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Illllll’il llllfillllll’llllllmllll 3 1293 01427 1732 This is to certify that the dissertation entitled The Effects of Plant Growth Regulators on Kentucky bluegrass (Poa pratensis L.) and Supina bluegrass (Poa supina Schrad.) in Reduced Light Conditions presented by John C. Stier has been accepted towards fulfillment of the requirements for Ph.D. degreein Crop & Soil Sciences O/fl / Major pro or Daté 5,42%»? /9‘9?’ MSU is an Affirmative Anion/Equal Opportuniti- Institution O~12771 LIBRARY Michigan State University PLACE IN RETURN BOX to remove this checkout from your record. TO AVOID FINE return on or before date due. mns DUE DATE ours I DATE DUE 1198 «lemmas-p.14 THE EFFECTS OF PLANT GROWTH REGULATORS ON KENTUCKY BLUEGRASS (POA PRA TENSIS L.) AND SUPINA BLUEGRASS (P. SUPINA SCHRAD.) IN REDUCED LIGHT CONDITIONS By John C. Stier A DISSERTATION Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Crop and Soil Sciences 1997 ABSTRACT THE EFFECTS OF PLANT GROWTH REGULATORS ON KENTUCKY BLUEGRASS (POA PRA TENSIS L.) AND SUPINA BLUEGRASS (P. SUPINA SCHRAD.) IN REDUCED LIGHT CONDITIONS By John C. Stier Turfgrass management in reduced light conditions (RLC; < 30% full sunlight) is difficult because turf growth is affected by lack of sufficient light energy. Turf plants in RLC are relatively weak and cannot withstand traffic or other damage due to excessive shoot elongation. reduced tillering, and reduced root growth. In normal light conditions plant growth regulators which inhibit gibberellic acid (GA) biosynthesis are occasionally used on turfgrass tO reduce mowing requirements by suppressing shoot growth. The Objective of the research was to determine the effects of two GA-inhibitors (flurprimidol and trinexapac-ethyl) on turfgrass in RLC. The primary reason for the research was to develop a set of management strategies to maintain turfgrass in RLC for athletic events, e.g. athletic fields in covered stadia, although the results should be applicable to many turf situations. Three studies were conducted. In the first set of experiments, the effects of flurprimidol were tested at three nitrogen (N) rates (24, 48, and 96 kg ha'1 month'l ) on Kentucky bluegrass, with and without traffic, at two levels of RLC (approximately 1-2 and 8 mol photosynthetically active radiation (PAR) day'l). A second study was undertaken to compare the relative shade tolerance of Supina bluegrass (Poa supina Schrad.) to Kentucky bluegrass (P. pratensis L.) with different combinations of trinexapac-ethyl and foliar-applied iron. In the third study the effects of trinexapac-ethyl on photosynthesis of Kentucky bluegrass and Supina bluegrass in RLC were assessed. The effects of trinexapac-ethyl on photosynthesis of Supina bluegrass maintained at low and high N rates (24 and 96 kg ha'1 month") were also evaluated. Both flurprimidol and trinexapac-ethyl effectively suppressed shoot growth and enhanced turf quality in RLC. Supina bluegrass was significantly more tolerant of RLC compared to Kentucky bluegrass although neither grass prospered at 1-2 mol PAR day'l. Supina bluegrass had greater rates of photosynthesis than Kentucky bluegrass on a turf area basis although this was related to the higher leaf area index (LAI) of Supina bluegrass. Trinexapac-ethyl did not affect photosynthetic rates in either Species. Nitrogen rate had little effect on photosynthesis in RLC but the high N rate did reduce LAI. Copyright by JOHN CLINTON STIER 1997 Dedicated to my wife, Valerie Ann Stier ACKNOWLEDGEMENTS I would like to extend my deepest appreciation to Dr. John (Trey) N. Rogers, III, for his guidance, support, and the belief in me he has always held. I will forever consider myself the most fortunate of graduate students for all the wonderful opportunites Dr. Rogers has provided. My graduate committee also deserves Special recognition for the time and efforts they have provided to me: Dr. James Flore, for the training and use of photosynthesis equipment and his lab space; Dr. Joseph Vargas, for his lab space and materials for identification of fungal pathogens as well as his insights on interacting with others; Dr. James Crum, for the special encouragement and wisdoms he shared with me. I am thankful for the help and friendship of T.J. Lawson, Richard Fogarsi, Thomas Nikolai, Mark Collins, and John C. Sorochan, as well as the myriad of student workers and golf course professionals with whom I have interacted, during the long, busy, and incredibly exciting days when we provided the world with the first indoor turfgrass athletic field at the Pontiac Silverdome for the 1993 US. Cup and 1994 World Cup. vi TABLE OF CONTENTS LIST OF TABLES ................................................................................................ ix LIST OF FIGURES .............................................................................................. xvii CHAPTER 1 INTERACTION OF NITROGEN AND FLURPRIMIDOL ON KENTUCKY BLUEGRASS (POA PRA TENSIS L.) IN REDUCED LIGHT CONDITIONS... 1 Introduction ............................................................................................... 1 Materials and Methods .............................................................................. 2 Results and Discussion .............................................................................. 8 Experiment I .................................................................................. 8 Turf Quality ....................................................................... ll Clipping Yields .................................................................. 15 Surface Characteristics ...................................................... 2O Experiment 11 ................................................................................. 25 Turf Quality ....................................................................... 27 Clipping Yields .................................................................. 33 Surface Characteristics ...................................................... 33 Plant Density ..................................................................... 41 Conclusions ............................................................................................... 41 CHAPTER 2 THE EFFECTS OF TRINEXAPAC-ETHYL AND F OLIAR IRON ON SUPINA BLUEGRASS (POA S UPINA SCRAD.) AND KENTUCKY BLUEGRASS (P. PRA T ENSIS L.) ....................................................................... 44 Introduction ............................................................................................... 44 Materials and Methods .............................................................................. 46 Experimental Environment ........................................................... 46 Plot Construction and Maintenance .............................................. 48 Data Collection .............................................................................. 50 Results and Discussion .............................................................................. 52 Turf Color and Quality .................................................................. 52 Experiment I: Turf Not Subjected to Traffic ..................... 52 Experiment 11: Turf Subjected to Traffic ........................... 60 Clipping Yields .............................................................................. 69 Experiment I: Turf Not Subjected to Traffic ..................... 69 VII Experiment 11: Turf Subjected to Traffic .......................... 76 Turf Shear Resistance .................................................................... 76 Experiment I: Turf Not Subjected to Traffic ..................... 76 Experiment 11: Turf Subjected to Traffic ........................... 82 Plant Density and Biomass ............................................................ 86 Experiment I: Turf Not Subjected to Traffic ..................... 86 Experiment 11: Turf Subjected to Traffic ........................... 91 Chlorophyll Concentration ............................................................ 91 Experiment I: Turf Not Subjected to Traffic ..................... 9] Experiment II: Turf Subjected to Traffic .......................... 98 Conclusions ............................................................................................... 98 CHAPTER 3 PHOTOSYNTHESIS OF SUPINA BLUEGRASS (POA SUPINA SCHRAD.) AND KENTUCKY BLUEGRASS (P. PRA T ENSIS L.) IN REDUCED LIGHT CONDITIONS AS AFFECTED BY NITROGEN AND TRINEXAPAC-ETHYL ....................................................................................... 1 01 Introduction ............................................................................................... 101 Materials and Methods .............................................................................. 104 Plot Establishment and Testing ..................................................... 104 Experiment 1: Species x PGR Study .................................. 104 Experiment 11: Nitrogen x PGR Study .............................. 107 Gas Exchange Measurements ........................................................ 108 Leaf Area Analysis ........................................................................ 110 Chlorophyll Analysis ..................................................................... l 10 Results ....................................................................................................... lll Experiment 1: Species x PGR Study .............................................. lll Experiment 11: Nitrogen x PGR Study ........................................... 111 Discussion .................................................................................................. 129 Conclusion ................................................................................................. l 3 5 APPENDIX ........................................................................................................... 137 LIST OF REFERENCES ...................................................................................... 138 viii LIST OF TABLES Table l - Photosynthetically active radiation (PAR) at the Hancock Turfgrass Research Center, East Lansing ......................................................................................... 9 Table 2 - Mean squares and treatment effects on the quality of Kentucky bluegrass maintained under ambient light conditions inside the Covered Stadium Simulator Facility ............................................................................................ 12 Table 3 - Main effects of nitrogen, flurprimidol, and traffic on the quality of Kentucky bluegrass maintained under ambient light conditions inside the Covered Stadium Simulator Facility, East Lansing, MI .......................................... 13 Table 4 - Quality rating values for the flurprimidol-by-nitrogen and nitrogen-by-traffic interactions in Kentucky bluegrass turf maintained in ambient light conditions in the Covered Stadium Simulator Facility, East Lansing, MI. .................................................................................................................. 16 Table 5 - Values for quality ratings of nitrogen-by-flurprimidol-by-traffic interaction of Kentucky bluegrass turf maintained under ambient light conditions in the Covered Stadium Simulator Facility, 22 Feb. 1994 ............................ 16 Table 6 - Mean squares and significance of treatment effects on clipping yields of Kentucky bluegrass maintained under ambient light conditions of the Covered Stadium Simulator Facility, East Lansing, MI. .............................................................. 17 Table 7 - Effect of nitrogen rate, flurprimidol, and traffic on clipping yields of Kentucky bluegrass maintained under ambient light conditions of the Covered Stadium Simulator Facility (CSSF), East Lansing, MI. ................................................. 18 Table 8 - Values for the significant nitrogen-by-flurprimidol interaction on clipping yields of Kentucky bluegrass maintained under ambient light conditions in the Covered Stadium Simulator Facility (CSSF), 1993-94. ....................................... 21 Table 9 - Values for the significant flurprimidol-by-traffic interactions on clipping yields of Kentucky bluegrass turf maintained under ambient light conditions of the Covered Stadium Simulator Facility (CSSF) ................................................................. 22 Table 10 - Mean squares and significance of treatment effects on the shear resistance of Kentucky bluegrass maintained under ambient light conditions inside the Covered Stadium Simulator Facility (CSSF) ............................................... 23 Table 11 - Main effects of nitrogen, flurprimidol, and traffic on the shear resistance of Kentucky bluegrass turf maintained under ambient light conditions inside the Covered Stadium Simulator Facility (CSSF) ............................................... 24 Table 12 - Mean squares and the significance of treatment effects on the surface hardness of Kentucky bluegrass turf maintained in ambient light conditions of the Covered Stadium Simulator Facility (CSSF) .................................... 26 Table 13 - Effects of traffic on Clegg Impact Values of Kentucky bluegrass turf maintained in reduced light conditions inside the Covered Stadium Simulator Facility (CSSF) ............................................................................................................. 26 Table 14 - Mean squares for the effects of nitrogen rate, flurprimidol, and traffic on the quality of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility, East Lansing, MI. ............................ 28 Table 15 - Main effects of nitrogen rate and flurprimidol on the quality of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility, East Lansing, MI. ........................................................... 29 Table 16 - Values for the significant interactions of nitrogen rate and flurprimidol treatment on the quality of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF), East Lansing, MI. ................................................................................................................ 31 Table 17 - Quality ratings for the significant nitrogen rate-by-flurprimidol-by-trafiic interactions on Kentucky bluegrass turf under supplemental light conditions ............. 32 Table 18 - Mean squares and significance of treatment effects on clipping yields of Kentucky bluegrass maintained under supplemental light conditions of the Covered Stadium Simulator Facility, East Lansing, MI. ............................................................ 34 Table 19 - Effect of nitrogen rate, flurprimidol, and traffic on clipping yields of Kentucky bluegrass maintained under supplemental light conditions of the Covered Stadium Simulator Facility (CSSF), East Lansing, MI. ............................................... 35 Table 20 - Clipping yields for the significant nitrogen-by-flurprimidol interactions in Kentucky bluegrass turf maintained under supplemental light conditions in the Covered Stadium Simulator Facility (CSSF) ............................................................... 37 Table 21 - Clipping yields for the significant flurprimidol-by-traffic interactions in Kentucky bluegrass turf maintained under supplemental light conditions in the Covered Stadium Simulator Facility (CSSF) .............................................................. 38 Table 22 - Mean squares and significance of treatment effects on the shear resistance of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF) .............................................................. 39 Table 23 - Main effects of nitrogen, flurprimidol, and traffic on the shear resistance of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF) .............................................................. 39 Table 24 - Mean squares and significance of treatment effects on the surface hardness of Kentucky bluegrass turf maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF) ............................. 40 Table 25 - Clegg Impact Values (gmax) for the flurprimidol-by-traffic interaction (3 Feb. 1993) and flurprimidol-by-nitrogen interaction (8 Apr. 1994) in Kentucky bluegrass turf maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF) ............................................................................. 40 Table 26 - Plant density, shoot density, and verdure weight of Kentucky bluegrass maintained under supplemental light in the Covered Stadium Simulator Facility, 10 Dec. 1993 to 23 August 1994 ................................................................................. 42 Table 27 - Photosynthetically active radiation (PAR) at the Hancock Turfgrass Research Center, East Lansing, MI. ........................................................................... 53 Table 28 - Mean squares and treatment effects on quality ratings of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ............................................................................................................... 54 Table 29 - Mean squares and treatment effects on color ratings of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ............................................................................................................... 55 Table 30 - Effect of species, trinexapac-ethyl, and iron on quality of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ............................................................................................................... 56 Table 31 - Effect of species, trinexapac-ethyl, and iron on color of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ............................................................................................................... 58 xi Table 32 - Quality rating values from selected dates for the significant species-by- trinexapac-ethyl interactions on non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ............................................ 61 Table 33 - Color ratings for the significant species-by-trinexapac-ethyl interaction on non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ........................................................................... 62 Table 34 - Mean square and treatment effects on quality of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................................................................................... 63 Table 35 - Mean square and treatment effects on color of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................................................................................... 64 Table 36 - Effect of species, trinexapac-ethyl, and iron on quality of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .......................................................................................... 65 Table 37 - Effect of species, trinexapac-ethyl, and iron on quality of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .......................................................................................... 67 Table 38 - Quality rating values for the significant species-by-trinexapac-ethyl interactions on turfgrass subjected to traffic in reduced light conditions ................... 70 Table 39 - Color rating values for the significant species-by-trinexapac-ethyl interactions on turfgrass subjected to traffic in reduced light conditions ................... 71 Table 40 - Mean squares and significance of treatment effects on clipping yields of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .......................................................................... 72 Table 41 - Effects of species, trinexapac-ethyl, and iron on clipping yields of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ......................................................................................... 73 Table 42 - Values for the significant interactions of trinexapac-ethyl and species on clipping yields of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ......................................................... 75 Table 43 - Mean squares and Significance of treatment effects on clipping yields of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ......................................................................... 77 xii Table 44 - Effects of species and trinexapac-ethyl on clipping yields of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................................................................... 78 Table 45 - Values from significant interactions of trinexapac-ethyl and species on clipping yields of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ........................................ 80 Table 46 - Mean squares and significance of treatment effects on shear resistance of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................................... 81 Table 47 - Effect of species and trinexapac-ethyl on shear resistance values (N cm) of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................................... 83 Table 48 - Shear resistance values (N am) for the significant species-by- trinexapac-ethyl interaction on non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ..................... 84 Table 49 - Mean square and Significance of treatment effects on shear resistance of turfgrass subjected to traffick in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ....................................... 84 Table 50 - Effect of species and trinexapac-ethyl on shear resistance values (N cm) of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ...................................................... 85 Table 51 - Shear resistance values (N om) for the Significant species-by- trinexapac-ethyl interactions on turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .................... 87 Table 52 - Mean squares and treatment effects on plant density and biomass of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ..................................................................................... 88 Table 53 - Effect of species, trinexapac-ethyl, and iron on plant density and biomass of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ..................................................... 89 Table 54 - Plant density and biomass values for the significant species-by- trinexapac-ethyl interactions on non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ................... 90 xiii Table 55 - Mean squares and treatment effects on plant density and biomass of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ..................................................................... 92 Table 56 - Effect of species, trinexapac-ethyl, and iron on plant density and biomass of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .................................................... 93 Table 57 - Plant density and biomass values for the species-by-trinexapac-ethyl interactions on turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ..................................... 94 Table 58 - Mean squares and treatment effects on chlorophyll of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ................................................................................................ 95 Table 59 - Effect of species, trinexapac-ethyl, and iron on chlorophyll of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. .................................................................... 96 Table 60 - Mean squares and treatment effects on chlorophyll of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. ................................................................................... 99 Table 61 - Effect of species, trinexapac-ethyl, and iron on chlorophyll content of turfgrass subjected to traffic in reduced light conditions ................................ 99 Table 62 - Photosynthetically active radiation (PAR) of plots in the Covered Stadium Simulator Facility (CSSF), Hancock Turfgrass Research Center, East Lansing, MI. .............................................................................................. 106 Table 63 - Mean squares and treatment effects of species and trinexapac-ethyl on photosynthetic parameters of turfgrass maintained in reduced light conditions (approximately 5 mol PAR day'l ), turf surface area basis, 23 November 1996 ............................................................................................ l 12 Table 64 - Photosynthetic differences between Supina bluegrass and Kentucky bluegrass in reduced light conditions (approximately 5 mol PAR day'l ), turf surface area basis, 23 November 1996 ........................................... 113 Table 65 - Mean squares and treatment effects of species and trinexapac-ethyl on photosynthetic parameters of turfgrass maintained in reduced light conditions (approximately 5 mol PAR day'l), leaf area basis, 23 November 1996 ............ 114 xiv Table 66 - Photosynthetic differences between Supina bluegrass and Kentucky bluegrass in reduced light conditions (approximately 5 mol PAR day'l), leaf area basis, 23 November 1996 ................................................................... 115 Table 67 - Mean squares and significance of treatment effects on leaf area index (LAI), fresh leaf weight, and chlorophyll concentration of turfgrasses in reduced light conditions (approximately 5 mol PAR clay'l ), 24 Nov. 1996.... 116 Table 68 - Effects of species and trinexapac-ethyl on foliage and chlorophyll of turfgrasses in reduced light conditions, approximately 5 mol PAR day", 24 Nov. 1996 ................................................................................................... 117 Table 69 - Mean squares and treatment effects of photosynthetic characteristics of Supina bluegrass affected by trinexapac-ethyl and nitrogen rate in reduced light conditions (approximately 5 mol PAR day'l), turf surface area basis ..... 118 Table 70 - Effects of nitrogen and trinexapac-ethyl on photosynthetic parameters of Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), turf surface area basis ............................................................ 121 Table 71 - Interaction of N rate and trinexapac-ethyl on transpiration (E) and stomatal conductance (gs) of Supina bluegrass maintained in reduced light conditions of approximately 5 mol PAR day"l ................................................ 123 Table 72 - Mean squares and treatment effects of photosynthetic characteristics of Supina bluegrass affected by trinexapac-ethyl and nitrogen rate in reduced light conditions (approximately 5 mol PAR day'l), leaf area basis....124 Table 73 - Effects of nitrogen and trinexapac-ethyl on photosynthetic parameters of Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), leaf area basis ..................................................................... 126 Table 74 - Mean squares and significance of treatment effects on leaf area index (LAI), fresh leaf weight, and chlorophyll concentration of Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), 17 Nov. 1996 ................................................................................................... 128 Table 75 - Effects of nitrogen rate and trinexapac-ethyl on leaf area index (LAI), fresh leaf weight, and chlorophyll concentration of Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), 17 Nov. 1996 ..... 130 Table 76 - Mean squares and treatment effects on pink snow mold (Microdochium nivale) effects on Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), 18 Nov. 1996 .......................... l3] XV Table 77 - Effects of nitrogen rate and trinexapac-ethyl on pink snow mold (Microdochium nivale) damage to Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), 18 Nov. 1996 ........................ 131 Table 78 - Particle size analysis of sand used in sandzpeat mixture (80:20) ............................................................................................................ 137 Table 79 - Particle size analysis of sand used in Experiment 11: Nitrogen x PGR study (Chapter 3) ................................................................................ 137 xvi LIST OF FIGURES Figure 1 - Photosynthetic photon flux density of sunlight, ambient light inside the Covered Stadium Simulator Facility (CSSF), and supplemental light inside the CSSF supplied by 400 W high pressure sodium lamps, 1515 h, 10 Feb. 1994 ......................... 10 xvii KEY TO SYMBOLS AND ABBREVIATIONS CER, carbon exchange rate; PGR, plant growth regulator; GA, gibberellic acid; gm“, peak deceleration; TE, trinexapac-ethyl; PAR, photosynthetically active radiation; PPFD, photosynthetic photon flux density; Ly, langley; E, transpiration; gs, stomatal conductance; gm, mesophyll conductance; WUE, water use efficiency; LAI, leaf area index; N, nitrogen; P, phosphorus; K, potassium; CSSF, Covered Stadium Simulator Facility. xviii Chapter 1 INTERACTION OF NITROGEN AND FLURPRIMIDOL ON KENTUCKY BLUEGRASS (POA PRA T ENSIS L.) IN REDUCED LIGHT CONDITIONS INTRODUCTION Turfgrasses in intense shade have weak tissues, reduced root to shoot ratio, and reduced tillering resulting in a reduced quality turf which cannot withstand traffic (Beard, 1973; Wilkinson and Beard, 1975). Minimal nitrogen inputs are recommended to maintain a balance between shoot and root growth and to minimize the growth of excessively succulent tissue. Traffic is to be minimized or avoided (Dudeck and Peacock, 1992). Consequently, current recommendations for turf in shade do not allow for management techniques in which traffic is a factor. Yet golf courses often have areas subjected to both shade and traffic. In addition, recent interest in the use of turfgrass systems for covered stadia requires the development of new management techniques for turf subjected to traffic under intensely shaded conditions (Anonymous, 1995; Kierle, 1995; Rogers, 1994; Tracinski, 1993). Plant growth regulators (PGRS) which are gibberellic acid biosynthesis-inhibitors (GA-inhibitors) have been used successfully to decrease vertical growth (i.e., clipping yields) (Dernoeden, 1984; Diesburg and Christians, 1989; Johnson, 1988) by inhibiting cell elongation (Kaufmann, 1986a). Side effects of GA-inhibitors include darker green 2 color and increased turf density due to enhanced tillering, often following a transient phytotoxic response resulting in tip die back (Demoeden, 1984; Watschke, 1981). In shade, excessive turfgrass shoot elongation leads to weak, traffic-intolerant turf (Beard, 1973). Preliminary research has proven the potential of GA-inhibitors to control shoot elongation and provide a higher quality turf compared to untreated turf in reduced light conditions although phytotoxicity can occur (Rogers et al., 1996; Rogers and Stier, 1993). In normal light situations multiple applications of GA-inhibitors increase the level of suppression but also increase the potential for phytotoxicity, especially when turf is grown under a stress condition (Demoeden, 1984; Johnson, 1988; Vitolo et al., 1990). Nitrogen fertilization in concert with PGR application has been reported to successfully minimize or overcome the Short-term deleterious effects of PGRS in normal light situations (Devitt and Morris, 1988). The effects of nitrogen rate on PGR-treated turf in reduced light conditions are unknown. The primary obj ective of this research was to determine the appropriate rate of nitrogen to apply to flurprimidol-treated Kentucky bluegrass turf in reduced light conditions. Secondary objectives included determining the amount of light required to maintain turf in an enclosed environment, the effects of traffic, and turf response to flurprimidol in reduced light conditions. MATERIALS AND METHODS The research was conducted inside the Covered Stadium Simulator Facility (CSSF) at the Hancock Turfgrass Research Center between Dec. 1992 and April 1994. The CSSF 3 was constructed to simulate the conditions inside the Pontiac Silverdome, a covered stadium (Stier et al., 1993). The CSSF was a 600 m2 air-supported structure constructed of Sheerfill II®, a fiberglass fabric (Chemical Fabrics Corporation, Buffalo, NY) which transmitted approximately 11 i 2% sunlight. Temperature and relative humidity were recorded daily with a sling psychrometer. Furnaces on the endwalls of the facility were used to maintain the temperature typically at 16.8 i 0.9 sd °C. Actual temperatures occasionally ranged from 3 to 23 ° C due to the poor insulating characteristics of the fiberglass fabric, the inability of the furnaces to compensate for excessively low outdoor temperatures (e.g., -10 C), and lack of an active cooling mechanism. Relative humidity (RH) averaged 44.8 i 6.2 Sd % with a range of 24-70% RH. Portable plots were established in wood boxes (1.2 x 1.2 x 0.15m depth) filled with a sandzpeat mix (80:20 v/v) (Table 78, Appendix). The pH was 7.3 with initial P and K levels of 63 kg ha'1 and 30 kg ha", respectively. Holes were drilled in the bottoms of the boxes for drainage. The sandzpeat mixture was compacted using hand-held tampers. Starter fertilizer (13-25-12) was applied to the soil which supplied 7.6 g N m'z, 6.4 g P m' 2, and 5.8 g K me. On 30 Sept. 1992 the plots were sodded with a washed Kentucky bluegrass blend (20% each of ‘Trenton’, ‘Midnight’, ‘Aspen’, ‘Rugby’, and ‘Kelly’). The plots were moved into the CSSF for testing on 7 Dec. 1992 through 10 April 1993. The experiments were repeated 3 second year (season). In 1993, plots were sodded 10 Sept. using a washed Kentucky bluegrass blend (20% each of ‘Trenton’, ‘Midnight’, ‘Aspen’, ‘Glade’, and ‘Parade’). These plots were moved into the CSSF on 10 Dec. 1993 and maintained until 8 April or 23 August 1994 depending on the experiment. In both years 4 plots were mowed once to twice weekly at 3.8 cm during establishment and irrigated as necessary to prevent moisture stress. Urea (2.4 g N m'z) was applied to aid establishment at three weeks after sodding in 1992 and at two and five weeks after sodding in 1993. Two experiments were conducted to assess the effects of nitrogen rate in PGR-treated turf. Experiment I was conducted in the ambient light conditions of the CSSF . Experiment 11 was conducted in the CSSF under supplemental lighting. Supplemental light was supplied by 430 W high pressure sodium (HPS) lamps suspended 2.7 m above the turf surface. An automatic timer controlled the lamps to provide a 12 hr photoperiod (0700 to 1900 hr). Reflective (metallic) mylar sheets were suspended in parallel along the two long sides of the rectangular plot area to separate the lighted plots from the unlighted plots and to reflect light from the lamps for increased uniformity of irradiance. Radiation data outside the CSSF were collected with a LI-PY14226 pyranometer and integrated daily. Radiometric units (Ly day") were converted to quantum units using the following equation based on conversion units from Thimijan and Heins (1983): Equation 1: ((Ly day”/1.05)*3600*24)/ 106 = mol PAR day'l. Radiation data for plots in ambient light conditions of the C SSF were estimated based on the percent transmission of photosynthetically active radiation (PAR) through the fabric, measured at the turf surface. Radiation data inside the CSSF from Dec. 1992 through April 1993 were determined at the turf surface weekly within one hour of the solar zenith using a hand-held photometer (Greenlee Inc., Rockford, IL). Occasionally a portable spectroradiometer (Li-Cor, Lincoln, NE) was used from Dec. 1992 to April 1993 to determine only photosynthetically active radiation (PAR), 400-700 nm. Starting 5 Dec. 1 993 radiation data inside the CSSF were collected weekly using only the spectroradiometer. For plots in supplemental light conditions in 1992-1993, photometeric units (lux) were converted to quantum units (umol rn'2 s") by multiplying against a conversion factor (0.2215) derived from data collected concurrently with the photometer and the spectroradiometer. Starting Dec. 1993 radiation data inside the CSSF were collected weekly using only the spectroradiometer. Based on measurements collected when ambient PAR inside the CSSF was low (e.g., 10 umol m'2 5'1 during rainstorms, pre- dawn, or evening), the HPS lamps supplied approximately 173 i 22 umol PAR m"2 s". The metallic mylar curtains on both sides of the plot area blocked much of the light transmitted into the CSSF. Measurements at different times of the day under a range of sunlight conditions (sunny, cloudy) Showed approximately 10% of the sunlight transmitted into the CSSF impinged on the plots under supplemental light in the morning and late afternoon; at midday approximately 80% of the light transmitted into the CSSF fell on the plots under supplemental light. Because ambient light levels peaked at midday, it was estimated that approximately 50% of the daily ambient PAR inside the C SSF contributed to the total daily PPF D of plots under supplemental light. The total daily photosynthetic photon flux density (PPFD) of plots in supplemental light conditions was estimated as follows using the average PPFD (umol m'2 s") from all plots: Equation 2: (((173 umol m’2 s’1 PAR*60 sec min’l)*60 min h‘l)*12 h)/1 x 106 = mol PAR day" from HPS lamps + 0.5 mol PAR day'I ambient light = mol PAR day", supplemental light plots The experiments were arranged as randomized complete block, Split plot designs with three replications. Treatments were arranged in a 3 x 2 x 2 factorial with nitrogen rate and flurprimidol as main plots. Simulated soccer traffic was applied as a sub-plot treatment split over the main plots. Nitrogen rates were 24, 48, and 96 kg ha'1 per treatment date. Urea nitrogen was applied with a drop spreader on the same dates as flurprimidol. Flurprimidol was applied at the label rate of 1.12 kg ha'1 in 1168 L H20 ha" I using a COz-powered backpack sprayer; control plots received no flurprimidol. Nitrogen and flurprimidol were applied on the following dates: 16 Dec., 21 Jan., and 26 Feb. (supplemental light study only) 1992-93, and 17 Dec., 4 F eb., and 21 Mar. 1993-94. The turf was irrigated with approximately 1.25 cm water immediately following fertilization and flurprimidol applications. Additional irrigation was supplied as necessary to prevent visible drought stress symptoms (blue-green turf color, footprinting, wilting). Plots in ambient light conditions received approximately 1.25 cm water at 14- 21 day intervals, while plots in supplemental light conditions received approximately 1.25 cm at seven to 10 day intervals. Traffic was applied to one-half of each plot by having a person (approximately 75-115 kg) walk 50 passes while wearing molded soccer cleats on each of the following dates: 29 Dec., 14 Jan., 21 Jan., 29 Jan., 6 Feb., 20 Feb., 10 Mar., and 24 Mar. (supplemental lighted study only on latter two dates) 1992-93, and 6 Jan., 11 Jan., 25 Jan., 1 Feb., 10 Feb., 22 Feb., 2 Mar., 17 Mar., 24 Mar., and 31 Mar. 1993-94. Traffic was applied immediately after mowing and prior to irrigation. Fungicides were applied at the onset of disease symptoms. On 9 Jan. 1993 and 14 Jan. 7 1994 chlorothalonil (tetrachloroisophthalonitrile; 16.5 kg ha") was applied to control leafspot and melting out diseases caused by Drechslera/Bipolaris spp. Plots were mowed once to twice weekly as needed to prevent removal of more than one-third of the leaf tissue. Clippings were collected, oven-dried at 60 °C for 48 h, and weighed. Turf quality was rated visually at five to 14 day intervals; ratings were conducted more frequently at the beginning of each year to assess rapid changes in turf quality and became less frequent as turf quality fluctuated less abruptly. Visual turf quality was based on a one to nine scale, with one representing completely necrotic turf or bare soil and nine representing dense, uniform turf with good color. A value of five was considered the minimum value for acceptable turf. Turf and rooting strength were evaluated periodically using an Eijkelkamp Shear vane apparatus (Eijkelkamp, Giesbeek, The Netherlands) (Rogers and Waddington, 1990). The amount of force (torque) required to tear the turf was collected in two locations from each plot on every measurement date. Treatment effects on the turf surface hardness were periodically evaluated using a Clegg Impact Soil Tester (CIT) (Lafayette Instrument Co., Lafayette, IN). The CIT provided surface hardness values by measuring maximum deceleration of a 2.25 kg hammer when dropped from a 0.46 m height (Rogers and Waddington, 1990). Impact values were collected from three locations in each plot on each measurement date. On 23 August 1994, one core (10 cm diam) was collected for plant biomass estimates from each plot which had received supplemental light. Plant density was evaluated by counting the number of live plants in each core. Verdure was removed from each core, and all living tissue was oven-dried at 60° C for 48 h then weighed. The number of 8 Shoots per plant was determined by averaging the number of shoots from five plants selected at random from each plot. Data were analyzed using MSTAT analysis of variance procedures for a 3-by-2 factorial experiment in a randomized complete block, split-plot design with three replications. The three nitrogen rates and two flurprimidol levels were split into trafficked and non-trafficked turf. RESULTS AND DISCUSSION Experiment I Turf quality, growth, and other attributes declined over time. To document the trends data are presented for individual dates throughout the course of the experiment. Data are presented for each year due to different results between years. Although some of the differences could have been due to different cultivars in the second year, the differences were probably due to a longer and more favorable establishment period during the autumn 1993. Average daily PAR values of ambient light in the CSSF increased steadily from December through April from, ranging from approximately 1 mol PAR day'1 in December to approximately 3 mol PAR day'l in April, but the turf did not respond (Table 1). Light quality transmitted through the fiberglass fabric of the CSSF mirrored the light quality of sunlight but light quantity was reduced approximately 90% (Figure 1). 9 Table l. Photosynthetically active radiation (PAR) at the Hancock Turfgrass Research Center, East Lansing, MI. Location Dec. 1992 Jan. 1993 Feb. 1993 Mar. 1993 Outside mol PAR day'1 I average 7.5 12.6 23.2 21.1 stnd deviation 3.9 5.4 3.1 12.2 CSSF, ambient light 1 average 0.8 1.4 2.5 2.3 stnd deviation 0.4 0.6 0.3 1.3 CSSF, Supplemental light § average 7.9 8.2 8.7 8.6 stnd deviation 1.2 1.2 1.1 1.6 Dec. 1993 Jan. 1994 Feb. 1994 Mar. 1994 Apr. 1994 Outside mol PAR clay'l average 9.5 10.8 19.3 24.3 31.7 stnd deviation 4.9 5.0 7.0 9.9 14.5 CSSF, ambient light average 1.0 1.2 2.1 2.7 3.5 stnd deviation 0.5 0.6 0.8 1.1 1.6 CSSF, Supplemental light average 8.0 8.1 8.5 8.8 9.2 stnd deviation 1.2 1.2 1.4 1.5 1.8 T PAR was collected with a pyranomter (Li-Cor, model PY 14226, Lincoln NE) and integrated daily. Radiation units (Ly clay'I ) were converted to quantum units (mol PAR day'I ) based on the conversion methods in Thimijan and Heins (1983). 1 CSSF = Covered Stadium Simulator Facility. Ambient PAR inside the CSSF was estimated by measuring the percent PAR transmitted into the CSSF at turf levels with a photometer (Greenlee Inc., Rockford IL) or a portable spectroradiometer (Li-Cor, Lincoln NE). § Supplemental lighting was supplied by 400 W high pressure sodium lamps. Because reflective mylar curtains on two sides of the plots blocked an estimated 50% of the ambient light from plots which received supplemental light, 50% of the total daily PAR was added to the total daily PAR supplied by the lamps (5.4 mol m"2 day'l ) to estimate the total daily PAR received by turf under the supplemental light. .32 .8“. A: .5 32 .859 $2qu 2385 :9; 3 8v 3 8:33 “Ammo 9: gas 29. 3285233 new .Aummov Esau. 56.355 5286 8.98 m5 mass 29. Essen .2923. Co :28 x3 86%. 285588;“. A 9%: AES Smco_o>m>> coo omm oom co». 10 I I ‘ll oo+woo.o roamooh oo+moo. r oo+womé oo+wood , oo+womN oo+moo.m oo+mom.m oo+moo.v oo+mom.v oo+woo.m A A A A 3 we .25 was 29. 323238 ammo ...... A E «.5 .85 8: 29. A8553 .ammo l A 3. We .85 OS: 2923 i i I (1'5 z_UJ mind) Ausuap xnu uoioud ogreutu/lsoioud 1111mm Significant treatment effects on Kentucky bluegrass quality are shown in Table 2. In 1992-93 there were no interactions between treatments. In 1993-94 occasional interactions occurred between nitrogen-by-flurprimidol and nitrogen-by-traffic; a three- way interaction occurred on 22 Feb. 1994. In both 1992-93 and 1993-94 the turf recovered from winter dormancy once placed inside the CSSF, but did not survive well. Quality declined to unacceptable values (< 5.0) within 49 days after installation in the CSSF in 1993 and within 82 days in 1994 regardless of treatment. The turf became nearly completely necrotic within 72 days in 1993 although better quality was sustained for the entire trial (105 days) in 1994. Traffic rapidly decreased turf quality and affected turf quality more often than nitrogen or flurprimidol (Table 2). Nitrogen rate did not affect turf quality in 1992-93; in 1993-94, the high rate (96 kg ha'l month'l) reduced turf quality within 74 days after installation in the CSSF (Table 3). High nitrogen rates are known to result in succulent tissues which render turf more susceptible to traffic and disease injury (Beard, 1973). High nitrogen rates have also been associated with decreased shoot density and root to shoot ratio in Shaded conditions (Burton et al., 1959; Eriksen and Whitney, 1981; Schmidt and Blaser, 1967); flurprimidol did not alter this response at this level of light. Flurprimidol increased turf quality on two dates only after the second application in both seasons. Traffic began to decrease turf quality after one application (50 passes) in 1992 and after four applications (200 passes) in 1994. In 1993, the flurprimidol-by-traffic interaction on 3 Feb. showed flurprimidol increased turf quality in a non-trafficked 12 .33 .5: 0 Es: 393m 8: 33 058,—. 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In the nitrogen-by-traffic interaction, traffic decreased turf quality at the medium and high nitrogen rates, but did not significantly affect turf quality at the low nitrogen rate. For the three-way interaction on turf quality, traffic did not affect quality at low or medium nitrogen rates when treated with flurprimidol, but did significantly reduce turf quality at the medium and high nitrogen rates in the absence of flurprimidol (Table 5). Weekly clipping yields were affected by flurprimidol and traffic in 1992-93 and by all three treatment groups individually in 1993-94 (Table 6). Data presented are intended to describe trends of main effects and interactions therefore interactions occurring only once were not discussed. Flurprimidol and traffic significantly reduced clipping yields beginning immediately after their first application (Table 7). The second application of flurprimidol inhibited vertical growth nearly completely, causing clipping yields to be at or near zero for the duration of the studies. Zero vertical growth is undesirable if it prevents turf recovery following damage. In this study flurprimidol did not appear to prevent recovery any more than untreated turf because the low light level was the limiting factor for growth. 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Traffic and flurprimidol also interacted in both seasons with flurprimidol decreasing clipping yields more than traffic (Table 9). We; Shear resistance of turf was most affected by traffic and only minimally affected by nitrogen or flurprimidol. No interactions occurred (Table 10). Traffic consistently decreased shear resistance values. High nitrogen rates decreased shear resistance compared to low nitrogen rates (Table l 1). Low shear resistance values due to traffic and high nitrogen rates were probably due to reduced turf cover and possibly reduced root structure although rooting was not measured. Flurprimidol had little effect, causing an increase on one date in 1994. 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A; 3 AS 3 AS 3 3 3 2 Z no A? 3233 2._ 8A: 2._ 8A: 2._ of: 2._ 3o 2._ 88 2 ._ A38 2._ 88 3.3.: «A :2 w: 3238:: 32 2 .%2 2 .32 N .83 mm .83 2 3.: CM .52 : 82 we do do do we do No A8233 A3 3 A; S o.o A; A; A; A3 to 2: no no A: a .33 S 2 A; 2: A3 3 2: 9o 3 2 E: 3 3o 3. :55? 2._ 88 2._ 88 2._ 8A. 2._ 88 2._ 88 2._ 8.0 2._ 3o 2.33 be”. 0. bum w ”A .2 9: 32332:: bum _ .53. am .5: mm .52. m_ .52 o mag A we no 2 23> 12 .wEmqu “mam .Emmov 326mm— SBEEE E:_3Sm 32050 23 5 22223280 Ew: 255:8 83:: 32:32.32: 33 mmfiwozz 32:33. :0 333A wcaazo no 2532225 oEab-zn-_03_E:E:: Emoc_:w_m 23 .8 mos—,3 .o 033% 23 Table 10. Mean squares and significance of treatment effects on the shear resistance of Kentucky bluegrass maintained under ambient light conditions inside the Covered Stadium Simulator Facility (CSSF), East Lansing, MI. Shear resistance (N-m) 1992-93 1993-94 Source of variation (if 22 Dec. 11 Jan. 3 Feb. 28 Dec. 8 Apr. Replication 2 22.028 2.507 9.299 9.528 1.021 N rate (N) 2 5.778 21.049" 5.132 0.778 37.646* Flurprimidol (F) 1 23.361 0.563 10.028 0.028 30250" N x F 2 0.444 5.063 1.799 8.778 3.771 Error 10 8.828 4.724 3.624 4.828 5.738 Traffic (T)T 1 ----- 25840" 40.1 1 1 ----- 20250" N x T 2 ----- 4.215 4.090 ----- 2.146 F x T 1 ----- 2.007 4.000 ----- 0.250 N x F x T 2 ----- 7.340 0.896 ----- 0.187 Error 12 ----- 3.993 1.681 ----- 2.507 CV, % 0.00 10.74 8.38 0.00 15.97 "‘ Significant at the 0.05 probability level. ‘1 Traffic applications were not started until 29 Dec. 1992 and 6 Jan. 1994. 24 Table 11. Main effects of nitrogen, flurprimidol, and traffic on the shear resistance of Kentucky bluegrass turf maintained under ambient light conditions inside the Covered Stadium Simulator Facility (CSSF), East Lansing, M1. Shear resistance (N-m) 1992-93 1993-94 Treatment 22 Dec. 1 1 Jan. 3 Feb. 28 Dec. 8 Apr. Nitrogen (kg ha'l ) ‘1 24 20.9 20.1 16.2 22.6 11.7 48 19.9 18.2 15.0 22.8 10.0 96 21.2 17.5 15.2 22.5 8.1 LSD (0.05) ns 2.0 ns ns 3.1 Flurprimidol (kg ha’1 ) none 19.9 18.7 15.0 22.5 9.0 1.12 i 21.5 18.5 16.0 22.6 108* Traffic without ---- 19.4 16.5 22.5 10.7 with § ---- 17.7* 14.4" 22.5 9.2“ "', ** Significant at the 0.05 and 0.01 probability levels, respectively. T Nitrogen was applied as urea on 16 Dec. 1992, 18 Jan., 17 Dec. 1993, 4 Feb., and 21 Mar. 1994. I F lurprimidol was applied on the same days as nitrogen. §Traffic was applied 29 Dec. 1992, 14 Jan., 21 Jan., 29 Jan., and 6 Feb. 1993, and 6 Jan., 11 Jan., 25 Jan., 1 Feb., 10 Feb., 22 Feb., 2 Mar., 17 Mar., and 24 Mar. 1994. 25 Treatment effects on surface hardness were not consistent between years (Table 12). Relative differences in gmax values between years were probably due to the use of different accelerometers in the CIT equipment following a repair in 1993. In the first season (1992-93) traffic treatments appeared to reduce surface hardness, while in the second season traffic treatments increased surface hardness (Table 13). Generally traffic will increase surface hardness by compaction and reduction of thatch and turf cover (Rogers and Waddington, 1990). Surface hardness is also affected by soil moisture with higher soil moisture providing lower gmax values (Rogers and Waddington, 1992), however, soil moisture was not determined in this study. In Feb. 1993 most turf in trafficked areas was dead although a thick (approximately 1.5 cm) mat layer remained which may have retained sufficient moisture to cause a decrease in gmax. Compaction in the mat layer by the traffic may also have caused decreased water infiltration. Non- trafficked areas may have had lower soil moisture values due to water uptake by the turf and increased infiltration rates. In Feb. and Apr. 1994 turf cover was higher than in 1993 and soil moisture values may have been more equivalent between trafficked and non- trafficked turf. Experiment 11 Supplemental lighting supplied approximately 8.4 i 1.4 mol PAR day'l (Table 1). The HPS lamps emitted a significant portion of their light in the yellow, orange, and red wavelengths (Figure 1). Ambient light in the CSSF was minimal and contributed little to the PAR on plots under supplemental light. 26 Table 12. Mean squares and the significance of treatment effects on the surface hardness of Kentucky bluegrass turf maintained in ambient light conditions of the Covered Stadium Simulator Facility (CSSF), East Lansing, MI. 1993 1994 Source of variation df 3 Feb. 3 Feb. 8 Apr. Replication 2 99.750 398.401 * 240.465 N rate (N) 2 1226.750 33.347 106.747 F lurprimidol (F) 1 205.444 31.923 30.988 NxF 2 196.861 143.191 37.814 Error 10 78.783 93.879 130.947 Traffic (T) 1 3211.111" 753.503" 1497.690" N x T 2 206.194 57.341 116.328 F x T 1 225.000 9.714 124.695 N x F x T 2 37.750 6.930 45.859 Error 12 157.333 18.056 50.820 CV, % 9.09 6.23 9.48 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. Table 13. Effects of traffic on Clegg Impact Values of Kentucky bluegrass turf maintained in reduced light conditions inside the Covered Stadium Simulator Facilty (CSSF), East Lansing, M1.f gmax Ambient light Supplemental light 3‘, 1993 1994 1993 1994 Treatment Feb. 3 3 Feb. 8 Apr. 3 Feb. 3 Feb. 8 Apr. Nitrogen (kg ha'l ) § 24 141.3 66.5 72.2 156.0 74.4 83.5 48 134.8 69.8 75.4 146.6 69.2 79.0 96 134.8 68.2 78.1 147.2 71.0 79.2 LSD (0.05) ns ns ns ns ns ns Flurprimidol (kg ha" ) 0.00 139.8 67.2 74.3 144.3 72.6 82.9 1.12 1] 134.1 69.1 76.2 155.6 70.5 78.2 Traffic without 147.4 63.6 68.8 154.9 66.4 73.8 with # 128.6" 72.8" 81.7" 145.0 76.7" 87.3" *, " Significant at the 0.05 and (YOI probability levels, respectively; ns = not significant at p=0.0§ T Plots were established outside during autumn of each year and moved into the CSSF on 7 Dec. 1992 and on 10 Dec. 1993. I Supplemental light was supplied from 400 W high pressure sodium lamps. § Nitrogen was applied as urea on 16 Dec. 1992, 18 Jan., 26 Feb., 21 Dec. 1993, and 4 Feb., 21 Mar. 1994. 11 F lurprimidol was applied on the same dates as nitrogen fertilizer. # Traffic was applied on 29 Dec. 1992, 14 Jan., 21 Jan., 29 Jan., 6 Feb., 20 Feb., 10 Mar., 24 Mar. 1993, and 6 Jan., 11 Jan., 25 Jan., 1 Feb., 10 Feb., 22 Feb., 2 Mar., 17 Mar., 24 Mar., and 31 Mar. 1994. 27 Turf quality remained relatively stable under the supplemental light conditions. Traffic had more of an effect on the turf in the first year than in the second year for probably two reasons: 1) shorter period of establishment in the first year, and 2) a heavier person (approximately 115 kg) applied the traffic the first year while a lighter person (approximately 75 kg) applied the traffic the second year. 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The nitrogen-by-flurprimidol interactions showed flurprimidol negated the effects of nitrogen on clipping yields while increasing nitrogen rates significantly increased clipping yields in the absence of flurprimidol (Table 20). The flurprimidol-by-traffic interaction showed traffic did not reduce clipping yields when flurprimidol was applied because flurprimidol acutely reduced clipping yields compared to traffic (Table 21). rf c h ra ri i s Treatments affected turf shear strength in both seasons (Table 22). No interactions occurred. Traffic and higher nitrogen rates decreased turf shear strength. Unlike the results in experiment one, the lower shear resistance values were probably due to reduced rooting and increased turf succulence as turf cover was not significantly diminished by either treatment. Flurprimidol did not affect shear strength in season one but caused a slight decline in season two (Table 23). 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Replication 2 50.361 * * 10.896 10.882 5.027 23 .083 5.090 N rate (N) 2 0.528 6.813 12.340 127.823M 5.250 281.757M Flurprimidol (F) 1 1.000 16.000 16.000 15.867 8.028 20.250" N x F 2 19.083 10.146 3.271 3.151 0.861 6.896 Error 10 5.828 6.929 8.315 3.643 9.083 3.599 Traffic (TN 1 ------ 12.250 4.000 125.814" ------ 20250" N x T 2 ------ 2.021 4.146 2.014 ------ 10.021 F x T 1 ------ 0.694 0.111 1.914 ------ 2.778 N x F x T 2 ------ 1.882 1.549 0.034 ------ 0.632 Error 12 ------ 4.604 7.271 2.537 ------ 3.764 CV, % 0.00 9.61 12.87 12.82 0.00 10.95 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. T Traffic was not started until 29 Dec. 1992 the first year and 6 Jan. 1994 the second year. Table 23. Effects of nitrogen, flurprimidol, and traffic on the shear resistance of Kentucky bluegrass maintained under supplemental light conditions inside the Covered Stadium Simulator Facility (CSSF), East Lansing, MI.“ Shear resistance (N -m) 1992-93 1993-94 Treatment 22 Dec. 1 1 Jan. 3 Feb. 20 May 28 Dec. 8 Apr. Nitrogen (kg ha'1 ) I 24 21.8 23.1 22.0 16.1 23.7 22.2 48 21.5 22.2 20.0 11.5 24.7 18.4 96 21.9 21.6 20.8 9.7 24.9 12.6 LSD (0.05) ns ns ns 1.7 ns 2.4 Flurprimidol (kg ha'l ) none 21.9 21.7 20.8 11.8 24.9 18.5 1.12§ 21.6 23.0 21.6 13.1 23.9 170* Traffic without -—-- 22.9 21.3 14.3 ---- 18.5 with 11 ---- 21.8 20.6 10.6 -—-- 17.0“ * Significant at the 0.05 probability level; ns=not significant at p=0.05. 1' Supplemental light (approximately 8.4 mol day'l ) was supplied by 400 W high pressure sodium lamps. I Nitrogen was applied as urea on 16 Dec. 1992, 18 Jan., 17 Dec. 1993, 4 Feb., and 21 Mar. 1994. § Flurprimidol was applied on the same days as nitrogen. 11 Traffic was applied 29 Dec. 1992, 14 Jan., 21 Jan., 29 Jan., and 6 Feb. 1993, and 6 Jan., 11 Jan., 25 Jan., 1 Feb., 10 Feb., 22 Feb., 2 Mar., 17 Mar., and 24 Mar. 1994. 40 Table 24. Mean squares and significance of treatment effects on the surface hardness of Kentucky bluegrass turf maintained under supplemental light conditions in the Covered Stadium Simulator Facility, East Lansing, MI. 1993 1993-94 Source (if 3 Feb. 3 Feb. 8 Apr. Replication 2 2762.861“ * 85.343 146.551* N rate (N) 2 331.361 84.010 77.048 Flurprimidol (F) 1 1156.000 37.414 198810“ N x F 2 245.583 109.471 214.666" Error 10 316.228 35.931 23.308 Traffic (T) 1 880.111 945.563" 1653.778” N x T 2 525.194 2.843 0.564 F x T 1 2177.778" 0.122 17.921 N x F x T 2 739.528 9.880 41.792 Error 12 195.889 19.469 30.086 CV, % 9.33 6.17 6.81 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. Table 25. Clegg Impact Values (gum) for the flurprimidol-by-traffic interaction (3 Feb. 1993) and flurprimidol-by-nitrogen interaction (8 Apr. 1994) in Kentucky bluegrass turf maintained under supplemental light conditions in the Covered Stadium Simulator Facilty, East Lansing, MI.* g max 3 Feb. 1993 8 Apr. 1994 flurprimidol (kg ha" )1; flurprimidol (kg ha'T )§ Trafficfl 0.00 1.12 Nitrogen (kg ha" )# 0.00 1.12 g —— g ————-- without 157.0 152.8 24 83.2 83.8 with 131.6 158.4 48 86.3 71.8 96 79.3 79.0 LSD (0.05) 14.4 6.2 T Supplemental light (approximately 8.4 mol day'I ) was supplied by 400 W high pressure sodium lamps. I Flurprimidol was applied 16 Dec. 1992 and 18 Jan. 1993 § F lurprimidol was applied 21 Dec. 1992 and 4 Feb. 1993. 1] Traffic was applied 29 Dec. 1992, 14 Jan., 21 Jan., and 29 Jan. 1993 # Nitrogen was applied on the same dates as flurprimidol. 41 hardness in the absence of flurprimidol; traffic did not affect surface hardness of turf treated with flurprimidol (Table 25). Untreated turf was flaccid and traffic caused a prostrate growth (grain), forming a cushion on the surface which absorbed the impact of the CIT hammer. Turf treated with flurprimidol remained rigid and had an upright growth which resulted in similar amounts of foliage removal during mowing, thus providing similar cushioning, regardless of traffic. The flurprimidol-by-nitrogen interaction was more difficult to decipher. CIT values were inconsistent among treatments and did not indicate an orderly or meaningful response (Table 25). BlanLdensjtx By August 1994 all turf was maintaining fair to excellent quality and had completely recovered from traffic. Plant biomass data showed flurprimidol significantly affected turf growth five months after the final treatment had been applied. While the number of plants per unit area was less in plots treated with flurprimidol compared to control plots, the number of shoots per plant was nearly double, and verdure mass was approximately 25% greater (Table 26). CONCLUSIONS Kentucky bluegrass turf in the early stages of winter dormancy recovered sufficiently within two weeks at approximately 1 mol PAR rn'2 day'1 and temporarily provided acceptable quality. However, this level of light was insufficient to maintain acceptable Kentucky bluegrass turf for periods of longer than eight weeks. At 1-—3 mol m'2 day '1 PAR the best nitrogen rates were a low or medium rate (24 and 48 kg ha'1 month'l). 42 Table 26. Plant density, shoot density, and verdure weight of Kentucky bluegrass maintained under supplemental light in the Covered Stadium Simulator Facility, 10 Dec. 1993 to 23 August 1994.’ No. of Treatment No. of plants m’2 I shoots plant'l § Verdure (g m‘2 )1] Nitrogen rate (kg ha'l ) # 24 8885 2.8 60.8 48 8144 3.2 60.7 96 7095 3.5 49.1 LSD (0.05) ns ns ns F lurprimidol rate (kg ha'1 ) Ti 0.00 9474 2.2 49.7 1.12 6609 ** 4.0 “ 64.1 *,** Significant at the 0.05 and 0.01 probability levels, respectively. 1‘ Supplemental light, approximately 8.4 mol PAR day", was supplied from 400 W high pressure sodium lamps. 1 Plants were counted from a 10 cm diam core extracted from each plot. § Five randomly selected plants from each plot were used for analysis. 1] Verdure was collected from a 10 cm diam core extracted from each plot and included all living above ground plant tissue. # Nitrogen was applied as urea on 21 Dec. 1993, 4 Feb. 1994, 21 March 1994. 11' Flurprimidol was applied on the same dates as nitrogen fertilizer. 43 Traffic and high nitrogen rates (e. g., 96 kg ha") hastened demise of the turf, while flurprimidol extended the period of acceptable quality for a short period (e. g., two weeks). Two or more full rate applications of flurprimidol at four to six weeks halted the turf vertical growth rate which may have reduced the potential for recovery from damage (Stier et al., 1994), although lack of sufficient light would probably have been the limiting factor for recovery. Kentucky bluegrass turf recovered from winter dormancy within two weeks when placed in supplemental light conditions. Reduced light of approximately 8.4 mol PAR m' 2 day'I was sufficient to maintain high quality turf indefinitely, even in trafficked conditions. The medium nitrogen rate (48 kg ha'1 mol) was considered optimal as it provided the most desirable combination of quality, yield, hardness, and shear resistance. Flurprimidol significantally improved turf quality throughout the study and was paramount for maintaining high turf quality. Timing of flurprimidol applications and rates need to be further assessed as turf vertical growth was nearly totally halted following the second application. Flurprimidol rates and application intervals should be determined that allow a steady suppression of growth without inhibiting turf recovery from traffic and other damages. Diesburg and Christians (1989) reported the combination of growth phase and season affected turf response to PGRS. Since the environment of indoor stadia is moderated, long-term or permanent use of turf in covered stadia or other reduced light conditions may require unique rates and application intervals due to the lack of seasonal changes. Chapter 2 THE EFFECTS OF TRINEXAPAC-ETHYL AND FOLIAR IRON ON SUPINA BLUEGRASS (POA S UPINA SCHRAD.) AND KENTUCKY BLUEGRASS (P. PRA T ENSIS L.) INTRODUCTION 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 with the exception of the fine fescues (e.g., F estuca rubra L., F. rubra var. commutata Gaud.). As a turf, fine fescues perform best in conditions of well-drained soil and low fertility but have poor traffic tolerance due in part to a slow recuperative rate (Beard, 1973). 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). Rough bluegrass (Poa trivialis L.) has better shade tolerance than Kentucky bluegrass but lacks traffic tolerance. A relatively shade and traffic tolerant cool-season turfgrass species is desirable for golf courses, lawns, and athletic fields. Supina bluegrass (P. supina Schrad.) has been cultivated as a cool-season turfgrass in Germany for over 20 years (Bemer, 1984). Supina bluegrass is a stoloniferous turfgrass capable of forming a dense turf at low mowing heights suitable for lawns, athletic fields. 44 45 and golf course fairways, tees, and putting greens (Berner, 1980; Nonn, 1994; Pietsch, 1989). The stolons are significantly more robust and have shorter intemodes compared to rough bluegrass (personal observation). Supina bluegrass is found naturally in high traffic areas (e. g., human and cattle paths) and in moist, shaded areas in woods near the Alps (Bemer, 1984; Pietsch, 1989). Supina bluegrass is well adapted to cold weather and is common even in the sub-alpine regions of the Alps (Bemer, 1984; Kéck and Walch, 1977; Skirde, 1971). In Germany, Supina bluegrass often encroaches and fills in high wear areas on sports fields (Kéck and Walch, 1977); subsequent testing documented the high wear tolerance which is at least partly due to a rapid recuperative rate (Bemer, 1980; Berner 1984). In addition, Supina bluegrass has been observed to have a high level of shade tolerance on golf courses, lawns, and in controlled tests in Germany although the actual data have not been reported (Pietsch, 1989). 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). Drawbacks to the production and use of Supina bluegrass are its poor seed yield (hence, high cost), poor drought tolerance, undefined management schemes, and light green leaf color (Berner, 1980; Leinauer et al., 1991). The development of management schemes requires controlled investigation. While seed yield and drought tolerance are characteristics not easily altered, leaf color is an adjustable parameter which could increase the acceptablity of Supina bluegrass if a darker color can be easily obtained. Plant growth regulators (PGRS) and foliar applications of iron have been used successfully to enhance (darken) turf foliage in normal field conditions (Brueninger et al., 46 1983; Freeborg, 1983; Glinski et al., 1992; Yust et al., 1984). F oliar applications of iron have also been useful to negate the transient phytotoxicity which can result from a PGR (Carrow & Johnson, 1990). Recent reports indicate PGRS can also significantly enhance turf color and quality in reduced light conditions (RLC) (< 30% full sunlight) (Rogers et al., 1996; Stier et al., 1994) although the effects of iron are relatively unknown. Although moderate RLC result in increased chlorophyll content, extreme RLC reduce chlorophyll content resulting in a lighter green color (Beard, 1973). In our research we have found chlorophyll levels in Kentucky bluegrass decline at less than approximately 10 mol photosynthetically active radiation (PAR) day'l, equivalent to approximately 20% full summer sunlight (unpublished data). The objectives of this research were to: 1) Compare the shade tolerance of Supina bluegrass and Kentucky bluegrass under a defined light regime, and 2) Determine the effects of multiple applications of trinexapac-ethyl (below label rates) and foliar applications of iron on the growth and quality of Supina bluegrass and Kentucky bluegrass in RLC. MATERIALS AND METHODS Experimental environment The research was conducted inside the Covered Stadium Simulator Facility (CSSF) at the Hancock Turfgrass Research Center from Dec. 1994 through May 1996. Constructed initially in 1992 with a fiberglass fabric (Sheerfill IV, Chemical Fabrics Corporation, Buffalo, NY) which transmitted 11 :t 2% sunlight, the fabric was replaced in late October 47 1994. The new fiberglass fabric, Sheerfill IV®, transmitted approximately 10.5 i 1.4% solar radiation from Nov. 1994 through April 1995. After being bleached by the sun in the spring and summer of 1995, the fabric transmitted approximately 15.5 i 3.0 % solar radiation from Dec. 1995 through May 1996. Quality of the light transmitted through the Sheerfill IV® was equivalent to that transmitted by Sheerfill II® (Figure 1, Chapter 1). Temperature and relative humidity were recorded daily with a sling psychrometer. Temperature was maintained typically at 16.6 C using furnaces. Actual temperatures ranged from 12.2 to 24.4 C due to the inability of the furnaces to compensate for extremely low outdoor temperatures (e. g., -10 C) and due to the lack of an active cooling system as outdoor air temperatures rose during the spring. Relative humidity averaged 45.6 i 12.5 % with a range of 28-63%. Daily totals of photosynthetically active radiation (PAR) in the CSSF were determined based on the percent of PAR transmitted through the fabric onto the turf surface inside the CSSF. To determine percentage of light transmission, data were collected weekly from each plot inside the CSSF within one hour of the solar zenith using a Li-Cor 1800 portable spectroradiometer (Li-Cor, Lincoln, NE). Two to four measurements were collected outside the CSSF with the spectroradiometer immediately before and immediately after collecting data inside the CSSF. Daily solar radiation data outside the CSSF were collected with a Li-Cor PY 14226 pyranometer (Li-Cor, Lincoln, NE) located approximately 15 m away from the CSSF. Pyranometer data were integrated hourly and daily through a Maxi weather station (Rain Bird Sales. Inc., Glendora, CA). Radiometric 48 units from the pyranometer were converted to quantum units using the following equation which was based on conversion units from Thimijan and Heins (1983): Equation 1: ((Ly day'l/l .05)*3600*24)/106 = mol PAR day". The average percentage of light transmitted into the CSSF was used to determine the daily PAR inside the C SSF based on the data recorded outside with the pyranometer. t i n m in n Portable plots were established in wood boxes (1.2 x 1.2 x 0.15m depth) filled with a sandzpeat mix (80:20 v/v) (Table 78, Appendix). The pH was 7.3 with initial P and K levels of 85 and 90 kg ha", respectively, in 1994. In 1995, the pH was 7.7 with initial P and K levels of 131 and 85 kg ha", respectively. Sixteen holes (0.6 cm diam) were drilled in the bottoms of each box to provide drainage. The sandzpeat mixture was compacted using hand-held tampers. Starter fertilizer (13-25-12 in 1994) was added to the sandzpeat mixture surface prior to sodding to supply 66 kg P ha'1 and 58 kg K ha'I in 1994. The plots were sodded 28 September 1994 and 28 August 1995. Additional fertilizer was applied twice in 1994 (24, 20, and 18.5 kg N, P, K ha'l, respectively, on 29 Sept. and 36, 30, and 28 kg N, P, K ha", respectively, 13 Oct.) and once in 1995 (24, 20, and 18.5 kg N, P, K ha", respectively, on 29 Sept.) prior to moving the plots into the CSSF. Supina bluegrass ‘Supranova’ and Kentucky bluegrass ‘Victa’l’Abbey’ (50:50 v/v) were used both years. In 1994 the Supina bluegrass sod had been raised in a woody yard waste compost media while in 1995 washed Supina bluegrass sod grown in a sandy loam soil was used for establishment (sod raised in the woody compost was not 49 available). In both 1994 and 1995 washed Kentucky bluegrass sod grown in an organic soil was used for establishment. Plots were mowed once to twice weekly depending on height of cut and growth rate. During establishment (approximately three weeks) plots were mowed with a rotary mower set at 5 cm height. The height was gradually lowered to 3 cm; a reel mower was used once a 3 cm cutting height was achieved. Plots were irrigated as necessary to prevent visible drought stress (bluish-green color, footprinting, wilting). Trinexapac-ethyl (0.19 kg ha], approximately two-thirds the full label rate for Kentucky bluegrass) was applied to six plots each of Supina bluegrass and Kentucky bluegrass on 3 Oct. 1994 and 9 Oct. 1995. A COz-powered backpack sprayer with 8002 flat fan nozzles was used to apply the trinexapac-ethyl in 896 L H20 ha". Plots were moved into the CSSF for testing from 12 Dec. 1994 through 12 April 1995 and from 8 Dec. 1995 through 11 June 1996. The plots were arranged in the CSSF in a completely randomized design with three replications per treatment. Two experiments were designed to determined treatment effects in both non-trafficked (Experiment I) and trafficked (Experiment 11) conditions. Traffic was applied by having a person (approximately 70-75 kg) jog 50 passes each week. Traffic was applied 28 Dec. 1994 through 16 Mar. 1995 (total of 144 passes) and 26 Jan. 1995 through 26 Apr. 1996 (total of 168 passes). Additional trinexapac-ethyl (0.08 kg ha", approximately one-quarter the full label rate for Kentucky bluegrass, diluted in 896 L H20 ha") was applied on 21 Dec. 1994, 20 Jan., 18 Feb., and 16 Mar. 1995 for the first year’s testing and on 31 Jan., 15 Mar., and 26 Apr. 1996 for the second year’s testing. Iron (1.14 kg ha'1 as FeSO4-7HZO) was applied to foliage using F erromec 50 AC (PBI Gordon Corp., Kansas City, M0) on the following dates: 10 Jan., 14 F eb., and 17 Mar. 1995; 28 Feb. and 13 May 1996. Plots were fertilized monthly with 24, 2, and 20 kg ha'l N, P, and K, respectively (1 8- 3-18). Approximately 1.25 cm water was applied immediately following fertilizer application. Additional irrigation was applied as necessary to prevent drought stress (approximately 1.25 cm at seven to 14 day intervals). Industrial fans were occasionally used for 24—72 h periods to dry the turf surface following irrigation to discourage fungal pathogen activity. Iprodione (3-(3,5-Dichlorophenyl)-N-(l-methylethyl)-2,4-dioxo-l- imidazolidinecarboximide) was applied to all plots on 23 Dec. 1994 (3 kg ha'l), 6 Mar. (6 kg ha") and 14 Apr. (6 kg ha'l), 1995 to control Microdochium patch (Microdochium nivale), primarily on the Supina bluegrass. Data eellectien 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. Mowing was always performed immediately preceding data collection or fertilizer, trinexapac-ethyl, or traffic application. Clippings were generally collected for clipping yield determination except occasionally when time limits precluded clipping collection. Clippings were collected from a 41 x 117 cm strip through the center of each trafficked and non- trafficked plot. Clippings were dried in a forced-air oven at 60 C for 48 h then weighed. Turf color and quality were evaluated visually on a one to nine scale. A one rating represented 100% necrotic turf/bare soil, while a nine rating represented dark green or ideal turf, respectively. A value of five was considered the minimum acceptable unit. 51 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 m quadrat(Skog1ey and Sawyer, 1992). 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. The leaf portion next to the shoot (< 5 mm distant) was avoided due to possible physiological differences compared to the more mature leaf region (Skinner and Nelson, 1995). Chlorophyll was extracted in three ml MN-dimethlyformamide (DMF) incubated 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. On 12 Apr. 1995 and 31 May 1996 samples of ten randomly selected plants were collected from each plot for biomass assessments. Average leaf number shoot’l, average shoot number plant", and average oven-dry weight plant'I were determined for each sample. Data were analyzed using MSTAT analysis of variance procedures. Data were analyzed as a 2x2x2 factorial in a completely randomized design with three replications. 52 RESULTS AND DISCUSSION Temperature inside the CSSF averaged 16.6 C i 1.5 C with a range of 12-24 C. Relative humidity averaged 46% i 12% with a range of 28-63%. Photosynthetically active radiation inside the CSSF ranged from approximately 1 mol PAR day'1 during December 1994 to approximately 5 mol PAR day'1 in May 1996 (Table 27). The Supina bluegrass responded to the increased PAR in the spring more than did Kentucky bluegrass. In general, quality of the Supina bluegrass in 1995-96 was superior to that in 1994-95, probably largely due to the higher light transmittance of the Sheerfill IV fabric due to bleaching by the sun during the summer of 1995. Infisnlnnandnualitx Ex r’me I: Tur n 4b '6 ' d1 tra zc Turf color and quality was affected by both species and trinexapac-ethyl as soon as observations began once inside the CSSF (Tables 28 and 29). The turf was dormant when it was brought into the CSSF and recovered quickly the first year (1994) but slowly the second year (1995). In 1995 the weather had become quite cold in early November without an appropriate transition (“hardening-oft”) period between growing and non- growing conditions which probably caused the delay in recovery inside the C SSF. The extra N application in autumn 1994 may also have contributed to faster green-up of turf inside the CSSF. Turf treated with trinexapac-ethyl was particularly slow to recover inside the CSSF during the second year (Tables 30 and 31). Once recovered from 53 Table 27. Photosynthetically active radiation (PAR) at the Hancock Turfgrass Research Center, East Lansing, MI. Location Dec. 1994 Jan. 1995 Feb. 1995 Mar. 1995 Apr. 1995 mol PAR day'l I average 8.4 9.4 19.2 24.8 26.9 stnd deviation 3.9 6.5 6.6 10.4 12.8 CSSF 1, Ambient light average 0.9 1.0 2.0 2.6 2.8 stnd deviation 0.4 0.7 0.7 1.1 1.4 Dec. 1995 Jan. 1996 Feb. 1996 Mar. 1996 Apr. 1996 May 1996 mol PAR day'I average 9.5 10.6 14.6 27.8 29.0 34.1 stnd deviation 4.4 4.7 7.6 10.2 14.6 14.5 CSSF, Ambient light average 1.5 1.6 2.3 4.3 4.5 5.3 stnd deviation 0.7 0.7 1.2 1.6 2.3 2.2 1' PAR was collected with a pyranomter (Li-Cor, model PY 14226, Lincoln NE) and integrated daily. Radiation units (Ly day'l ) were converted to quantum units (mol PAR m'2 day'I ) based on the conversion methods in Thimijan and Heins (1983). i CSSF = Covered Stadium Simulator Facility. Ambient PAR inside the CSSF was estimated by measuring the percent PAR transmitted into the CSSF at turf level with a photometer (Greenlee Inc., Rockford IL) or a portable spectroradiometer (Li-Cor, model L1-1800, Lincoln NE). 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Turf density, uniformity, and overall appearance contributed to the quality ratings. Kentucky bluegrass turf density declined over time and the turf died in 1996 due in part to powdery mildew (Erysiphe gramim's). No powdery mildew was observed on Supina bluegrass although Microdochium patch (Microdochium nivale) occasionally occurred. The Microdochium patch was controlled by using fans to dry the turf and with fungicide. Supina bluegrass had a lighter green color than Kentucky bluegrass except towards the end of the second year when the Kentucky bluegrass became necrotic (Table 31). Iron had negligible effect on either color or quality of turf in either year except for minor, transient (< 4 wks) increases in turf color which, while statistically significant, were not as dramatic as those caused by trinexapac-ethyl. Interactions between species and trinexapac-ethyl occurred frequently (Tables 32 and 33). Supina bluegrass was more sensitive to trinexapac-ethyl than Kentucky bluegrass. Trinexapac-ethyl usually increased the turf quality and enhanced the color of Supina bluegrass compared to Kentucky bluegrass except at the beginning of the second year when trinexapac-ethyl delayed recovery from winter dormancy. xr'mnI: r 'cd r 6 Treatment effects on the quality and color of turf subjected to traffic were similar to those of untrafficked turf although actual values differed (Tables 34 and 35). Traffic treatments resulted in unacceptable turf quality and color (rating values < 5) two to three weeks after traffic applications began in 1994-1995 and three to five weeks after traffic applications began in 1996 (Tables 36 and 37). 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Ed the 3&ng 303:3— o; 2 _.m m.m we o6 $.3ng «Emzm $6on .E< c .82 S bum vm be“. a .22 cm .53. o EQEEEH moo— .=2 .9553 5mm £850 5883* mmfiwfish x828: .mcoEucoo Em: 3262 E oEwb 9 380.33 mmetB .3 .875 so so: can .Efioémmmonb .88on .3 80km Sm Bomb 68 .82 $2 2 Ba .6“ mm 8 ,0QO a 33% m2: .2 E 3.: 8: n .328 some 72 wx modv cog .a< cm :82 2 :5: _m A 72 wx 2.8 32 .80 0 ”385553 E :32? 8a v.88 $.23 w:_3o__8 of so 32%? 83 35983on; F .32 .32 2 Ba :2 3 :52 2 so .0QO a 33% a: 72 3 3.: so: a .326 :23 Ta; wx woe 39 .52 m: rnom w. :5: om A 72 wx a. .8 vmfl .80 m ”885:0th E :32: Rm 822 $83 wage—.8 2: co Big 33 350-8355; H EB 52w fimvnoE: $3 03885th 058020 o\ooo_ u 28 ”2QO 2:: 2 28 a co xzmsmg $38 was 8:5 + .bo>uuoam2 .m_o>o_ 323395 56 can mod 8 Emu—.tcwfi t...» m.m _.m wN 9m .5; ad ad v.0 two» m.m ON ”N v... m._ m6 v6 9o o: :o: 3.253 2 2E 69 eight to nine weeks in both years. Supina bluegrass showed signs of recovery, however, within three weeks after traffic treatments were ended; Kentucky bluegrass did not recover. Trinexapac-ethyl treatments resulted in superior recovery of Supina bluegrass compared to untreated turf (Tables 38 and 39) while Kentucky bluegrass was unaffected. As with non-trafficked turf, iron had little or no effect. 1' in i x r'ment I: Tur n u ' d Ira z Clipping yields of Kentucky bluegrass were significantly different compared to Supina bluegrass yields throughout the study (Table 40). Kentucky bluegrass clipping yields were greater than those of Supina bluegrass for the first two to three months inside the CSSF after which they were either no different (1995) or significantly less as the Kentucky bluegrass died (1996) (Table 41). In this study clipping yield data was only partly indicative of the turf’ 5 response to RLC; Supina bluegrass has a creeping growth habit which can be expected to result in less clipping yield compared to Kentucky bluegrass in even normal sunlight (Bemer, 1980). Trinexapac-ethyl treatments significantly reduced clippings of both species on most dates. As with the effect on color and quality, Supina bluegrass was more sensitive to trinexapac-ethyl than Kentucky bluegrass in terms of clipping yield reduction (Table 42). Lack of PGR efficacy after the first application and the rapid growth flushes following growth suppression commonly documented in non-RLC were not observed (Cooper et al., 1985; Shearing and Batch, 1982). The potential decline in turf color and quality observed due to the potential for retention of senescent foliage/suppression of new growth 7O .ABmv some 72 wx no.8 oaa_ ..a< cm :32 m. :5: _m A 7m: wx 2.8 maa_ .80 a “$855.8 5 :39? 2a moat $83 wEBo:£ 05 co 8:35 83 350-8355; m .328 some 7m: wx no.8 maa_ .82 2 snow 2 :52. cm A 7m: wx a_ .8 vaa. .80 m ”885:0qu E :39? 2a 8:: ”82% w530=£ 05 :o 3:95 83 350-3355; H £8 22388 “8 o:_m> E2558 05 mm; m ”f2 ESE: .omcouna .28 23%.: Eat face—u— mofiom 05: 9 one a co >=m=£> 8:2 33 £330 .r _._ ad ad md 0.0 $98 Dwa— o._ N._ o._ o._ N... m._ _ _ o._ m._ m._ 8.2ng 325:3— : Z 3. 2 S «M 3 2 nm E 889:3 2&5 mo» 0: mo» o: mo» 0: mo» 0: 8» o: 363m m 350-333,?“ 2:: : 52 am a: 2 .a< em .32 am caa_ ad ad wd Amodv Om; a.N _.m _.m 9v a.m w.m $.2ng 5.03:3— w.m m._ ad 9m 3. oh $8323 «Eqsm me» 0: mo» 0: me» o: 860nm H 350-8355: ho..— a .53. cm .5: o maa_ A2 .wfimfifi “mam .6800 5.808% 383.5% x085: £53628 Em: 3262 E oEmb 9 380.33 mmfiwta so 3258an Ecsémamonb$9m28am EmoE=w_m 05 com mwcufi >530 .mm 033. 71 A33 :08 7a: m: no.8 eaa_ .a< cm :32 2 £2 3“ A 72: w: a. .8 maa_ .80 a “88558“: E 5.6% 2a 822 $83 wEBo=£ 05 so 3:9? 83 Thfioomaaxofic. a .38: :08 _.m: w: no.8 maa_ .82 2 ran: a: $8.. om A 7a: m: a: .8 Va: .60 m 68055qu 5 =265. 2m 8:2 3.3% wage—:8 05 co Bias mm? 350-8%55; H “ta :0on faunas: f3 0:88:th cacao—:0 caco— n 25 6.an 2:: 9 28 m :o 2:253 :88 83 .830 + Nd N.N _._ 5o 9o A38 amu— o._ 3 3 3 I 3 2 3 n. 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E Va 3‘ Nam o.~ 0.. N.N N.N o.m vd NM fin We o: no: QM *ON EL.— ..imd 1.6.— _.:._N.~ .I.N._ ER.— ..:..m.N *m.N **m.v .50» Nm Tm _.m w.m Em m.m wN NM NV Qm v.0 0: 350-8355; _.m m6 0N m.N N.m *...o.m **o.m _.:..N.m 3&6 **N.v “aged games—n \0:93:02 3 3 3 3 2 Z I E 3 ma 3 magmas «59% E m 363m .82 mm .32 2 .82 m .32 _ bum mm how 2 how w bum _ .92. mm .53. M2 .5: 2 Enchant. moa— ..=z $5284 “mam c850 ganged mmfiwtsh x8283 £53650 Em: 3262 E 38th uoonabéo: («a £22» wzaazo no so: new .Rfioémamxofib .882? .3 Sootm .3. £an 74 .82 52 2 as non mm .5 6QO a 33%. a: 72 9. 3.: .5: _r A23 :08 Tu: wx no.8 com. .E< cm :52 2 :22. _m A 7m: wx 2 .8 32 .80 a ”835:0th E 550% 8a 3:2 ”8:3 wEBoze 2: so Bin? 83 ifioémqnxoafi. a .32 .32 2 2a :9; z .5. 2 8 Smog 8 33% 83 A .2 wx 3.: S: H A88 some 7m: 3 8.8 32 .52 2 spam M: :5: om A 72 wx E .8 32 .80 m ”$855.3 5 53% 8a 822 38% wEBozfl 2: :0 Rings 33 SSoéQOfiEfi. + 52883. .232 £3305 :3 Ba 36 a Escewa t... M: 2 .3 as 3 5 .5 to: 3 7% M: N.N nm E E 3 n: S: 3 2. .8: 23:58 3» 03m... 75 A83 some 7m; 9. 3.8 09$ .E< em 9.82 2 £5. 5 A was wx 2 .8 .32 .80 o ”momoficoaa E :32? 2a 3%... $83 336:8 2: co coins was 350-8355; H A83 :28 7mg wx no.8 mag .32 E rnom M: :52. om A was wv. 2 .8 go— .80 m ”$855.8,“ 5 Esonm 2m v.88 $.83 w53o=8 05 co com—mam £3 350-835:th + ad 5o v4 ad m.m m.m ad ”6 6o 3: .1. Na 5.2 ”.2 NE NE mmfiwozz 308:3. m._ 3. v; 9v _.m 2. Em 2: M: E: N.N ow 9v odm ms ad. mmfiwosz «Easm 0 we c me» 0: mo» 0: 8» o: 8% o: 8» 0: me» c: me» 0: 3» o: 83on «SSuémqaxofih 32 2 $2 x Q: m .a< om .a< 2 .a< _ .52 «N .52 E 009 3 3 mo 5. 3 3. _.~ 3 M: 2. mm 3 E 3 mm 3 383:3 Agassi ed 9N ed a; o6 To. Nd Wm 5o m._ o._ Wm o._ N.N NM— w.~ mwfiwozfi «Easm ~-E m me» c: we» on mm» o: 3% c: me» 0: me» o: 8% o: 8» o: 860% Lzfioémamxofih .32 w .32 _ bum mm bum 2 bum w bon— _ .52 mm .5: M: 32 dz .wfimamq “mam £850 :ohwomom mmSth. x0855 £83628 Em: 8262 E $8th noonmbéoc .8 $33 quEB so 86on can Efioégmxofib no 328885 Emoficwv. how 32m> .9 03m... 76 resulting from long-term PGR applications was also not observed (Watschke, 1976; Kaufmann, 1986b). Apparently the successive low rate (0.08 kg ha") applications of trinexapac-ethyl suppressed growth only to the point where carbohydrates may have been shifted to enhance tillering without causing growth cessation (Hanson and Branham, 1987). For practical use it may be important to be able to monitor the level of active PGR in the turf if PGRS are to be used on a continuous basis to maintain turf in RLC. Immunological techniques may prove to be the most expedient method for determining the amount of active ingredient in the plant. Ex "rim nt [1: ur '11!) 'e ' ’ tra I Treatment effects on the clipping yields of trafficked turf were similar to non- trafficked turf (Table 43) but actual yields were much lower than those from non- trafficked turf (Table 44). Trinexapac-ethyl continued to affect Supina bluegrass more significantly compared to Kentucky bluegrass (Table 45). In 1995 yields of Supina bluegrass were zero after Feb. 1 although turf yields in 1996 indicated continued growth throughout the trial. Since total inhibition of growth will slow or eliminate turf recovery it is important to match timing and rates of application of PGRS to maintain acceptable turf cover. W E r' ' t ' r l Turf species and trinexapac-ethyl consistently affected turf shear resistance while other treatment effects were rare and inconsistent (Table 46). Shear resistance declined 77 50380nt .m_o>o_ 325305 _o.o EB mod 2: E EmoEcwE 3.. .... on; 3.3 2.: 3.2. :d. 8.9. 8.8 a. _ m 3.: x .>o a: .o 32. $3 :3 3: ._ $3 N3; :3 a _ S 2 Sum 83 3.3 :3 $2“ 32 La: 5.8 $2.3 o: ._ _ m... x mp x m 32. 83 S: $3 33 we; _ _ S a _ .o Red _ m: x m: 83. 83 a: 83 ES 255 2.3 2 S 33 _ mm x m ~86 2 3 2 m... 83; 33 83 23 E _ .: 2 do _ ab .5: :83 83 38.2 SN.» :33 S 3. ~22: 33 o: .o _ me x m :33 83. an: 83 :Ene :89: :33? age 2.3 _ mp .33 :os .: cm: 33 92.2 80. _ N $3. :330 83 _ @ macaw 3: t a: a a: m .a< am .22 2 .23. _ a: «N a: 2 .32 m 0% EN. 032 $23 8.2: 2.3 6.2. 3.8 mm: 3.2 NS». Sam .x. .>0 8? mm; :3 83 83V 53 £3 32 82 8: £3 2 Sum 89o $2 .355 23 83 RS. 33 23 :50 2: .o :30: _ m: x E x m 23 m2: 08¢ 83. mm; 89 v8.0 33 :56 E: 22 _ m: x E. 3; ._ cam .53 £8 £2. 82 23 83 89¢ 22. £3 _ m: x m as; 3.2 Sod Sod 83. 83 03¢ 83 83 $2. $8; _ 33 .5: onto :2 .136 .22 :33. £356 .33 :§._ :5; wood :2: _ me x m :3 Sad :83 :ofle :83 :3 no :82 :82 :53 :20: :Equ _ me $3 $3 82 .22 :33 .8? :NSo :c8.§ :03: £38 :mflafl _ av 860% .52 mm .52 2 .52 w .32 _ .fia mm a: 2 .fia m sea _ .5: mm .5: M: .52 2 :c 853 82 :2 $2355 “mam .3300 5.880% mmflwta. x828: £52280 Em: 30sz 5 uEmb 9 c8833 38th mo mac—2» wcaazo :o 303% 82585 mo ougoc_:w_m 38 83:3. :82 .3 03m... 78 .390 {ad 0; ”M _.:._w._ f..m.N 3.9% w.v m._ am?» m._ w; QM Em m.m .6 ad Ev _.m o: 35063305; .2 :3 3 mm mm 3 E to E 383:3 5.323. m2 QN o.m N6 52 vd 3‘ _.m Nu— mmfiwosfi 2:96 E w museum 32 2 b3 u bwE m a< am .5. 2 .a< _ .52 E. .32 2 .52 w 2258: 32 2 ad Yo Yo wd ed 5o m._ o.m wd fie H3» 5o Yo Yo Yo 5o wd 9o m2 o.m 5m 9m 0: so: We v.0 .3." _ .o trod ..:..N.o *.._N.o frmd .I,w.o firm.— **O.N .256 .2. m2» 2 ad 5o Mg m._ N; ad 2 "mm m.m ms 0: 35868305; m..— o._ 9o :06 11 *2 3.. _ ._ :NN film Ioé 33.x 383:3 .9835! 2 no 3 mo 3 S 3 do no I 2 $3.33 2&3 N.E w 363m .32 mm .82 2 .32 m .32 _ bon— NN how 2 bum m be; _ .52 mm .5: M: 5:. 2 Hanged. 32 ._2 $5234 “mam .5250 53832 mmfiwtsh x085: .mcoEncou Em: 8262 E oEmb 8 380.33 38th go .633 wcaazo co 350-3355: can 860% .3 38.2.5 .3 033 79 .82 32 2 Ba .8“. mm :0 6QO 8 33% a: :2 mx 3.: no: _r A23 some 72 wx M33 03— .E< cm £22 m. :5: _m A 7a: wx S .8 32 .80 o ”momoficobwa E 5505 2a mesh 68% wEBo:£ 2: so 32%? mm? Efioéamaxofifi a .82 .52 t 23 so; 2 :52 2 8 6%“. 8 33% as A :2 w; 2.: :2. H A88 some 72 wx wodv 39 .32 2 rnom M: :5: cm A 7m: wx 2.8 go. .80 m ”885:0th E 5505. 2a 82$ 38% wage—.8 05 co 3:qu mmB ifiobamxocth + 52:89.2 .232 £3305 :3 as 8d a Eocewa :9. ad 2 3 3 E ”A E E M: F 3» 3 3 _.~ 3 X .3 mm 3 E 2. :8. 3:53 3 22¢ 80 A33 none 72 wx no.8 82 ..a< on :82 m.— ..5: 1m A 72 wx o. .8 32 .80 o “momoficoaa E 5597. 2a 8E ”menu 9.50:8 2: co 3:an 83 350-3355; H Aofiu some 72 wx no.8 3E .32 3 room M: :5: cm A 7s... wx 2.8 $2 .80 m ”8855.22. 5 56% 2m 8:: $23 wEBo—é 2: co v3.3a 33 150-8355; + v.0 ad ozm 8 ad 3 m: m: Amodv Om: We Yo wd We a; 04 Q». 9N N.m m.m QM 9v 9m 2. :. Qm 389:3 5.08:3. 2 3 3 3 3 we 5 3 no 3 2 cm «A no 3 5 383:3 2&3 #8 w no» 0: mo» 0: me» 0: me» o: 8» o: 8% 0: we» on me» o: mfluoam H 35983555 $2 2 32 w 32 m .a< om .a< a .a< _ .52 a .32 N. coa— 2 2. no 3 no to 9o 3 A38 mm; _.o m._ od m._ vd Wm Yo Om 5o 0.. o; w.m Nam 3V Qm 0A: 3.3ng 533:3— o.o md ed to ad No od _.o od _.o o.o md md 5o 9m 5v mmfiwoan Siam E w NO 8» 0: mo» 0: 3» 0: mm» 0: me» o: 8» o: 8% 0: 8» o: 86on * Efioémamxofih .32 w .32 _ do“. mm bon— m. .nom w bum _ .23. mm .5: 2 E2535. moo— .=2 .wfimcmq “mam c850 5883* $8th x085: .mcoEnzoo Em: 3268 E stab 2 380.33 323:: mo 33% wEaE—u so 860% new 350-0333?“ .3 328885 EmoEcwv. Eat mo=_m> .mv 2an 81 Table 46. Mean square and significance of treatment effects on shear resistance of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 1994-1995 Source of variation df 20 Dec. 20 Jan. 22 Feb. 31 Mar. Species (S) 1 369.094" 2.344 546.260" 240.667" Trinexapac-ethyl (TE) 1 0.010 94.010" * 0.260 20167“ S x TE 1 3.760 10.010 1.760 3.375 Iron (Fe) 1 3.760 15.844 3.760 2.042 S x Fe 1 0.010 17.510 0.260 2.667 TE x Fe 1 0.010 36.260 1.260 32.667" S x TE x Fe 1 7.594“ 0.094 0.010 15042" Error 16 1.042 17.760 2.271 3.219 CV, 0/o 5.57 25.77 10.17 14.08 1996 Source of variation (if 17 Jan. 12 Mar. 31 May Species (S) l 121.500M 48.167* 28.711” Trinexapac-ethyl (TE) 1 l .500 18.375 5.753"‘ S x TE 1 7.042 15.042 6773* Iron (Fe) 1 4.167 5.042 0.023 S x Fe 1 2.042 30.375 0.315 TE x Fe 1 7.042 16.667 0.003 S x TE x Fe 1 0.667 32.667 0.065 Error 16 2.281 8.000 0.815 CV, % 7.29 16.80 10.31 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. 82 over time in both years regardless of treatment (Table 47). The decline in shear resistance indicated a lack of sufficient turf growth and rooting probably due to insufficient light energy to sustain turf permanently. In the first year Kentucky bluegrass had higher shear resistance than Supina bluegrass probably due to the presence of rhizomes in Kentucky bluegrass which added stability to the turf (McNitt, 1994). Shear resistance values of Kentucky bluegrass were significantly lower than those of Supina bluegrass in 1996, perhaps due to increased root growth of Supina bluegrass during the longer establishment period in autumn of 1995 compared to autumn 1994. However the practical significance of such a difference in shear resistance may not be important as values were still relatively close. In addition, a desirable value for shear resistance using the Eij kelkamp shear vane has not been established despite previous attempts (Liesecke and Schmidt, 1978) . Shear resistance values are generally important for their ability to indicate relative turf strength and rooting differences. In 1996 the shear resistance of Supina bluegrass was significantly increased by trinexapac-ethyl while Kentucky bluegrass shear resistance was unchanged (Table 48). Trinexapac-ethyl enhanced Supina bluegrass growth and development more than Kentucky bluegrass, which resulted in more biomass aboveground which enhanced the shear resistance of the Supina bluegrass. E r' ' r ' t r Treatment effects on turf shear resistance were similar to those on non-trafficked turf (Table 49). Shear values declined over time due to loss of turf density and actual values were lower than for non-trafficked turf (Table 50). Supina bluegrass continued to be 83 Table 47. Effect of species, trinexapac-ethyl, and iron on shear resistance values (N'm) of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 1994-1995 Treatment 20 Dec. 20 Jan. 22 Feb. 31 Mar. Species N m Supina bluegrass 14.2 16.0 10.0 9.0 Kentucky bluegrass 22.4” 16.7 19.6M 15.3M Trinexapac-ethyl no 18.3 14.4 14.7 11.2 yesl' 18.3 18.3" 14.9 130* Iron no 18.7 15.5 15.2 12.4 yes: 17.9 17.2 14.4 11.8 1996 Treatment 17 Jan. 12 Mar. 31 May Species N m Supina bluegrass 23.0 18.2 9.9 Kentucky bluegrass 18.5” 15.4“ 7.7M Trinexapac-ethyl no 20.5 16.0 8.3 yes§ 21.0 17.7* 92* Iron no 20.3 16.4 8.7 yes1l 21.1 17.3 8.8 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. TTrinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (o. 19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). 1' Iron (1.14 kg ha'l ) was applied as FeSO4 on 10 Jan., 14 Feb., 17 Mar 1995. § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha'l ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha'l each date). 1| Iron (1.14 kg ha'l ) was applied as FeSO4 on 28 Feb. and 13 May 1996. 84 Table 48. Shear resistance values (N-m) for the significant species-by-trinexapac-ethyl interaction on non- trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 31 May 1996 trinecapac-ethylt Species no yes N m Supina bluegrass 8.8 10.9 Kentucky bluegrass 7.7 7.6 LSD (0.05) 1.1 l“ Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). Table 49. Mean square and significance of treatment effects on shear resistance of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 1994-1995 Source of variation df 20 Jan. 22 Feb. 31 Mar. Species (S) 1 8.167 546.260M 152.510" Trinexapac-ethyl (TE) 1 60.167 10.010 15.844* * S x TE 1 12.042 1.260 8760* Iron (Fe) 1 4.167 8.760 10010“ S x Fe 1 18.375 6.510 4.594 TE x Fe 1 22.042 1.760 0.844 S x TE x Fe 1 1.500 0.260 25.010" Error 16 15.010 3.083 1.948 CV, % 25.62 13.57 15.47 1996 Source of variation (if 12 Mar. 31 May Species (S) 1 45.375* 19.260" Trinexapac-ethyl (TE) 1 5.042 16.667" S x TE 1 2.667 24.000" Iron (Fe) 1 5.042 2.344 S x Fe 1 6.000 0.094 TE x Fe 1 6.000 0.667 S x TE x Fe 1 15.042 0.167 Error 16 6.667 1.255 CV, % 18.33 15.86 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. 85 Table 50. Effect of species, trinexapac-ethyl, and iron on shear resistance values (N -m) of turfgrass subjected to trafficT in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 1994-1995 Treatment 20 Jan. 22 Feb. 31 Mar. Species N m Supina bluegrass 14.5 8.2 6.5 Kentucky bluegrass 15.7 17.7“ 11.5” Trinexapac-ethyl no 13.5 12.3 8.2 yesi 16.7* 13.6 9.8" Iron no 14.7 12.3 9.7 yes§ 15.5 13.5 8.4* 1996 Treatment 12 Mar. 31 May Species N m ----------------- Supina bluegrass 15.5 8.0 Kentucky bluegrass 12.7 6.2" Trinexapac-ethyl no 13.6 6.2 yesfl 14.5 7.9” lron no 13.6 7.4 yes# 14.5 6.8 *,** Significant at the 0.05 and 0.01 probability levels, respectively. T Traffic was applied by persons who jogged on the turf 50 passess each week while wearing molded soccer cleats on the following dates: 28 Dec. 1994 through 16 Mar. 1995 and 26 Jan. through 26 Apr. 1996. I Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha'1 ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.03 kg ha" each date). § Iron (1.14 kg ha'I ) was applied as FeSO4 on 10 Jan., 14 Feb., 17 Mar 1995. 11 Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (o. 19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha‘1 each date). # Iron (1.14 kg ha'l ) was applied as FeSO4 on 28 Feb. and 13 May 1996. 86 more sensitive to trinexapac-ethyl applications in both 1995 and 1996 compared to Kentucky bluegrass (Table 51). Blantdensjtyjnihiomass Ex r' n ' T r ' 1 Species, trinexapac-ethyl, and species-by-trinexapac-ethyl interactions existed for specific plant weight, plant density, tiller, and leaf counts (Table 52). The turf density of Supina bluegrass was much greater than Kentucky bluegrass and positively influenced quality ratings. Supina bluegrass had more tillers and leaves per plant plus a higher specific plant weight (Table 53). Plant density (number plants per unit area) was either the same or greater than Kentucky bluegrass. Trinexapac-ethyl increased turf density by increasing the plant density, number of tillers and leaves per plant, and specific plant weight. Apparently trinexapac-ethyl was effective at repartitioning carbohydrates in the plant to produce axillary tillers and more leaves per plant similar to the effects of paclobutrazol and fiurprimidol (Hanson and Branham, 1987). Turf left untreated, 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 increased leaf area index (LAI) resulting from the trinexapac-ethyl application. As with color, quality, and clipping yields, Supina bluegrass was more sensitive to trinexapac-ethyl than Kentucky bluegrass and exhibited a strong positive response while Kentucky bluegrass was largely unaffected (Table 54). 87 Table 51. Shear resistance values (N -m) for the significant species-by-trinexapac-ethyl interactions on turfgrass subjected to traffic’( in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 31 March 1995 31 May 1996 trinexapac-ethyl' Species no yes: no yes§ N m Supina bluegrass 5.1 7.9 6.1 9.8 Kentucky bluegrass 1 1.3 1 1.8 6.3 6.0 LSD (0.05) 1.6 1.0 1' Traffic was applied by persons who jogged on the turf 50 passess each week while wearing molded soccer cleats on the following dates: 28 Dec. 1994 through 16 Mar. 1995 and 26 Jan. through 26 Apr. 1996. I Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha'l ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha‘1 each date). § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha'I ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha'1 each date). 88 Table 52. Mean squares and treatment effects on plant density and biomass of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 12 April 1995 Plant density Specific plant Source of variation df (No. plants m'z) Tillers plant'l Leaves plant’I weight (mg) Species (S) 1 1695697630 15.360” 180950" 94.169“ Trinexapac-ethyl (TE) 1 13834140784" 2.667* 34.800" 143.277" S x TE 1 749137.231 3.227* 33.844“ 48.053" Iron (Fe) 1 4924.957 0.202 3.300 1.242 S x Fe 1 60.805 0.135 1.170 0.814 TE x Fe 1 102207.627 0.375 13.650 14.291 S x TE x Fe 1 572082.701 0.482 3.920 2.857 Error 16 610266.41 1 0.400 4.872 7.807 CV, % 13.71 33.27 30.82 29.72 31 May 1996 Plant density Specific plant Source of variation df (No. plants m'z) Tillers plant'l Leaves plant'l weight (mg) Species (S) 1 20665817941“ 10.402M 388.815" 2061.277“ Trinexapac-ethyl (TE) 1 832314.080" 0.807” 12.042" 10.010 S x TE 1 1208798011“ 0.807" 1.500 2.306 Iron (Fe) 1 26813.528 0.240 2.282 58.033 S x Fe 1 32164.085 0.240 8.167 191.648 TE x Fe 1 1520.039 0.082 1.500 198.490 S x TE x Fe 1 547.217 0.082 0.015 220.584 Error 16 28759.193 0.087 2.187 66.212 CV, % 15.79 17.84 20.17 33.78 *,** Significant at the 0.05 and 0.01 probability levels, respectively. 89 Table 53. Effect of species, trinexapac-ethyl, and iron on plant density and biomass of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 12 April 1995 Plant density Specific plant Treatment (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species Supina bluegrass 5430.8 2.7 9.9 1 1.4 Kentucky bluegrass 5962.4 1.1" 4.4* 7.4" Trinexapac-ethyl no 4937.4 1.6 6.0 7.0 yesl‘ 6455.8" 2.2* 8.4" 11.9" lron no 5710.9 1.8 6.8 9.2 yes: 5682.2 2.0 7.5 9.6 31 May 1996 Plant density Specific plant Treatment (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species Supina bluegrass 1990. 2.3 1 1.4 33.4 Kentucky bluegrass 134" 1.0" 3.3" 14.8" Trinexapac-ethyl no 875 1.5 6.6 23.4 yes§ 1248'” 1.8" 8.0“ 24.8 lron no 1028 1.6 7.0 22.5 yesfil 1095 1.8 7.6 25.6 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. TTrinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). 1 Iron (1.14 kg ha'1 ) was applied as FeSO, on 10 Jan., 14 Feb., 17 Mar 1995. § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). 11 Iron (1.14 kg ha" ) was applied as FeSO4 on 28 Feb. and 13 May 1996. 90 Table 54. Plant density and biomass values for the significant species-by-trinexapac-ethyl interactions on non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, Ml. 12 April 1995 Plant density Specific plant (no. plants m'2 ) Tillers plant" Leaves plant" weight (mg) trinexapac-ethylf Species no yes no yes no yes no yes Supina bluegrass 49370 6366 2.0 3.4 7.5 12.3 7.5 15.2 Kentucky bluegrass 5379 6544 1.1 1.1 4.4 4.4 6.4 8.4 LSD (0.05) ns 0.8 2.7 3.4 31 May 1996 Plant density Specific plant (no. plants m'2 ) Tillers plant" Leaves plant" weight (mg) trinexapac-ethylz Species no yes no yes no yes no yes Supina bluegrass 1579 2400 2.0 2.7 10.4 12.3 33.0 33.7 Kentucky bluegrass 172 96 1.0 1.0 2.8 3.8 13.9 15.8 LSD (0.05) 208 0.4 ns ns 1' Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (o. 19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). I Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). 91 Trinexapac-ethyl had less effect on the biomass of turf subjected to traffic compared to non-trafficked turf (Table 55). Supina bluegrass continued to exhibit a higher plant density, more tillers and leaves per plant, and higher specific plant weight compared to Kentucky bluegrass (Table 56). In addition, both species exhibited a higher specific plant weight compared to untrafficked turf apparently due to less competition for light and perhaps water and nutrients. Unlike non-trafficked turf, trinexapac-ethyl had little effect on response to traffic between species (Table 57). Chlamhxflflmemraticn xr'mn'Tlrn u' t Species and trinexapac-ethyl significantly affected chlorophyll a and b levels and total chlorophyll (Table 58). The greater levels of chlorophyll a, b, and total chlorophyll in Kentucky bluegrass compared to Supina bluegrass were consistent with the darker green color of Kentucky bluegrass but were not effective in providing superior shade tolerance (Table 59). Trinexapac-ethyl also enhanced chlorophyll concentration (leaf area basis) in both turf species probably due to decreased cell enlargement. Trinexapac-ethyl cannot be expected to affect chlorophyll synthesis directly since its known modes of action have been described as blocking gibberellic acid (GA) biosynthesis only at the end of the pathway, primarily by preventing 3-13 hydroxylation of the biologically inactive GAZO to the biologically active GA, (Rademacher, 1991). Since carotenoids and GA, have a common precursor, geranylgemalypyrophosphate (GGPP), it is possible GA inhibition 92 Table 55. Mean squares and treatment effects on plant density and biomass of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Resarch Center, East Lansing, MI. 12 April 1995 Plant density Specific plant Source of variation df (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species (S) 1 1368037471 15.844" 301.750" 193.007" Trinexapac-ethyl (TE) 1 51 13419.885 * 0.260 9.004 9.805 S x TE 1 107254.135 0.304 3.450 1.540 Iron (Fe) 1 8755.448 0.000 0.120 4.200 S x Fe 1 190673.985 0.020 2.100 1.101 TE x Fe 1 1 17712.025 0.004 0.454 1.224 S x TE x Fe 1 78798.975 0.000 2.220 1.325 Error 16 689430.079 0.293 4.807 8.944 CV, % 39.94 26.78 25.16 21.78 31 May 1996 Plant density Specific plant Source of variation df (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species (S) 1 5545355292" 6.934" 320.470“ 2058.869“ Trinexapac-ethyl (TE) 1 818147.252M 0.350 6.720 110.039 S x TE 1 934886.447" 0.510 4.420 98.537 Iron (Fe) 1 41101.925 0.070 1.450 5.482 S x Fe 1 107254.140 0.020 0.400 31.763 TE x Fe 1 35021.756 0.034 0.634 0.196 S x TE x Fe 1 0.000 0.004 0.034 224.298 Error 16 94729.003 0.125 2.390 125.984 CV, % 50.62 22.59 22.95 39.29 ‘2" Significant at the 0.05 and 0.01 probability levels, respectively. 93 Table 56. Effect of species, trinexapac-ethyl, and iron on plant density and biomass of turfgrass subjected to trafficl in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 12 April 1995 Plant density Specific plant Treatment (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species Supina bluegrass 1840 2.8 12.3 16.6 Kentucky bluegrass 2317 1.2“ 5.2” 10.9” Trinexapac-ethyl no 1617 1.9 8.1 13.1 yes§; 2540* 2.1 9.3 14.4 lron no 2098 2.0 8.8 14.1 yes§ 2060 2.0 8.6 13.3 31 May 1996 Plant density Specific plant Treatment (No. plnts m") Tillers plant" Leaves plant" weight (mg) Species Supina bluegrass 1089 2.1 10.4 37.8 Kentucky bluegrass 127* * 1.0M 3.1" 19.3" Trinexapac-ethyl no 423 1.4 6.2 26.4 yesfil 793" 1.7 7.3 30.7 lron no 649 1.5 6.5 28.1 yes# 567 1.6 7.0 29.0 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. ‘1 Traffic was applied by persons who jogged on the turf 50 passes each week while wearing molded soccer cleats on the following dates: 28 Dec. 1994 through 16 Mar. 1995 and 26 Jan. through 26 Apr. 1996. TTrinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). I Iron (1.14 kg ha" ) was applied as FeSO4 on 10 Jan., 14 Feb., 17 Mar 1995. § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). 11 Iron (1.14 kg ha" ) was applied as FeSO4 on 28 Feb. and 13 May 1996. 94 Table 57. Plant density and biomass values for the species-by-trinexapac-ethyl interactions on turfgrass subjected to traffic* in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 12 April 1995 Plant density Specific plant (no. plants m'2 ) Tillers plant" Leaves plant" weight (mg) trinexapac-ethylI Species no yes no yes no yes no yes Supina bluegrass 1445 2234 2.6 3.0 11.3 13.2 16.2 17.0 Kentucky bluegrass 1789 2846 1.2 1.2 4.9 5.4 10.0 1 1.8 LSD (0.05) ns ns ns ns 31 May 1996 Plant density Specific plant (no. plants m'2 ) Tillers plant" Leaves plant" weight (mg) trinexapac-ethyl§ Species no yes no yes no yes no yes Supina bluegrass 707 1471 1.8 2.4 9.4 1 1.4 33.7 42.0 Kentucky bluegrass 140 1 14 1.0 1.0 3.0 3.2 19.2 19.4 LSD (0.05) 377 ns ns ns ‘1 Traffic was applied by persons who jogged on the turf 50 passes each week while wearing molded soccer cleats on the following dates: 28 Dec. 1994 through 16 Mar. 1995 and 26 Jan. through 26 Apr. 1996. I Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (o. 19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). 95 Table 58. Mean squares and treatment effects on chlorophyll of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 4 April 1995 Total Chlorophyll Source of variation df Chlorophyll a Chlorophyll b chlorophyll asb Species (S) 1 238.644" 30.173" 435.968" 0.029 Trinexapac-ethyl (TE) 1 314.071" 47.124" 601.301" 0.163* S x TE 1 1.815 0.008 2.227 0.000 Iron (Fe) 1 5.587 0.980 10.921 0.000 S x Fe 1 4.267 0.022 5.125 0.035 TE x Fe 1 1 1.289 1.701 21.263 0.001 S x TE x Fe 1 19.802 1.071 30.759 0.009 Error 16 13.265 2.285 26.225 0.027 CV, % 15.21 19.45 16.16 5.25 29 May 1996 Total Chlorophyll Source of variation df Chlorophyll a Chlorophyll b chlorophyll asb Species (S) 1 32.109 2.734 55.937“ 0.029 Trinexapac-ethyl (TE) 1 121.590” 10.935M 200.797 *" 0.020 S x TE 1 8.592 0.680 15.360 0.029 Iron (Fe) 1 19.91 1 2.257 33.844 0.032 S x Fe 1 13.681 1.344 22.042 0.000 TE x Fe 1 0.341 0.070 1.033 0.022 S x TE x Fe 1 0.928 0.109 1.288 0.034 Error 16 4.710 0.470 7.887 0.043 CV, % 10.03 9.78 9.79 6.70 *,** Significant at the 0.05 and 0.01 probability levels, respectively. 96 Table 59. Effect of species, trinexapac—ethyl, and iron on chlorophyll content of non-trafficked turfgrass in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 4 April 1995 Total Chlorophyll Treatment Chlorophyll a Chlorophyll b chlorophyll axb 11g cm'2 leaf tissue Species Supina bluegrass 20.8 6.7 27.4 3.1 Kentucky bluegrass 27.1" 8.9" 36.0" 3.1 Trinexapac-ethyl no 20.3 6.4 26.7 3.2 yesl 27.6” 9.2" 36.7" 30* Iron no 24.4 8.0 32.4 3.1 yes§: 23.5 7.6 31.0 3.1 29 May 1996 Total Chlorophyll Treatment Chlorophyll a Chlorophyll b chlorophyll asb pg cm'2 leaf tissue Species Supina bluegrass 20.5 6.7 27.1 3.1 Kentucky bluegrass 228* 7.3 30.2" 3.1 Trinexapac-ethyl no 19.4 6.3 25.8 3.1 yes§ 23.9" 7.7** 31.6M 3.1 Iron no 20.7 6.7 27.5 3.1 yes1l 22.5 7.3* 29.9 3.1 ‘, " Significant at the 0.05 and 0.01 probability levels, respectively. TTrinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha'1 each date). 1 Iron (1.14 kg ha'l ) was applied as FeSO4 on 10 Jan., 14 Feb., 17 Mar 1995. § Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 9 Oct. 1995 (0.19 kg ha" ), 31 Jan., 15 Mar., 26 Apr. 1996 (0.08 kg ha" each date). 11 Iron (1.14 kg ha" ) was applied as FeSO4 on 28 Feb. and 13 May 1996. 97 could cause a feedback mechanism to shunt additional GGPP to carotenoid production. Current evidence does not support the existence of such a feedback mechanism as typically non-active gibberellins(GA1 precursors) continue to accumulate in the presence of a (GA) biosynthetic inhibitor (Rademacher, 1991). Iron application caused only minor, temporary darker green turf color. Chlorophyll levels were relatively unaffected within two weeks following an application of iron. This result suggests the plants were already at their maximum capacity for using iron for chlorophyll production or else energy levels within the turfgrass plants were too low to utilize the iron. Auxiliary studies showed that while foliar applications of iron failed to enhance turf color or chlorophyll levels in Kentucky bluegrass, iron levels in plant tissues were increased threefold following an application of iron sulfate at the same rate used in the current study (unpublished data). Another possible explanation for the inconsistency between color enhancement and lack of effect on chlorophyll content is that the leaves used for chlorophyll analysis might have been partially unexpanded at the time of the iron application thus the effect would have been seen particularly on older leaves. Chlorophyll asb ratios were relatively unaffected by any treatment. Chlorophyll asb ratios were approximately 3:1 which is equivalent to the ratio of approximately 3:1 observed for sun plants (Nobel, 1991). This indicates Supina bluegrass is not a “shade” plant per se but apparently has mechanisms for shade tolerance which are lacking in Kentucky bluegrass. 98 E r'm ' r Treatment effects on chlorophyll quality and quantity were similar between trafficked and untrafficked turf in 1995 (Table 60). As expected, data from 1995 indicated traffic did not affect chlorophyll content (Table 61). Data were not collected in 1996 due to insufficient plant material (the youngest fully matured leaves were consistently too short from mowing to use in analysis). CONCLUSIONS Supina bluegrass was more tolerant of RLC than Kentucky bluegrass. The light conditions tested were too low to sustain Supina bluegrass permanently in a trafficked conditions and were marginal for non-trafficked conditions. The enhanced growth of Supina bluegrass due to increasing light levels and photoperiod in the spring indicated the actual light requirement to sustain Supina bluegrass under traffic was greater than the test conditions generally provided. Iron had negligible effect on any characteristic of either turfgrass species. Trinexapac- ethyl treatments provided superior enhancement of color and quality compared to iron. In addition, iron did not provide the enhanced biomass associated with trinexapac-ethyl treatments. Supina bluegrass was consistently more responsive to trinexapac-ethyl than Kentucky bluegrass. Trinexapac-ethyl may have helped to increase the turf quality in RLC by shifting carbohydrate partitioning away from primary shoots to stimulate tillering with a subsequent increase in the LAI in addition to promoting a more prostrate 99 Table 60. Mean squares and treatment effects on chlorophyll of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, MI. 4 April 1995 Total Chlorophyll Source of variation df Chlorophyll a Chlorophyll b chlorophyll a:b Species (S) 1 104.125M 14.774" 278.734" 0.077" Trinexapac-ethyl (TE) 1 179.252* * 23. 108* * 328486" * 0.027 S x TE 1 0.017 0.134 4.708 0.010 Iron (Fe) 1 5.636 0.485 12.600 0.007 S x Fe 1 0.956 0.002 5.180 0.020 TE x Fe 1 24.990 3.190 52.896* 0.000 S x TE x Fe 1 13.395 3.046 4.797 0.056 Error 16 7.701 1.048 11.416 0.013 CV, % 10.98 12.94 10.18 3.54 *,** Significant at the 0.05 and 0.01 probability levels, respectively. Table 61. Effect of species, trinexapac-ethyl, and iron on chlorophyll content of turfgrass subjected to traffic in reduced light conditions, Hancock Turfgrass Research Center, East Lansing, M1.T 4 April 1995 Total Chlorophyll Treatment Chlorophyll a Chlorophyll b chlorophyll asb pg cm'2 leaf tissue Species Supina bluegrass 23.2 7.1 29.8 3.3 Kentucky bluegrass 27.4* * 8.7* * 36.6* * 3 .2* Trinexapac-ethyl no 22.5 6.9 29.5 3.3 yes: 28.0" 8.9M 36.7” 3.2 Iron no 24.8 7.8 32.5 3.2 yes§ 25.8 8.1 33.9 3.2 *, ** Significant at the 0.05 and 0.01 probability levels, respectively. T Traffic was applied by persons who jogged on the turf 50 passes each week while waring molded soccer cleats from 28 Dec. 1994 through 16 May 1995. I Trinexapac-ethyl was applied on the following dates; rates are shown in parentheses: 3 Oct. 1994 (0.19 kg ha" ), 20 Jan., 18 Feb., 16 Mar. 1995 (0.08 kg ha" each date). §1ron (1.14 kg ha" )was applied as FeSO4 on 10 Jan., 14 Feb., 17 Mar 1995. 100 and compact growth. These attributes could be expected to enhance net photoassimilation on a turf area basis, creating a favorable cycle as more carbohydrates can be produced to regenerate tissues damaged by traffic or disease. Supina bluegrass was superior to Kentucky bluegrass in RLC due in part to an apparent resistance to powdery mildew. Supina bluegrass was more susceptible to pink snow mold in RLC compared to Kentucky bluegrass. This problem was partly controlled by providing wind movement over the turf with portable fans although occasional fungicide applications were still necessary to prevent noticeable disease damage. Chapter 3 PHOTOSYNTHESIS OF SUPINA BLUEGRASS (POA SUPINA SCHRAD.) AND KENTUCKY BLUEGRASS (P. PRA T ENSIS L.) IN REDUCED LIGHT CONDITIONS AS AFFECTED BY NITROGEN AND TRINEXAPAC-ETHYL INTRODUCTION Turfgrass performance in reduced light conditions (RLC; < 30% sunlight) is often poor due to insufficient light for photosynthesis and normal turf growth. Turfgrass species and cultivars may vary widely in their tolerance to RLC although all may exhibit reduced tillering, reduced rooting, and an upright spindly growth (Beard, 1973). Supina bluegrass (Poa supina Schrad.), a stoloniferous grass native to the sub-alpine regions of Europe, has been developed in Germany as a turfgrass with purportedly good to excellent shade and traffic tolerance (Bemer, 1984; Nonn, 1994; Pietsch, 1989; Skirde, 1971). Preliminary research supports the hypothesis that Supina bluegrass is more tolerant of RLC than Kentucky bluegrass (P. pratensis L.) (Stier and Rogers, 1995) which is commonly used in the United States but has poor shade tolerance (Beard, 1973). The mechanism(s) for the apparent shade tolerance of Supina bluegrass is/are unknown. In addition to the use of shade-tolerant turfgrasses proper management techniques are also important for turf performance in RLC. Previous research has indicated the potential for plant growth regulators (PGRS) which inhibit gibberellic acid (GA) biosynthesis to improve turf quality in reduced light conditions (Rogers et al., 1996; Stier and Rogers, 101 102 1995). Turf treated with GA-inhibitors in RLC was more uniform with darker color and increased density compared to untreated turf. While the GA-inhibitors effectively suppressed shoot elongation, other mechanisms by which the GA-inhibitors improved turf quality were unknown. Possibilities range from enhanced photosynthetic rates (Gausman et al., 1991), increased carbohydrate production or partitioning (Hanson and Branham, 1987; Wang et a1. 1985), increased chlorophyll levels (Wang et a1 1985, Archbold and Houtz, 1988), increased protein/enzyme levels and/or activity (Wang et al. 1985), altered hormonal levels affecting foliar production (Gausman et al., 1991), to gene expression (Gausman et al., 1991). Conversely, Archbold and Houtz (1988) reported flurprimidol and paclobutrazol decreased photosynthetic rates and Rubisco activities in strawberry plants. DeJong and Doyle (1984) found paclobutrazol reduced shoot growth of nectarine trees but did not affect photosynthesis. Mefluidide, generally considered a mitotic inhibitor which also may inhibit GA-biosynthesis (Wilkenson, 1982), consistently reduced photosynthetic rates of ‘Baron’ Kentucky bluegrass while amidochlor occasionally enhanced photosynthesis (Spokas and Cooper, 1991). In the early 1990’s a new GA-inhibitior, trinexapac-ethyl (TE), was labeled for use on turfgrasses, primarily to decrease mowing requirements by suppressing shoot growth (Vitolo et al., 1990). The potential side effects of TE on plant physiology are relatively unknown due to its recent release but may be different than other turf GA-inhibitors. TE apparently blocks 3-B oxidation of the biologically inactive GAzo to form the biologically active GA, as opposed to flurprimidol and paclobutrazol which inhibit ent-kaurene 103 oxidation oxidative steps earlier in the biosynthetic pathway (Coolbaugh et al., 1982; Rademacher, 1991). In normal (full sun) conditions GA-inhibitor effects on turfgrass can vary with nitrogen (N) rate and turf species or cultivars. Watschke (1981) found differences in responses of two Kentucky bluegrass culitivars (‘Merion’ and ‘Pennstar’) to paclobutrazol and flurprimidol. Other studies showed high N rates reduced the effects of flurprimidol on common bermudagrass [C ynodon dactylon (L.) Pers.] (Devitt and Morris, 1988) but not on ‘Tifway’ hybrid bermudagrass [Cynodon transvaalensis Burtt-Davy x C. dactylon (L.) Pers.] (Johnson, 1988). Johnson (1994) corroborated the differences in response to trinexapac-ethyl between common bermudagrass and ‘Tifway’ hybrid bermudagrass. In RLC of approximately 5-6 mol photosynthetically active radiation (PAR) day", medium to high N rates (48 and 96 kg ha" at four to six week intervals) resulted in significantly better quality Kentucky bluegrass compared to low N rates (24 kg ha" at four to six week intervals) when flurprimidol was applied, although low and medium N rates provided superior turf in the absence of flurprimidol (Chapter 1). Due to the demand for improved turfgrasses and management schemes for turf in RLC, studies were initiated to examine the effects of N rate, trinexapac-ethyl, and species on turf photosynthesis in RLC. Two hypotheses were tested: 1) Supina bluegrass was more tolerant of RLC compared to Kentucky bluegrass due to a greater carbon exchange rate (CER), i.e., enhanced photosynthetic rate, and 2) Trinexapac-ethyl improved turfgrass quality in RLC by enhancing CER. The objectives of this research were to determine if differences in CER existed between Supina bluegrass and Kentucky 104 bluegrass and to determine the effects of trinexapac-ethyl on CER of the two species. A second set of objectives were to determine the influence of nitrogen rate and trinexapac- ethyl on the CER of Supina bluegrass. MATERIALS AND METHODS £11 111'! | 111' x e ' ' 1 Portable plots were established outside in full sun conditions. Wooden boxes (1.2 x 1.2 x 0.15 m depth) were filled with a sandzpeat mixture (80:20 v/v) (Table 78, Appendix). The pH was 7.8 with initial P and K levels of 85 kg ha" and 90 kg ha", respectively. Sixteen holes (0.6 cm diam) were drilled on approximately 23 cm spacings in the bottom of each box to provide drainage. Starter fertilizer (13-25-12) was applied to the soil and supplied 76 kg N ha", 64 kg P ha", and 58 kg K ha". Ten plots each were sodded in Sept. 1995 with Supina bluegrass ‘Supra’ or Kentucky bluegrass ‘Blacksburg’. The sod had been grown in a composted wood mulch on polyethylene sheeting during the summer of 1995 (Cairol and Chevallier, 1981). Plots were mowed two to three times weekly to 3 cm height and irrigated as necessary to prevent moisture stress. Plots were fertilized bimonthly with 48 kg ha'l N, 3 kg ha'l P, and 40 kg ha" K. To prepare plots for testing in reduced light conditions (RLC), plots were fertilized with 48 kg N ha", 41 kg P ha", and 38 kg K ha" on 26 Aug. 1996. Plots were fertilized thereafter on a biweekly basis with 37 kg N ha", 3 kg P ha", and 30 kg K ha". 105 On 18 Sept. 1996 trinexapac-ethyl (0.19 kg ha") was applied to five plots each of Supina bluegrass and Kentucky bluegrass. Plots were moved into the Covered Stadium Simulator Facility (CSSF) on 4 October 1996 and arranged in a randomized complete block (RCB) design with five replications. Air temperature was maintained at 15.9 C i 2.9 C (range was 10-20 C). Relative humidity was 55.4 i 8.7%. High pressure sodium lamps (400 W), suspended 2.7 m above the turf surface, provided a steady but reduced light condition of approximately 100 i 9 umol m'2 s" on a 12 h photoperiod (ppd) and provided approximately 4.3 mol PAR m'2 day", not including ambient light (Table 62). Iprodione (3-(3,5-dichlorophenyl)-N-(1-methethyl)-2,4-dioxo- 1-imidazolidinecarboximide), 5.93 kg ha", was applied with a COz-powered backpack sprayer on 2 November 1996 to control Microdochium patch (Microdochium nivale). An open gas exchange system was used to determine photosynthetic rates (Sams and Flore, 1982) using a polycarbonate chamber (4.9 cm2, approximately 24 cm3) secured over the turf surface. Gas exchange measurements, foliar characteristics, and chlorophyll concentrations were determined 23-25 Nov. 1996 approximately seven weeks after the turf was moved into the CSSF. Carbon exchange rates (CERs) were collected on 23 Nov. between 1200-1600 h after C02 levels in ambient air had stabilized following large fluctuations earlier in the day. Data were analyzed as a 2 x 2 factorial in a RCB design with two species (Supina bluegrass and Kentucky bluegrass) and TE treatments (0.00 and 0.19 kg ha" trinexapac-ethyl) as main plots with five replications. 106 Table 62. Photosynthetically active radiation (PAR) of plots in the Covered Stadium Simulator Facility (CSSF), Hancock Turfgrass Research Center, East Lansing, MI. 1996 Location October November mol PAR day" Outside)r average 1 5.5 9.8 standard deviation 9.4 4.3 CSSF, ambient light 1 average 2.4 1.5 standard deviation 1.5 0.7 CSSF, supplemental light § average 5.5 5.1 standard deviation 1.1 0.7 1' PAR was integrated daily using a pyranometer (Li-Cor, model PY 14226, Lincoln NE). Radiometric units (Ly day'l ) were converted to quantum units (mol PAR rr1'2 day" ) based on the conversion units published by Thimijan and Heins (1983). I PAR inside the CSSF was determined by measuring the percent of PAR transmitted into the CSSF at turf level with a portable spectroradiometer (Li-Cor, model LI-1800, Lincoln NE). § Supplemental light (approximately 100 umol m'2 s"; 12 h photoperiod) was supplied with 400 W high pressure sodium lamps. 107 E . II' 1 1‘ E SE 1 Portable plots were established outside in full sun conditions. Wooden boxes (1.2 x 1.2 x 0.15 m depth) were filled with sand (Table 79, Appendix). Sixteen holes (0.6 cm diam) were drilled on approximately 23 cm spacings in the bottom of each box to provide drainage. Starter fertilizer (13-25-12) was raked into the upper 2 cm of the sand surface to provide 76 kg N ha", 64 kg P ha", and 58 kg K ha". Sixteen plots were sodded 29 August 1996 with Supina bluegrass ‘Supranova’ washed sod. Plots were irrigated as necessary to prevent moisture stress. Plots were mowed at 5 cm height at seven day intervals for the first 14 days, after which mowing height was gradually reduced to 3 cm height during the following 21 days. Thereafter, plots were mowed two to three times weekly to 3 cm height. Beginning 18 Sept. 1996 plots were fertilized with either a low N rate, 24 kg N ha" month", or a high N rate, 96 kg N ha" month" applied in split applications biweekly at 48 kg N ha". Potassium was applied biweekly at 48 kg K ha" to all plots. On 18 Sept. 1996 trinexapac-ethyl (0.19 kg ha") was applied to four plots each fertilized with low or high N rates. Plots were moved into the CSSF on 4 October 1996 and arranged in a randomized complete block (RCB) design with four replications. High pressure sodium lamps (400 W) were suspended 2.7 m above the turf surface and provided a steady but reduced light condition of approximately 100 i 9 pmol 1n‘2 5". The lamps were on a 12 h photoperiod (ppd) which totalled approximately 5 mol PAR rn'2 day", including ambient light (Table 62). Iprodione (3-(3,5-dichlorophenyl)-N-(1- methethyl)-2,4-dioxo-1-imidazolidinecarboximide), 5.93 kg ha", was applied with a 108 COz-powered backpack sprayer on 2 November 1996 to control Microdochium patch (Microdochium nivale). Carbon exchange rates were determined 15 and 16 Nov. 1996 (one and two days after mowing, respectively) using an open gas exchange system with a polycarbonate chamber (surface area = 27 cm2; volume approximately 200 cm3). The same location on each plot was assayed on both dates. Leaf areas from CER sampling areas were determined 16 Nov. 1996. Samples for chlorophyll analysis were collected 18 Nov. and analyzed 20 Nov. 1996. Carbon exchange rates were determined again on 26 Nov. (1 day after mowing) using a smaller polycarbonate chamber (surface area = 4.9 cm2; volume approximately 24 cm3) to determine the effects of a greater flow ratezsurface area on the CER. Leaf areas were determined from the sample areas the same day. Samples for chlorophyll analysis were collected 25 Nov. and analyzed 27 Nov. Photosynthetic measurements were collected between 0900-1200 h on all dates. Gas exchange and foliar data were analyzed as a 2 x 2 factorial with N rate (24 and 96 kg ha" month") and TE (0.00 and 0.19 kg ha") as main plots with four replications. Gas esehange measurements Carbon dioxide assimilation and related parameters were measured using an open system. The system was comprised of an ADC LCA2 infrared gas analyzer (IRGA), an air supply unit (ASU) capable of delivering up to 600 ml min" flow, a Parkinson leaf chamber (PLC) (Analytical Development Co. Ltd., Hoddesdon, England), and a polycarbonate assimilation chamber (PLC). Semi-flexible polyethylene tubing was used to connect the system components. Ambient air inside the CSSF was used as the air source and was drawn from a distance of at least 4 m away from the experimental site to 109 minimize C02 fluctuations due to the investigator. Air was drawn from approximately 0.3 m above the asphalt floor from a comer of the facility subject to little air movement which minimized C02 fluctuation. Air drawn from a height of 4 m inside the facility, or from outside the facility, had serious C02 fluctuations due apparently to fumace-emitted (heated) air. The COZ fluctuations prevented accurate measurements even when the air was passed through containers up to 250 L in attempts to dampen the C02 fluctuations. Ambient air (approximately C02 = 349 i 5 11L L" except on 26 Nov. when C02 = 427 i 1 1 11L L") was passed into the ASU which pumped at a flow rate of 500 ml min". The air was passed into a dome-shaped polycarbonate chamber (either 4.91 cm2 opening, volume approximately 24 cm3, or 27.3 cm2 opening, volume approximately 200 cm3) through in inlet port midway at or slightly below the turf surface. The chamber was secured over the turf surface using wire which was hooked over bolts at the chamber base and inserted into the turf. An exit port near the top of the chamber passed air into the IRGA for analysis of C02 concentration. Steady readings of C02 differential between the chamber and ambient air were achieved within one to two minutes. Immediately following gas exchange determination, a PLC was connected between the outlet port of the assimilation chamber and the IRGA for temperature and relative humidity measurements. The temperature and relative humidity of the ambient air were then measured. The photosynthetic photon flux density (PPFD) was determined during assimilation using a Li-Cor 19OS quantum sensor (LiCor, Lincoln, NE). Photosynthetic parameters were calculated on both a turf surface area and leaf area basis using a BASIC l 10 computer program (Moon and Flore, 1986). No attempt was made to inhibit the effects of soil respiration on CER. Leafareaanalxsis Leaf area of the turf was determined using a Li-Cor 300 leaf area meter (Li-Cor, Lincoln, NE). Leaf blades were excised from shoots and placed flat on sheet of clear contact paper. The contact paper was taped inside a folded piece of transparency paper; the sheets were then passed through the leaf area meter. The average of three readings were collected for each sample and the “blank” area of the contact paper plus tape was subtracted. Chlamphxfljnalxsis Sections (1 cm length) were collected from the middle of the youngest, fully expanded leaf blades of 10 plants per plot. Leaf widths were measured for leaf area determination. The mass of each 10 segment sample was determined to evaluate fresh leaf weight. Chlorophyll was extracted according to the methods of Moran and Porath (1980) using the extinction coefficients and formulae determined by Inskeep and Bloom (1985). Chlorophyll was extracted from each 10 leaf segment sample in 3 ml N,N- Dimethylformamide during incubation in the dark for 48 h at 4 °C. Absorbance values were measured using a UV-Vis spectrophotometer to determine chlorophyll a, chlorophyll b, and total chlorophyll concentrations. ll 1 RESULTS Ex 'm n Species significantly affected CER, E, and g5 on a turf surface basis while trinexapac- ethyl did not affect gas exchange parameters (Table 63). No interactions occurred between species and TE on any gas exchange parameters. Supina bluegrass CER on a turf surface area basis was over 50% greater than CER of Kentucky bluegrass and significantly different at p=0.05 (Table 64). Higher transpiration rate and stomatal conductance of Supina bluegrass corresponded with the greater CER compared to lower values observed from Kentucky bluegrass. Trinexapac-ethyl did not significantly enhance CER although CER was 36% greater in treated versus control plots. On a leaf area basis neither species or TE affected CER (Table 65). Values of gas exchange parameters were quite similar between the two species on a leaf area basis (Table 66). Species and TE both significantly affected LAI, fresh leaf weight, and chlorophyll levels (Table 67). There were no significant interactions between species and TE. Supina bluegrass turf had a greater LAI and lower fresh leaf weight but less chlorophyll compared to Kentucky bluegrass (Table 68). Trinexapac-ethyl resulted in greater LA] and increased chlorophyll levels in both species. Chlorophyll azb was not affected by any treatment. Experiment 11: Nitregen x PGR study Nitrogen and trinexapac-ethyl did not have a significant effect on CER or other gas exchange parameters of Supina bluegrass when evaluated on a turf area basis (Table 69). A higher than normal (approximately 350 11L L" COZ) ambient C02 level and decreased 112 .xocomoao own .833 MS? 60:90:28 £3088 . aw . N00 .5qu BEBE .5 ”8:826:00 .858on am ”coumufimgb .m ”88 owcwnoxo c880 JED + 66153365 85 65 a 38%me .. 8.8 3.; 3A: 3. G 3.5. 8.9. .x. .>o m2: at? ESQ: 3:: 83 ES ccem owed $2 82% 6%.: $0.0 9.2 m: x m $2 $3 $32 9.2M 33 Red Am: 35668965: 22 $2: 3552 52.3” as? .. _ m2 A5 66ch E: 83 32.82 macs em; 22 eccecsacm A To”: BE Now 35 A em as 655 A moo as Now .255 A em as 35.5 A _.m as .655 A we N.E N8 .655 856m a: a _ .MSB em 6 .w m 5.5 .003 con8o>oz mm £23 38 85.35 E A 18¢ add 38 m bofigxoaamv mcoEwcoo Em: 3268 E coESEmE 3.5th: 50 .Amhoonqu osofiimoaosa so Efiobamxosg 98 86on mo gumbo 5:253: 28 88:3 S82 .8 £th 113 .82 .acm 2 BE? 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Small and large chamber sizes (flow rate approximately 500 ml min") resulted in similar values when compared between the species x PGR and nitrogen x PGR studies. Carbon exchange rates on a turf area basis were lower at the high N rate compared to the low N rate although treatment effects were not significant at p=0.05 (Table 70). Carbon exchange rates 48 h after mowing were slightly greater compared to 24 h after mowing but there were still no significant differences among treatments. An interaction occurred on 15 Nov. 1996 between species and TE on E and g8 when gas exchange parameters were determined on a turf area basis. Twenty-four hours after mowing, E and gS of turf maintained at high N and treated with TB were significantly greater than untreated, high N turf or low N turf regardless of treatment (Table 71). This interaction was not significant on a leaf area basis and was not observed when the experiment was repeated on 26 Nov. 1996. On a leaf area basis nitrogen was the only treatment effect to produce any significant effects (Table 72). TE had no effect and there were no interactions. The high nitrogen rate increased CER, E, gs, and gm although the results were only significant for CER (p=0.10), E, and gS on one date 24 h after mowing and were not significant 48 h after mowing (Table 73). Both nitrogen rate and TE significantly affected LAI and chlorophyll content of Supina bluegrass (Table 74). 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Interaction of N rate and trinexapac-ethyl on transpiration (E) and stomatal conductance (gs) of Supina bluegrass maintained in reduced light conditions of approximately 5 mol PAR day". ambient co2 = 346 i 3 uL L" E gs trinexapac-ethyl (kg ha'I ) T N rate (kg ha'1 ) 0.00 0.19 0.00 0.19 24 0.22 0.18 7.62 6.15 96 0.13 0.27 4.92 9.57 LSD (0.05) 0.10 4.2 1 Applied 18 Sept. 1996. 124 RS ooo.o Koo $5.3 ooo.o ooo.o Sum ooo.o E _.o ooo.o 2 S; ooo.o too me x z “:2” 83 Show oooém ooo.o ooo.o Amt 3538985 sumo 52o om: ooo.ofl ooo.o ooo.o Azo Sweoz ooo.o ago 24.2 35:2 ooo.o ooo.o Snoozoom m5 7... we 6:3 2 a of a8 2225 L .3 : a 5. .woosoe 8% ; om .82 .32 om hogan—.9 gag—Emma «:8 ad 59. oo.ov ommo No.3 too oo. 8 so .>u :3 $2 Goo oo _ .23. ooo.o ooo.o Sum 33 ooo.o zoo ooo.o»? ooo.o ooo.o m: x z ooo.o ooo.o ooo.o ooo.o ooo.o oo.o Amt assoaoxoofi $3: 82 wow: 33%: Soo o_o.o 9o Emoaz ooo.o 2 2 oo: ooo.oomm ooo.o Goo 8:8:on m5 7... we as: 2 a of a8 Basso L .3 N a to. .mosse 5% o oo .32 .82 2 oo.om oo.o: oo.o... 3.2 a2 2.3 so .>o ow: ooo.o ooo.o £35 ooo.o foo Sum 32 ooo.o oo: $2989. ooo.o oo.o E x z ooo.o ooo.o ooo.o Smoom ooo.o ooo.o at 350-8385. ooo.o ooo.oo .53 84.8 :mooo Kmoo 9o 38:2 $3 5.2 ooo.o *ooito ooo.o .mooo oooogaom mi _.m we as: M: a o: .60 2285 L .3 m .+. so .waaoe 5% o om 32 5oz 2 3:82? 2523.33 «Eu MKN ACNE LoE NOD 3.5 :35 38.5 ALE: .255 A12: .063 A m E .255 A _.m N.E 3&3 830m M.oo x _ MS? em mw d _- N. m #50 Emma ooo.o won: A38 ~35 BE m boommeoaamv mcoEwcoo Em: 8968 E 88 somehow: EB ifioéamxoqtu 3 wooootw mmmcmoin ocmmsm mo moumtooogfio ouofiqmmopona mo 38b... 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I .... g A383 m 23 128 53838“ .282 353% So as ,mod .26 2: a 88%.me .1 .* cod $6 mos 36 3d mmém o\o .>U wood :06 526 3.0:». wmod vomd 8th 29¢ 35% mnvd mnmd mowd owed m; x Z mood * *mmo.R * * _ mwxm 3mm A .5“ *mvoé 30 A .N Am: 350-8355; Cod :Novdfl “1.036 ivomdo mid ibmfim AZV 5&0me wood *moAdm “AME: 33.2 0:; mg; :oswozmom em EU BE .5 9 :5 a 20 f5 we :5 850m A ~-Eo wlv $238220 is (32 amour,— 602 .>oZ : .A 19% Mag 38 m bofifimxoaamv mafia—Eco Em: 3038 E mmfiwoin «fiasm mo 2058:5800 :Emouoio cam ”£303 use. 23¢ .AFJV x09: moan awo— co 38am “58:85 mo oogomemm 98 38:3 :32 .3 2an 129 chlorophyll content. The high nitrogen rate significantly reduced LAI although chlorophyll content was increased (Table 75). Trinexapac-ethyl significantly increased LAI, fresh leaf weight, and chlorophyll concentration. The ratio of chlorophyll asb was not affected by any treatment. Although it was not a planned component of the study, N rate was observed to affect Microdochium patch (Microdochium nivale) (Table 76). Microdochium patch severely damaged turf maintained at high N rates while turf at low N rates sustained significantly less damage (Table 77). DISCUSSION Carbon exchange rates (approximately 1 umol C02 m'2 s", turf area basis) were comparable to results obtained using an open system to determine CER of Kentucky bluegrass during sod establishment in similarly reduced light conditions of 150 umol m"2 s'1 PAR (Kamok and Augustin, 1981). Kamok and Augustin (1981) reported increasing assimilation rates on a sward area basis with increasing days after mowing which corresponded to increased shoot height. Since fine turf is normally mowed frequently (e.g., one or two day intervals) the photosynthetic rate within one to two days following mowing was deemed important in the current study. Morgan and Brown (1983) concluded the optimal LAI of bermudagrass for photosynthesis was approximately 4.7 at 1600-2000 umol m'2 s'1 PAR while lesser LAls resulted in significantly lower CER. The optimal LAI for cool-season turfgrasses in reduced light conditions is unknown but the higher LAI of Supina bluegrass was 130 32 .aom M: 3%: reggae .232 Essoa So as ,2; .26 05 a :aéeam r. u or v0.“ $9.5m 100$ imodm *3: .I.m Home Nm.m Nm.mm Ks Ewm mo 0m oo.o A 72 mos Efioomqwxoctb com €523 1.3.x imndm 33 1:” 00 end Exam m2. No.3 3.0 m.m vm A 75:2: Tm: we: cowobmz as 20 33 Eu a Eu a :5 f5 we :5 access: a? L32 5on A $80 was =EmoHoEU .ooofi .>oZ C ,ATAg ~33 BE m £03meonva £82980 Em: 3262 E mmfiwofin «Emsm mo 2235:8200 233820 98 Jammy? awo— nmofi .AZd x09: 33 .32 so “Afioommmxocg 98 88 come“? we mgoobm .m5 2an 131 Table 76. Mean squares and treatment effects on Microdochium patch (Microdochium nivale) effects on Supina bluegrass in reduced light conditions (approximately 5 mol PAR day"), 18 Nov. 1996. Turfgrass Source of variation Color Density Quality Replication 0.099 421 .229 1.307 Nitrogen rate (N) 0.391 6123.063" 62.106" Trinexapac-ethyl (TE) 8.266” 742.563 8.266 N x TE 0.391 60.063 2.641 Error 0.488 175.229 5.307 CV, % 9.35 18.690 45.23 ** Significant at the 0.01 probability level. Table 77. Effects of nitrogen rate and trinexapac-ethyl on Microdochium patch (Microdochium nivale) damage to Supina bluegrass in reduced light conditions (approximately 5 mol PAR day'l), 18 Nov. 1996. Turfgrass Treatment Color;r Density: Quality§ Nitrogen rate (kg ha’l month)‘ 24 7.3 90.4 7.1 48 7,6 512” 3.1" Trinexapac-ethyl (kg ha'1 ) 0.00 6.8 64.0 4-4 0.19 H 8.2“ 77.6 5.8 ** Significant at the 0.01 probability level. 1‘ Color was rated visually on a one to nine scale, one = chlorotic, yellow turf, nine: dark green turf color with 5 a minimum rating for acceptable color. 1 Percent turf cover, visual estimate. § Quality was rated visually on a one to nine scale, one=100% necrotic turf, nine=dense, uniform, ideal turf with 5 a minimum rating for acceptable turf. 11 Nitrogen was applied as urea T'l‘ Applied 18 Sept. 1996. 132 apparently responsible for most or all of the difference in CER between the two species a sward area basis. There were no significant gas exchange differences between species on a leaf area basis. Supina bluegrass plants have a prostrate growth habit and stolons with short intemodes and numerous tillers which apparently provided a greater leaf area for photon capture and gas exchange compared to Kentucky bluegrass which exhibits an increasingly more vertical growth habit as PPFD declines (Wilkinson and Beard, 1973). The high N rate did not increase photosynthesis on a turf area basis because the amount of foliage was significantly decreased. High disease incidence associated with the high N rate may have caused a reduction in foliage although areas which appeared to be relatively unaffected by disease were chosen for CER measurements. The direct relationship between N rate and photosynthesis in non light-limiting situations appears to be largely dependent on the increased leaf biomass stimulated by higher N rates which affect carbon partitioning (Belanger et al., 1994; Gastal and Belanger, 1993; Nelson et al., 1993; Walker and Ward, 1973). In the current study, the high N rate may have stimulated excessive shoot growth early on after being placed in the reduced light conditions and depleted the carbohydrate pool necessary to sustain foliar growth and development. The high N rate may also have stimulated respiration which would have depleted the pool of nonstructural carbohydrates and resulted in reduced tillering. On a leaf area basis, the high N rate had a tendency to increase photosynthesis although this was significant only on one of the three dates. This result concurs with Walker and Ward (1973) who reported photosynthetic rates of centipedegrass [Eremochloa ophiuroides (Munro) Hack.] were directly dependent on N rate. The higher 133 N rate may have resulted in greater enzyme, particularly Rubisco, production (Ogata et al., 1983; Stitt and Schulze, 1994) and/or greater mesophyll conductance (gm) (Bolton and Brown, 1980). The lack of significant effect of TE on photosynthesis in RLC is not surprising. GA- inhibitors (paclobutrazol, flurprimidol) which act to inhibit ent—kaurene oxidation to ent- kaurenoic acid have been associated with both increases and decreases in photosynthetic rates in strawberries (Archbold and Houtz, 1988). Trinexapac-ethyl, however, inhibits the latter stages of GA biosynthesis, primarily by inhibiting hydroxylation at the 38 position of GAZOto produce a biologically active GAl (Rademacher, 1991). Several other differences exist between trinexapac-ethyl and other GA-inhibitors commonly used on turf which may influence their effects on plant physiology: 1) trinexapac-ethyl is foliar- absorbed (Vitolo et al., 1990), while paclobutrazol and flurprimidol are drenched into the ground for root uptake (Watschke et al., 1992), and 2) trinexapac—ethyl may be less phytotoxic than paclobutrazol and flurprimidol (Watschke and DiPaola, 1995). It is important to understand the mechanism(s) by which trinexapac-ethyl affects turfgrass growth and physiology in order to successfully use trinexapac-ethyl to maintain high quality turf in RLC. Green et a1. (1990) reported flurprimidol significantly reduced the ET rate of St. Augustinegrass for 5 weeks after application. Although the ET components were not split into the respective components of evaporation and transpiration, it was implied the reduced leaf extension rate was responsible for lowering the ET. Such data are important as decreased transpiration in RLC will further inhibit photosynthate production, an undesirable effect. 134 TE did significantly enhance leaf area and chlorophyll concentrations of both species and of Supina bluegrass across nitrogen rates but did not signficantly affect CER, even on a sward area basis. The effects of TE on photosynthesis may have been complicated by reduced senescence and increased LAI since increased leaf age and greater canopy development have been reported to reduce individual leaf photosynthetic rate (Morgan and Brown 1983). The improved turf quality associated with TE on turf in RLC may be related only to darker green leaf color and increased leaf area and/or tillering. Reduced leaf senescence rate and additional tillering could have been stimulated by TE side effects on other hormones or by TE altering carbohydrate levels and partitioning in the plants. GA- inhibitors have been shown to affect levels of other hormones such as abscisic acid in wheat (Buta and Spaulding, 1990) but their effects on hormones in turf is not known. Research on PGR effects on carbohydrate partitioning in turf is scarce. The key publication in the area, produced prior to the release of TE, indicates even GA-inhibitors with similar modes of action (paclobutrazol and flurprimidol) vary in their effect on assimilate partitioning (Hanson and Branham 1987). It is interesting to note that both paclobutrazol and flurprimidol did significantly decrease photoassimilate partitioning to roots four weeks after treatment (Hanson and Branham, 1987) although this may have been a transient response and not resulted in long-term effects. In the long term, reduced photoassimilate partitioning to roots could decrease turf quality and growth due to reduced root production. Studies on root growth of turf treated with flurprimidol or TE indicated these compounds had either no effect or had a beneficial effect on root growth 135 (Demoeden, 1984; Elam, 1993; McCarty etal., 1990). Studies on the effects of GA- inhibitors on photosynthate partitioning and hormone levels in turfgrass in RLC are warranted. Chlorophyll concentration did not affect photosynthetic rates (r2 = 0.07). Differences in chlorophyll concentration were often statistically significant at p=0.05 when analyzed between species, between N rates, and between TE and untreated plots, but were not great enough to result in different photosynthetic rates. The quantity of photosynthetically active radiation, not chlorophyll, limited the CER. Species and TE did have a significant role in turf color (Ch. 2), however, and for practical reasons species and TE must be considered when managing turf in reduced light conditions. Chlorophyll a:b ratios were typical of “sun” plants, approximately 3 (Nobel, 1991), and were not affected by any treatments. CONCLUSION The relative shade tolerance of Supina bluegrass compared to Kentucky bluegrass appeared to be related to a greater LAI and not to superior gas exchange properties (e. g. CER, transpiration, stomatal resistance). The high N rate did not sufficiently enhance Supina bluegrass photosynthetic rates to offset problems associated with the lower LAI compared to the low N rate or the problem of the increased Microdochium patch incidence (Microdochium nivale). Trinexapac-ethyl did not seem to affect gas exchange parameters of photosynthesis. It is likely TE improved turf quality in RLC by 136 mechanisms other than enhanced photosynthesis, possibly by altering photosynthate partitioning. APPENDIX 137 Table 78. Particle size analysis of sand used in sandzpeat mixture (80:20) . Description Very Gravel coarse Coarse Medium Fine Very fine Silt & clay diameter (m) > 2 2-1 1-0.5 0.5-0.25 0.25-0.l 0.1-0.05 < 0.05 Percent (%) 1.5 1.3 11.6 70.8 14.4 0.35 0.05 2.8 82.4 14.8 Table 79. Particle size analysis of sand used in Experiment 11: Nitrogen x PGR study (Chapter 3). Description Very Gravel coarse Coarse Medium Fine Very fine Silt & clay diameter (m) > 2 2-1 1-0.5 0.5-0.25 0.25-0.1 0.1-0.05 < 0.05 Percent (%) 0.9 6.7 32.0 40.0 16.2 1.6 0.5 7.6 72.0 18.3 LIST OF REFERENCES LIST OF REFERENCES Anonymous, 1995. The Kohlerdome. Panstadia International 2(2):70-71. Archbold, DD, and KL. Houtz. 1988. Photosynthetic characteristics of strawberry plants treated with paclobutrazol or flurprimidol. HortScience 23(1):200-202. Beard, J .B. 1973. Turfgrass: Science and culture. Prentice-Hall, Englewood Cliffs, NJ. Belanger, G., F. Gastal, and RR. Warembourg. 1994. Carbon balance of tall fescue (F esuca arundinacea Schreb.): Effects of nitrogen fertilization and the growing season. Annals of Bot. 74(6): 653-659. Bemer, P. 1980. Characteristics, breeding methods, and seed production of Poa supina Schrad. p. 409-412. In J .B. Beard (ed.) Proc. 3rd Int. Turfgrass Res. Conf., Munich, Germany. 11-13 July 1977. Int. Turfgrass Soc. and ASA, CSSA, and SSSA, Madison, WI. 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