Chapter 2. Effects of Exogenous Fructose and Adjuvant Applications on Leaf Injury of. Supina Bluegrass under Reduced Light Conditions.
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1 - Chapter 2 Effects f Exgenus Fructse and Adjuvant Applicatins n Leaf Injury f Supina Bluegrass under Reduced Light Cnditins. ABSTRACT Turfgrass management in reduced light cnditins «30 % full sunlight) is difficult because turf grwth is affected by a lack f sufficient light energy. In turn, plant carbhydrates are reduced, due t insufficient light inhibiting phtsynthesis. T cmbat this prblem, a simple study was initiated t investigate the ptentials fr exgenus fructse applicatins t the leaves f turfgrass grwing under reduced cnditins. Thus, the bjective f this study was t determine whether a physilgical respnse ccurs with exgenus fructse applicatins t turfgrass grwn under reduced light cnditins. Fructse applied t supina bluegrass under reduced light cnditins five times per week fr eight cnsecutive weeks causes t much leaf injury unless the applicatin rate is 1.25% weight per vlume fructse at the 815-L ha' rate with 0.1 % weight per vlume Break Thru adjuvant. Weekly fructse applicatins (ne time per week) yielded acceptable levels f leaf tissue injury fr the 1,2,4,6, and 8 times applicatin rates. The 1x rate prvided the least amunt f leaf tissue injury while demnstrating psitive physilgical respnses cmpared t the cntrl. 24
2 INTRODUCTION Turfgrass is used fr athletics, recreatin, aesthetics, and utility. Each f these situatins requires turf t perfrm a different rle in a different setting. As a result f this dynamic utilizatin, turfgrass is ften managed under sub-ptimal grwing cnditins. One such cnditin is shade (reduced light cnditins; RLC). Turfgrass lcated in RLC makes up abut 20-25% f all managed turfgrass (Beard, 1973). Turfgrass management in RLC «30 % full sunlight) is difficult because turf grwth is affected by a lack f sufficient light energy (Stier and Rgers, 2001). Light intensity can be reduced by as much as 95% frm surrunding plant material, cluds, r ther shading structures (Beard, 1973). As a result, the light, quality that reaches the turf surface is altered, creating an envirnment nncnducive t nrmal plant grwth and develpment (Wilsn, 1997). Plants need light energy frm the sun fr phtsynthesis t prvide energy fr grwth, develpment, and reprductin (Taiz and Zeiger, 1991). EnergYlight+ 24H C02 > C12H H20 Phtsynthesis is driven by the visible prtin f the light spectrum (400 t 700 nm), called phtsynthetically active radiatin (PAR; Lawlr, 1987), and cmprises up t 50% f the earth's direct radiatin (Salisbury and Rss, 1992). When plant carbhydrates are limiting, ften due t insufficient light frm inhibited phtsynthesis, exgenus applicatins f sugars have shwn ptential t becme a management technique fr turf under RLC (unpublished data). 25
3 Early research has fcused n the ptential fr exgenus sugar applicatins t be taken up by a myriad f plant species. Exgenus sucrse applicatins (10 % in slutin) were fund t cause an increase in the dry weight f tmat plants (Lycpersicum esculentum Mill., Went, 1944; Went and Carter, 1948). In additin, Berrie (1960) determined a greater increase in tmat dry weight with exgenus sucrse applicatins (10 % in slutin) particularly in cnditins where carbhydrate synthesis is limited, and especially when respiratin is prceeding rapidly and phtsynthesis slwly. Other exgenus sugar absrptin research has lked at plant uptake using leaf disks. Weatherley (1954) attempted t determine that sucrse absrptin was an active transprt when the water was absrbed thrugh passive absrptin using Atrpa belladnna leaf disks. Hwever, under aerbic and anaerbic cnditins, evidence fr the active uptake f sucrse is far frm cnclusive. Using carrt leaf disks, Grant and Beevers (1963) determined the ptimal absrptin times and cnditins fr several sugars. The greatest time fr sugar absrptin fr carrt leaf disks was when the disks were respiring. Glucse uptake als increased ten fld when temperatures increased frm 3-25 "C. Finally, withhlding xygen depressed the uptake f glucse, fructse, galactse and xylse (Grant and Beevers, 1963). Mixed results have been fund when exgenus applicatins f sucrse have been applied t plants as a spray mixed with urea (46-0-0). Eatn and Ergle (1952) fund that spraying the upper leaf surface f cttn plants (Gssypium hirsutum var. Missdel x Acala) daily thrugh the fruiting perid with 26
4 20% sucrse, 1% nitrgen (urea) and the tw in cmbinatin failed t imprve plant grwth and resulted in significant decreases in the number f blls that were set. In a related study, Alvim (1960) determined that 10 % sucrse and gibberellic acid were effective in prtecting against injury by 20/0 urea spray n kidney bean plants (Phase/us vu/garus). In additin, 10 % sucrse sprays prevented a reductin in rt dry weight as a result f gibberellic applicatins (Alvim, 1957). In his research n clver (Triflium repens L.), Van Schreven (1959) fund that cncentratins f 0.5 and 1% glucse stimulated ndule frmatin, bth in the absence and in the presence f additinal light. ndules were frmed in the presence f 2% glucse. The highest numbers f Van Schreven als fund that in the absence f additinal light, ndulatin was stimulated by 0.5, 1, and 20/0 sucrse and in the presence f additinal light, by 0.5 and 1 10 sucrse. In bth cases, 0.5% sucrse effected the greatest stimulatin. Exgenus sugar applicatins fr plants in light and dark cnditins have als been studied using sybean (G/ycine max) seedlings. In the light, translcatin f sucrse-c" r glucse-c 14 ut f the treated leaf was very slw. Only 1% f the C 14 frm sucrse was translcated after 14 hurs. In the dark, 100/0 f the C 14 frm glucse was translcated t all parts f the seedling in 3 hurs. The bulk f the translcated C 14 accumulated in the stem between the treated leaf and the rt (Nelsn and Grham, 1957). Althugh, this research was cnducted n a legume, the findings between the effects f sugar 27
5 p:s applicatins in the light versus dark supprt reasn fr studying the effects f sugar applicatins t turfgrass under reduced light cnditins. While mst f the exgenus sugar applicatin research has fcused n nn-grass (Pacea) crps, sme research has lked at the grass family. Exgenus applicatins t crn (Zea maize) rts cncluded that entering glucse and fructse mixed readily with the endgenus pls (Grant and Beevers, 1963). Sucrse uptake als appears t depend n extracellular hydrlysis in yung crn rts (Giaquinta, et al., 1983; Lin, et al., 1984; Singh and Maclachlan, 1986). Unfrtunately, because turfgrass grws in a cntiguus cmmunity, the practicality f the applied sugars reaching the rts befre interceptin frm shts, thatch, sil, insects and micrbes is nt likely. Cmpsitins incrprating a pstemergence herbicide and a sugar, particularly fructse, as a ptentiatr f the herbicide against weeds withut decreasing the tlerance f a crp plant t the herbicide has becme a new methd fr killing weeds. Fructse in slutin can be used fr plant uptake at a range f % weight per vlume, but 1.25% is bestjpenner and Rggenbuck, 1999). Althugh this research is fcusing n the use f exgenus fructse applicatins t increase the efficacy f herbicides, subsequent findings have determined that the leaves f bth bradleaf and grassy weeds readily take up exgenus fructse applicatins. readily absrbed an rgansilicne Hwever, in rder fr the fructse t be is necessary as an adjuvant (Penner and Rggenbuck, 1999). 28
6 Based n the afrementined findings with exgenus sugar applicatins t plants, an investigatin was warranted t determine if exgenus sugar applicatins can cunter the effects f reduced light cnditins (shade) fr turfgrass situatins. In this experiment ur bjective was t determine whether a physilgical respnse ccurs with exgenus fructse applicatins t turfgrass grwn under reduced light cnditins. Fructse applicatin frequencies and rates were investigated t determine the psitive and/r negative physilgical respnses when applied t turfgrass under reduced light cnditins. 29
7 MATERIALS AND METHODS On 20 September 1999 prtable plts were filled with a sand rt zne and sdded with ne year ld supina bluegrass sd (Pa supina Schrad.; Manderley Sd Inc., Napean, ON, Canada) using six 1.2 x 1.2 m high-density plyethylene ITM mdules (GreenTech, Inc., Richmnd, VA, USA) at the Hancck Turfgrass Research Center (HTRC) at Michigan State University, East Lansing, MI, USA. Beginning in May January 2001, the plts were fertilized tw times per mnth at 0.5-g N m- 2 applicatin" using Lebann Cuntry Club fertilizer (Lebann Seabard Crp., Lebann, PA, USA). On 15 September 2000 the established ITM mdules were mved inside the indr research facility at the HTRC. The indr research facility is a 400 m 2 air-supprted structure cnstructed f Ultralux" (Chemical Fabrics Crpratin, Buffal, NY, USA), a fiberglass fabric which transmits apprximately 20% +/- 2% phtsynthetically active radiatin (Figure 5). Tw studies investigating the effects f exgenus fructse (ISOCLEAR 55, Cargill, Inc., Daytn, OH, USA) applicatins t supina bluegrass under reduced light cnditins were cnducted n 4 December 2000 thrugh 26 January 2001 and 4 December 2000 thrugh 12 January 2001 fr studies ne and tw, respectively. The experimental design fr these studies was a cmpletely randmized blck design with three replicatins. Statistical analysis was dne using Agriculture Research Manager, versin 6.18 (Gylling Data Management, Inc. Brkings, SO, USA). 30
8 paz (1) "en "5 c 0 I I L- "'0 C I ur w w w w iu w ur co t-- CD u-) C'0 N (_SZ_W [um) l!suap xnu UOl04d ::>!Ja4Jusl4d + W (l)..c.... (l) "'0 -Ci5 C......c _Q>.... c (l) :.c EO r "'0 (l) C C.... r..., ::J "5,m := C ::J..c (/')0 '+-('1').... -Ci5 c (l)n "'0 c x CO ::J i;:: - c..c 0- Q.(/') (,)C r t) c r w (/') c 0(3 a.. $ l,(j..c (l) L- L- r ::J (l) O>(/') tr 31
9 Study ne investigated the effects f fructse at 1.25% weight per vlume and tw different adjuvants (Break Thru - rgansilicne, Aquatrls Inc. Cherry Hill, NJ, USA; Apsa 80 - Alkylaryl alkxylate, Amway Crp., Ada, MI, USA) at bth 0.1 and 0.25% weight per vlume applied five times per week fr eight weeks. Study tw cmpared the afrementined treatments using weekly applicatins fr six weeks. Fur stck slutins cntaining the different adjuvant treatments (Break Thru and Apsa 80 at bth 0.1 and 0.25% weight per vlume) were made with reverse smsis water and 1.25% weight per vlume fructse. The slutins were applied at 1, 2, 4, 6 and 8 times the rate f 815 L ha'. Treatments were applied t the leaves as a spray using a 500-ml hand trigger plastic spray bttle. Bth f the studies used individual ITM mdules as a replicatin. Plt sizes fr treatments were 225 crrr', Data cllectin cnsisted f visually assessing physilgical cmparisns f injury and txicity effects. Turfgrass leaf injury was rated was rated using a 1-9 scale, with 1 being all green with n leaf damage, and 9 being all brwn r dead leaf tissue. 32
10 RESULTS AND DISCUSSION Study ne - Five times per week applicatins Significant differences ccurred thrughut the eight weeks f data cllectin fr bth treatments (adjuvants and fructse rate; Table 2). The effects f the tw adjuvants (Apsa 80 and Break Thru ) at applicatin rates f 0.1 and 0.25% weight per vlume shwed significant differences fr injury n turfgrass leaf tissue when applied five times per week fr eight weeks regardless f the rate f fructse applied (Figure 6). All treatments yielded unacceptable turfgrass leaf tissue quality with the exceptin f the 0.1 % weight per vlume rate using the adjuvant Break Thru. In additin, different applicatin rates (1,2,4,6, and 8 times 815-L ha") f fructse at 1.25% weight per vlume, regardless f the adjuvant treatment, shwed significant differences fr injury t turfgrass leaf tissue when applied five times perweek (Figure 7). Only the 1x rate f fructse (815-L ha') shwed very little injury thrughut the duratin f the study, suggesting that a rate f 1.25% weight per vlume fructse applied at 815-L ha' is the limit fr preventing unacceptable levels f leaf tissue injury (Figure 7). 33
11 Table 2. Study ne - analysis f variance fr leaf tissue injury frm exgenus fructse t applicatins with an adjuvanf applied five times per week n supina bluegrass under reduced light cnditins. E. Lansing, MI December January Mean Square Errr SOURCE DF 8 Dec. 15 Dec. 30 Dec. 7 Jan. 12 Jan. 28 Jan. Ttal 59 R Adjuvant (A) * 7.26* 6.95* 21.29* * 71.48* Fructse rate (B) * * 9.98* 17.10* * 35.68* AB * ERROR t The tw adjuvants tested were Break Thru - rgansilicne, Aquatrls Inc. Cherry Hill, NJ, USA; Apsa 80 - Alkylaryl alkxylate, Amway Crp., Ada, MI, USA) at bth 0.1 and 0.25% weight per vlume :t: Fructse applicatin rates were 1, 2, 4, 6, and 8 times 815 L ha' using 1.25% weight per vlume fructse - ISOCLEAR 55, Cargill, Inc., Daytn, OH, USA. 34
12 M It) ci O (j) -J It) ci O (j) -J CD It) ci (j) -J CD L.O (6-) A.infu!:leal ssej6jjnl N N.:::t:. '--. en 0) E C :,j:j en C t+=...j > r.... C.... r ""C en > r ::J==W...,0. ""C. r r en.... C C C 0 :,j:j '-.c.- :>""C :> C ""C '+-c.... O.c 2...Ci5 ""C 0 W O)""C CD '- '- I '- ::J ::J._ O)'-""C.c C u..... ::J 35
13 >< >< >< >< >< T"""NCOcx) t + f - + M a l!) c:i C/) -...l l!) c:i C/) -...l l!) c:i C/) -...l c s T"""..:::s:. L.. J2 L.. +J 0. C (J) E u:.;::i ::.= 0.> 0..- c'- (J) l!) c:i C/) -...l.... 'c..c...j L{) T""" 00..c +J (J) (1) E :.;::i C :e "0 C +J..c.2> f"- a l!) c:i C/) -...l f"- c M (6-) Jtlnfu! jeal ssej6j.jnl N 36
14 Study tw - One time per week applicatins After six weeks f testing, significant differences in turfgrass leaf tissue injury ccurred between weekly applicatins with the tw adjuvants (Table 3). Regardless f the adjuvant, the rate f 0.1 % weight per vlume adjuvant in slutin caused significantly less tissue injury than the rate f 0.25% weight per vlume in slutin (Figure 8). Significant differences in turfgrass leaf tissue injury was bserved with different applicatin rates f fructse (Table 3). The 1x rate f fructse (815-L ha') shwed very little injury thrughut the duratin f the study, suggesting that a rate f 1.25% weight per vlume fructse applied at 815-L ha' is ideal fr preventing leaf tissue injury (Figure 9). In additin, 1.25% weight per vlume fructse applied at 2, 4, 6, and 8 times the 815-L ha' yielded acceptable turfgrass leaf injury levels. These results suggest that weekly (ne time per week) exgenus fructse applicatins are mre practical than daily (five times per week) applicatins fr turfgrass under reduced light cnditins. Fr supina bluegrass grwing under reduced light cnditins the rate f 1.25% weight per vlume fructse applied at 1, 2, 4, 6, and 8 times 815-L ha' is acceptable in terms f injury and triggers psitive physilgical changes t ccur cmpared t the cntrl plants where n fructse applicatins were applied (Figure 9). These bservatinal cmparisns suggest that the plant is readily absrbing exgenus fructse applicatins and is using it fr metablic prcesses. 37
15 Table 3. Study tw - analysis f variance fr leaf tissue injury frm exgenus fructse t applicatins with an adjuvant! applied ne time per week n supina bluegrass under reduced light cnditins. E. Lansing, MI December January Mean Square Errr SOURCE OF 8 Dec. 15 Dec. 30 Dec. 7 Jan. 12 Jan. Ttal 59 R Adjuvant (A) * * 2.55* Fructse rate (B) * * 1.43* 2.60* AB ERROR t The tw adjuvants tested were Break Thru - rgansilicne, Aquatrls Inc. Cherry Hill, NJ, USA; Apsa 80 - Alkylaryl alkxylate, Amway Crp., Ada, MI, USA) at bth 0.1 and 0.25% weight per vlume + Fructse applicatin rates were 1, 2, 4, 6, and 8 times 815 L ha' using 1.25% weight per vlume fructse - ISOCLEAR 55, Cargill, Inc., Daytn, OH, USA. 38
16 Ii) (f)...j I CX) E:J r en>o 0 I «ID O '-" Q.N Q. 0 en c C >.c r.- E >. "O r 0 0 CX)..... Ii) (f)...j <0 N.:: S Ii) (f)...j N CX)... :J 0> :J...._..c U-1-39
17 I"-- Ii) c:i (j) L{) Ii) c:i c:r (j) L{) Ii) c:i c:r (j) L{) Ii) c:i c:r (j) L{) s L- 0) 0)-0 CJ) E 0) -- () () 0) 20 '+- 0) c E >c Ci5 2 a.-.r: _ 0')0') "(5.s 3: CJ) c r u.,-j N(j) r L-W O) 0) r 0) L- 3: c :.;::i r a. a. (1) r E i= -J L() c 0).r: -- L- 0) a. 0) c CJ) c :E "'0 C Ii) c:i (j) s c I"-- L{) M (6- )AJnrU! :Jeal ssej6jjnl N 40
18 Figure 10. Digital illustratin demnstrating physilgical differences between tw supina bluegrass plants grwn in reduced light cnditins. The plant n the left has received weekly exgenus fructse applicatins fr six weeks and the plant n the right has nt received any fructse, E. Lansing, MI 12 Jan
19 CONCLUSIONS Fructse applied t supina bluegrass under reduced light cnditins five times per week fr eight cnsecutive weeks causes t much leaf injury unless the applicatin rate is 1.250/0 weight per vlume fructse at the 815-L ha' rate with 0.1 % weight per vlume Break Thru adjuvant. Weekly fructse applicatins (ne time per week) yielded acceptable levels f leaf tissue injury fr the 1,2,4,6, and 8 times applicatin rates. The 1x rate prvided the least amunt f leaf tissue injury while demnstrating psitive physilgical respnses cmpared t the cntrl. Overall, this study was successful as a preliminary study t warrant mre in-depth research fr understanding the uptake and translcatin f exgenus fructse applicatins t turfgrass under reduced light cnditins. 42
20 REFERENCES Alvim, P Net assimilatin rate and grwth behavir f beans as affected by gibberellic acid urea and sugar sprays. Plant Physi/. 35: Beard, J.B Tutigrass: Science and Culture. Englewd Cliffs, New Jersey: Prentice Hal/. Berrie, A.M The Effect f Sucrse Sprays n the Grwth f Tmat. Plant Physilgia Plantarum 13: Eatn, F.M. and D.R. Ergle Relatinship f seasnal trends in carbhydrate and nitrgen levels and effects f girdling and spraying with sucrse and urea t the nutritinal interpretatin f bll shedding in cttn. Plant Physi/. 27: Giaquinta, R.T., W. Lin, N.L. Sadler, and V.R. Franceschi Pathway f phlem unlading f sucrse in crn rts. Plant Physi/. 72: Grant, B.R. and H. Beevers Absrptin f sugars by plant tissues. Plant Physi/. 38: Lawlr, D.W Phtsynthesis: metablism, cntrl and physilgy. Jhn Wiley & Sns, Inc., New Yrk, NY. Lin, W., M.R. Schmitt, W.O. Hitz, and R.T. Giaquinta Sugar transprt in islated crn rt prtplasts. Plant Physi/. 76: Nelsn, C.D. and P.R. Grham Uptake and translcatin f C 14 -labelled sugars applied t primary leaves f sybean seedlings. Can. J. Bt. 35: Penner, D. and F.C. Rggenbuck Cmpsitins cntaining herbicide and mnsaccharides and methd f use theref. U.S. Patent Date issued: 31 August. Salisbury, F.B., and C.W. Rss Plant Physilgy, furth editin. Belmnt, Califrnia: Wadswrth Publishing Cmpany. 43
21 - Singh, R. and G. Maclachlan Differential rates f uptake f glucse, fructse and sucrse by pea stem prtplasts. J. Exp. Bt. 37(181): Stier, J.C. and J.N. Rgers, I Trinexapac-Ethyl and irn effects n supina and Kentucky bluegrasses under lw irradiance. Crp Sci. 41 : Taiz, L., and E. Zeiger Plant Physil. Redwd City, Califrnia: The Benjamin/Cummings Publishign Cmpany, Inc. Van Schreven, D.A Effects f added sugars and nitrgen n ndulatin f legumes. Plant and Sil. XI(2): Weatherley, P.E On the uptake f sucrse and water by flating leaf disks under aerbic and anaerbic cnditins. New Phytl. 55: Went, F.W Plant grwth under cntrlled cnditins I. Crrelatin between varius physilgical prcesses and grwth in the tmat plant. Amer. J. Bt. 31: 597. Went, F.W. and M. Carter Grwth respnses f tmat t applied sucrse. Amer. J. Bt. 35: Wilsn, J.R Adaptive respnses f grasses t shade: Relevance t turfgrass fr lw light envirnments. Int. Tutfgrass Sc. Res. J. 8:
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