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Quantifying a Daily Light Integral Requirement of a ‘TifEagle’ Bermudagrass Golf Green

^a Dep. of Horticulture, Clemson Univ., Clemson, SC 29634-0375
^b Dep. of Experimental Statistics, Clemson Univ., Clemson, SC 29634-0375

^* Corresponding author (toddb{at}sepro.com)

	ABSTRACT

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

 
‘TifEagle’ bermudagrass [Cynodon dactylon (L.) Pers. x C. transvaalensis Burtt-Davy] is a fine textured hybrid bermudagrass used for golf course putting greens in the subtropical and tropical regions of the world. The growth and performance of TifEagle and other bermudagrass often decline in reduced light environments (RLEs). This study quantified a daily light integral (DLI; mol m^–2 d^–1) required to maintain commercially acceptable TifEagle bermudagrass maintained under golf green conditions. Three levels of shade, none, low, and high (0, 41, and 92%), were applied to mature TifEagle bermudagrass during morning (sunrise to 1100 h) and afternoon (1500 h to sunset) hours, allowing full irradiance from 1100 to 1500 h to mimic a golf green situation where trees or structures are adjacent to, but rarely directly overhead. The study was performed for 8 wk in June to August 2001 and 2002. An average minimal DLI of 32.6 mol m^–2 d^–1 was needed to maintain commercially acceptable TifEagle turf quality (TQ 7). Other plant responses measured included percentage lateral regrowth (RG), total shoot chlorophyll, and total nonstructural carbohydrates (TNC). These responses declined significantly when the DLI 32.6 mol m^–2 d^–1. Differences in diurnal shade exposure occurred. High afternoon shade reduced TifEagle bermudagrass TQ, percentage lateral RG, shoot chlorophyll, and TNC by 3.0 rating units and 17, 39, and 27%, respectively, compared with no afternoon shade (NSA). High morning shade reduced TQ, percentage lateral RG, and shoot chlorophyll by 1.5 rating units and 11 and 16%, respectively, compared with no morning shade (NSM). Overall, afternoon shade applications were more detrimental to TifEagle bermudagrass growth and performance compared with morning shade.

Abbreviations: AH, afternoon high shade • AL, afternoon low shade • DLI, daily light integral • MH, morning high shade • ML, morning low shade • NSA, no afternoon shade • NSM, no morning shade • PPF, photosynthetic photon flux • RE, regrowth • RLE, reduced light environment • TQ, turf quality • WAS, weeks after shade initiation

	INTRODUCTION

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

 
DWARF-TYPE BERMUDAGRASSES such as TifEagle are becoming an increasingly popular choice for golf putting greens in the southern USA. These grasses provide an excellent putting surface when exposed to sufficient irradiance. Surrounding trees and other tall objects cause a RLE, which lowers TQ of shade-sensitive bermudagrass. Diurnal variations in irradiance and resulting turfgrass quality have been long observed by turfgrass managers; however, little published results are available. The majority of research on C₄ grasses and shade has been performed by applying continuous shade, allowing various percentages of photosynthetic photon flux (PPF) (Kephart et al., 1992; Gaussoin et al., 1988; Miller and Edenfield, 2002; Ervin et al., 2002).

Continuous shade studies are more applicable to turfgrass areas maintained directly below a tree canopy or closely adjacent to a building structure when only diffuse irradiance reaches the turf. In nature, golf greens rarely have continuous overhead shade; therefore, direct and diffuse irradiance reaches turfgrass leaves at various intervals surrounding solar noon hours. In the northern hemisphere, existing trees to the east and/or west reduces irradiance during morning and afternoon hours, respectively. However, no reports exist examining the importance of morning and afternoon irradiance to the growth and development of bermudagrass putting greens. Morning irradiance has been long associated with healthy and high performing turfgrass, especially in cool-season grasses. Creeping bentgrass [Agrostis stolonifera var. palustris (Huds.) Farw.] surrounded by trees to the east and southeast have been observed by turfgrass managers to decline during summer months due to lack of morning irradiance (Bell and Danneberger, 1999). In addition to reduced light, areas shaded during morning hours have higher relative humidity and prolonged periods of dew and leaf wetness, both of which encourages disease and added stress (Bell and Danneberger, 1999). A study performed in Ohio applied 6 h of morning and/or afternoon shade to creeping bentgrass with 80 and 100% shade cloth. Morning or afternoon shade at either level had no detrimental effects on bentgrass quality (Bell and Danneberger, 1999). Therefore, in cool-season grasses, morning shade did not prove more physiological detrimental than afternoon shade.

Unlike cool-season grasses, which reach light saturation at approximately 1/2 full sunlight, C₄ grasses such as bermudagrass require full sunlight to reach maximum photosynthetic capacity (McCarty, 2001). Therefore, previous research with cool-season grasses is not directly applicable to dwarf-type bermudagrasses. Limited RLE research exists on a bermudagrass golf putting green where trees are positioned adjacent and rarely directly above. The objective of this research was to quantify the DLI requirement of a dwarf bermudagrass golf green by exposure to various shade applications during morning and afternoon hours.

	MATERIALS AND METHODS

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

 
The study was conducted during summers of 2001 and 2002 on a TifEagle bermudagrass research green at Clemson, SC (34°40' N, 82°49' W). The Year 1 study was conducted from 26 June to 22 August 2001 and Year 2 from 19 June to 15 August 2002. The green was sprigged in June 1999 with 2.6 m³ ha^–1 of certified TifEagle bermudagrass. Soil profile construction and soil physical properties followed United States Golf Association recommendations with a 85:15 (v/v) sand/peat mix (United States Golf Association Green Section Staff, 1993).

Three levels of shade, none (0%), low (41%), and high (92%), were applied during morning hours and afternoon hours daily for 8 wk with treatments arranged in a 3-by-3 factorial design (Table 1). Morning hours were deemed sunrise to 1100 h and afternoon hours from 1500 h to sunset. Morning and afternoon shade applications were spaced equally approximately 1 to 2 h from solar noon, which ranges from 1300 to 1400 h during summer months in Clemson, SC. This was performed to ensure all plots received direct irradiance when intensity was greatest. Low and high shade applications used a neutral density, polyfiber black shade cloth allowing 59 and 8% full irradiance, respectively (model no. SC-BL40 and SC-BL90; International Greenhouse, Sidel, IL). Light quality measurements were made with a spectroradiometer (LI-1800; LiCor, Inc., Lincoln, NE) under shade tents. No differences in red/far-red were detected between shade treatments. Percentage shade of each shade cloth was determined by comparing PPF (µmol m^–2 s^–1) under shade cloths at the turf canopy to full-irradiance measurements with a LI-190SA quantum sensor (LiCor, Inc.). Measurements were taken twice during each year on a clear, cloud-free day before morning shade tent removal and following afternoon tent application. All shade treatments were removed at 1100 h and reapplied at 1500 h, therefore allowing direct irradiance from 1100 to 1500 h.

Measurements
Hourly light integrals were recorded in a LI-1000 (LiCor, Inc.) datalogger fitted with a LI-190SA (LiCor, Inc.) quantum sensor programmed to collect readings every minute. Daily light integrals for shade treatments (sunrise to 1100 h, and 1500 h to sunset) were calculated by taking 8% (high shade) and 59% (low shade) of PPF for the shade period. Daily light integrals (mol m^–2 d^–1) were averaged across the growing season for both years. Standard errors were calculated for yearly and 2-yr DLI averages for each treatment.

Visual ratings of TQ, assessing color, density, uniformity, and aesthetic appeal, were determined weekly. Turf quality was rated on a 1-to-9 scale with 9 = best TQ. Unacceptable TQ was deemed <7.

Percentage lateral RG was evaluated by removing 10.8-cm-diam. TifEagle bermudagrass plugs at the initiation of each study from each replicate. Holes were backfilled with a similar sand media. A wire mesh grid was constructed equal to the dimension of the original hole. The grid contained 230 square holes, 0.4 cm² in area. A green shoot present in a 0.4-cm² square denoted a point. Percentage lateral RG was calculated weekly by [green shoot points/total squares (230)]. Weekly percentage lateral RG and TQ data were averaged to yield overall RG and TQ ratings during the 8-wk shade application.

Total shoot chlorophyll concentration was measured 4 and 8 weeks after shade initiation (WAS) for both years. Fresh clippings were harvested and collected during mowing from individual replicates and chlorophyll extracted using dimethyl sulfoxide (DMSO) (Hiscox and Israelstam, 1979). The DMSO technique extracts chlorophyll from shoot tissue without grinding or maceration (Hiscox and Israelstam, 1979). Total shoot chlorophyll concentration (mg g^–1 fresh) was determined using spectrophotometer (Beckman DU-64, Beckman Instruments, Inc., Fullerton, CA) absorbance values at 645 and 663 nm in the equation proposed by Arnon (1949).

The TNC (mg g^–1) of below ground tissue, including rhizomes and adjacent roots, which are the primary carbohydrate storage organs for bermudagrasses, were measured at the end of the study (8 WAS) for both years. Below ground tissues were harvested using a 5-cm-diam. plugger to a depth of 6.5 cm. Two samples were taken per individual replicate before sunrise to minimize diurnal fluctuations in carbohydrates (Westhafer et al., 1982). Invertase (β-D-fructofuranoside fructohydrolase, 433 units mg^–1; Sigma I-4753) and amyloglucosidase (1,4--D-glucan glucohydrolase, 23000 units g^–1; Sigma A-7255) (Sigma Chemical Co., St. Louis, MO) were added to convert sucrose to glucose and fructose moieties and starch to glucose, respectively. The TNC was measured by using Nelson's Assay, which quantifies the reducing sugars, glucose and fructose, in plants (Nelson, 1944; Somogyi, 1945).

Statistical Design and Analysis
The study was a RCB design with treatments arranged in a 3-by-3 factorial treatment design. Morning and afternoon shade represented the two factors with three levels of shade. Experiments were conducted during the summers of 2001 and 2002. Each yearly run had three replications of each treatment combination. Plots were 1 by 1.5 m in dimension with a 1-m buffer zone between plots to prevent shade overlap.

Data were analyzed using ANOVA. The general linear model procedure (GLM) of SAS (SAS Institute, 1987) was used for all these calculations. SigmaPlot program (SPSS Inc., Chicago, IL) was used for data subjected to regression analysis. Treatment x year interactions were not significant; therefore, data were pooled across the 2 yr. Mean separations were performed using Fisher's LSD at the = 0.05 level.

	RESULTS AND DISCUSSION

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

 
Plant responses (TQ, lateral RG, chlorophyll, and TNC) are reported as the treatment combinations of the levels of morning and afternoon shade. Main effect means of morning or afternoon shade were also analyzed.

Daily Light Integral
Yearly and 2-yr combined totals are shown in Table 2. Two-year averages ranged from 41.6 to 22.1 mol m^–2 d^–1, depending on shade application. Main effect DLI means were calculated to determine light quantity based on diurnal shade applications (Table 3). Main effect morning DLI ranged from 35.8 to 28.3 mol m^–2 d^–1 in no-shade and high-shade plots, respectively. Main effect afternoon DLI ranged from 37.8 to 26.0 mol m^–2 d^–1 in no-shade and high-shade plots, respectively (Table 3).

Main effect means of morning and afternoon shade applications demonstrated differences between diurnal light interruptions. Low and high morning shade reduced TQ by 0.5 and 1.5 rating units, respectively, compared with NSM (Table 3). Plots receiving high morning shade maintained an unacceptable TQ rating of 5.8. During afternoon hours, TQ was reduced by 1.0 and 3.0 rating units with low and high shade applications, respectively, compared with NSA. High afternoon shade received an overall TQ rating of 4.9 (Table 3). High shade applications on TifEagle bermudagrass during both morning and/or afternoon hours were extremely detrimental to TQ. Because of a lower afternoon main effect TQ mean (4.9) compared with high morning shade (5.8), it can be concluded that afternoon shade is potentially more detrimental (or afternoon irradiance is more important) to growing commercially acceptable TifEagle bermudagrass under golf green conditions.

TifEagle bermudagrass TQ ratings were plotted against 2-yr DLI to quantify a DLI requirement (Fig. 1) . According to the model, a DLI point estimate of 32.6 mol m^–2 d^–1 was required for TifEagle bermudagrass mowed at 3.2 mm in Clemson, SC, during the months of June to August to maintain a TQ 7 (Fig. 1). The 95% confidence interval for the point estimate was 31.9 to 33.9 mol m^–2 d^–1.

View larger version (14K): [in this window] [in a new window]   Fig. 1. Turf quality response of TifEagle bermudagrass to various daily light integrals during the summers of 2001 and 2002.

Percentage lateral RG main effect means again demonstrated the negative effects of high shade application to TifEagle bermudagrass. High shade in the morning decreased percentage lateral RG by 11% compared with NSM (Table 3). Likewise, high afternoon shade decreased percentage lateral RG by 17% compared with NSA or low afternoon shade. Percentage lateral RG differences did not occur between no shade and low shade in both morning and afternoon applications.

Total Shoot Chlorophyll
Total shoot chlorophyll was measured at 4 and 8 WAS during both years. A treatment x year interaction occurred (P = 0.001) at 8 WAS; therefore, data is presented in separate years. No treatment x year interaction occurred at 4 WAS.

At 4 WAS, highest total shoot chlorophyll concentration of 3.12, 3.06, and 2.93 g kg^–1 occurred with ML/NSA, NSM/NSA, and MH/NSA, respectively (Table 2). Many studies have demonstrated increased chlorophyll concentration with reduced irradiance (Beard, 1997). Low shade applications (35% shade) on cool-season or C₃ turf species, such as perennial ryegrass (Lolium perenne L.), have shown an increase in chlorophyll concentration (Van Huylenbroeck and Van Bockstaele, 2001). Additionally, a study performed on Kentucky bluegrass (Poa pratensis L.) and red fescue (Festuca rubra subsp. rubra) noted an increase in chlorophyll as light intensity decreased (Wilkinson and Beard, 1973). These studies indicate increased chlorophyll with low shade applications to cool-season turfgrass; however, in bermudagrass, chlorophyll increases may not occur in shade because of greater sunlight requirements of warm-season turfgrasses.

In this study, low or high shade applications during morning hours combined with NSA did not decrease chlorophyll concentration compared with NSM/NSA plots. An increase in shoot chlorophyll with reduced light intensity did not occur in TifEagle bermudagrass. The NSM/AH, ML/AL, ML/AH, MH/AL, and MH/AH reduced chlorophyll by 31, 13, 39, 29, and 48%, respectively, compared with NSM/NSA (Table 2). When comparing chlorophyll means against DLI, any plots receiving 32.9 mol m^–2 d^–1 responded with a decrease in chlorophyll compared with NSM/NSA.

Total shoot chlorophyll main effect means at 4 WAS indicate negative afternoon shade effects on TifEagle bermudagrass. Low and high afternoon shade applications reduced total shoot chlorophyll concentration by 16 and 39%, respectively, compared with NSA (Table 3). Additionally, all plots receiving an afternoon shade application, except NSM/AL, contained less chlorophyll than NSM/NSA (Table 3). High morning shade reduced total shoot chlorophyll by 16% compared with NSM (Table 2). Therefore, afternoon shade applications to TifEagle were more detrimental to shoot chlorophyll concentration compared with morning shade.

At 8 WAS in 2001, similar results as 4 WAS occurred with treatment means. The NSM/AH, ML/AL, ML/AH, MH/AL, and MH/AH reduced chlorophyll by 48, 19, 65, 58, and 81%, respectively, compared with NSM/NSA (Table 2). In 2001, treatments receiving a DLI 32.0 mol m^–2 d^–1 maintained less total chlorophyll compared with NSM/NSA. In 2002, at 8 WAS, MH/AH reduced total shoot chlorophyll concentration by 35% reduction compared with NSM/NSA (Table 2).

Shoot chlorophyll main effect means at 8 WAS during 2001 indicated a reduction in total shoot chlorophyll at both levels of shade in morning and afternoon durations. Low and high morning shade reduced chlorophyll by 13 and 40%, respectively, compared with NSM (Table 3). Also, high morning shade decreased TifEagle chlorophyll by 31% compared with low morning shade. Greater chlorophyll reductions occurred with afternoon shade applications. Low and high afternoon shade decreased chlorophyll by 24 and 65%, respectively, compared with NSA (Table 3). High afternoon shade decreased chlorophyll by 54% compared with low afternoon shade.

In 2002, high morning shade reduced chlorophyll concentration by 12% compared with NSM (Table 3). No differences occurred between NSM and ML. High afternoon shade reduced chlorophyll concentration by 19% compared with NSM. Greater differences in chlorophyll concentration at 8 WAS were observed in 2001 compared with 2002. The difference in DLI may have attributed to the yearly variation in total shoot chlorophyll. Generally, during 2001, plots received approximately 1 to 2 mol m^–2 d^–1 less PAR than received in 2002. This slight difference in DLI may have altered total shoot chlorophyll concentration.

Total Nonstructural Carbohydrates
Morning and afternoon shade application influenced TNC concentration of TifEagle bermudagrass (Table 2). Reserve carbohydrate storage are the result of photosynthate production rate exceeding allocation and mobilization of nonstructural carbohydrates (fructose, glucose, sucrose, and starch) for energy and ATP production. In this study, highest TNC concentration of 40.9 mg g^–1 resulted from NSM/NSA, or maximum irradiance exposure (Table 2). Treatments receiving no shade either during morning and/or afternoon hours (NSM/AH and MH/NSA) or no application of high shade in the morning or afternoon (NSM/AL, ML/NSA, and ML/AL) did not differ from NSM/NSA. Total nonstructural carbohydrates were reduced by 37, 30, and 43% with ML/AH, MH/AL, and MH/AH, respectively, compared with NSM/NSA (Table 2). These plots received a DLI 28.8 mol m^–2 d^–1. Overall, a downward linear trend in TNC occurred with a corresponding decrease in DLI (Fig. 2) .

View larger version (15K): [in this window] [in a new window]   Fig. 2. Total nonstructural carbohydrates (TNC) of TifEagle bermudagrass below ground tissue to various daily light integrals during the summers of 2001 and 2002.

	CONCLUSIONS

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

 
A 2-yr study in Clemson, SC, quantified a DLI necessary for commercially acceptable TifEagle bermudagrass maintained to golf course standards during the months of June to August. According to the model, a DLI of 32.6 mol m^–2 d^–1 was required to provide commercially acceptable TifEagle TQ (Fig. 1). Other plant responses showed significant declinations from plots receiving full irradiance throughout the day. Percentage lateral RG differed from NSM/NSA plots when the DLI 26.2 mol m^–2 d^–1. Total shoot chlorophyll differed from NSM/NSA when the DLI 32.9 mol m^–2 d^–1. Finally, TifEagle bermudagrass TNC concentration differed from NSM/NSA with a DLI 28.8 mol m^–2 d^–1. Therefore, from this study it can be concluded a decline in TifEagle bermudagrass quality, growth, and metabolic responses can be expected when the average DLI falls below approximately 33 mol m^–2 d^–1.

Main effect means suggest afternoon shade can be extremely detrimental to TifEagle bermudagrass growth. When comparing no-shade with high-shade means in the morning and afternoon, greater reductions followed high afternoon shade applications in every plant response measured. This was probably because of the 2 mol m^–2 d^–1 greater reduction in the main effect DLI from high afternoon shade (26.0 mol m^–2 d^–1) compared with high morning shade (28.3 mol m^–2 d^–1) (Table 3). High afternoon shade applied 1500 h to sunset reduced TifEagle bermudagrass TQ, percentage lateral RG, shoot chlorophyll, and TNC by 39, 17, 39, and 27%, respectively, compared with NSA. High morning shade reduced TQ, percentage lateral RG, and shoot chlorophyll by 21, 11, and 16%, respectively, compared with NSM. These data can be applied to directional positioning of trees or other irradiance-blocking structures.

	NOTES

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

 
Research conducted at Clemson Univ., Clemson, SC. Technical Contribution No. 5042 of the Clemson University Experiment Station. Mention of a product does not constitute a guarantee or warranty of the product by Clemson University or the authors and does not imply its approval to the exclusion of other products that may also be suitable.

Received for publication January 5, 2004.

	REFERENCES

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

 

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This Article Abstract Figures Only Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Similar articles in this journal Similar articles in ISI Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via HighWire Citing Articles via ISI Web of Science (3) Citing Articles via Google Scholar Google Scholar Articles by Bunnell, B. T. Articles by Rajapakse, N. C. Search for Related Content PubMed Articles by Bunnell, B. T. Articles by Rajapakse, N. C. Agricola Articles by Bunnell, B. T. Articles by Rajapakse, N. C. Related Collections Turfgrass Management Turfgrass