Effects of Plant Density and Number of Shoots on Yield and Fruit Characteristics of Peppers Grown in Glasshouses

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Turk J Agric For 27 (3) 29-35 TÜB TAK Effects of Plnt Density n Numer of Shoots on Yiel n Fruit Chrcteristics of Peppers Grown in Glsshouses H. Y l z DAfiGAN*, Kz m ABAK Deprtment of Horticulturl Science, Fculty of Agriculture, University of Çukurov, 133 An - TURKEY Receive: 19.9.2 Astrct: Plnt ensity n pruning systems ply key role in the effective use of the re insie the greenhouse. Pepper (Cpsicum nnuum L.) cultivrs, Amzon-long green n Blo ell-shpe type, were grown in the winter cultivtion perio in glsshouse. A constnt spce of 8 cm etween rows with ifferent within row spcings (45 cm, 3 cm n 15 cm) n shoot numers (etween one n four shoots per plnt) were pplie to optimize plnt ensity n numer of shoots. Wier within row spcing n higher shoot numers per plnt increse the numer of leves. However, the iniviul lef le re ws higher for nrrower withinrow spcings with fewer shoots. While higher plnt ensities with greter numer of shoots reuce photosyntheticlly ctive rition, they increse the lef re inex t fruiting level. In orer to otin high yiels n ssumption of 8 x 15 cm with two shoots per plnt is suggeste for peppers. When expensive see is use then 8 x 3 cm spcing with three shoots per plnt might e more economicl. Plnt ensity n the numer of shoots i not ffect fruit qulity chrcteristics, such s fruit weight, length, imeter, volume, ry mtter, totl solule solis n the ph of the flesh in either cultivr. Key Wors: Cpsicum nnuum L., plnt spcing, pruning, yiel, fruit qulity Bitki Yo unlu u ve Sürgün Sy s n n Ser Bier Yetifltiricili ine Verim ve Meyve Özellikleri Üzerine Etkileri Özet: Ser içerisineki ln n etkin kulln lms konusun, itki yo unlu u ve um sistemi nhtr ir rol oynmkt r. Bier (Cpsicum nnuum L.) çeflitlerinen Amzon-(uzun yeflil meyveli) ve Blo (olml k meyveli) çeflitleri k fl önemine ser yetifltirilmifltir. Bitki yo unlu u ve sürgün sy s n optimize etmek için, s r rs mesfe tüm uygulmlr sit olrk 8 cm kulln l rken, frkl s r üzeri mesfeler (45 cm, 3 cm ve 15 cm) ile sürgün sy lr (1, 2, 3 ve 4 sürgün/itki) kulln lm flt r. Bitkilere s r üzeri mesfe rl kç sürgün sy s zlt lm flt r, zln sürgün sy s n yprk sy s zl rken her ypr n ln rtm flt r. Yüksek itki yo unlu u ve fzl sürgün sy s, çiçek ve meyve seviyesine PAR (Fotosentetiksel ktif fl n m) e erlerinin zlms n neen olurken yn itkilere LAI (Yprk ln ineksi) in rtm fl olu u tespit eilmifltir. Bu çl flm sonun, ser ierlerinen yüksek verim lilmek için 8 x 15 cm itki yo unlu u ve 2 sürgünlü um önerilmekteir. Anck, tohum mliyeti yüksek olu u urumlr tohumun ekonomik kulln m n s lmk için 8 x 3 cm ve 3 sürgünlü um önerileilir. Bitki yo unlu u ve sürgün sy s n n, ier meyvelerine rl k, oy, çp, hcim, meyve suyun SÇKM ve ph gii klite özelliklerini önemli üzeye etkilemei i tespit eilmifltir. Anhtr Sözcükler: Cpsicum nnuum L., itki s kl, um, verim, meyve klitesi Introuction The min pepper groups grown in the Meiterrnen sin re consist of ell-shpe, chrleston, conic n long-green fruit types of cultivrs. Growers o no pruning in open fiel pepper cultivtion. However, in greenhouse prouction, pruning n plnt ensity re very importnt for the optimiztion of plnt spcing per unit re. In greenhouse cultivtion of ell-shpe n long-green peppers, the effects of plnt ensity n pruning system re unknown n growers o not pply regulr pruning systems epening on plnt ensity. There re mny stuies on plnt ensity for ifferent types of pepper: ell or sweet pepper (Ahme, 1984; Stoffell n Bryn, 1988; Ceul, 1995; Jolliffe n Gye, 1995), cyenne pepper (Decoteu n Grhm, 1994), pepperoncini pepper (Motsenocker, 1996) n pprik pepper (Cvero et l., 1). Plnt ensity n plnt rrngement hve pronounce influence on plnt evelopment, growth n the mrketle yiel of mny vegetle crops (Stoffell n Bryn, 1988). A numer of stuies hve inicte liner increse in fruit yiel when plnt ensity is increse (Decoteu n Grhm, 1994; * Corresponence to: sgn@mil.cu.eu.tr 29

Effects of Plnt Density n Numer of Shoots on Yiel n Fruit Chrcteristics of Peppers Grown in Glsshouses Jolliffe n Gye, 1995; Motsenocker, 1996; Stoffell n Bryn, 1988). Porter n Etzel (1982) n Ahme (1984) reporte tht incresing plnt ensity resulte in greter yiel of ell pepper fruit. Mrketle ell pepper yiel increse with greter plnt ensity (Btl n Smittle, 1981; Everett n Surmny, 1983; Locscio n Stll, 1982). Sunstrom et l. (1984) reporte n increse in mechniclly hrveste tsco pepper (Cpsicum frutescens L.) yiel when within-row spcing ws ecrese. An increse in yiel with higher plnt ensity ws result of increse numers of fruit per hectre in irect-seee pprik pepper (Cvero et l., 1). Although most uthors hve reporte some informtion on the plnt ensity of ifferent types of peppers in open fiel prouction, stuies on greenhouse pepper prouction re limite. The optimiztion of plnt n shoot spcing in the greenhouse prouction of ell pepper ws investigte y Ceul (1995); plnts re usully trine to two shoots n the spcing of plnts is 8 x 3 cm. The present stuy ws crrie out to clrify the optimum plnt ensity n pruning system for ellshpe n long-green peppers grown in greenhouses. Mterils n Methos The experiment ws conucte t the greenhouse of the Horticulturl Deprtment of the Agriculturl Fculty in Çukurov University, An, Turkey uring the - 1 winter cultivtion perio. The Amzon F 1 cv (long-green type) hs 15- cm long fruit 1.5-2. cm in with n with intermeite pungency, n the Blo F 1 cv (ell-shpe type) hs fruit 7-8 cm long n 6-7 cm wie with sweet tste. Sees of the cultivrs, ought from Dutch compnies, Blo from Rijk Zwn n Amzon from Royl Sluis, were sown on August 7,. Seelings were plnte in the greenhouse t the 5-6 leves growth stge on Septemer,. The constnt istnce etween rows ws 8 cm. Within-row spcings use were 45 cm, 3 cm n 15 cm. Ech plot ws 4.5 m long n.8 m in with (3.6 m 2 ). The numers of plnts grown in the ech plot were 1, 15 n 3 epening on the within-row spcings (45, 3 n 15 cm, respectively). Two ifferent pruning methos (one n two shoots for 15 cm; two n three shoots for 3 cm n three n four shoots for 45 cm) were pplie on ech within-row spcing. Depening on the shoot numers (etween one n four shoots) the pruning methos pplie t the plnt evelopment stges were first, secon, thir n fourth emrnchment stges. During the cultivtion perio, every 1 or 15 ys, ll sie shoots were remove on the min shoots n fruiting occurre on trine min shoots only. The optimiztion of plnt spcing n shoot numers in greenhouse ell pepper prouction (Ceul 1995) were chosen s the min moels for moifying n pting ell-shpe n long-green peppers. The following tretments were rrnge in split plot esign with four replictions, The greenhouse ws hete t night to mintin minimum temperture of 16 ºC. Mximum tempertures (ytime) insie the greenhouse epene on the outsie ir tempertures n vrie from ºC to 34 ºC uring the cultivtion perio. One umle ee colony ws plce in the greenhouse for pollintion. A rip irrigtion system ws use for irrigtion n fertiliztion. During the experiment 68 kg N, 2 kg P 2 O 5 n 75 kg K 2 O were pplie per hectre. The numer of leves per plnt ws recore in the ifferent plnt evelopment stges: 35, 74 n 148 ys fter trnsplnting (DAT). The le re of the lef locte on the closest noe to the first rnching point ws mesure once, on 74 DAT. At the en of the experiment, the lef re of the plnts ws recore n the lef re inex (LAI) ws clculte, i.e. lef re ivie y spcing re. Mesurements of single-lef les n whole plnt lef res were me with 1 plnts from ech repliction (4 plnts for ech tretment) using igitl Hitchi t mrk lef re meter evice. The fruit hrvest strte on Novemer 11, (52 DAT) n continue for 215 ys with weekly hrvests, ening on June 13, 1 (267 DAT). The first 11 hrvests (from Novemer 11 to Ferury 28) were trete s the erly hrvest perio. On April 4, 1 (197 DAT), fruits from ech repliction were smple for to investigte fruit qulity chrcteristics such s fruit weight, length, imeter, volume, ry mtter, totl solule solis (TSS) n the ph of the flesh in oth cultivrs. Photosyntheticlly Active Rition (PAR) vlues were otine, using igitl quntum meter (BQM moel, Spectrum Technologies, Inc.), from three levels on 3

H. Y. DAfiGAN, K. ABAK Decemer 11, (82 DAT) n Ferury 2, 1 (135 DAT). The levels were ove the plnt level (), fruit setting level () n fruiting level (c). The percentge vlues otine from n c were clculte with respect to. Results The numer of leves ws increse y wier withinrow spcing n higher shoot numers in oth cultivrs (Figure 1). However, the iniviul lef le re ws generlly higher in nrrower within-row spcings (in higher plnt ensity) with fewer shoots (Figure 2). A higher plnt ensity with higher numer of shoots increse LAI (Tle 1). The increse in plnt ensity ws the importnt fctor in the more effective coverge of the soil surfce y leves. The cultivr Blo prouce higher numer of leves thn the cultivr Amzon, lthough the opposite pplies with regr to LAI (Tle 1). In oth cultivrs, PAR t the fruit setting n fruiting levels ws similrly ffecte y within-row spcing n 1 AMAZON 1 BALO 14 1 1 st mesurement 35 DAT 14 1 st mesurement 35 DAT 1 1 1 8 8 4 c c 4 c c 1 1 Numer of leves per plnt 14 1 1 8 4 2 n mesurement 74 DAT c c Numer of leves per plnt 14 1 1 8 4 2 n mesurement 74 DAT c c c Figure 1. 1 14 1 1 8 4 3 r mesurement 148 DAT c c Tretments 1 14 1 1 8 4 3 r mesurement 148 DAT e c Tretments Effects of spcing/numer of shoots n cultivrs on numer of leves per plnt t ifferent mesurement tes. The sme letters on the rs of the sme grph re not significntly ifferent ccoring to Duncn s Multiple Rnge test. DAT: Dys fter trnsplnting. Tretments: I. Spcing 8 x 15 cm (8.3 plnts/m 2 ), pruning to one shoot, 8.3 shoots/m 2, II. Spcing 8 x 15 cm (8.3 plnts/m 2 ), pruning to two shoots, 16.7 shoots/m 2, III: Spcing 8 x 3cm (4.2 plnts/m 2 ), pruning to two shoots, 8.3 shoots/ m 2, IV. Spcing 8 x 3 cm (4.2 plnts/m 2 ), pruning to three shoots, 12.5 shoots/m 2, V. Spcing 8 x 45 cm (2.3 plnts/m 2 ), pruning to three shoots, 8.3 shoots/m 2, VI. Spcing 8 x 45 cm (2.3 plnts/m 2 ), pruning to four shoots, 11.1 shoots/m 2 c 31

Effects of Plnt Density n Numer of Shoots on Yiel n Fruit Chrcteristics of Peppers Grown in Glsshouses AMAZON 18 1 14 1 1 8 4 Lef Ble Are (cm 2 ) I II III IV V VI Tretments 18 BALO 1 14 1 1 c c c 8 4 I II III IV V VI Tretments Figure 2. Effects of spcing/numer of shoots n cultivrs on lef le re mesure on 74 DAT. The sme letters on the rs of the sme grph re not significntly ifferent ccoring to Duncn s Multiple Rnge test. See the legen of Fig. 1 for efinition of the tretments numer of shoots. The plnts on the nrrowest withinrow spcing (15 cm) with two shoots h the lowest PAR vlues t fruiting level, ut the highest LAI (Tle 1). The results concerning yiels re given in Tle 2. Both cultivrs prouce similr erly n totl yiels. As within-row spcing ecrese n the numer of shoots increse, oth erly n totl yiels per m 2 increse. The fruit numer per plnt ws correlte positively with the within-row spcing (r =.93** Amzon, r =.92** Blo) n the numer of shoots per plnt (r =.98**for oth cultivrs). The highest totl yiel otine ws 11.15 kg/m 2 (men of oth cvs) from the highest plnt ensity (8 x 15 cm) with two shoots per plnt (Tle 2). The secon highest yiel, which ws from n 8 x 3 cm 32

H. Y. DAfiGAN, K. ABAK Tle 1. Tretments Effects of spcing/numer of shoots n cultivrs on the photosyntheticlly ctive rition (PAR) in percentges of rition mesure over the plnts n LAI. 11. 12. 2. 2. 1 (82 DAT*) (135 DAT) FSL** FL*** FSL FL LAI Cultivr Amzon 42 33 39 35 8. Blo 43 29 41 39 5.16 F vlue NS++ NS NS NS *** Spcing/numer of shoots I. 8 x 15cm/1 shoot 55 4 4 36 7.88 II. 8 x 15cm/2 shoots 41 19 41 31 8.35 III: 8 x 3cm/2 shoots 38 31 41 4 6.53 c IV. 8 x 3cm/3 shoots 4 33 38 37 6.95 c V. 8 x 45cm/3 shoots 41 32 44 39 4.81 e VI. 8 x 45cm/4 shoots 4 32 38 38 5.56 e F vlue NS NS NS NS *** Interctions Cultivrs x no. of shoots NS NS NS NS NS * DAT: Dys After Trnsplnting, ++NS: Nonsignificnt ** Fruit Setting Level *** Fruiting Level Men seprtion within colums one y Duncn s Multiple Rnge test. Vlues with the sme letter re not significntly ifferent. Tle 2. Effects of spcing/numer of shoots n cultivrs on totl n erly yiels of ellshpe n long-green types of peppers. Totl Totl Erly Erly Fruit Fruit Tretments yiel yiel yiel yiel no. per no. per (kg/m 2 ) (kg/plnt) (kg/m 2 ) (kg/plnt) plnt m 2 Cultivrs Amzon 8.15 1.88 3.39.78 74.2 3.24 Blo 8. 1.82 3..73 29.6 129.79 F vlue NS++ NS NS NS *** Spcing/numer of shoots I. 8 x 15cm/1 shoot 7.29 c.87 e 3.15 c.38 e 23.9 e 199. c II. 8 x 15cm/2 shoots 11.15 1.34 4.39.53 36.9 38.6 III: 8 x 3cm/2 shoots 7.19 c 1.73 c 2.97 c.71 c 48. c. c IV. 8 x 3cm/3 shoots 8.72 2.9 3.53.85 59.7 249.7 V. 8 x 45cm/3 shoots 6.42 c 2.31 2.59 c.93 64.1 174.1 VI. 8 x 45cm/4 shoots 7.7 c 2.77 3.14 c 1.13 78.7 218.7 c F vlue *** *** *** *** *** *** Interctions Cultivrs x no. of shoots NS NS NS NS *** *** ++ NS: Nonsignificnt * ** *** Significnt t P.5,.1 n.1, respectively. Men seprtion within colums one y Duncn s Multiple Rnge test. Vlues with the sme letter re not significntly ifferent. 33

Effects of Plnt Density n Numer of Shoots on Yiel n Fruit Chrcteristics of Peppers Grown in Glsshouses Tle 3. Effects of spcing/numer of shoots n cultivrs on some fruit qulity chrcteristics of peppers. Weight Length Dimeter Volume Dry Solule Tretments (g) (cm) (mm) (cm 3 ) mtter solis ph (%) (%) Cultivrs Amzon 25.5.8.1 31.1 6.9 5.1 5.7 Blo 61.1 7.5.8 11. 5. 4.8 5.7 F vlue *** *** *** *** *** NS NS Spcing/numer of shoots 8 x 15cm/1shoot 42.4 14. 41.2 71.7 6.4 5. 5.8 8 x 15cm/2 shoots 42.4 13.7 38. 71.5 6.3 4.9 5.7 8 x 3cm/2 shoots 55. 14.3 41.3 66.5 6.5 5. 5.7 8 x 3cm/3 shoots 44.4 14.8 41.2 73.4 6.4 4.9 5.7 8 x 45cm/3 shoots 44.4 13.7 41.3 69.8 6.4 5. 5.7 8 x 45cm/4 shoots 42.1 14.3 39.8 7.7 6.5 4.8 5.7 F vlue NS NS NS NS NS NS NS Interctions Cultivrs x no. of shootsns NS NS NS NS NS NS ++ NS: Nonsignificnt* ** *** Significnt t P.5,.1,.1 respectively. Men seprtion within colums one y Duncn s Multiple Rnge test. spcing n three shoots per plnt, ws 8.72 kg/m 2. The lrgest within-row spcing (8 x 45 cm) with three shoots prouce the lowest yiel (6.42 kg/m 2 ). The yiel per m 2 ecrese when wier within-row spcing n lower numer of shoots per plnt were teste (Tle 2). The cultivr Amzon prouce higher numer of fruits (74.2 fruits/plnt) thn the cultivr Blo (29.6 fruits/plnt) (Tle 2). However, the ifference etween the fruit numers per plnt i not ffect the yiel (oth cultivrs h similr erly n totl yiels) ecuse Amzon h lower fruit weight (25.5 g) thn Blo (61.1 g) (Tle 3). Within-row spcing n the numer of shoots i not ffect the fruit qulity chrcteristics. Depening on the tretment, s men of oth cultivrs, fruit weight vrie from 42.1 g to 55. g, fruit length from 13.7 cm to 14.8 cm, fruit imeter from 38. mm to 41.3 mm, volume from 66.5 to 71.7 cm 3, ry mtter from 6.3% to 6.5%, solule solis from % 4.8 to 5.% n flesh ph from 5.7 to 5.8. As expecte, significnt ifferences in some fruit chrcteristics etween the two cultivrs teste were oserve ue to the fruit types eing ellshpe n long-green (Tle 3). Discussion Decresing the numer of shoots reuce the numer of leves on the plnts. However, the surfce of single lef ws extene (Figure 2). Vnn et l. (1986) n Ceul (1995) oserve similr compenstory growth properties in sweet peppers. Reucing the shoot numer while incresing plnt ensity cuse more effective coverge of the soil y the lef cnopy (Tle 1). While higher plnt ensity with higher numer of shoots (8 x 15 cm, two shoots) reuce PAR t fruiting level, it increse LAI (Tle 1). Stoffell n Bryn (1988) n Ceul (1995) suggeste tht t higher plnt ensity, plnt efficiency woul e increse y higher LAI n high net ssimiltion rte in ell peppers. Motsenocker (1996) in pepperoncini peppers n Jolliffe n Gye (1995) in ell peppers reporte tht lef re per plnt ecrese while LAI increse s the plnt ensity of the peppers increse. At higher plnt ensities of the ell pepper, greter rition interception ue to higher LAI in resulte higher iomss n fruit yiel (Ceul, 1995; Lorenzo n Cstill 1995). As plnt ensity increse, erly n totl yiel per m 2 in the oth cultivrs increse while erly n totl yiel per plnt ecrese. This is proly ecuse less spcing crete higher interplnt competition. Similr finings 34

H. Y. DAfiGAN, K. ABAK hve een reporte from open fiel experiments for ell peppers (Stoffell n Bryn, 1988; Btl n Smittle, 1981; Everelt n Surmny, 1983; Jolliffe n Gye, 1995) cyenne peppers (Decoteu n Grhm, 1994) pepperoncini peppers (Motsenocker, 1996) n pprik pepper (Khn et l., 1997; Cvero et l., 1). Ceul (1995) reporte tht the gretest yiel of sweet peppers (8.37 kg/m 2 ) ws otine from single-shoot plnts grown t highensity, 8 x 15.6 cm, in greenhouse experiment. In our greenhouse experiment, with ellshpe (Blo) n long-green (Amzon) peppers, the gretest yiel per m 2 ws otine from the highest plnt ensity, 8 x 15 cm, with two shoots prune tretment (11.53 kg/m 2 from Blo n 1.76 kg/m 2 from Amzon). Although Ceul (1995) reporte tht the limittion of shoot numers ffecte fruit weight (the fewer shoots, the hevier fruits), in our experiment fruit size prmeters such s weight, length n imeter were not significntly ffecte y within-row spcing or the numer of shoots (Tle 3). This coul e ue to the fct tht fruit ws hrveste t pproximtely the sme stge of evelopment throughout the experiment (Stoffell n Bryn, 1988; Jolliffe n Gye, 1995). In conclusion, within-row spcing n the numer of shoots ffect the prouctivity of pepper plnts. The prouctivity of the plnt t high plnt ensity cn vry epening on the numer of shoots per plnt. In orer to otin high yiel from the unit re in the greenhouse, we suggest proucing these cultivrs with 8 x 15cm spcing n two shoots per plnt (8.33 plnt/m 2 n 16.66 shoots/m 2 ). If the see cost is high n economicl use of the see is consiere, secon suggestion 8 x 3 cm with three shoots (4.17 plnts/m 2 n 12.51 shoots/m 2 ) might e cceptle. Plnt ensity n numer of shoots o not ffect the fruit chrcteristics when the fruit is hrveste t the sme stge of evelopment. Acknowlegements The uthors wishes to thnk Prof. S. Ceul for vlule iscussions on visit to Turkey. The uthors lso re grteful to MSc stuents Bulut Ekici n Siel Koç for their consierle ssistnce in the greenhouse work, n to Dr. Set Büyüklc for her help with the English text. References Ahme, M.K., 1984. Optimum plnt spcing n nitrogen fertiliztion of sweet pepper in the Sun Gezir. Act Hortic. 143: 35-31. Btl, K.M. n D.A. Smittle. 1981. Reponses of ell pepper to irrigtion, nitrogen n plnt popultion. J. Amer. Soc. Hort. Sci. 16: 259-262. Cvero, J., R. Orteg Gil n M. Gutierrez. 1. Plnt ensity ffects yiel, yiel components n color of irect-seee pprik pepper. HortScience 36(1): 76-79. Ceul, S., 1995. Optimiztion of plnt n shoot spcing in glsshouse prouction of sweet pepper. Act Horticulture 412: 321-329. Decoteu, D.R. n H.A.H. Grhm. 1994. Plnt sptil rrngement ffects growth, yiel n po istriution of cyenne peppers. HortScience 29: 149-151. Everett, P.H. n R. Surmny. 1983. Pepper prouction s influence y plnt spcing n nitrogen-potssium rtes. Proc. Fl. Stte Hort. Soc. 96: 79-82. Jolliffe, P.A. n M.M. Gye. 1995. Dynmics of growth n yiel component responses of ell peppers (Cpsicum nnuum L.) to row covers n popultion ensity. Scienti Hortic. 62: 153-164. Khn, B.A., J.R. Cooksey n J.E. Motes. 1997. Within-row spcing effects on trits of importnce to mechnicl hrvest in ppriktype peppers. Scienti Hortic. 69: 31-39. Locscio, S.J. n W.M. Stll. 1982. Plnt rrngement for increse ell pepper yiel. Proc. Fl. Stte Hort. Soc. 95: 333-335. Lorenzo, P. n N. Cstill. 1995. Bell pepper yiel response to plnt ensity n rition in unhete plstic glsshouse. Act Hortic. 412: 33-334. Motsenocker, C.E., 1996. In-row plnt spcing ffects growth n yiel of pepperoncini pepper. HortScience 31: 198-. Porter, W.C. n W.W. Etzel. 1982. Effects of luminum-pinte n lck polyethylene mulches on ell pepper, Cpsicum nnuum L. HortScience 17: 942-943. Stoffell, P.J. n H.H. Bryn. 1988. Plnt popultion influences growth n yiels of ell pepper. J. Amer. Soc. Hort. Sci. 113: 835-839. Sunstrom, F.J., C.H. Thoms, R.L. Ewrs n G.R. Bskins 1984. Influence of N n plnt spcing on mechniclly hrveste Tsco peppers. J. Amer. Soc. Hort. Sci. 19: 642-645. Vnn, D.R., J.S. Fletcher, N.R. Acchirey n L. Beevers. 1986. Influence of prtil efolition of green pepper on the senescence, growth, n nitrte reuctse of remining lef. Plnt n Soil 91: 357-361. 35