Partial D e f o l i a t i o n of Vitis vinifera L. cv.

C a b e r n e t Sauvignon/99 Richter: Effect on Root Growth, C a n o p y Efficiency; Grape C o m p o s i t i o n , and Wine Quality
J. J. H U N T E R ~*, H. P. R U F F N E R 2, C. G. V O L S C H E N K 3, a n d D. J. LE R O U X 4
vitis vinifera L. cv. Cabernet Sauvignon/99 Richter was grown under field conditions. The effect of partial defoliation (33%) in the lower half of the canopy at berry set stage, and thereafter at pea-size and veraison, respectively, on root development, distribution, and composition as well as on canopy efficiency, yield, grape composition, and wine quality was investigated. Defoliation evidently stimulated occurrence of fine and extension roots, which may have increased the absorptive capacity of the root system. Root number decreased with increasing depth and roots occurred predominantly in the top 800 mm of the soil profile. Starch was the principal carbohydrate storage form in the roots, irrespective of root size. Starch synthesis appeared not affected by root age. Sucrose and organic acid patterns were similar. Citric and tartaric acids were the main organic acids in roots, followed by malic acid. Elevated sugar and organic acid levels were found in roots of treated vines. The results demonstrate that the remaining leaves of partially defoliated vines were able to sustain normal metabolic functions in the roots. Canopy density was efficiently reduced by partial defoliation, leading to increased light penetration, fruit exposure, and photosynthetic activity of mature and old leaves. Although partially defoliated vines had much less leaf area per gram fresh berry mass at ripeness, yield increased considerably with defoliation at pea-size and veraison. Root density, yield, and cane mass were related. Grape total soluble sugar content was unaffected, but titratable acidity increased and the pH of the must decreased with partial defoliation. Ostensible increases in wine constituents (anthocyanins, phenolics), color density, cultivar character intensity, and overall wine quality were found in wines from treated vines.

KEY WORDS: Vitis vinifera, partial defoliation, root growth, root composition, canopy efficiency, grape composition, wine quality

Excessive vegetative growth and dense canopies of grapevines occur to some extent in all grape growing regions of the world. This is due primarily to the use of propagation material free of harmful viruses and the indiscriminate use of fertilizers, notably nitrogen, as well as to improvements in viticultural practices: e.g., soil management, irrigation, cultivation, and pest and disease control. Long term choices regarding rootstockscion combinations, training and trellising systems, and plant spacing greatly affect canopy density. In South Africa, a favorable climate, especially high temperature, contribute to vigorous growth. Canopy microclimate and source:sink relationships in grapevines are detrimentally affected by excessive growth, reducing photosynthetic activity of leaves (20,21, 22,23,37,39,60,62). Yield (59,63), grape composition, and wine quality (7,16,17,61,64,65) are also negatively affected. High humidity and low air flow in a dense
1,3.4Plant Physiologist, Plant Physiologist and Agricultural Research Technician, respectively, Nietvoorbij Institute for Viticulture and Oenology, Private Bag X5026, 7599 Stellenbosch, Republic of South Africa; 2Enzymologist, Institute of Plant Biology, Mycology and Phytochemistry, University of ZQrich, Zollikerstrasse 107, CH-8008 ZL~rich, Switzerland. *Corresponding author. This research was conducted at the Nietvoorbij Institute for Viticulture and Oenology.

Acknowledgements: Valuable technical contributions by A. J. Heyns, E. Burger, W. J. Hendricks, L. M. Paulse, and R. Skrivan are appreciated.
*Presented in part at the IV International Symposium on Grapevine Physiology, 11-15 May 1992, Italy, and at the InternationalSymposium on Table G rape Production, 28-29 June 1994, California, USA. Manuscript submitted for publication 20 June 1994. Copyright 1995 by the American Society for Enology and Viticulture. All rights reserved.

canopy-interior (24) promote bunch rot as reported by Smart et al. (64). Excess foliage further impedes effective pest and disease control (67). Against this background, excessive vigor is of major concern to producers striving to obtain prolonged maximum production of quality grapes. It is known that grapevine leaf photosynthesis is influenced by various factors (22 and references therein) and photoassimilate supply to the various sinks comprises a complex system of diversion and balance (20,21,37,39,52). Minimizing vegetative dominance will, therefore, require careful plant manipulation to prevent physiological imbalances and ensure that both sources and sinks function to full capacity. Partial defoliation is widely recognized as an invaluable practice to counteract the deleterious effects of excessive growth and plays a beneficial role in grapevine production (35,37,38,64). However, in many experiments with partial defoliation, leaves were indiscriminately removed and plants severely stressed. While focusing on a single problem, short- and long-term effects on leaf, fruit, and root physiology were frequently neglected. Therefore, the effects of different degrees of partial defoliation (33% and 66%) over the whole canopy, commencing at different developmental stages of the vine (budburst, berry set, pea-size, and veraison), on various physiological aspects were examined extensively (16,17,18,21,22,23,24,25). Based on this study, a method to defoliate grapevines discriminatively and with practical applicability was suggested. The objec306

A m . J. E n o l . V i t i c . , V o h 46, No. 3, 1 9 9 5

PARTIAL DEFOLIATION

-- 307

tive of the present investigation was to test this selective defoliation method by m e a s u r i n g some key parameters, including canopy efficiency, yield parameters, m u s t composition, and wine quality. Effects on development, distribution, and carbohydrate and organic acid contents of the root system are emphasized.

Materials

and Methods

E x p e r i m e n t a l v i n e y a r d : An l 1-year-old vineyard, situated in the Western Cape at Nietvoorbij, Stellenbosch, was used. Vitis vinifera L. cv. Cabernet Sauvignon (clone CS46), grafted onto 99 Richter (clone RY30), was spaced 3.0 1.5 m on a Glenrosa soil (Series 13, Kanonkop) (43) and trained to a 1.5-m slanting trellis described by Zeeman (77). Shoots not situated on two-bud spurs were suckered at approximately 30-cm shoot length. Soil c h a r a c t e r i s t i c s : The soil was double deepploughed in two directions to a depth of 800 m m prior to planting of the vines. Chemical characteristics, clay, silt, and sand contents in the different soil layers were reported by H u n t e r and Le Roux (18). Bulk density and water content in four soil layers, i.e., 0 cm to 30 cm, 30 cm to 60 cm, 60 cm to 90 cm, and 90 cm to 120 cm were determined during the winter according to s t a n d a r d methods. Phylloxera, margarodes, and nematode occurrence in the soil was determined with methods described by De Klerk (8,9) and Loubser (42). D e f o l i a t i o n t r e a t m e n t s : Three t r e a t m e n t s were applied: 0% (non-defoliated), and two respective 33% defoliation t r e a t m e n t s (consecutive removal of one out of every three leaves) applied evenly on main and lateral shoots from side to side in the canopy. The first 33% defoliation on both these t r e a t m e n t s was applied in the zone opposite and below bunches at berry set stage. The remaining part of the lower half of the canopy was subsequently 33% defoliated at either pea-size (treatment two) or veraison (treatment three). R o o t study. Root distribution: To determine root distribution, the profile wall method of BShm (3) was used, as modified by H u n t e r and Le Roux (18). Roots were plotted in each of six soil depths (0 - 20 cm, 20 - 40 cm, 40- 60 cm, 60- 80 cm, 80 - 100 cm, and 100 - 120 cm) in five root diameter classes: i.e., <0.5 mm, 0.5 m m to 2 mm, 2 mm to 5 mm, 5 mm to 10 mm, and >10 m m (66). Roots were categorized as fine (<0.5 mm), extension (0.5 - 2 mm), p e r m a n e n t (2 - 5 mm), and framework (5 - 10 mm and > 10 mm) roots according to Richards (54).

enized for 45 seconds using an Ultra-Turrax macerator operating at 20 500 rpm. The homogenate was transferred to a 0.45-~tm filter (Millipore Co.) and extraction repeated with 2 25 mL 80% ethanol. The filter residue was frozen at -20C, freeze-dried, and kept for starch analysis. Filtrates were combined, dried in a rotary evaporator at 35C and the residue redissolved in 5 mL 50% aqueous acetonitrile. The extract was passed through a column with intermediate base anion exchange resin (Bio-Rex 5, Bio-Rad Laboratories) and the organic acids subsequently desorbed from the resin with 10% H2SO 4. Both neutral sugar and organic acid fractions were passed through Sep-pak Cls cartridges and stored at-4C prior to analyses by HPLC, using the equipment and conditions described previously (26).

Extraction a n d analysis o f starch: A 50-mg sample of the freeze-dried, insoluble root material was transferred to an Eppendorf vial. One mL 80% aqueous acetone was added, followed by vortexing (10 sec) and sonication (10 min). The suspension was left at-4C for six hours, centrifuged (10 min) at full speed in an Eppendorf centrifuge, and the s u p e r n a t a n t decanted. The residue was then t a k e n up in 1 mL ethanol and treated as above, except t h a t the time lapse between sonication and centrifugation was omitted. After addition of 1 mL H20, the sample was washed again, the sediment frozen at 20C and freeze-dried overnight. The lyophilized material was t a k e n up in 550 ~tL H20 , followed by vortexing (10 sec) and sonication (10 min), whereafter it was left at -4C for 60 minutes and centrifuged. Immediately after centrifugation (10 min), a 50-~L aliquot was removed as control.
Starch was then gelatinized by incubating the sample in a boiling water bath (5 min with open caps and 55 min closed). After allowing the material to cool, 500 ~tL of an enzyme mix containing 5 U (z-amylase (Sigma A6380) and 2 U amyloglucosidase (Sigma A-7255) in 0.1 M Na-acetate (acetic acid/Na-acetate) buffer (pH 5.0) was added, the mixture vortexed for 10 seconds and incubated at 40C with constant shaking at 35 rpm, to allow hydrolyzation of starch [vials were removed and vortexed (10 sec) every 30 minutes]. After three hours, the samples were centrifuged (10 min) and diluted (1:39) with water. Glucose generated from starch was determined by using the ABTS [2,2' azino-di(3 ethylbenzthiazoline)-6'sulfonate] reagent, which consisted of 3.45 g Na2HPO 4 2H20, 1.6 g NaH2PO 4 H20 , 2350 U glucose oxidase (Boehringer no. 646423), 375 U peroxidase (Boehringer no. 127361) and 125 mg ABTS (Boehringer no. 102946), dissolved in 250 mL H20. A 50-~tL aliquot of the diluted sample was mixed With 950 ~tL of the above reagent. Absorbancy was read at 436 nm after 30 minutes. The blank consisted of a mixture of water and reagent. To obtain a glucose s t a n d a r d curve, seven standards were prepared: i.e., O, 5, 10, 20, 30, 40, and 50 mg glucose/100 mL. Results are expressed in mg starch after multiplication with a factor of 0.9, which allows for the reduced molecular weight of glucose in the polymer.

Root sampling: Roots from each of the above five classes were sampled randomly in the whole profile. They were then frozen a t - 2 0 C prior to freeze-drying with a Christ freeze-drying unit. Root samples were ground using a Cyclotec 1093 Sample Mill and stored at room temperature. Extraction a n d analyses o f sucrose, hexoses a n d organic acids: A modified method ofRuffner et al.
(57) was used for extraction: I g dry material from each root class was suspended in 50 mL MeOH-CHC13 - 0.2 M HCO2H (12:5:3 v/v) (pH approximately 4.2) and homog-

Am. J. Enol. Vitic., Vol. 46, No. 3, 1995

308 m HUNTER

et al.

Canopy m e a s u r e m e n t s : The potential of vines to produce quality winegrapes was assessed by scoring canopies with the vineyard score card outlined by S m a r t et al. (64). Light intensity just above the vine cordon was determined in the late morning with a LI-COR Line Q u a n t u m Sensor and expressed as a percentage of ambient light intensity. Photosynthetic activity (mg COz/dm2/hr) of basal (just above bunches) and middle leaves was measured with a portable photosynthesis meter (ADC), as described by H u n t e r and Visser (22). V e g e t a t i v e growth, yield and grape composition: Budding, bud fertility, yield, leaf area/g fresh berry mass, cane mass, total soluble solids, titratable acidity, and m u s t pH were determined as described previously (16,24,25). Wine c o m p o s i t i o n and quality: Wines were made as reported by H u n t e r et al. (17). Anthocyanin content, color density and phenolic content of wines were measured spectrophotometrically at 530 nm, 530 + 420 nm, and 280 nm, respectively, in a Varian UV/VIS spectrophotometer (Model 2200) using 10 m m quartz cells. Wines were evaluated sensorially for cultivar character intensity and overall wine quality. E x p e r i m e n t a l d e s i g n a n d statistical analyses: The experiment was laid out as a completely randomized design. For the root study, five uniform, healthy vines (replications) per t r e a t m e n t were randomly selected. The study was conducted in the fourth growth season after partial defoliation was commenced. Data were collected in winter (July/August). Canopy, growth, yield, and grape composition measurements were made on 14 replications (one vine plots) for each of the three treatments. Canopy scores of each t r e a t m e n t were done on five vines by three judges. T r e a t m e n t s were applied for three consecutive years. Data were collected at ripeness. Except for canopy scores, means obtained over the last two years are presented. Wine quality determinations and evaluations were made on duplicate wines after three years. Wines were evaluated sensorially for cultivar character intensity by 17 judges and for overall wine quality by 22 judges, using a nine-point scale. The scores of each judge were expressed as a percentage of m a x i m u m score. The n a t u r e of the data was such t h a t it could not be statistically analyzed. Where possible, a one-way analysis of variance was performed on the raw data; differences between treatment means were determined using Student's t-LSD test.

3025o~
t._

- 2.0

1.5 20 15 1.0 = (9 "o

i

(9

n 10 0 5 0 I 0-30 , , 30-60 60-90 Soil layer (cm)

0.5

m i

' 0.0 90-120

Fig. 1. Soil water content and bulk density of a Glenrosa soil (Series 13, Kanonkop) at Nietvoorbij, Stellenbosch, measured in winter.

tion index (bulk density) was higher in the top 60 cm, exceeding the critical value of 1.5 g/cm 3for root penetration reported by Richards (54) (Fig. 1). Growth conditions with respect to soil water and bulk density were similar for non-defoliated and partially defoliated vines (data not shown). The numbers of nematode species and phylloxera were insignificant r e g a r d i n g grapevine health (data not shown). Margarodes were not found.

Results and Discussion

Soil characteristics: As reported earlier (18), clay and silt content of the soil generally increased with depth, while the percentage of sand in the different sand classes decreased. Resistance and pH of the soil as well as P, K, and Ca content decreased with increasing depth, while Mg content increased. Percentage soil water increased with depth, whereas the soil compac-

Root distribution: Root density was not significantly affected by partial defoliation as applied in this study. However, an a p p a r e n t stimulation of root density (number/m 2)was evident after partially defoliating vines at pea-size [397] and veraison [409], respectively, compared to non-defoliated vines [383]. This, as well as the ostensible increase in root density the later defoliation was applied, correspond to previous findings on fieldgrown Cabernet Sauvignon (18), but do not support the decrease in root dry mass found when leaf area of potgrown Muscat d'Alexandrie and Thompson Seedless was reduced (5,33). However, it must be emphasized t h a t vines used in the latter studies were only one year old and leaf area severely reduced, whereas vines used in the present study were older t h a n 10 years at the time defoliation commenced. Fine roots (<0.5 mm) dominated the soil profile, while framework roots of more t h a n 10 mm were present in lowest numbers (Table 1). Evidently, growth of fine and extension (<0.5 - 2 mm) roots was stimulated by partial defoliation. Growth of p e r m a n e n t (2 - 5 mm) and framework (5 - 10 mm) roots was reduced, compared to t h a t of non-defoliated vines. It is apparent that grapevines and other plants absorb a large proportion of water and nutrients through roots t h a t have undergone secondary thickening (2,50,70). However, it stands to reason t h a t occurrence of higher numbers of fine and extension roots which are rapidly growing, would not only increase the absorptive capacity and activity of the root system and allow more efficient utilization of nutrients and water from the soil, but also enhance the production of growth regulators (cytokinin, gibberellin, abscisic acid) which regulate shoot and fruit development (10,54). Notwithstanding the findings of Van Zyl and Van Huyssteen (72) t h a t thicker roots of Chenin

Am. J. Enol. Vitic., Vol. 46, No. 3, 1995

PARTIAL DEFOLIATION - - 309

Table 1. Effect of 33% defoliation of grapevines on the total number of roots per diameter class.
Root d i a m e t e r class (mm) Treatment

their numbers decreased by a factor of four in the 100 cm to 120 cm soil layer. Interestingly, root occurrence (Table 2) decreased with decrease in soil compaction (Fig. 1), indicating t h a t soil compaction was not restricting root penetration in this study (cf. also 50). The slightly higher root numbers of partially defoliated vines in the 40 cm to 120 cm soil layer suggest an improved penetration and utilization of deeper soil layers, which may have positive implications regarding performance under non-irrigated and/ or drought conditions. It was also found by H u n t e r and Visser (22) that, although the rate of photosynthesis increased when vines were partially defoliated, a lower transpiration:photosynthesis ratio was needed by their leaves.

<0.5 Control 631 *Pea size defoliation 659 *Veraison defoliation 674 Mean 655 a

0.5 - 2 62 77 78 72 b

2-5 29 16 23 23 bc

5 - 10 11 9 8 9 bc

>10 1 1 2 1c

*Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Values designated bythe same letter do not differ significantly (p< 0.05).

blanc/99 Richter had a better regenerative ability t h a n thinner roots, regeneration was clearly not impeded by defoliation. It would even seem as if the root system adjusted its growth rate by initiating new roots, as was found for apple trees (44). Regardless of treatment, the n u m b e r of roots decreased with increasing depth (cf. also 51) (Table 2). As is generally found for grapevines (18,68,71), roots were predominantly located in the top 80 cm, beyond which

Root composition: Starch, sugar, and organic acid concentrations correspond to data of Winkler and Williams (75), indicating t h a t starch is the main assimilate storage form in the roots, exceeding 25% of root dry weight and representing 90% of the carbohydrates analyzed, followed by sucrose and almost identical glucose and fructose concentrations (Fig. 2 and 3). Root classes differed in their ability to function as starch stores, p e r m a n e n t roots having the highest concentration. The results nevertheless indicate t h a t the activity of starch-synthesizing enzymes was not limited by Table 2. Effect of 33% defoliation of grapevines on the total number of roots per soil layer. root age. Since no evident conversion Soil layer (cm) of starch to sugar during the winter Treatment 0 - 20 20 - 40 40 - 60 60 - 80 80 - 100 100 - 120 dormancy period was found (75), invertase-mediated hydrolysis of suControl 177 151 145 144 78 40 crose probably continues to cater for *Pea size defoliation 168 139 164 157 89 44 maintenance metabolism (respirato*Ve raison defoliation 177 151 156 153 106 43 ry losses), albeit at a low rate. Invertase is known to be abundant in grapeMean 174 a 147 b 155 ab 151 b 91 c 42 d vines (57). This, however, raises the question of the carbon source which is * Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. able to sustain accumulation of suValues designated by the same letter do not differ significantly (p < 0.05). crose along a concentration gradient in the roots. Given the higher relative 330 -20 BB Starch hexose concentrations of fine, extenab 300 sion, and p e r m a n e n t roots, they seem Sucrose b to be playing a dual role in assimilate Glucose a 270 m metabolism, namely utilizing sugar c ab -15 E A 240and storing starch, whereas the framework roots tended to be more storageab E 210o') orientated.
-A

~' 1 8 0 "ID "~ 1 5 0 E 120J~

a__

NJ
ri.N
~N

a_~
bc

E -10 ~
. w o x

o "m

9060300

-5
-

~
L._

o ~

< 0.5

"" 2-5 5-10 0.5-2 Root diameter class (mm)

>10

Fig. 2. Sugar content of roots of different diameter sizes. Values designated by the same letter do not differ significantly (p < 0.05) for each sugar.

Sucrose and organic acid levels in the different root classes behaved similarly. As was found by Kliewer (28,29), tartaric and citric acids were predominant, with malic acid present in smaller amounts. A reverse pattern was found in xylem exudates of one-year-old spurs of Chardonnayjust before budburst (11). The intermediates of the tricarboxylic acid cycle, citric and malic acid, may be utilized as a n a p l e r o t i c sources of e n e r g y (34,53,56) in maintenance reactions.

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310-

HUNTER et aL

30 t
~'w 2 . 5 E p, "o 2.0

BB T a r t a r i c a c i d I--] C i t r i c a c i d ~
Malic acid

ti
~

ab

~ 1.5
-o

"g m
.2 1.0
O) o 0.5
era

ab

a b
bc

b ab
i

ab

0,0

< 0.5

0.5-2 2-5 5-10 Root diameter class (mm)

>10

Fig. 3. Organic acid content of roots of different diameter sizes. Values designated by the same letter do not differ significantly (p < 0.05) for each organic acid.
300 a

-15 BB Control 1 [ ~ *Pea size defoliation

*Veraison defoliation

A W W m

{4 W

1=
-/" a

10~" "o

"o

E
ID 0

E
J O L_

or)

o
0

~ Starch

~%~ Sucrose

~ Glucose

P~ Fructose

Fig. 4. Effect of 33% defoliation of the grapevine on root starch and sugar content. *Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Values designated by the same letter do not differ significantly (p< 0.05) for each sugar or for starch.

reserves (5,6,24,25,30,32,33,45,49), whereas little negative or even beneficial effects are observed when partial defoliation is applied after veraison (24,25,46,58). The results on root distribution and composition in the present study clearly demonstrate that neither assimilate sources, nor size and activity of the root system were depleted/affected by partial defoliation. Root systems of established, field-grown vines, therefore, do have the capacity to withstand the assumed stress conditions induced by partial defoliation. Photosynthates supplied by the remaining leaf area during the growth season were apparently still meeting/exceeding the demand for assimilates needed for growth, storage and respiration, processes which are critically important for survival of perennial plants and sustained productivity (41). Although the levels of root reserves do not fluctuate substantially during the dormancy period (27,29,75), the data do not indicate whether higher proportions of carbohydrates were mobilized from storage regions to support growth in spring upon partial defoliation in the previous season; if so, it was not detrimental. It is possible that the root system may find itself in a position of reduced sink strength due to apical dominance (cf. 73) under vigorous foliar growth conditions during the vegetative season. As soon as the above-ground vegetative dominance and the normal assimilate distribution p a t t e r n between leaves and bunches (20,21,36,52,74) are changed by partial defoliation, the roots may have access to assimilatory reserves and recently produced photosynthates in the phloem which effectively neutralize possible stress conditions imposed by partial defoliation.
BB Control

However, the physiological significance of tartaric acid remains enigmatic (55). After defoliating vines from budburst onwards, Marangoni et al. (47) suggested that c a r b o h y d r a t e s and organic acids occurring in the xylem sap in the pre-bloom growth phase were derived largely from plant reserves. Whether tartaric acid is transported to the new growth areas in spring and whether it is available for accumulation in the berries are open to speculation. Although none of the sugars or organic acid levels were significantly affected by partially defoliating vines elevated concentrations were found for most compounds (Fig. 4 and 5). Premature and severe defoliation of grapevines causes mobilization of stored carbohydrate reserves from roots, trunks, and canes and induces stress conditions, as indicated by decreases in yield, lower bud fertility in the following season, delayed ripening, and reduction in dry weight and carbohydrate

~, 2.0- ][---1*Pea size defoliation w *Veraison defoliation I .................................... E a ~ 1.5~ ~ 1.0O

a __

a a

o "8 0.5 m o
Tartaric acid Citric acid Malic acid Fig. 5. Effect of 33% defoliation of the grapevine on root organic acid content. *Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Values designated by the same letter do not differ significantly (p < 0.05) for each acid.
0.0

Am.

J. Enol. Vitic., Vol. 46, No. 3, 1995

PARTIAL D E F O L I A T I O N - 311

Table 3. Effect of 33% defoliation of grapevines on canopy characteristics and vine performance.

BB Control *Pea size defoliation i! *Veraison defoliation ~i

Treatment Parameter
2Canopy gaps (%) 2Canopy density index 2Fruit exposure (%) Light intensity (% ambient) 3Bud fertility index Budding(%) Leaf area (cm2)/g fresh berry mass Yield (kg/vine) Cane mass (kg/vine)

Control
10 - 20 >2 30 13.00 b 1.30 a 91.50b 17.27 a 3.48 b 1.30 ab

~Pea-size defoliation
50 <2 50 26.00 a 1.38 a 98.40a 13.23 b 3.87 b 1.02 b

~Veraison defoliation
45 2 55 32.00 a 1.42 a 94.10ab 9.73 c 5.55 a 1.61 a >,,

6-

c4
Jo o Jo
L_

<

~Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. 2Scored with the vineyard score card according to Smart et aL (66). 3Bud fertility = number of bunches/number of shoots originating from buds allocated during pruning. Values designated bythe same letter do not differ significantly (p < 0.05) for each parameter.

Anthocyanin content
(As3o)

Color density
(A4,o+s3o)

Phenolic content
(6280)

Fig. 7. Effect of 33% defoliation on wine composition. *Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set.

.c7-

BB Control
[ ~ *Pea size defoliation I! { ~ *Veraison defoliation
~.....,..~.................~ ..-.-o-.~,~.~...~-~ .-,.~.-~............... ~ ~,~.,..o~......... ~ , - ~ . ~ .,,-~ ~ . , . . . ~

80

BB Control *Pea size defoliation *Veraison defoliation

-80

~60 0 5 .
ol

E ~4-

a A
o

i
' v

o~ =
m

~3,,C e~
0

~ 2 - I

40!._ ,

-40
"==

"

01.

I
m ~ m

= ._>
,1~ =-

I
"

~=
t_.

Basal leaves Leaf position

Middle leaves

tO 20-

> -~n 0

Fig. 6. Effect of 33% defoliation on photosynthetic activity of basal and middle leaves. *Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Values designated by the same letter do not differ significantly (p < 0.05) for each leaf position.

0
Table 4. Effect of 33% defoliation of grapevines on grape composition at ripeness. Fig. 8. Effect of 33% defoliation on wine quality. *Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Canopy efficiency: Canopy gaps, fruit light exposure, and number of leaf layers of partially defoliated vines (Table 3) were in accordance with parameters recommended for the production of higher quality grapes (62,64). The results also imply that the incidence of pests and diseases would be reduced and chemical control would benefit from partial defoliation (4,38,67,76). Both defoliation treatments significantly improved light intensity in the interior of the canopy (Table 3). This certainly contributed to the higher photosynthetic activities of middle and basal leaves (Fig. 6), confirming

Treatment Parameter
Total soluble solids (Brix) Titratable acidity (g/L) pH

Control
24.77 ab 7.31 c 3.24 a

*Pea-size defoliation
25,15 a 7.63 b 3,21 a

*Veraison defoliation
24.63 b 8.18 a 3.16 b

*Defoliation in the lower half of the canopy; preceded by defoliation in the bunch zone at berry set. Values designated bythe same letter do not differ significantly (p < 0.05) for each parameter.

Am. J. Enol. Vitic., Vol. 46, No. 3, 1995

312 - - H U N T E R et aL

the stimulating effect of partial defoliation (15,22,39) and the deleterious effect of interior-canopy shade on photosynthetic response (22,37,48,62).

Yield parameters: Since the period just prior to bloom is critical in determining the number of bunches per bud (1), it is remarkable that defoliation at pea-size and veraison (and concomitant higher light intensity received by the basal buds) can apparently still increase bud fertility (Table 3), unless the initial defoliation in the bunch zone at berry set stage benefited conversion of anlagen to inflorescences instead of tendril primordia. The more shaded interior of the non-defoliated vines probably also impeded nutrient supply from the leaves to the buds, decreasing bud fertility and budding capacity (Table 3). Notwithstanding the 23% and 44% less leaf area/g fresh berry mass at ripeness (Table 3), partially defoliated vines produced approximately 11% and 59% higher yields after defoliation at pea-size and veraison, respectively (Table 3). This is in contrast to previous results with severe defoliation (24,30) and further proves that enhanced mobilization of carbohydrate reserves as a result of partial defoliation as suggested earlier (30), is very unlikely under the conditions of this experiment. Lower and more favorable source:sink ratios created by partial defoliation as well as the improved canopy microclimate favoring metabolic activity of both leaves and grapes are a more obvious explanation. The stimulation of yield and photosynthetic activity is all the more remarkable because the major part of defoliation took place during the time of decreasing canopy photosynthesis (19,40) and passive root growth (71). In line with previous results (25), cane mass was little affected by partial defoliation; however, early defoliation reduced shoot growth, compared to later defoliation (Table 3). In general, the results coincide with data for various rootstocks under different cultural practices that root density, yield, and cane mass are closely related (18,68,71). Grape composition and wine quality: Partial defoliation had no effect on total soluble solid accumulation in the fruit, but increased titratable acidity and reduced must pH (Table 4). Apparently, meaningful increases in both wine composition (Fig. 7) and wine sensory data (Fig. 8) were found when vines were partially defoliated. General: Reasons for the improved performance of partially defoliated vines were discussed in previous papers (16,17) and were mostly related to the fact that an inferior canopy microclimate caused by excessive foliage is in a direct and indirect way detrimental to the general metabolism of vines. Supply (via photosynthesis) and demand [via respiration, cell division and/or enlargement, and storage (14)] under warm and cool climates (31,35,37,62,64,65) are affected, resulting in vines functioning below their individual maximum potential/efficiency. Conclusions
Although the root system was relatively insensitive

to defoliation as applied in this study, the tendentious stimulation of root growth and root system efficiency indicate a fine balance between above-ground and subterranean growth. Considering the much lower leaf area per gram fresh berry mass of partially defoliated vines as compared to that of non-defoliated vines, it is evident that performance of leaves in supplying photosynthates for growth and development depend to a large extent on microclimate and source:sink ratio. This indicates that the generally observed relationship between above-ground and subterranean growth of grapevines is very important in the control of general metabolism, soil utilization, yield, and wine quality and can be positively manipulated by careful canopy management. It is evident that the increase in photosynthetic activity of remaining leaves after partial defoliation was sufficient to sustain or even increase normal growth of the root system. The homeostatic mechanism by which the balance between shoot and root output (size X activity) is maintained (44,54) was obviously not disrupted. The results indicate that higher carbon fixation rates ensured high export from sources and utilization in sinks and vice versa (12,13,14,69). Growth regulators may play an important role in their control of this relationship (74). It would seem as if both the root system and the canopy increased their efficiency in response to partial defoliation. As suggested earlier (23), senescence of remaining leaves on partially defoliated vines may be inhibited, which is important in the light of the elevated sucrose production in leaves and their presumed contribution to storage pools, particularly after harvest (19). All factors investigated point to the fact that discriminative defoliation as applied in this study does not impede grapevines in any way. On the contrary, plant performance was improved to such an extent that partial defoliation must be considered a necessity for vigorously growing grapevines in a given situation. The manipulations have no deleterious effects on the environment and the vines retain normal allocation patterns, which is important for continued health and longevity particularly of perennial plants such as the grapevine.

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