Introduction

Grapevines are regularly subjected to environmental constraints, which are mainly soil water availability limitations and supra-optimal temperatures during summer. Even in cool to temperate climates, conditions of water stress may periodically arise during summer months, depending on vineyard topography and rainfall amounts. The pedoclimatic situation (soil type, soil available water reserves, rainfall) and climate evolution both play a major role in the development of edaphic water deficits. Global warming and heat waves, observed over the last few decades, have further accentuated abiotic stress. The grapevine, originating from the Mediterranean region, is traditionally considered a drought-resistant plant (Carbonneau, 1998). Its ability to endure water scarcity comes from its capacity to reach down to great depths, thanks to a large and deep well-developed root system (Bauerle et al., 2008; Barrios-Masias et al., 2015). Furthermore, the vine’s water needs are relatively modest compared with other crops (Pereira et al., 1998). Controlled vine yields and progressive, moderate water deficit are needed to obtain quality wines (Seguin, 1975; van Leeuwen and Seguin, 1994).

Vine sensitivity to water deficit depends on factors that are intrinsic to each site (soil type, mesoclimate) and varies according to genetic (rootstock/scion association) (Spring, 1997; Lovisolo et al., 2016), agronomic (cover crops, leaf-fruit ratio, irrigation) and other factors. Therefore, the environmental constraints imposed by topography, climate and its evolution sometimes require adaptations of management techniques. These adaptations are mainly irrigation to ensure yield and maintain or improve wine quality (Dry and Loveys, 1998; Chaves et al., 2007) in association with other practices (e.g., soil cover, grass management, leaf-fruit ratio; Celette and Gary, 2013).

Numerous studies have shown that limiting vine water availability affects photosynthesis (A) and leaf transpiration (E) (Chaves et al., 2010), plant (Stevens et al., 1995) and root development (Dry et al., 2000b), and leaf and berry mineral nutrition (Gonzalez-Dugo et al., 2010). Stomatal control of leaf transpiration constitutes a physiological drought-avoiding mechanism by optimizing crop water use while preventing embolism events (Lovisolo et al., 2002). High temperatures together with water deficit conditions can lead to leaf fall, in turn leading to a source-sink imbalance, a decrease in the plant canopy’s capacity for photosynthesis, and incomplete ripening of berries due to limited carbohydrate availability (Chaves et al., 2007). It is also recognized that not only the severity of a drought but also the timing and duration influence the final berry size and composition (Deloire et al., 2004). Generally, moderate water stress during the ripening period is favorable for sugar accumulation (van Leeuwen et al., 2009) and increases the anthocyanin and tannin contents in berries (Matthews and Anderson, 1989; Koundouras et al., 2006). In temperate climates, conditions of water deficit are favorable for producing high-quality red wines (van Leeuwen et al., 2009).

Physiological indicators (leaf and stem water potentials, relative carbon isotope composition) that allow direct evaluations of vine water status during the season have been the object of much research (Choné et al., 2001). These indicators were also used in the present study to create different vine water regimes during the growing season by imposing deficit irrigation (DI) from flowering to ripening (irrigated vines with respect to maximum leaf water potential) and water stress without irrigation (non-irrigated vines). The effects of water supply on leaf gas exchange (A, E), water use efficiency (WUE), leaf stomatal conductance, petiole hydraulic conductivity, plant vigor, yield components, and Pinot noir grape and wine quality were analyzed in a field experiment on adult vines (> 10 years) under relatively dry and warm alpine valley conditions (Valais) in Switzerland.

Materials and Methods

Figure 1. Evolution of annual rainfall (A) and mean temperatures (B) recorded from May to September from 1864 to 2015. Data from the meteorological station in Sion, Switzerland (46°13’ N, 7°21’ E; 510 m asl; MétéoSuisse database).

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Results

Plant water status

Seasonal courses of Ψ_PD, measured between 2009 and 2015, highlighted significant differences between irrigated and non-irrigated vines (Figure 2). No water stress was observed in well-watered vines, with Ψ_PD values between -0.05 and -0.25 MPa throughout the season. On the other hand, non-irrigated vines (rain-fed) presented Ψ_PD values that became increasingly negative from the end of July (DOY 210). These values ranged between -0.3 and -0.5 MPa, reflecting moderate water stress from veraison (DOY 220) to the end of the ripening period, except for 2014, when the water deficit was small. Water stress was accentuated even more in non-irrigated vines where the soil was covered by a plastic waterproof sheet. In this case, water stress generally developed from veraison, with Ψ_PD values < -0.5 MPa, especially during the hot, dry summers of 2009, 2011, 2012 and 2015. In contrast, water stress was moderate in 2010, 2013 and 2014 due to cooler temperatures and greater rainfall (Tables 1 and 2), thus occasioning a lower ETP and a less rapid depletion of water soil resources.

Figure 2. Seasonal evolution of predawn leaf water potential (ΨPD) for different irrigation treatments. Arrows indicate irrigation onset. Means +/- SE for eight leaves. Letters: statistically significant at the 5% level of probability. Pinot Noir, Leytron (Switzerland), 2009-2015.

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The Ψ_PD and Ψ_STEM values measured during the veraison-harvest period correlated well (R² = 0.86, P<0.01 and R² = 0.87, P<0.01, respectively) with the relative carbon isotope composition (δ¹³C) of must sugars at harvest (Figure 3). δC¹³ values were between -23 ‰ (high water stress) in non-irrigated vines and -27 ‰ (no water stress) in irrigated vines during the study period. In addition, a correlation was observed between Ψ_PD values and midday Ψ_STEM values (R² = 0.80, P<0.01), measured at solar noon, and a vapor pressure deficit (VPD) between 2.5 and 3.5 kPa during the 7-year study period (results not presented).

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Figure 3. Relationship between predawn leaf water potential (Ψ_PD, A) and midday stem water potential (Ψ_STEM, B) measured during the veraison-harvest period and relative carbon isotope composition (δ¹³C) in must sugars at harvest for different irrigation treatments. Means +/- SE. Pinot noir, Leytron (Switzerland), 2009-2015.

Water and gas exchange relationship

Figure 4 illustrates the impacts of the different water regimes, estimated by Ψ_PD during the night or Ψ_STEM at midday (Figures 4A, B), on gs (Figure 4C), mesophyll resistance (r_m; Figure 4D), net photosynthesis A (Figure 4E) and leaf transpiration E (Figure 4G) measured at midday during the 2009 season. Non-irrigated vines presented lower values of gs, A and E than irrigated vines from the end of July (DOY 210) and particularly in September, with a decrease in gas exchange of nearly 40% compared with irrigated vines. There was a progressive increase in r_m during the season, with an important rise as water stress increased in non-irrigated vines. In water-stressed vines where the soil was covered with an impermeable plastic sheet, major reductions in gas exchange (A, E) and gs were measured from the end of July. Very low levels of gs, A and E were recorded during the grape ripening phase (August-September), generally corresponding to residual conductance g_res (~30 mmol m^-2s^-1) and to rates of photosynthesis and leaf transpiration of approximately 2 µmol CO₂ m^-2s^-1 and 1 mmol H₂O m^-2s^-1, respectively. Gas exchange (A, E) and gs in adult non-senescent leaves, measured at midday during periods covering bloom through to mid-ripening, generally correlated with Ψ_PD values: correlation coefficients (R²) rose to approximately 0.92 (P<0.01) for A, 0.90 (P<0.01) for E and 0.91 (P<0.01) for gs.

Figure 4. Changes in predawn leaf water potential (ΨPD, A), stem water potential (ΨSTEM, B), stomatal conductance (gs, C), mesophyll resistance (rm, D), photosynthetic rate (A, E), intrinsic water use efficiency (WUEi, F), transpiration rate (E, G) and instantaneous water use efficiency (WUEinst, H) during the 2009 season. Means +/- SE for eight leaves. Pinot noir, Leytron (Switzerland).

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WUE_i (A/gs) was found to increase progressively during the growing season (Figure 4F) and was greatest in vines where water deficit was highest (non-irrigated vines with a covered soil) compared to irrigated vines. WUE_inst (A/E), however, tended to decrease during the growing phase (Figure 4H) and was found to be slightly lower in non-irrigated vines, which presented the greatest water deficit at the end of the season. A correlation between WUE_i and δ¹³C in must sugars was noted during the seven years of observation (Figure 5). The increase in WUE_i was associated with that in δ¹³C.

Daily rates of leaf transpiration E (Figure 6B) and stem sap flow (global shoot transpiration; Figure 6C) were greatly dependent on vine water status (Ψ_PD values) and daytime climatic conditions, such as solar radiation, air temperature and relative humidity (Figure 6A). E and sap flow values rose rapidly in the morning as solar radiation was intercepted and air temperatures increased. Nevertheless, a lapse was observed in the early hours of the morning (lag phase) between sap flow values, which remained very low despite light conditions, and leaf transpiration on the external part of the canopy. The maximum sap flow values (0.17 L/h/m²) were recorded during the afternoon, when air temperatures were highest and air humidity was lowest (greatest VPD), in well-watered vines with Ψ_PD of -0.06 MPa. In non-irrigated vines, the maximum E and sap flow values were lower, particularly in the afternoon at the hottest times. During most of the daytime, sap flow rates measured in water-stressed vines (Ψ_PD = -0.55 MPa) represented approximately 30 to 40% of values recorded in well-watered vines.

Figure 5. Relationship between intrinsic water use efficiency (WUEi) and relative carbon isotope composition (δ13C) in must sugars at harvest for different irrigation treatments. Means +/- SE. Pinot Noir, Leytron (Switzerland), 2009-2015.

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Figure 6. Diurnal course of solar radiation (A), mean air temperature (A), relative humidity (A), leaf transpiration rate (B) and sap flow rate (C) through the shoots for different irrigation treatments during the day of 26 July 2009. Means +/- SE of eight leaves (leaf transpiration) and four vines (sap flow rate). Pinot Noir, Leytron (Switzerland).

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Cavitation vulnerability

The specific hydraulic conductivity in petioles (K_petiole) was found to be highest when Ψ_STEM values were above -0.5 MPa (Figure 7A), in the absence of water stress. There was a large decrease in K_petiole as water deficit increased, and notably with Ψ_STEM values less than 1.2 MPa, observed in non-irrigated vines with plastic-covered soils. A cavitation vulnerability curve (Figure 7B) was constructed by plotting PLC in relation to Ψ_STEM. At Ψ_STEM = -1.0 MPa, a PLC of approximately 20% was noted. The 50% PLC threshold was reached when Ψ_STEM was approximately -1.25 MPa, according to the fitted function parameters. Non-irrigated vines with plastic-covered soils showed PLC values > 60% in petioles when Ψ_STEM was less than -1.3 MPa. Irrigated vines presented very low vulnerability to cavitation with Ψ_STEM values > -0.7 MPa. An intermediate situation was observed in non-irrigated vines (rain-fed vines).

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Figure 7. Specific hydraulic conductivity in petioles (K_petiole, A) and vulnerability to cavitation (PLC, percent loss of conductivity, B) versus stem water potential (Ψ_STEM) for grapevine leaf petioles measured in situ for different irrigation treatments. Pinot Noir, Leytron (Switzerland), 2013.

Leaf area and plant vigor (pruning weight)

The seasonal evolution of total leaf area (LA) per vine, in relation to the different irrigation treatments, is presented in Figure 8. An increase in LA through the end of July (~DOY 210) was observed in all three irrigation treatments. After this date (one week before veraison), a progressive decrease was observed initially, followed by a sharp drop in LA in vines subjected to major water stress (non-irrigated vines with plastic-covered soils). In these vines, loss of leaves in the basal zone of shoots, due to drought, largely explains the fall in LA, which went from ~2.5 m²/vine (end of July) to ~1.2 m²/vine (end of September). An intermediate situation was again found in non-irrigated vines where a slight decrease in LA/vine was observed at the very end of the season.

Vine water status also influenced plant vigor, which was estimated from the pruning weight obtained in winter (Table 4). Moderate to high water stress (non-irrigated vines with plastic-covered soils) diminished plant vigor (lower pruning weights), compared with well-watered vines (irrigated vines). On average, over the years of experimentation (2009-2015), plant vigor decreased as vines were subjected to increasing water stress. Compared with irrigated vines, a decrease of nearly 20% in pruning weight was observed in non-irrigated vines with plastic-covered soils, and of 7% in non-irrigated vines (rain-fed).

Figure 8. Seasonal evolution of leaf area per vine for different irrigation treatments. Means +/- SE. Letters: statistically significant at the 5% level of probability. Pinot Noir, Leytron (Switzerland) 2009.

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Table 4. Pruning weight (g/vine) per year and the 2009-2015 mean. Letters: statistically significant at the 5% level of probability. Leytron (Switzerland).

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	Pruning weight (g/vine)							Mean 2009-2015
	2009	2010	2011	2012	2013	2014	2015
Irrigated vines	701a	750a	720a	903a	807a	717a	788a	770a
Non-irrigated vines	722a	735a	592b	763b	819a	671a	700b	715ab
Non-irrigated vines + plastic covered	689a	696a	423c	727b	681b	575b	622b	630b

Phenolic compounds and wine tasting

Analyses of total phenol index (DO 280) and anthocyanins were performed according to Ribéreau-Gayon et al. (1998). Figure 9 illustrates that the wines from non-irrigated vines were distinguished by their deeper colors and were thus richer in anthocyanins (Figure 9A) and phenolic compounds (Figure 9B) than wines from well-watered vines (irrigated vines). The main anthocyanins analyzed in wines were malvidin 3-0- glucoside and peonidin 3-0- glucoside, which represented approximately 80% of the proportion of the five anthocyanidins present in wines (results not presented).

Figure 10 shows the wine tasting results, performed a few weeks after bottling, for the four following key characteristics: bouquet (quality of aroma), wine structure, quality of tannins, and overall wine appreciation. Globally, the wines from non-irrigated vines were more appreciated for their superior tannin quality (smooth and supple), stronger structure and more asserted varietal aromas than wines from irrigated vines. The latter were judged to be of lower quality during 2009-2013, and again in 2015, but in a less pronounced way (no significant difference). In 2014, high summer rainfall and the absence of water stress in all three irrigation treatments resulted in wines with highly similar taste characteristics.

Figure 9. Anthocyanin content (A) and total polyphenol index (PI, B) in wines from different irrigation treatments. Means +/- SE. Letters: statistically significant at the 5% level of probability. Pinot Noir, Leytron (Switzerland) 2009-2015.

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Figure 10. Evaluation of wine quality at tasting according to different sensory variables: olfactive (bouquet), gustatory (structure, tannin quality) and overall impression for the different irrigation treatments. Perception scale from 1 (poor) to 7 (high). Letters: statistically significant at the 5% level of probability. Pinot Noir, Leytron (Switzerland) 2009-2015.

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Discussion

References

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