RESEARCH NOTE I N F L U E N C E OF W I N D B R E A K S A N D CLIMATIC R E G I O N ON D I U R N A L F L U C T U A T I O N OF LEAF W A T E R P O T E N T I A L , S T O M A T A L C O N D U C T A N C E , A N D LEAF T E M P E R A T U R E OF GRAPEVINES

Brian M. Freeman, W. M. Kliewer, and Peter Stern

Respectively, Graduate Student and Professor of Viticulture, Department of Viticulture and Enology, University of California, Davis; and Winemaker, Turgeon & Lohr Winery, San Jose, CA. Manuscript submitted 9 November 1981. Revised manuscript received 22 February 1982. Accepted for publication 24 May 1982.

ABSTRACT Reduced stomatal conductance of Chardonnay grapevines grown near Greenfield in the Salinas Valley was associated with both dense fog during most of the morning and then with wind in the afternoon. After 13:00 hours, when wind speed markedly increased in the Greenfield area, vines grown in the absence of a windbreak had markedly lower stomatal conductance than vines protected by a windbreak. Wind did not appear to increase water stress of non-sheltered vines, which were less stressed (less negative leaf water potential) than sheltered vines (more negative leaf water potential), indicating that the reduced stomatal conductance of non-sheltered vines was not due to water stress. Leaf stomatal conductance remained high at Davis for about 12 hours per day compared to four and eight hours for non-sheltered and sheltered vines grown near Greenfield. Since stomatal conductance, CO2 assimilation, and rate of photosynthesis are directly related, low stomatal conductance indicates that C02 assimilation and photosynthesis are being curtailed. Consequently, non-sheltered vines located in the more windy areas of Salinas Valley may be photosynthetically active for as little as four hours per day, whereas the photosynthetic active period of vines grown in the presence of windbreaks may be nearly doubled. The effects of reduced photosynthetic activity on grape quality is discussed.

Stomatal regulation is a complex interaction of external and internal factors (1,5). In many plants wind can cause stomatal closure and consequently limit CO2 uptake and photosynthesis, even though adequate soil moisture is available. During most of the summer months, the Salinas Valley is characterized by daily winds that begin blowing in from the ocean during the late morning or early afternoon hours and generally continue late into the night. These winds coupled with dense morning fog very likely reduce the daily period for active CO2 uptake and assimilation in grapevines growing in parts of the Salinas Valley as well as other areas with similar climatic characteristics. To investigate this climatic condition, a series of measurements were made on 11 August 1981 to compare changes in stomatal conductance and leaf water relations of Chardonnay grapevines growing in sheltered (behind windbreaks) and non-sheltered sites near Greenfield, California. Similar meausrements were made on Carignane vines grown at the University of California Experimental Vineyard at Davis on 19 August 1981. Temperature conditions were mild at Davis on this day, with a daytime maximum temperature of 27C compared to 26C at Greenfield on August 11. The Greenfield area is a Region II and Davis is a Region IV according to the Winkler-Amerine degree day system of classifying climat233

ic regions in California. The main objective of this investigation was to evaluate the relative importance of windbreaks on diurnal fluctuations of leaf temperature, stomatal conductance, and leaf water potential of grapevines. MATERIALS AND METHODS Stomatal conductance and temperature were measured on the abaxial surface of leaves at one to two hour intervals from about 06:00 to 20:00 hours with a Li-Cor Li-65 Autoporometer (Li-Cor Instrument Corp., Lincoln, Nebraska). Leaf water potential was measured with a portable pressure chamber, Model 600 (PMS Instrument Co., Corvallis, Oregon) using 30 second interval readings, and average wind speed was measured with a handheld anomometer. Data presented in Figures 1 to 4 are the means of three measurements. Exposed, fully expanded leaves, about the 15th leaf from the base of the shoot, were selected for measurements. The Greenfield site was located on the Turgeon & Lohr Vineyard, two kilometers northwest of Greenfield. The sheltered site vines were located 70 meters from an approximately 20 meter high eucalyptas tree windbreak. The non-sheltered site (no windbreak trees) was in the same block of vines, but about 400 meters east of the sheltered site and 160 meters from the north edge of the vineyard. The variety at both these sites was Chardon-

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nay, which was head-trained and cane-pruned on a threewire vertical trellis consisting of a cane wire 107 cm from the ground and two foliage supporting wires 30 and 60 cm above the cane wire. The Davis site was located at the University of California Experimental Vineyard and consisted of eightyear-old Carignane vines cordon-trained and spur pruned on a three-wire trellis consisting of a cordon wire 107 cm from the ground and two foliage supporting wires on a 90 cm wide crossarm 45 cm above the cordon wire. RESULTS Leaf temperatures of vines grown at Davis and sheltered vines at Greenfield (Fig. 1) did not differ significantly at 08:00 hours; however, at Greenfield ter~peratures of leaves from non-sheltered vines were 0.8C lower than sheltered leaves at 08:00 hours (P <0.01). Maximum leaf temperatures at Greenfield and Davis were reached at 13:00 and 15:00 hours, respectively, and did not differ significantly (Fig. 1). Maximum temperatures of nonsheltered vine leaves at Greenfield were about 1C lower than sheltered leaves, not significantly different at the 5% level. Late afternoon (17:00 to 18:00 hours) nonsheltered leaf temperatures at Greenfield were significantly (P <0.001) lower than leaf temperatures at Davis measured at the same time of the day. Also, late afternoon exposed leaf temperatures of sheltered vines at Greenfield were about 2C warmer than non-sheltered leaves (P<0.01) (Fig. 1).
30

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0.4

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Figure 2. Diurnal changes in stomatal conductance of Chardonnay vines grown at Greenfield under sheltered (B) and nonsheltered (FI) conditions and Carignane vines grown at Davis (A).

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/
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The s t o m a t a l conductance of the non-sheltered Greenfield vines increased over four-fold between 11:00 and 12:00 hours and was significantly (P<.05) greater than that of the sheltered vines. However, it then declined rapidly again when the wind velocity reached rates in excess of 3 m sec -1 (6.7 mph) beginning shortly after 12:00 hours. This indicated that the stomata were again beginning to close (Fig. 2). The stomatal conductance of the sheltered vines, on the other hand, increased more slowly than the non-sheltered vines. However, the stomatal conductance of sheltered vines continued to increase until about 15:00 hours and was significantly (P<.05) greater than that of the non-sheltered vines from 15:00 to 18:00 hours. This provides indirect evidence that the sheltered vines were photosynthesizing for three to four hours longer than the non-sheltered vines (Fig. 2). The eucalyptus windbreak at Greenfield reduced the windspeed from 4 to 6 m sec -1 to 1.5 m sec -1 (Fig. 3). At Davis there was no detectable wind on 19 August. The

1S

10 E00
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r~ o

TIME

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Figure 1. Diurnal changes in leaf temperatures of Chardonnay vines grown at Greenfield under sheltered (n) and non-sheltered (FI) conditions and Carignane vines grown at Davis (A).

Stomatal conductance of leaves at Davis was usually less than at Greenfield (Fig. 2). This may be due to varietal differences (1,3), but other factors such as leaf age and preconditioning to stress cannot be ruled out. Thus, only relative comparisons or trends can be made between measurements made at Davis and Greenfield (Fig. 2). The delay in the increase in stomatal conductance at Greenfield was closely associated with the dense fog that limited the amount of light available for photosynthesis, which did not clear off until about 11:00 hours.

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Figure 3. Diurnal changes in wind velocity of Chardonnay vines grown at Greenfield under sheltered (B) and non-sheltered (FI) conditions.

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&

-9

-12
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Figure 4. Diurnal changes in leaf water potential of Chardonnay vines grown at Greenfield under sheltered (11) and non-sheltered (El) conditions, and Carignane vines grown at Davis (A).

data in Figure 4 reveals that the wind did not cause stomatal closure indirectly through increased transpiration, and hence water stress in the vines, since the leaf water potential of the non-sheltered vines was higher (less negative) than the sheltered vines. The water potential of vines in both sites declined as the day progressed, as is normally the case (6). When the stomata of the nonsheltered vines were nearly closed (about 14:00 hr) the leaf water potential remained nearly constant at about - 8 bars until around 17:00 hours. Between 15:00 and 18:00 hours the leaf water potential of the sheltered vines was significantly (P<.05) more negative than the non-sheltered vines. A leaf water potential o f - 1 3 to - 1 5 bars is generally considered necessary to cause stomata of grapevines to close (4,6). The water potential of the sheltered vines continued to decline until about 16:00 hours, reaching - 1 1 bars, at which time their stomata began to close; i.e. stomatal conductance declined (Figs. 2 and 4). Thus, stomatal closure in the non-sheltered vines was not attributable to high leaf water potential since the - 1 3 bar threshold level for stomatal closure was never reached. Therefore, it is concluded that stomatal closure was a direct effect of wind and not moisture stress. DISCUSSION Stomatal conductance for water vapor in leaves (which is what the autoporometer measures) is also an estimation of CO2 assimilation into leaves (5). Conductance of water vapor and CO2 into the leaf involves identical pathways and diffusion mechanisms. Water vapor, because of its lower molecular weight diffuses 1.6 times faster than C02. The sensitivity of assimilation and transpiration rates remains constant when stomatal conductance is varied by changes in temperature and humidity (2). However, the ratio of assimilation to transpiration is not constant under all conditions (2), consequently stomatal conductance is only a guide to potential CO2 uptake. The heat summation for both Davis and Greenfield on the dates that mesurements were taken were similar, so according to the Winkler-Amerine degree day system the

assimilation should have been similar. The stomatal conductance data suggest that non-sheltered vines at Greenfield were assimilating at this maximum rate for only about four hours per day, whereas, vines at Davis continued at a maximum rate for 12 to 13 hours. The maximum stomatal conductance at Davis was less than at Greenfield, but this may be due to varietal differences (1,3) or leaf age and preconditioning effects. Fog limited stomatal conductance of the Greenfield vines during most of the morning hours, whereas, wind appeared to be the dominant factor in the afternoon. The presence of a windbreak at Greenfield increased the period of maximal stomatal conductance to about eight hours compared to four hours for non-sheltered vines. Increased leaf to air vapor pressure deficit reduced stomatal conductance and a linear relationship was established between stomatal conductance and leaf to air vapor pressure deficit for apples (8). This response is attributed to epidermal water loss and not a decrease in bulk leaf water potential. Vapor pressure deficit was not measured for the days reported in this experiment, but wind would reduce vapor pressure deficit and may have been the reason for the reduced stomatal conductance of the non-sheltered vines. The late time in the year when grapes mature in the northern part of the Salinas Valley may be in large measure due to both the light limitations of fog and wind factors that greatly limit the daily active period of photosynthesis needed to manufacture the sugar for fruits. The reduced photosynthetic activity of vines grown in the Salinas Valley may also indirectly affect potassium levels in grape berries. In an experiment with Ricinus communis L., Smith and Milburn (7) studied phloem loading of potassium and sucrose. Potassium was loaded into the phloem when sucrose was not available due to an extended dark period. Potassium concentration in the phloem was increased by 30% and 60% after 12 and 24 hour dark periods, respectively. The 20 hour period in which C02 was not actively assimilated in non-sheltered vines at Greenfield may increase the potassium level in the phloem, which would then be available for loading into grape berries. It is unknown if grapevines behave similarly to Ricinus, as attempts to obtain uncontaminated phloem exodate samples from grapevines have been unsuccessful. Work is in progress to determine the importance of wind movement and light levels on photosynthesis, stomatal conductance and potassium accumulation in fruits of grapevines. LITERATURE CITED 1. Dtiring, H. Studies on environmentally controlled stomatal transpiration in grapevines. I. Effects of light intensity and air humidity. Vitis 15:82-7 (1976). 2. Hall, A. E. and E. D. Schulze. Stomatal response to environment and a possible interrelation between stomatal effects on transpiration and CO2 assimilation. Plant, Cell and Environment 3:46774 (1980). 3. HoF~icker,W. Investigations on the influence of changing soil water supply on the photosynthesis intensity and the diffusive resistance of vine leaves. Vitis 15:171-82 (1976).

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4. Kriedemann, P . E . and R . E . Smart. Effects of irradiance, temperature and leaf water potential on photosynthesis of vine leaves. Photosynthetica 5:6-15 (1971). 5. Raschke, K. Stomatal action. Ann. Rev. Plant Physiol. 26:30940 (1975). 6. Smart, R. E. Aspects of water relations of the grapevine (Vitis

vinifera). Am. J. Enol. Vitic. 25:84-91 (1974).

7. Smith, J. A. C. and J . A . Milburn. Osmoregulation and the control of phloem-sap composition in Ricinus communis L. Planta 148:28-34 (1980). 8. Warrit, B., J. J. Landsberg and M. R. Thorpe. Responses of apple leaf stomata to environmental factors. Plant, Cell and Environment 3:13-22 (1980).

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