of Photosystem I to Photosystem II, causing a decrease in the chlorophyll a:b ratio since Photosystem II has a high proportion of chlorophyll b compared with Photosystem I. Apparently none of the treatments increased granal stacking to a degree that would dramatically enhance shade tolerance.

Clipping Yields
Species, TE, and species x TE interaction resulted in significant differences in clipping yields on more than half the dates (Tables 3 and 4). Trinexapac-ethyl reduced yields of both turfgrasses in the first year but reduced yields of only supina bluegrass in the second year (data not shown). Averaged over both years, TE reduced supina bluegrass yields 61% in non-trafficked turf (99.7 g cm-2 total yield for untreated turf vs. 38.7 g cm-2 for TE-treated turf) and 56% in trafficked turf (39.1 g m-2 total yield for untreated turf vs. 17.4 g cm-2 for TE-treated turf). Kentucky bluegrass yields were reduced 20% (105.2 g m-2 total yield for untreated turf vs. 83.9 g cm-2 for TE-treated turf) and 14% (50.1 g m-2 total yield for untreated turf vs. 38.0 g cm-2 for TE-treated turf) for non-trafficked and trafficked turf, respectively. Some of the difference between species yields was due to the prostrate growth habit of supina bluegrass, which was enhanced by TE.

During the first year, TE decreased supina bluegrass yields to zero (data not shown) approximately 10 wk after installation in the CSSF when irradiance ranged from approximately 1 mol m-2 d-1 during December and January to 2.6 mol m-2 d-1 in March. In the trafficked plots, supina bluegrass yields declined to zero within 4 wk after traffic began (approximately 6 wk after installation in the CSSF). Although acceptable turf quality was maintained on the non-trafficked plots during the period of non-yield, quality of the trafficked plots plummeted due partly to excessively suppressed growth. During the second year when irradiance was greater, a reasonable amount of growth was maintained even in the trafficked plots, with oven-dry weights ranging from approximately 0.5 to 4.3 g m-2.

Plant Density and Biomass
Experiment I: Turf Not Subjected to Traffic
The two-way interaction between species and TE was significant for plant density, tillering, leaf number, and specific weight (Table 6). Trinexapac-ethyl significantly enhanced plant density, tillering, leaf number, and specific weight of supina bluegrass but not of Kentucky bluegrass. Even without TE, supina bluegrass had approximately two to three times the number of tillers and leaves per plant compared with Kentucky bluegrass (Table 6). In 1996 supina bluegrass had significantly greater plant density.


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Table 6. Treatment effects on turf density and development of non-trafficked supina (SB) and Kentucky (KB) bluegrasses in reduced-irradiance conditions, East Lansing, MI

 
The developmental response in supina bluegrass may have been partly due to a redistribution of carbohydrates (Hanson and Branham, 1987) which increased tillering (Foreman and Marshall, 1990). Trinexapac-ethyl likely increased the number of leaves by reducing the leaf-extension period and plastochron, thereby accelerating leaf production (Foreman and Marshall, 1990). Foreman and Marshall (1990) reported an experimental GA inhibitor caused an increase in leaf width, a result also observed with TE under reduced irradiance (Stier, unpublished data, 1996). The increased leaf area index (LAI) resulting from increased leaf width and number would increase carbohydrate production (Quian and Engelke, 1999) and growth. The reaction was likely greater in supina bluegrass partly due to its prostrate growth habit, which would allow more tillers and leaves to form without being removed by mowing. In our study, untreated turf, particularly Kentucky bluegrass, exhibited a spindlier, more upright growth habit and may have exhausted carbohydrate reserves by cell and shoot elongation without benefitting from an LAI resulting from the TE application.

A powdery mildew epidemic greatly reduced Kentucky bluegrass turf density the second year of the study. Turf density of both species was much lower the second year compared with the first year since plant counts were conducted 24 wk after installation of the plots in the CSSF; in the first year, counts were conducted at 15 wk. The decline in plant population over the longer time period indicates a lack of sufficient irradiance for long-term growth of either species.

Experiment II: Turf Subjected to Traffic
Similar to non-trafficked turf, the main species effect showed supina bluegrass had more tillers, leaves, and a greater specific plant weight compared with Kentucky bluegrass (Table 7). Unlike the results seen in non-trafficked turf, the main effects of TE did not affect tillering, leaf number, and specific plant weights of either species when turf was subjected to traffic. Trinexapac-ethyl did increase plant density. The only species x TE interaction was an increase in the density of TE-treated supina bluegrass in 1996 but not in Kentucky bluegrass turf.


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Table 7. Treatment effects on turf density and development of supina (SB) and Kentucky (KB) bluegrasses when subjected to traffic{dagger} in reduced-irradiance conditions, East Lansing, MI

 
The visible effects of TE on tillering, leaf number, and specific plant weights may have been nullified as emerging tillers and leaves were destroyed by routine traffic. However, the greater plant density of TE-treated turf may have been due to increased wear tolerance, which slowed the destructive force of traffic. Shearman and Beard (1975) showed total cell wall content was correlated to turfgrass wear tolerance. Foreman and Marshall (1990) showed N-t-butylcarbonyl-N-(4'-chlorophenyl methyl) aminopyrazine, another GA inhibitor, reduced leaf length of perennial ryegrass (Lolium perenne L.) without altering mesophyll cell number. Since there is no suggested or reported evidence that TE reduces cell wall deposition, suppression of cell enlargement by TE (King et al., 1997) may have increased the relative total cell wall content. This phenomenon could enhance wear tolerance and account for the increased rigidity of TE-treated turf compared with untreated turf (unpublished data, 1993–1996). An unexpected benefit of such a phenomenon was the higher quality of cut we observed in TE-treated turf, while untreated turf was flaccid and was often bent, torn, or cut nonuniformly (unpublished data, 1993–1996). The lack of turf-density response to TE treatments by Kentucky bluegrass was due to the overwhelming effects of powdery mildew, which destroyed much of the Kentucky bluegrass turf.

Turf Shear Resistance
There were significant main effects on shear resistance due both to species and TE but no meaningful interactions (Table 8). Kentucky bluegrass exhibited greater shear resistance than supina bluegrass during the first year; supina bluegrass had greater resistance the second year, although the shear resistance of Kentucky bluegrass was comparable in January and March of both years. Trinexapac-ethyl significantly enhanced shear resistance on three of seven test dates but the magnitude of response was much less than that observed between species.


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Table 8. Main effects of turf species and trinexapac-ethyl on shear resistance (Nm) of non-trafficked turfgrass in reduced-irradiance conditions, East Lansing, MI

 
Rogers and Waddington (1990) attributed the relatively high shear resistance of Kentucky bluegrass to the presence of rhizomes, a feature not shared by supina bluegrass, which helps explain the greater values achieved by Kentucky bluegrass during 1994–1995. During the second year, two events occurred that favored the shear resistance of supina bluegrass: (i) The severe powdery mildew infection and concomitant decline in Kentucky bluegrass turf density probably reduced shearing resistance. (ii) The supina bluegrass was able to take advantage of the higher irradiance in the second year to produce a thicker turf with more stolons during the months of January and March. Regardless of treatment, however, shear strength of both species declined over time in both years. The decline in shear resistance indicated a lack of sufficient turf growth and rooting, probably due to insufficient irradiance to sustain turf permanently. The occasionally greater shear resistance produced by TE may have been due to enhanced root growth (Rogers, unpublished data, 1998). Trafficked turf had similar shear strength compared with non-trafficked turf (data not shown).

Adequate shear strength is important to minimize the amount of tearing and size of divots removed during play. Appropriate shear strength is critical for maximum playability and player safety since too little shear strength can cause athletes to slip and perhaps injure muscles; too much shear strength can result in turf toe. Unfortunately, only anecdotal data are available to indicate acceptable and unacceptable shear values, and these can depend on the type of shear vane used to measure the turf. In our trials, we designated a value of 10 Nm as the minimum acceptable value since the turf was easily torn from the soil at values <10 Nm. Values between 10 to 15 were considered fair, values 15 to 20 were considered good, and values >20 were exceptional. With these criteria we can make the following observations. In the first year when transmission of solar irradiance was restricted to approximately 10%, supina bluegrass provided only acceptable shear strength for approximately 30 d. Kentucky bluegrass' shear strength was acceptable for at least 90 d the first year, although turf quality was not acceptable for more than 30 d (20 d when trafficked). In the second year, when the fabric transmitted approximately 15% of solar irradiance, supina bluegrass provided acceptable shear strength for at least 90 d, which was similar on a practical basis to Kentucky bluegrass.

Effects of Iron and Interactions with Species and Trinexapac-Ethyl
Interactions between iron and species, iron and TE, and three-way interactions were rare and deemed insignificant, based on their relative effect in each situation. The response due to iron was significant with respect to quality, color, and yield on only a few dates (Tables 3 and 4). Iron did not affect turf density, growth, or development. Contrary to expectations and results obtained under normal irradiance, iron did not affect total chlorophyll level. Iron treatment did increase chlorophyll b from 6.7 to 7.3 µg cm-2 in the non-trafficked experiment on 29 May 1996, but the lack of effect on chlorophyll a:b and chlorophyll b levels of both the non-trafficked and trafficked (data not shown) turf in 1995 suggest iron had negligible effect. The lack of iron effect on chlorophyll and the chlorophyll a:b was unexpected since previous work under normal irradiance showed iron was capable of increasing chlorophyll and decreasing chlorophyll a:b in turf (Lee et al., 1996). The lack of effect on chlorophyll content indicates either the iron did not enter the foliage or was not utilized by the turf for chlorophyll production. The extremely low irradiance in the current study may have compromised the ability of the turf to utilize the foliar iron.

Received for publication September 21, 1999.


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