This action might not be possible to undo. Are you sure you want to continue?
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 27°C compared to 26°C 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-
Am. J. Enol. Vitic., Vol. 33, No. 4, 1982
This indicated that the stomata were again beginning to close (Fig.2 nay.4 0. 1). however. The eucalyptus windbreak at Greenfield reduced the windspeed from 4 to 6 m sec -1 to 1. 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. This may be due to varietal differences (1. Vol. Also. 30 0. The 1S 10 E00 1000 1 SO0 2000 r~ o TIME (hrs) l@ Figure 1. Stomatal conductance of leaves at Davis was usually less than at Greenfield (Fig. Diurnal changes in wind velocity of Chardonnay vines grown at Greenfield under sheltered (B) and non-sheltered (FI) conditions. increased more slowly than the non-sheltered vines. No.8 0. 33.01) (Fig. Am.5 m sec -1 (Fig. 1) did not differ significantly at 08:00 hours. RESULTS Leaf temperatures of vines grown at Davis and sheltered vines at Greenfield (Fig. J. only relative comparisons or trends can be made between measurements made at Davis and Greenfield (Fig. 2). However. 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. Maximum leaf temperatures at Greenfield and Davis were reached at 13:00 and 15:00 hours. 4.0 500 I000 1500 2000 TTME (hrs) Figure 2.01). 1982 . This provides indirect evidence that the sheltered vines were photosynthesizing for three to four hours longer than the non-sheltered vines (Fig. which did not clear off until about 11:00 hours.INFLUENCE OF WINDBREAKS 1.3). Enol. not significantly different at the 5% level. 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 stomatal conductance of the sheltered vines. and did not differ significantly (Fig. 25 / 20 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<. 2). However.. the stomatal conductance of sheltered vines continued to increase until about 15:00 hours and was significantly (P<.234 . late afternoon exposed leaf temperatures of sheltered vines at Greenfield were about 2°C warmer than non-sheltered leaves (P<0. it then declined rapidly again when the wind velocity reached rates in excess of 3 m sec -1 (6..05) greater than that of the non-sheltered vines from 15:00 to 18:00 hours. Vitic. Maximum temperatures of nonsheltered vine leaves at Greenfield were about 1°C lower than sheltered leaves. on the other hand.8°C lower than sheltered leaves at 08:00 hours (P <0.7 mph) beginning shortly after 12:00 hours. Late afternoon (17:00 to 18:00 hours) nonsheltered leaf temperatures at Greenfield were significantly (P <0. respectively. 500 I000 1500 2000 TIME (hrs) Figure 3.001) lower than leaf temperatures at Davis measured at the same time of the day.05) greater than that of the sheltered vines. At Davis there was no detectable wind on 19 August. but other factors such as leaf age and preconditioning to stress cannot be ruled out. 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). 1). Thus. 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). at Greenfield ter~peratures of leaves from non-sheltered vines were 0. 3). 2). 2).
The water potential of vines in both sites declined as the day progressed.. it is concluded that stomatal closure was a direct effect of wind and not moisture stress.235 0 & -9 -12 500 1000 IS00 2000 TIME (hrs) Figure 4. Fog limited stomatal conductance of the Greenfield vines during most of the morning hours. Vitis 15:171-82 (1976).. whereas. 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. as is normally the case (6). E. I.1 1 bars. stomatal conductance declined (Figs. The heat summation for both Davis and Greenfield on the dates that mesurements were taken were similar. It is unknown if grapevines behave similarly to Ricinus. Thus.W. Studies on environmentally controlled stomatal transpiration in grapevines. However. HoF~icker.05) more negative than the non-sheltered vines. the ratio of assimilation to transpiration is not constant under all conditions (2).6). Vapor pressure deficit was not measured for the days reported in this experiment. 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). stomatal closure in the non-sheltered vines was not attributable to high leaf water potential since the . A leaf water potential o f . 3. Enol. stomatal conductance and potassium accumulation in fruits of grapevines.. so according to the Winkler-Amerine degree day system the assimilation should have been similar. whereas. and Carignane vines grown at Davis (A). consequently stomatal conductance is only a guide to potential CO2 uptake. Potassium was loaded into the phloem when sucrose was not available due to an extended dark period. The water potential of the sheltered vines continued to decline until about 16:00 hours. This response is attributed to epidermal water loss and not a decrease in bulk leaf water potential. Vitic.3) or leaf age and preconditioning effects. Hall. The stomatal conductance data suggest that non-sheltered vines at Greenfield were assimilating at this maximum rate for only about four hours per day. A. which would then be available for loading into grape berries. respectively. Plant. Water vapor. reaching . Vitis 15:82-7 (1976).1 5 bars is generally considered necessary to cause stomata of grapevines to close (4.1 3 to . The maximum stomatal conductance at Davis was less than at Greenfield. Potassium concentration in the phloem was increased by 30% and 60% after 12 and 24 hour dark periods. Therefore. 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. 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. but wind would reduce vapor pressure deficit and may have been the reason for the reduced stomatal conductance of the non-sheltered vines.I N F L U E N C E OF W I N D B R E A K S . because of its lower molecular weight diffuses 1. The reduced photosynthetic activity of vines grown in the Salinas Valley may also indirectly affect potassium levels in grape berries. D. The sensitivity of assimilation and transpiration rates remains constant when stomatal conductance is varied by changes in temperature and humidity (2). but this may be due to varietal differences (1. LITERATURE CITED 1. H. 33.e. i. Conductance of water vapor and CO2 into the leaf involves identical pathways and diffusion mechanisms. 4. When the stomata of the nonsheltered vines were nearly closed (about 14:00 hr) the leaf water potential remained nearly constant at about .6 times faster than C02. Vol. Cell and Environment 3:46774 (1980). wind appeared to be the dominant factor in the afternoon. 2. since the leaf water potential of the non-sheltered vines was higher (less negative) than the sheltered vines. 1982 . and hence water stress in the vines. vines at Davis continued at a maximum rate for 12 to 13 hours. Schulze. Diurnal changes in leaf water potential of Chardonnay vines grown at Greenfield under sheltered (11) and non-sheltered (El) conditions. at which time their stomata began to close. Effects of light intensity and air humidity. No. In an experiment with Ricinus communis L.1 3 bar threshold level for stomatal closure was never reached. and E.8 bars until around 17:00 hours. as attempts to obtain uncontaminated phloem exodate samples from grapevines have been unsuccessful. DISCUSSION Stomatal conductance for water vapor in leaves (which is what the autoporometer measures) is also an estimation of CO2 assimilation into leaves (5). Dtiring. Smith and Milburn (7) studied phloem loading of potassium and sucrose. Am. Investigations on the influence of changing soil water supply on the photosynthesis intensity and the diffusive resistance of vine leaves. Stomatal response to environment and a possible interrelation between stomatal effects on transpiration and CO2 assimilation. data in Figure 4 reveals that the wind did not cause stomatal closure indirectly through increased transpiration. 2 and 4). Work is in progress to determine the importance of wind movement and light levels on photosynthesis. J. Between 15:00 and 18:00 hours the leaf water potential of the sheltered vines was significantly (P<.
Raschke. B. Stomatal action. Responses of apple leaf stomata to environmental factors. Plant. 5. Ann. Thorpe. E . 4. E. Am.. J. 8. Smart. and J . J. J. Effects of irradiance. Kriedemann.. A . 33. E . A. 25:84-91 (1974). Vitic.236 m I N F L U E N C E O F W I N D B R E A K S 4. Planta 148:28-34 (1980). R. 7. Vol. R. Enol. Smith. Cell and Environment 3:13-22 (1980). Landsberg and M. 6. Aspects of water relations of the grapevine (Vitis vinifera). Photosynthetica 5:6-15 (1971). Enol. Rev. Warrit. No. K. J. temperature and leaf water potential on photosynthesis of vine leaves. and R . Milburn. P . C. 1982 . Osmoregulation and the control of phloem-sap composition in Ricinus communis L. Am. J. 26:30940 (1975). Plant Physiol. Smart. Vitic.
This action might not be possible to undo. Are you sure you want to continue?