- Email: [email protected]
Effect of irrigation frequency and water distribution pattern on leaf gas exchange of cv. ‘Syrah’ grown on a clay soil at two levels of water availability
Agricultural Water Management 177 (2016) 410–418
Contents lists available at ScienceDirect
Agricultural Water Management journal homepage: www.elsevier.com/locate/agwat
Effect of irrigation frequency and water distribution pattern on leaf gas exchange of cv. ‘Syrah’ grown on a clay soil at two levels of water availability Bárbara Sebastian a,∗ , José R. Lissarrague a , Luis G. Santesteban b , Rubén Linares a , Pedro Junquera a , Pilar Baeza a a Grupo de Investigación en Viticultura, Departamento de Producción Vegetal: Fitotecnia, Escuela Técnica Superior de Ingenieros Agrónomos, Universidad Politécnica de Madrid, C/Senda del Rey s/n, 28040 Madrid, Spain b Departamento de Producción Agraria, Universidad Pública de Navarra, Campus de Arrosadia, 31006 Pamplona, Spain
a r t i c l e
i n f o
Article history: Received 5 April 2016 Received in revised form 24 August 2016 Accepted 26 August 2016 Keywords: Drip irrigation Leaf water potential Grapevine Vitis vinifera L.
a b s t r a c t The implications of water availability in grapevine physiology have been widely studied before. However, for a given irrigation water amount, the effect of other aspects such as application frequency, or emitter spacing and flow rate (i.e.: distribution pattern) has been scarcely studied, with nearly no previous research on their implications on leaf gas exchange. The aim of this work was to evaluate the physiological response of grapevine to two irrigation frequencies (IrrF, every 2 and 4 days) and two water distribution patterns (DisP, 2 L h−1 emitters every 0.6 m vs. 4 L h−1 emitters every 1.2 m). The experiment was carried out in a cv. Syrah vineyard with a clay soil in central Spain, and the two factors were evaluated under two water availability conditions (low and medium). IrrF and DisP promoted changes in leaf gas exchange. Under low WA conditions, plants irrigated every 4 days had higher average net assimilation than plants irrigated every 2 days. Under medium WA conditions leaf gas exchange depended on the day of measurement with respect to irrigation. Water distribution pattern effect was less evident, but plants with closer emitters performed better under medium WA. The results obtained suggest that variations in irrigation frequency and water availability promote plant acclimation to water deficit conditions, more intense as irrigation dose was lower and as irrigation frequency was higher. © 2016 Elsevier B.V. All rights reserved.
1. Introduction Growers from arid and semi-arid areas are experiencing unprecedented pressure for water resources, likely to increase by the perspective of global warming (Fereres and Evans, 2006; IPCC, 2008). Therefore, the application of irrigation has to be optimized in those areas in order to maximize water use efficiency. Previous research has shown that grapevine yield and quality are greatly limited in the Mediterranean area by water deficit, temperature and evaporative demand during summer (Chaves et al., 2007; Escalona et al., 1999; Santesteban and Royo, 2006). In this context, proper irrigation management is known to play a key role in vineyard sustainability in both economic and ecological terms. In the last decades, several studies have been carried out to understand grapevine gas exchange and carbon assimilation under
∗ Corresponding author. E-mail address: [email protected] (B. Sebastian). http://dx.doi.org/10.1016/j.agwat.2016.08.032 0378-3774/© 2016 Elsevier B.V. All rights reserved.
different irrigation strategies, since the mechanisms that regulate carbon assimilation and partitioning play an important role in the balance between yield and quality (Cifre et al., 2005). Most of those studies focused either on analyzing the changes in leaf gas exchange due to different levels of water availability (Acevedo-Opazo et al., 2010; Medrano et al., 2002; Romero et al., 2010), or on comparing how different genotypes cope with water stress (Bahar et al., 2011; Padgett-Johnson et al., 2003; Schultz, 2003; Vandeleur et al., 2009). Other authors focused on the physiological response to sustained water deficit and subsequent rewatering (Gomez-del-Campo et al., 2007; Hochberg et al., 2013; Pou et al., 2012; Santesteban et al., 2009; Tomas et al., 2014), reporting that water deficit effects are maintained after rewatering, as acclimation mechanisms appear. However, most of the latter studies were performed on pots, but acclimation dynamics are probably more complex under field conditions, as water stress develops much more gradually (Flexas et al., 2006, 2009). In regions such as the Mediterranean area, where rainfall during summer is scarce, irrigation frequency, emitter spacing and flow
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
411
Table 1 Irrigation water amounts applied during the experiment and average leaf water potential at noon. Irrigation frequency
Low WA: 20% ETo Med WA: 40% ETo
Every 2 days (2d) Every 4 days (4d) Every 2 days (2d) Every 4 days (4d)
Irrigation water amount per irrigation event (mm)
1.85–3.70 3.70–7.40 3.70–7.40 7.40–14.80
rate can play an important role regulating water loss and water uptake. Maintaining the water status of the plants under fluctuating water supply can improve water use efficiency through acclimation (Cornic and Massacci, 1996). Irrigation frequency, emitter spacing and flow rate also determine, for a given soil, the wetted volume, which can result on losses of water trough evaporation or deep percolation, thus affecting water use efficiency (Goldberg et al., 1971; Levin et al., 1979; Smasjtrla et al., 1985; Wang et al., 2006). Earlier research has shown that irrigation frequency, emitter spacing and flow rate affect yield and grape composition (Goldberg et al., 1971; Myburgh 2012; Sebastian et al., 2015; Selles et al., 2004;), however, their effect on leaf gas exchange has not yet been studied on grapevine to our knowledge. The aim of this study was to evaluate the effect that irrigation frequency and water distribution pattern (i.e.: emitter spacing and flow rate) has on leaf gas exchange of cv. Syrah grown on a clay soil at two levels of water availability.
Total annual irrigation water amount (mm)
Average noon leaf water potential (MPa)
Fruitset-Veraison
Veraison-Harvest
105
−1.21
−1.55
204
−1.15
−1.43
2. Material and methods 2.1. Vineyard characteristics and experimental design The experiment was carried out in a 6-years old commercial cv. ‘Syrah/SO4 vineyard located in Malpica de Tajo, Toledo, Spain (39◦ 52 N, 4◦ 39 W, 493 m above sea level) during 2006. This region has a Mediterranean climate (P = 450 mm; ETP Penman = 1225 mm), and there are severe water deficits in the summer. Site characteristics and experimental features were explained in more detail in Sebastian et al. (2015), where the agronomical effects of irrigation frequency and water distribution patterns throughout four consecutive seasons in this vineyard were presented. Vineyard soil was classified as Typic Haploxeralf according to the Soil Survey Staff (2013), and the topsoil (0–25 cm) had 38% clay and the underlying soil layer (25–60 cm) had 64% clay. There was a firm soil layer that limited grapevine growth 0.6 m below the soil surface. Soil
Fig. 1. Influence of irrigation frequency (IrrF) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Low water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
412
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 2. Influence of irrigation frequency (IrrF) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Medium water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
bulk density was 1.67 g cm−3 (topsoil) and 1.44 g cm−3 (subsoil). The vineyard was trained as a bilateral cordon, spur-pruned, and shoots were vertically positioned. Row orientation was NW-SE, plant spacing was 2.7 m between rows and 1.2 m within the row. The experimental design was set up independently under two water availability conditions, labelled as low (Low WA, 20% of ETo ) and medium (Med WA, 40% of ETo ) at two adjacent fields. The irrigation doses (Dose) were calculated according to Eq. (1) on a weekly basis Dose =
(IrrCoefxETo − Re) 0, 9
(1)
ETo and Re being, respectively, the reference evapotranspiration and effective rainfall (>7 mm) of the previous week, and IrriCoef the irrigation coefficient defined for each water availability conditions (0.2 for Low WA and 0.4 for Med WA), and 0.9 the correction factor considering the efficiency of the irrigation system. The performance of the irrigation system and the dose applied were checked twice a week with flow meters. Two factors were considered in the experiment, namely irrigation frequency (IrrF) and water distribution pattern (DisP). The IrrF were established by watering every 2 and every 4 days. For DisP, there were two emitter distance and flow rate combinations −that resulted in the same amount of water applied per row meter at each irrigation event- were tested. The combinations were 2 L h−1 drip emitters every 0.6 m, and 4 L h−1 drip emitters every 1.2 m. For the sake of clarity, hereon the two DisP treatments will be referred
solely as 0.6 m and 1.2 m. These irrigation treatments had been applied the three previous seasons to the same plots. The experiment was laid out following a split-plot design, IrrF being the main factor. Experimental plots comprised two rows of 40 grapevines each, with 3 buffer grapevines at each end and a buffer row on each side. Each experimental plot covered 600 m2 . The amount of water applied throughout the season was 105 mm for Low WA treatments and 204 mm for Med WA (Table 1). Irrigation started ten days after fruitset (DOY 159), and stopped shortly after harvest. Both low and medium WA doses resulted in moderate to severe water stress between veraison and harvest (Table 1) according to the scale proposed by Van Leeuwen et al. (2009), and mimic the most commonly used irrigation dose range in this area, limited due to water availability restrictions. 2.2. Experimental measurements 2.2.1. Plant water status Plant water status was determined measuring leaf water potential on well-exposed, mature leaves. At fruitset, before irrigation treatments commenced, measurements were taken at predawn (PD ) and noon (n ). After having initiated irrigation treatments, in order to characterize the irrigation cycles, those measurements were taken one day after irrigation had been applied to both irrigation frequencies, and two days later at 3 phenological stages (pea-size, veraison, and end of ripening). Leaf water potential was determined using a Scholander type pressure chamber (PMS, Port-
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
413
Fig. 3. Influence of water distribution pattern (Disp) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Low water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
land, Oregon, USA), taking into account the considerations given by Turner and Long (1980) and Turner (1988). Briefly, leaf blades were covered with a plastic bag prior to severing the petiole, gas flow was limited to 0.2 bar s−1 and the measurement was performed within the 1-1.5 min after detaching the leaf from the plant. Measurements were carried out on 5 leaves per treatment (2 leaves in replicates 1 and 2, and 1 leaf in replicate 3). 2.2.2. Leaf gas exchange Net CO2 assimilation rate (AN ) and stomatal conductance (gs ) were measured at mid-morning, at noon, and in the afternoon with a portable IRGA system (Li-6200, LI-COR Inc., CITY, USA), on the same leaves used to determine leaf water potential, just before covering them with the plastic bag and severing the petiole. 2.3. Statistical analysis The results obtained for each water availability situation were analyzed separately through a two-way ANOVA. The relationships between leaf gas exchange parameters and leaf water potential and atmospheric conditions were analyzed through linear and curvilinear regression analysis. The equality of those regression equations (slopes and intercepts) were tested using pairwise Student tests following the methods described in Kleinbaum et al. (2008), log transformation being used for exponential functions. All calculations and statistical analyses were performed with R (R Development Core Team, 2016).
3. Results and discussion The effect of IrrF and DisP on daily evolution of leaf water potential and leaf gas exchange parameters was studied through two-way ANOVA. Interaction between factors was non-significant for nearly all dates and times in the day (for >90% of the variabledate-time combinations PIrrF∗DisP > 0.10), so their effect can be assumed to be independent enough to allow the study of each factor separately. Grapevine response to IrrF differed depending on water availability and phenological stage. To illustrate this effect, the daily evolution patterns of leaf water potential, net assimilation and stomatal conductance were compared for the 2- and 4-day irrigation cycles (Figs. 1 and 2). To ease the explanation, the day both 2d and 4d treatments had been irrigated has been labelled as the ‘day 1’ in the irrigation cycles. Under low WA, grapevines showed much reduced leaf gas exchange rates from veraison onwards. Irrespective of the time lag between irrigation and measurement, grapevines irrigated every 2 days tended to have lower leaf water potential, whereas leaf gas exchange parameters showed different behaviour trends depending on the phenological stage and the time elapsed since the last irrigation (Fig. 1). Thus, in general, the day after both treatments (2d, 4d) had been irrigated (‘day 2’), net assimilation and stomatal conductance of 4d grapevines were significantly higher across the whole day. This result is a direct consequence of the fact 4d grapevines had received twice the amount of water than 2d
414
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 4. Influence of water distribution pattern (Disp) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Medium water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
grapevines. On ‘day 4’ (one day after only 2d grapevines had been irrigated), grapevines responded differently depending on the time of the season. At pea-size and veraison, AN and gs of 2d grapevines tended to be higher at midmorning, although differences were not significant and attenuated along the day. However, at the end of ripening, AN and gs of 4d grapevines tended to be higher across the whole day, differences being significant only in the afternoon. Under medium WA (Fig. 2), leaf water potential (leaf ), AN and gs behaved differently depending on the time lag between irrigation and measurement. In general, PD was significantly higher for the treatments that had received more water the day before, the differences being higher on ‘day 4’, when only 2d grapevines had been irrigated the previous day. The AN and gs daily evolution patterns were also different depending on the time elapsed between last irrigation and leaf gas exchange measurement. The greatest statistical differences were found at peasize, when treatments that had received more water the day before showed higher leaf gas exchange (4d grapevines on ‘day 2’ and 2d grapevines on ‘day 4’). At veraison and at the end of ripening, no differences on leaf gas exchange were found on ‘day 2’ despite 4d grapevines had received twice the amount of water than 2d grapevines. On ‘day 4’ when 4d grapevines had been three days without irrigation, 2d grapevines showed higher leaf gas exchange rates. The low values of leaf water potential observed point out that in our experiment, the amount of irrigation water applied was not enough to replace the water extracted by the grapevines and, thus,
all the treatments suffered a sustained water stress. In fact, under Low WA, values of leaf < −1.5 MPa were reached. This value is close to the threshold for severe cavitation described for grapevine (Lovisolo et al., 2008; Salleo and Lo Gullo 1989) and gives an idea of the severity of the water stress imposed. Even for medium WA, stomatal conductance rates were low, but in agreement with those reported by other authors for similar leaf water potential levels (Kriedemann and Smart, 1971; Loveys and Kriedeman, 1973; Tramontini et al., 2013). These low values of stomatal conductance shown even at mid-morning can also be related to soil characteristics, as certain compaction occurred according to the bulk density observed (Dallas and Lewandowski, 2003). Plants growing in relatively compacted soils usually show low stomatal conductance even when soil water availability is high, as water uptake is less efficient (Tardieu 1989; Tardieu and Simonneau, 1998). In our study, leaf water potential daily dynamics may also be reflecting this fact, since its values dropped abruptly between predawn and midmorning, showing that water uptake did not balance water losses through transpiration even when evaporative demand was lower. The effect of water distribution pattern on leaf water potential and gas exchange was much smaller than that of irrigation frequency. In general, not significant differences were found between 0.6 m and 1.2 m grapevines under Low WA (Fig. 3), whereas under Med WA (Fig. 4), significant differences were more frequently found, pointing out that closer emitter spacing resulted in a slightly better water status and in higher AN and gs .
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
415
Fig. 5. Relationships between net assimilation and leaf water potential (a), stomatal conductance and leaf water potential (b), net assimilation and VPD (c) and stomatal conductance and VPD (d) under Low water availability conditions. Fitted lines equations for the relationships are presented. Slope and intercept values of log transformed equations followed by different letters correspond with significant differences at p < 0.10 according to pairwise Student tests.
In general terms, the differences in gas exchange parameters that were observed across the experiment are not as substantial as those reported by other researchers when comparing water availability levels (De Souza et al., 2005; Romero et al., 2010), which agrees with the fact the effect of IrrF and DisP on plant water status was relatively small. The fact that the effects observed were small agrees with other authors who found that in severe water stress conditions the effects of irrigation treatments in grapevine physiological performance are attenuated (Flexas et al., 1998), and that the effect of water distribution pattern can be reduced in the case of fine textured soils (Chaves et al., 2007; Marsal et al., 2008). However, the degree to which IrrF has been shown to affect AN and gs under Low
WA was enough to affect yield significantly (Sebastian et al., 2015). The analysis of the diurnal evolution of AN and gs (Figs. 1 and 2) allowed us to explain this behaviour: on ‘Day 2’, 4d grapevines were photosynthetically more active than 2d grapevines, since they had received twice the amount of water the previous day and, although on ‘Day 4’ 2d grapevines were expected to be more active than 4d grapevines that had not been irrigated for three days, differences in gas exchange rates appeared seldom, their magnitude was smaller, and did not balance those observed on ‘Day 2’. The different daily evolution patterns of leaf gas exchange observed depending on the amount of available water and the time elapsed since irrigation could be explained because, addition-
416
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 6. Relationships between net assimilation and leaf water potential (a), stomatal conductance and leaf water potential (b), net assimilation and VPD (c) and stomatal conductance and VPD (d) under Medium water availability conditions. Fitted lines equations for the relationships are presented. Slope and intercept values of log transformed equations followed by different letters correspond with significant differences at p < 0.10 according to pairwise Student tests.
ally to the effects IrrF may have on irrigation efficiency, the fact some grapevines received irrigation water every 2 days whereas others did every 4 days results in a different dynamic of stressrecovering that can result in a different degree of acclimation (Flexas et al., 2006, 2009; Pou et al., 2012). In order to evaluate to which extent differences in irrigation frequency caused acclimation, leaf gas exchange data measured along the day were plotted against water status and evaporative demand (Figs. 5 and 6). Relationships were fitted to exponential functions and log transformed into linear models to be compared statistically. The AN and gs were shown to be related to plant water status and vapour pressure deficit (Figs. 5 and 6), showing similar trends: a
decline of gs and AN for decreasing leaf and increasing VPD, which agrees earlier literature (Chaves et al., 2010; Lovisolo et al., 2010; Rogiers et al., 2012; Tomas et al., 2014). As expected, the differences between the situations considered were bigger when comparing leaf gas exchange parameters against VPD than against leaf water potential, which is logical as in the latter responses at the same level of water status are compared. When the effect of the WA and the time elapsed since the last irrigation was compared (Figs. 5 and 6), water availability conditions were observed to influence the impact of irrigation frequency on leaf gas exchange response to changes in VPD and leaf water potential. Under low WA (Fig. 5), gas exchange parameters of 2d
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
grapevines showed a steeper decline in response to increments in VPD and to reductions of leaf than 4d grapevines (higher slope), and hence, presented the lowest AN and gs values under high evaporative demands and low leaf water potentials, i.e.: under the most frequent conditions during the experiment. 4d grapevines showed the same response of AN and gs to changes in leaf and VPD irrespective of the time elapsed since irrigation. Significant differences between fitted lines only appeared when comparing 2d vs 4d for VPD, although for leaf water potential fitted lines followed the same pattern. Under medium WA (Fig. 6), 4d grapevines responded differently to changes in leaf and VPD depending on the time elapsed since irrigation. Thus, on ‘day 2’ of the irrigation cycle, AN and gs response to changes in VPD and leaf of 4d grapevines was the mildest, behaving as quite “optimistic” (Jones et al., 1980), maintaining high levels of leaf gas exchange at low values of leaf . However, on ‘day 4’, these grapevines, that had not been irrigated for three days, showed a steep decline of AN and gs to decreasing leaf and increasing VPD. Grapevines irrigated every 2 days under medium WA showed an intermediate response. The relationships between leaf gas exchange parameters, leaf water potential and the atmospheric demand showed that low WA treatments presented tighter stomata control than medium WA treatments. The fact that 4d grapevines under low WA conditions had a very similar response of leaf gas exchange to changes in leaf water potential irrespective the time elapsed since last irrigation suggests that some kind of acclimation occurred. This phenomenon would be the result of the interaction between hydraulic and rootmediated hormonal information (Lovisolo et al., 2010; Soar et al., 2006; Tardieu and Simonneau, 1998; Vandeleur et al., 2009). Under medium WA, grapevines irrigated every 4 days appeared not to acclimate, as after three days without irrigation when climatic conditions worsened they decreased leaf gas exchange. Thus, the grapevines irrigated every 4 days with the higher irrigation dose (medium WA) seemed to be less protected against drought stress than the grapevines irrigated with the lower irrigation dose. This could explain their physiological behaviour at the end of ripening, showing very low stomatal conductance after being three days without irrigation, even lower than 4d grapevines under low WA at the same stage (Fig. 2).
4. Conclusions Irrigation frequency and water distribution pattern affected leaf gas exchange of cv. Syrah grown in a clay soil, the effect of irrigation frequency being much more relevant than that of water distribution pattern. Irrigation intervals of two days or shorter should be avoided in clay soils under low water availability conditions, as they result in an irrigation efficiency loss. However, when water availability is higher, for the irrigation frequencies tested, this effect fades away, which makes that under these conditions, frequency could be chosen considering other management issues such as water scheduling, design, operational costs, etc. Water distribution pattern, a nearly unstudied factor, appeared not to be a very critical issue under the conditions of this study, although the closer spacing seemed to be slightly more efficient, especially for medium water availability conditions. Irrigation frequency clearly affected the relationship between leaf gas exchange parameters, plant water status and atmospheric conditions, and together with the amount of water applied had implications in the development of acclimation mechanisms that affected plant physiological response, thus affecting irrigation efficiency. Therefore, more importance should be given to this factor when implementing an irrigation strategy.
417
Acknowledgements The authors would like to thank Osborne Malpica Winery, for allowing us to carry out this research on their vineyards, and for their financial support of the study. The authors would also like to thank the staff in the Viticulture Research Group of the Polytechnic University of Madrid for their support in taking measurements.
References Acevedo-Opazo, C., Ortega-Farias, S., Fuentes, S., 2010. Effects of grapevine (Vitis vinifera L.) water status on water consumption, vegetative growth and grape quality: an irrigation scheduling application to achieve regulated deficit irrigation. Agric. Water Manage. 97, 956–964. Bahar, E., Carbonneau, A., Korkutal, I., 2011. The effect of extreme water stress on leaf drying limits and possibilities of recovering in three grapevine (Vitis vinifera L.) cultivars. Afr. J. Agric. Res. 6 (5), 1151–1160. ˜ M.F., Rodrigues, M.L., Lopes, M., Chaves, M.M., Santos, T.P., Souza, C.R., Ortuno, Maroco, J.P., Pereira, J.S., 2007. Deficit irrigation in grapevine improves water-use efficiency while controlling vigour and production quality. Ann. Appl. Biol. 150, 237–252, http://dx.doi.org/10.1111/j.1744-7348.2006.00123.x. Chaves, M.M., Zarrouk, O., Francisco, R., Costa, J.M., Santos, T., Regalado, A.P., Rodrigues, L., Lopes, C.M., 2010. Grapevine under deficit irrigation: hints from physiological and molecular data. Ann. Bot. 105, 661–676. Cifre, J., Bota, J., Escalona, J.M., Medrano, H., Flexas, J., 2005. Physiological tools for irrigation scheduling in grapevine (Vitis vinifera L.). An open gate to improve water-use efficiency? Agric. Ecosyst. Environ. 106, 159–170. Cornic, G., Massacci, A., 1996. Leaf photosynthesis under drought stress. In: Photosynthesis and the Environment. Springer, Netherlands, pp. 347–366. Dallas, H., Lewandowski, A., 2003. Protecting Urban Soil Quality: Examples for Landscape Codes and Specifications. USDA Natural Resources Conservation Services. De Souza, C.R., Maroco, J.P., dos Santos, T.P., Rodrigues, M.L., Lopes, C., Pereira, J.S., Chaves, M.M., 2005. Control of stomatal aperture and carbon uptake by déficit irrigation in two grapevine cultivars. Agric. Ecosyst. Environ. 106, 261–274. Escalona, J.M., Flexas, J., Medrano, H., 1999. Stomatal and non-stomatal limitations of photosynthesis under water stress in field-grown grapevines. Aust. J. Plant Physiol. 26, 421–433. Fereres, E., Evans, R.E., 2006. Irrigation of fruit trees and vines: an introduction. Irrig. Sci. 24, 55–57. Flexas, J., Escalona, J.M., Medrano, H., 1998. Down-regulation of photosynthesis by drought under field conditions in grapevine leaves. Aust. J. Plant Physiol. 25, 893–900. Flexas, J., Ribas-Carbo, M., Hanson, D.T., Bota, J., Otto, B., Cifre, J., McDowell, N., Medrano, H., Kaldenhoff, R., 2006. Tobacco aquaporin NtAQP1 is involved in mesophyll conductance to CO2 in vivo. Plant J. 48, 427–439. Flexas, J., Barón, M., Bota, J., Ducruet, J.M., Gallé, A., Galmés, J., Jiménez, M., Pou, A., Ribas-Carbó, M., Sajnani, C., Tomàs, M., Medrano, M., 2009. Photosynthesis limitations during water stress acclimation and recovery in the drought adapted Vitis hybrid Richter-110 (V. berlandieri × V. rupestris). J. Exp. Bot. 60, 2361–2377. Goldberg, S.D., Gornat, B., Bar, Y., 1971. The distribution of roots, water and minerals as a result of trickle irrigation. J. Am. Soc. Hortic. Sci. 96 (51), 645–648. Gomez-del-Campo, M., Baeza, P., Ruiz, C., Sotes, V., Lissarrague, J.R., 2007. Effect of previous water conditions on vine response to rewatering. Vitis 46 (2), 51–55. Hochberg, U., Degu, A., Fait, A., Rachmilevitch, S., 2013. Near isohydric grapevine cultivar displays higher photosynthetic efficiency and photorespiration rates under drought stress as compared with near anisohydric grapevine cultivar. Physiol. Plant. 147 (4), 443–452. IPCC, 2008. Climate change and water. In: B.C. Bates, B.C., Kundzewicz, Z.W., Wu, S., Palutikof, J.P. (Eds.), IPCC Tech. Paper VI. IPCC Secretariat, Geneva. Jones, H.G., Turner, N.C., Kramer, P.J., 1980. Interaction and integration of adaptive responses to water stress: the implications of an unpredictable environment. Adaptation of plants to water and high temperature stress, 353–365. Kleinbaum, D.G., Kupper, L., Muller, K.E., Nizam, A., 2008. Applied Regression Analysis and Other Multivariable Methods, 4th ed. Duxbury Applied, Thomsom Co, Belmont, USA. Kriedemann, P.E., Smart, R.E., 1971. Effects of irradiance, temperature, and leaf water potential on photosynthesis of vine leaves. Photosynthetica 5, 15–19. Levin, I., Van Pooyen, P., Van Rooyen, F., 1979. The effect of discharge rate and intermittent water application by point source irrigation on the soil moisture distribution pattern. Soil Sci. Soc. Am. J. 43, 8–16. Loveys, B.R., Kriedeman, P.E., 1973. Rapid changes in abscisic acid-like inhibitors following alterations in vine leaf water potential. Physiol. Plant. 28, 476–479. Lovisolo, C., Perrone, I., Hartung, W., Schubert, A., 2008. An abscisic acid-related reduced transpiration promotes gradual embolism repair when grapevines are rehydrated after drought. New Phytol. 180 (3), 642–651. Lovisolo, C., Perrone, I., Carra, A., Ferrandino, A., Flexas, J., Medrano, H., Schubert, A., 2010. Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Funct. Plant Biol. 37, 98–116.
418
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Marsal, J., Mata, M., Del Campo, J., Arbones, A., Vallverdú, X., Girona, J., Olivo, N., 2008. Evaluation of partial root-zone drying for potential field use as a deficit irrigation technique in commercial vineyards according to two different pipeline layouts. Irrig. Sci. 26 (4), 347–356. Medrano, H., Escalona, J.M., Bota, J., Gulías, J., Flexas, J., 2002. Regulation of photosynthesis of C3 plants in response to progressive drought: stomatal conductance as a reference parameter. Ann. Bot. 89, 895–905, http://dx.doi. org/10.1093/aob/mcf079. Myburgh, P.A., 2012. Comparing irrigation systems and strategies for table grapes in the weathered granite-gneiss soils of the Lower Orange River Region. S. Afr. J. Enol. Vitic. 33, 184–197. Padgett-Johnson, M., Williams, L.E., Walker, M.A., 2003. Vine water relations, gas exchange, and vegetative growth of seventeen Vitis species grown under irrigated and non-irrigated conditions in California. J. Am. Soc. Hortic. Sci. 128, 269–276. Pou, A., Medrano, H., Tomás, M., Martorell, S., Ribas-Carbó, M., Flexas, J., 2012. An anisohydric grapevine variety performs better under moderate water stress and recovery than isohydric varieties. Plant Soil 359, 335–349, http://dx.doi. org/10.1007/s11104-012-1206-7. R Core Team, 2016. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL http://www.R-project.org/. Rogiers, S.Y., Greer, D.H., Hatfield, J.M., Hutton, R.J., Clarke, S.J., Hutchinson, P.A., Somers, A., 2012. Stomatal response of an anisohydric grapevine cultivar to evaporative demand, available soil moisture and abscisic acid. Tree Physiol. 32 (3), 249–261. Romero, P., Fernández-Fernández, J.I., Cutillas, A., 2010. Physiological thresholds for efficient regulated deficit-irrigation management in winegrapes grown under semiarid conditions. Am. J. Enol. Vitic. 61, 300–312. Salleo, S., Lo Gullo, M.A., 1989. Xylem cavitation in nodes and internodes of Vitis vinifera L. plants subjected to water stress. Limits of restoration of water conduction in cavitated xylem conduits. In: Kreeb, K.H., Richter, H., Hinckley, T.M. (Eds.), Structural and Functional Responses to Environmental Stresses. SPB Academic Publishing, The Hague,The Netherlands, pp. 33–42. Santesteban, L.G., Royo, J.B., 2006. Water status, leaf area and fruit load influence on berry weight and sugar accumulation of cv.‘Tempranillo’ under semiarid conditions. Sci. Hortic. 109 (1), 60–65. Santesteban, L.G., Miranda, C., Royo, J.B., 2009. Effect of water deficit and rewatering on leaf gas exchange and transpiration decline of excised leaves of four grapevine (Vitis vinifera L.) cultivars. Sci. Hortic. 121 (4), 434–439. Schultz, H.R., 2003. Differences in hydraulic architecture account for near isohydric and anisohydric behaviours of two field-grown Vitis vinifera L. cultivars during drought. Plant Cell Environ. 26, 1393–1405, http://dx.doi.org/10.1046/j.13653040.2003.01064.x. Sebastian, B., Baeza, P., Santesteban, L.G., Sanchez de Miguel, P., De La Fuente, M., Lissarrague, J.R., 2015. Response of grapevine cv Syrah to irrigation frequency
and water distribution pattern in a clay soil. Agric. Water Manage. 148, 269–279. Selles, G., Ferreyra, R., Contreras, G., Ahumada, R., Valenzuela, J., Bravo, R., 2004. Effect of three irrigation frequencies applied by drip irrigation over table grapes (Vitis vinifera L. cv Thompson Seedless) located in the Aconcagua Valley (Chile). Acta Hortic. 646, 175–181. Smasjtrla, A.G., Harrison, D.S., Clark, G.A., 1985. Trickle Irrigation Scheduling I. Duration of Water Application IFAS Bulletin 204. University of Florida, Gainesville. Soar, C.J., Dry, P.R., Loveys, B.R., 2006. Scion photosynthesis and leaf gas exchange in Vitis vinifera L. cv. Shiraz: mediation of rootstock effect via xylem sap ABA. Aust. J. Grape Wine Res. 12, 82–96. Soil Survey Staff, 2013. Simplified Guide to Soil Taxonomy. USDA, Natural Resources Conservation Service. National Soil Survey Center, Lincoln, NE http://www.nrcs.usda.gov/wps/portal/nrcs/detail/soils/survey/class/ ?cid=nrcs142p2 053580. Tardieu, F., Simonneau, T., 1998. Variability among species of stomatal control under fluctuating soil water status and evaporative demand: modelling isohydric and anisohydric behaviours. J. Exp. Bot. 49 (Secial Issue), 419–432. Tardieu, F., 1989. Root system responses to soil structural properties: micro-and macro-Scale. In: Mechanics and Related Processes in Structured Agricultural Soils. Springer, Netherlands, pp. 153–172. Tomas, M., Medrano, H., Brugnoli, E., Escalona, J.M., Martorell, S., Pou, A., Ribas-Carbó, M., Flexas, J., 2014. Variability of mesophyll conductance in grapevine cultivars under water stress conditions in relation to leaf anatomy and water use efficiency. Aust. J. Grape Wine Res. 20, 272–280. Tramontini, S., Van Leeuwen, C., Domec, J.-C., Destrac-Irvine, A., Basteau, C., Vitali, M., Mosbach-Schulz, O., Lovisolo, C., 2013. Impact of soil texture and water availability on the hydraulic control of plant and grape-berry development. Plant Soil 368, 215–230. Turner, N.C., Long, M.J., 1980. Errors arising from rapid water loss in the measurements of leaf water potential by the pressure chamber technique. Aust. J. Plant Physiol. 7, 527–537. Turner, N.C., 1988. Measurement of plant water status by the pressure chamber technique. Irrig. Sci. 9, 289–308. Van Leeuwen, C., Tregoat, O., Chone, X., Bois, B., Pernet, D., Gaudillere, J.P., 2009. Vine water status is a key factor in grape ripening and vintage quality for red Bordeaux wine. How can it be assessed for vineyard management purposes? J. Int. Sci. Vigne Vin 43, 121–134. Vandeleur, R.K., Mayo, G., Shelden, M.C., Gilliham, M., Kaiser, B.N., Tyerman, S.D., 2009. The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine. Plant Physiol. 149, 445–460. Wang, F.X., Kang, Y., Liu, S.P., 2006. Effects of drip irrigation frequency on soil wetting pattern and potato growth in North China Plain. Agric. Water Manage. 79 (3), 248–264.
Contents lists available at ScienceDirect
Agricultural Water Management journal homepage: www.elsevier.com/locate/agwat
Effect of irrigation frequency and water distribution pattern on leaf gas exchange of cv. ‘Syrah’ grown on a clay soil at two levels of water availability Bárbara Sebastian a,∗ , José R. Lissarrague a , Luis G. Santesteban b , Rubén Linares a , Pedro Junquera a , Pilar Baeza a a Grupo de Investigación en Viticultura, Departamento de Producción Vegetal: Fitotecnia, Escuela Técnica Superior de Ingenieros Agrónomos, Universidad Politécnica de Madrid, C/Senda del Rey s/n, 28040 Madrid, Spain b Departamento de Producción Agraria, Universidad Pública de Navarra, Campus de Arrosadia, 31006 Pamplona, Spain
a r t i c l e
i n f o
Article history: Received 5 April 2016 Received in revised form 24 August 2016 Accepted 26 August 2016 Keywords: Drip irrigation Leaf water potential Grapevine Vitis vinifera L.
a b s t r a c t The implications of water availability in grapevine physiology have been widely studied before. However, for a given irrigation water amount, the effect of other aspects such as application frequency, or emitter spacing and flow rate (i.e.: distribution pattern) has been scarcely studied, with nearly no previous research on their implications on leaf gas exchange. The aim of this work was to evaluate the physiological response of grapevine to two irrigation frequencies (IrrF, every 2 and 4 days) and two water distribution patterns (DisP, 2 L h−1 emitters every 0.6 m vs. 4 L h−1 emitters every 1.2 m). The experiment was carried out in a cv. Syrah vineyard with a clay soil in central Spain, and the two factors were evaluated under two water availability conditions (low and medium). IrrF and DisP promoted changes in leaf gas exchange. Under low WA conditions, plants irrigated every 4 days had higher average net assimilation than plants irrigated every 2 days. Under medium WA conditions leaf gas exchange depended on the day of measurement with respect to irrigation. Water distribution pattern effect was less evident, but plants with closer emitters performed better under medium WA. The results obtained suggest that variations in irrigation frequency and water availability promote plant acclimation to water deficit conditions, more intense as irrigation dose was lower and as irrigation frequency was higher. © 2016 Elsevier B.V. All rights reserved.
1. Introduction Growers from arid and semi-arid areas are experiencing unprecedented pressure for water resources, likely to increase by the perspective of global warming (Fereres and Evans, 2006; IPCC, 2008). Therefore, the application of irrigation has to be optimized in those areas in order to maximize water use efficiency. Previous research has shown that grapevine yield and quality are greatly limited in the Mediterranean area by water deficit, temperature and evaporative demand during summer (Chaves et al., 2007; Escalona et al., 1999; Santesteban and Royo, 2006). In this context, proper irrigation management is known to play a key role in vineyard sustainability in both economic and ecological terms. In the last decades, several studies have been carried out to understand grapevine gas exchange and carbon assimilation under
∗ Corresponding author. E-mail address: [email protected] (B. Sebastian). http://dx.doi.org/10.1016/j.agwat.2016.08.032 0378-3774/© 2016 Elsevier B.V. All rights reserved.
different irrigation strategies, since the mechanisms that regulate carbon assimilation and partitioning play an important role in the balance between yield and quality (Cifre et al., 2005). Most of those studies focused either on analyzing the changes in leaf gas exchange due to different levels of water availability (Acevedo-Opazo et al., 2010; Medrano et al., 2002; Romero et al., 2010), or on comparing how different genotypes cope with water stress (Bahar et al., 2011; Padgett-Johnson et al., 2003; Schultz, 2003; Vandeleur et al., 2009). Other authors focused on the physiological response to sustained water deficit and subsequent rewatering (Gomez-del-Campo et al., 2007; Hochberg et al., 2013; Pou et al., 2012; Santesteban et al., 2009; Tomas et al., 2014), reporting that water deficit effects are maintained after rewatering, as acclimation mechanisms appear. However, most of the latter studies were performed on pots, but acclimation dynamics are probably more complex under field conditions, as water stress develops much more gradually (Flexas et al., 2006, 2009). In regions such as the Mediterranean area, where rainfall during summer is scarce, irrigation frequency, emitter spacing and flow
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
411
Table 1 Irrigation water amounts applied during the experiment and average leaf water potential at noon. Irrigation frequency
Low WA: 20% ETo Med WA: 40% ETo
Every 2 days (2d) Every 4 days (4d) Every 2 days (2d) Every 4 days (4d)
Irrigation water amount per irrigation event (mm)
1.85–3.70 3.70–7.40 3.70–7.40 7.40–14.80
rate can play an important role regulating water loss and water uptake. Maintaining the water status of the plants under fluctuating water supply can improve water use efficiency through acclimation (Cornic and Massacci, 1996). Irrigation frequency, emitter spacing and flow rate also determine, for a given soil, the wetted volume, which can result on losses of water trough evaporation or deep percolation, thus affecting water use efficiency (Goldberg et al., 1971; Levin et al., 1979; Smasjtrla et al., 1985; Wang et al., 2006). Earlier research has shown that irrigation frequency, emitter spacing and flow rate affect yield and grape composition (Goldberg et al., 1971; Myburgh 2012; Sebastian et al., 2015; Selles et al., 2004;), however, their effect on leaf gas exchange has not yet been studied on grapevine to our knowledge. The aim of this study was to evaluate the effect that irrigation frequency and water distribution pattern (i.e.: emitter spacing and flow rate) has on leaf gas exchange of cv. Syrah grown on a clay soil at two levels of water availability.
Total annual irrigation water amount (mm)
Average noon leaf water potential (MPa)
Fruitset-Veraison
Veraison-Harvest
105
−1.21
−1.55
204
−1.15
−1.43
2. Material and methods 2.1. Vineyard characteristics and experimental design The experiment was carried out in a 6-years old commercial cv. ‘Syrah/SO4 vineyard located in Malpica de Tajo, Toledo, Spain (39◦ 52 N, 4◦ 39 W, 493 m above sea level) during 2006. This region has a Mediterranean climate (P = 450 mm; ETP Penman = 1225 mm), and there are severe water deficits in the summer. Site characteristics and experimental features were explained in more detail in Sebastian et al. (2015), where the agronomical effects of irrigation frequency and water distribution patterns throughout four consecutive seasons in this vineyard were presented. Vineyard soil was classified as Typic Haploxeralf according to the Soil Survey Staff (2013), and the topsoil (0–25 cm) had 38% clay and the underlying soil layer (25–60 cm) had 64% clay. There was a firm soil layer that limited grapevine growth 0.6 m below the soil surface. Soil
Fig. 1. Influence of irrigation frequency (IrrF) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Low water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
412
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 2. Influence of irrigation frequency (IrrF) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Medium water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
bulk density was 1.67 g cm−3 (topsoil) and 1.44 g cm−3 (subsoil). The vineyard was trained as a bilateral cordon, spur-pruned, and shoots were vertically positioned. Row orientation was NW-SE, plant spacing was 2.7 m between rows and 1.2 m within the row. The experimental design was set up independently under two water availability conditions, labelled as low (Low WA, 20% of ETo ) and medium (Med WA, 40% of ETo ) at two adjacent fields. The irrigation doses (Dose) were calculated according to Eq. (1) on a weekly basis Dose =
(IrrCoefxETo − Re) 0, 9
(1)
ETo and Re being, respectively, the reference evapotranspiration and effective rainfall (>7 mm) of the previous week, and IrriCoef the irrigation coefficient defined for each water availability conditions (0.2 for Low WA and 0.4 for Med WA), and 0.9 the correction factor considering the efficiency of the irrigation system. The performance of the irrigation system and the dose applied were checked twice a week with flow meters. Two factors were considered in the experiment, namely irrigation frequency (IrrF) and water distribution pattern (DisP). The IrrF were established by watering every 2 and every 4 days. For DisP, there were two emitter distance and flow rate combinations −that resulted in the same amount of water applied per row meter at each irrigation event- were tested. The combinations were 2 L h−1 drip emitters every 0.6 m, and 4 L h−1 drip emitters every 1.2 m. For the sake of clarity, hereon the two DisP treatments will be referred
solely as 0.6 m and 1.2 m. These irrigation treatments had been applied the three previous seasons to the same plots. The experiment was laid out following a split-plot design, IrrF being the main factor. Experimental plots comprised two rows of 40 grapevines each, with 3 buffer grapevines at each end and a buffer row on each side. Each experimental plot covered 600 m2 . The amount of water applied throughout the season was 105 mm for Low WA treatments and 204 mm for Med WA (Table 1). Irrigation started ten days after fruitset (DOY 159), and stopped shortly after harvest. Both low and medium WA doses resulted in moderate to severe water stress between veraison and harvest (Table 1) according to the scale proposed by Van Leeuwen et al. (2009), and mimic the most commonly used irrigation dose range in this area, limited due to water availability restrictions. 2.2. Experimental measurements 2.2.1. Plant water status Plant water status was determined measuring leaf water potential on well-exposed, mature leaves. At fruitset, before irrigation treatments commenced, measurements were taken at predawn (PD ) and noon (n ). After having initiated irrigation treatments, in order to characterize the irrigation cycles, those measurements were taken one day after irrigation had been applied to both irrigation frequencies, and two days later at 3 phenological stages (pea-size, veraison, and end of ripening). Leaf water potential was determined using a Scholander type pressure chamber (PMS, Port-
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
413
Fig. 3. Influence of water distribution pattern (Disp) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Low water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
land, Oregon, USA), taking into account the considerations given by Turner and Long (1980) and Turner (1988). Briefly, leaf blades were covered with a plastic bag prior to severing the petiole, gas flow was limited to 0.2 bar s−1 and the measurement was performed within the 1-1.5 min after detaching the leaf from the plant. Measurements were carried out on 5 leaves per treatment (2 leaves in replicates 1 and 2, and 1 leaf in replicate 3). 2.2.2. Leaf gas exchange Net CO2 assimilation rate (AN ) and stomatal conductance (gs ) were measured at mid-morning, at noon, and in the afternoon with a portable IRGA system (Li-6200, LI-COR Inc., CITY, USA), on the same leaves used to determine leaf water potential, just before covering them with the plastic bag and severing the petiole. 2.3. Statistical analysis The results obtained for each water availability situation were analyzed separately through a two-way ANOVA. The relationships between leaf gas exchange parameters and leaf water potential and atmospheric conditions were analyzed through linear and curvilinear regression analysis. The equality of those regression equations (slopes and intercepts) were tested using pairwise Student tests following the methods described in Kleinbaum et al. (2008), log transformation being used for exponential functions. All calculations and statistical analyses were performed with R (R Development Core Team, 2016).
3. Results and discussion The effect of IrrF and DisP on daily evolution of leaf water potential and leaf gas exchange parameters was studied through two-way ANOVA. Interaction between factors was non-significant for nearly all dates and times in the day (for >90% of the variabledate-time combinations PIrrF∗DisP > 0.10), so their effect can be assumed to be independent enough to allow the study of each factor separately. Grapevine response to IrrF differed depending on water availability and phenological stage. To illustrate this effect, the daily evolution patterns of leaf water potential, net assimilation and stomatal conductance were compared for the 2- and 4-day irrigation cycles (Figs. 1 and 2). To ease the explanation, the day both 2d and 4d treatments had been irrigated has been labelled as the ‘day 1’ in the irrigation cycles. Under low WA, grapevines showed much reduced leaf gas exchange rates from veraison onwards. Irrespective of the time lag between irrigation and measurement, grapevines irrigated every 2 days tended to have lower leaf water potential, whereas leaf gas exchange parameters showed different behaviour trends depending on the phenological stage and the time elapsed since the last irrigation (Fig. 1). Thus, in general, the day after both treatments (2d, 4d) had been irrigated (‘day 2’), net assimilation and stomatal conductance of 4d grapevines were significantly higher across the whole day. This result is a direct consequence of the fact 4d grapevines had received twice the amount of water than 2d
414
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 4. Influence of water distribution pattern (Disp) on daily leaf water potential (), net assimilation (AN ) and stomatal conductance (gs ) evolution under the 2- and 4-day irrigation cycles for Medium water availability conditions at Pea-size (a, d, g), veraison (b, e, h), and end of ripening (c, f, i). Irrigation events for 2d- and 4d- treatments are indicated by black and white arrows, respectively. Daily VPD evolution is also shown as a grey line (a, b, c). Phenological stages dates: Peazize: 5/07/2006, Veraison: 27/07/2006; End of Ripening: 29/08/2006 (dd/m/yy). Statistical significance of differences between treatments are given as * (p < 0.1), ** (p < 0.05) and *** (p < 0.01) and ns (not significant).
grapevines. On ‘day 4’ (one day after only 2d grapevines had been irrigated), grapevines responded differently depending on the time of the season. At pea-size and veraison, AN and gs of 2d grapevines tended to be higher at midmorning, although differences were not significant and attenuated along the day. However, at the end of ripening, AN and gs of 4d grapevines tended to be higher across the whole day, differences being significant only in the afternoon. Under medium WA (Fig. 2), leaf water potential (leaf ), AN and gs behaved differently depending on the time lag between irrigation and measurement. In general, PD was significantly higher for the treatments that had received more water the day before, the differences being higher on ‘day 4’, when only 2d grapevines had been irrigated the previous day. The AN and gs daily evolution patterns were also different depending on the time elapsed between last irrigation and leaf gas exchange measurement. The greatest statistical differences were found at peasize, when treatments that had received more water the day before showed higher leaf gas exchange (4d grapevines on ‘day 2’ and 2d grapevines on ‘day 4’). At veraison and at the end of ripening, no differences on leaf gas exchange were found on ‘day 2’ despite 4d grapevines had received twice the amount of water than 2d grapevines. On ‘day 4’ when 4d grapevines had been three days without irrigation, 2d grapevines showed higher leaf gas exchange rates. The low values of leaf water potential observed point out that in our experiment, the amount of irrigation water applied was not enough to replace the water extracted by the grapevines and, thus,
all the treatments suffered a sustained water stress. In fact, under Low WA, values of leaf < −1.5 MPa were reached. This value is close to the threshold for severe cavitation described for grapevine (Lovisolo et al., 2008; Salleo and Lo Gullo 1989) and gives an idea of the severity of the water stress imposed. Even for medium WA, stomatal conductance rates were low, but in agreement with those reported by other authors for similar leaf water potential levels (Kriedemann and Smart, 1971; Loveys and Kriedeman, 1973; Tramontini et al., 2013). These low values of stomatal conductance shown even at mid-morning can also be related to soil characteristics, as certain compaction occurred according to the bulk density observed (Dallas and Lewandowski, 2003). Plants growing in relatively compacted soils usually show low stomatal conductance even when soil water availability is high, as water uptake is less efficient (Tardieu 1989; Tardieu and Simonneau, 1998). In our study, leaf water potential daily dynamics may also be reflecting this fact, since its values dropped abruptly between predawn and midmorning, showing that water uptake did not balance water losses through transpiration even when evaporative demand was lower. The effect of water distribution pattern on leaf water potential and gas exchange was much smaller than that of irrigation frequency. In general, not significant differences were found between 0.6 m and 1.2 m grapevines under Low WA (Fig. 3), whereas under Med WA (Fig. 4), significant differences were more frequently found, pointing out that closer emitter spacing resulted in a slightly better water status and in higher AN and gs .
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
415
Fig. 5. Relationships between net assimilation and leaf water potential (a), stomatal conductance and leaf water potential (b), net assimilation and VPD (c) and stomatal conductance and VPD (d) under Low water availability conditions. Fitted lines equations for the relationships are presented. Slope and intercept values of log transformed equations followed by different letters correspond with significant differences at p < 0.10 according to pairwise Student tests.
In general terms, the differences in gas exchange parameters that were observed across the experiment are not as substantial as those reported by other researchers when comparing water availability levels (De Souza et al., 2005; Romero et al., 2010), which agrees with the fact the effect of IrrF and DisP on plant water status was relatively small. The fact that the effects observed were small agrees with other authors who found that in severe water stress conditions the effects of irrigation treatments in grapevine physiological performance are attenuated (Flexas et al., 1998), and that the effect of water distribution pattern can be reduced in the case of fine textured soils (Chaves et al., 2007; Marsal et al., 2008). However, the degree to which IrrF has been shown to affect AN and gs under Low
WA was enough to affect yield significantly (Sebastian et al., 2015). The analysis of the diurnal evolution of AN and gs (Figs. 1 and 2) allowed us to explain this behaviour: on ‘Day 2’, 4d grapevines were photosynthetically more active than 2d grapevines, since they had received twice the amount of water the previous day and, although on ‘Day 4’ 2d grapevines were expected to be more active than 4d grapevines that had not been irrigated for three days, differences in gas exchange rates appeared seldom, their magnitude was smaller, and did not balance those observed on ‘Day 2’. The different daily evolution patterns of leaf gas exchange observed depending on the amount of available water and the time elapsed since irrigation could be explained because, addition-
416
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Fig. 6. Relationships between net assimilation and leaf water potential (a), stomatal conductance and leaf water potential (b), net assimilation and VPD (c) and stomatal conductance and VPD (d) under Medium water availability conditions. Fitted lines equations for the relationships are presented. Slope and intercept values of log transformed equations followed by different letters correspond with significant differences at p < 0.10 according to pairwise Student tests.
ally to the effects IrrF may have on irrigation efficiency, the fact some grapevines received irrigation water every 2 days whereas others did every 4 days results in a different dynamic of stressrecovering that can result in a different degree of acclimation (Flexas et al., 2006, 2009; Pou et al., 2012). In order to evaluate to which extent differences in irrigation frequency caused acclimation, leaf gas exchange data measured along the day were plotted against water status and evaporative demand (Figs. 5 and 6). Relationships were fitted to exponential functions and log transformed into linear models to be compared statistically. The AN and gs were shown to be related to plant water status and vapour pressure deficit (Figs. 5 and 6), showing similar trends: a
decline of gs and AN for decreasing leaf and increasing VPD, which agrees earlier literature (Chaves et al., 2010; Lovisolo et al., 2010; Rogiers et al., 2012; Tomas et al., 2014). As expected, the differences between the situations considered were bigger when comparing leaf gas exchange parameters against VPD than against leaf water potential, which is logical as in the latter responses at the same level of water status are compared. When the effect of the WA and the time elapsed since the last irrigation was compared (Figs. 5 and 6), water availability conditions were observed to influence the impact of irrigation frequency on leaf gas exchange response to changes in VPD and leaf water potential. Under low WA (Fig. 5), gas exchange parameters of 2d
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
grapevines showed a steeper decline in response to increments in VPD and to reductions of leaf than 4d grapevines (higher slope), and hence, presented the lowest AN and gs values under high evaporative demands and low leaf water potentials, i.e.: under the most frequent conditions during the experiment. 4d grapevines showed the same response of AN and gs to changes in leaf and VPD irrespective of the time elapsed since irrigation. Significant differences between fitted lines only appeared when comparing 2d vs 4d for VPD, although for leaf water potential fitted lines followed the same pattern. Under medium WA (Fig. 6), 4d grapevines responded differently to changes in leaf and VPD depending on the time elapsed since irrigation. Thus, on ‘day 2’ of the irrigation cycle, AN and gs response to changes in VPD and leaf of 4d grapevines was the mildest, behaving as quite “optimistic” (Jones et al., 1980), maintaining high levels of leaf gas exchange at low values of leaf . However, on ‘day 4’, these grapevines, that had not been irrigated for three days, showed a steep decline of AN and gs to decreasing leaf and increasing VPD. Grapevines irrigated every 2 days under medium WA showed an intermediate response. The relationships between leaf gas exchange parameters, leaf water potential and the atmospheric demand showed that low WA treatments presented tighter stomata control than medium WA treatments. The fact that 4d grapevines under low WA conditions had a very similar response of leaf gas exchange to changes in leaf water potential irrespective the time elapsed since last irrigation suggests that some kind of acclimation occurred. This phenomenon would be the result of the interaction between hydraulic and rootmediated hormonal information (Lovisolo et al., 2010; Soar et al., 2006; Tardieu and Simonneau, 1998; Vandeleur et al., 2009). Under medium WA, grapevines irrigated every 4 days appeared not to acclimate, as after three days without irrigation when climatic conditions worsened they decreased leaf gas exchange. Thus, the grapevines irrigated every 4 days with the higher irrigation dose (medium WA) seemed to be less protected against drought stress than the grapevines irrigated with the lower irrigation dose. This could explain their physiological behaviour at the end of ripening, showing very low stomatal conductance after being three days without irrigation, even lower than 4d grapevines under low WA at the same stage (Fig. 2).
4. Conclusions Irrigation frequency and water distribution pattern affected leaf gas exchange of cv. Syrah grown in a clay soil, the effect of irrigation frequency being much more relevant than that of water distribution pattern. Irrigation intervals of two days or shorter should be avoided in clay soils under low water availability conditions, as they result in an irrigation efficiency loss. However, when water availability is higher, for the irrigation frequencies tested, this effect fades away, which makes that under these conditions, frequency could be chosen considering other management issues such as water scheduling, design, operational costs, etc. Water distribution pattern, a nearly unstudied factor, appeared not to be a very critical issue under the conditions of this study, although the closer spacing seemed to be slightly more efficient, especially for medium water availability conditions. Irrigation frequency clearly affected the relationship between leaf gas exchange parameters, plant water status and atmospheric conditions, and together with the amount of water applied had implications in the development of acclimation mechanisms that affected plant physiological response, thus affecting irrigation efficiency. Therefore, more importance should be given to this factor when implementing an irrigation strategy.
417
Acknowledgements The authors would like to thank Osborne Malpica Winery, for allowing us to carry out this research on their vineyards, and for their financial support of the study. The authors would also like to thank the staff in the Viticulture Research Group of the Polytechnic University of Madrid for their support in taking measurements.
References Acevedo-Opazo, C., Ortega-Farias, S., Fuentes, S., 2010. Effects of grapevine (Vitis vinifera L.) water status on water consumption, vegetative growth and grape quality: an irrigation scheduling application to achieve regulated deficit irrigation. Agric. Water Manage. 97, 956–964. Bahar, E., Carbonneau, A., Korkutal, I., 2011. The effect of extreme water stress on leaf drying limits and possibilities of recovering in three grapevine (Vitis vinifera L.) cultivars. Afr. J. Agric. Res. 6 (5), 1151–1160. ˜ M.F., Rodrigues, M.L., Lopes, M., Chaves, M.M., Santos, T.P., Souza, C.R., Ortuno, Maroco, J.P., Pereira, J.S., 2007. Deficit irrigation in grapevine improves water-use efficiency while controlling vigour and production quality. Ann. Appl. Biol. 150, 237–252, http://dx.doi.org/10.1111/j.1744-7348.2006.00123.x. Chaves, M.M., Zarrouk, O., Francisco, R., Costa, J.M., Santos, T., Regalado, A.P., Rodrigues, L., Lopes, C.M., 2010. Grapevine under deficit irrigation: hints from physiological and molecular data. Ann. Bot. 105, 661–676. Cifre, J., Bota, J., Escalona, J.M., Medrano, H., Flexas, J., 2005. Physiological tools for irrigation scheduling in grapevine (Vitis vinifera L.). An open gate to improve water-use efficiency? Agric. Ecosyst. Environ. 106, 159–170. Cornic, G., Massacci, A., 1996. Leaf photosynthesis under drought stress. In: Photosynthesis and the Environment. Springer, Netherlands, pp. 347–366. Dallas, H., Lewandowski, A., 2003. Protecting Urban Soil Quality: Examples for Landscape Codes and Specifications. USDA Natural Resources Conservation Services. De Souza, C.R., Maroco, J.P., dos Santos, T.P., Rodrigues, M.L., Lopes, C., Pereira, J.S., Chaves, M.M., 2005. Control of stomatal aperture and carbon uptake by déficit irrigation in two grapevine cultivars. Agric. Ecosyst. Environ. 106, 261–274. Escalona, J.M., Flexas, J., Medrano, H., 1999. Stomatal and non-stomatal limitations of photosynthesis under water stress in field-grown grapevines. Aust. J. Plant Physiol. 26, 421–433. Fereres, E., Evans, R.E., 2006. Irrigation of fruit trees and vines: an introduction. Irrig. Sci. 24, 55–57. Flexas, J., Escalona, J.M., Medrano, H., 1998. Down-regulation of photosynthesis by drought under field conditions in grapevine leaves. Aust. J. Plant Physiol. 25, 893–900. Flexas, J., Ribas-Carbo, M., Hanson, D.T., Bota, J., Otto, B., Cifre, J., McDowell, N., Medrano, H., Kaldenhoff, R., 2006. Tobacco aquaporin NtAQP1 is involved in mesophyll conductance to CO2 in vivo. Plant J. 48, 427–439. Flexas, J., Barón, M., Bota, J., Ducruet, J.M., Gallé, A., Galmés, J., Jiménez, M., Pou, A., Ribas-Carbó, M., Sajnani, C., Tomàs, M., Medrano, M., 2009. Photosynthesis limitations during water stress acclimation and recovery in the drought adapted Vitis hybrid Richter-110 (V. berlandieri × V. rupestris). J. Exp. Bot. 60, 2361–2377. Goldberg, S.D., Gornat, B., Bar, Y., 1971. The distribution of roots, water and minerals as a result of trickle irrigation. J. Am. Soc. Hortic. Sci. 96 (51), 645–648. Gomez-del-Campo, M., Baeza, P., Ruiz, C., Sotes, V., Lissarrague, J.R., 2007. Effect of previous water conditions on vine response to rewatering. Vitis 46 (2), 51–55. Hochberg, U., Degu, A., Fait, A., Rachmilevitch, S., 2013. Near isohydric grapevine cultivar displays higher photosynthetic efficiency and photorespiration rates under drought stress as compared with near anisohydric grapevine cultivar. Physiol. Plant. 147 (4), 443–452. IPCC, 2008. Climate change and water. In: B.C. Bates, B.C., Kundzewicz, Z.W., Wu, S., Palutikof, J.P. (Eds.), IPCC Tech. Paper VI. IPCC Secretariat, Geneva. Jones, H.G., Turner, N.C., Kramer, P.J., 1980. Interaction and integration of adaptive responses to water stress: the implications of an unpredictable environment. Adaptation of plants to water and high temperature stress, 353–365. Kleinbaum, D.G., Kupper, L., Muller, K.E., Nizam, A., 2008. Applied Regression Analysis and Other Multivariable Methods, 4th ed. Duxbury Applied, Thomsom Co, Belmont, USA. Kriedemann, P.E., Smart, R.E., 1971. Effects of irradiance, temperature, and leaf water potential on photosynthesis of vine leaves. Photosynthetica 5, 15–19. Levin, I., Van Pooyen, P., Van Rooyen, F., 1979. The effect of discharge rate and intermittent water application by point source irrigation on the soil moisture distribution pattern. Soil Sci. Soc. Am. J. 43, 8–16. Loveys, B.R., Kriedeman, P.E., 1973. Rapid changes in abscisic acid-like inhibitors following alterations in vine leaf water potential. Physiol. Plant. 28, 476–479. Lovisolo, C., Perrone, I., Hartung, W., Schubert, A., 2008. An abscisic acid-related reduced transpiration promotes gradual embolism repair when grapevines are rehydrated after drought. New Phytol. 180 (3), 642–651. Lovisolo, C., Perrone, I., Carra, A., Ferrandino, A., Flexas, J., Medrano, H., Schubert, A., 2010. Drought-induced changes in development and function of grapevine (Vitis spp.) organs and in their hydraulic and non-hydraulic interactions at the whole-plant level: a physiological and molecular update. Funct. Plant Biol. 37, 98–116.
418
B. Sebastian et al. / Agricultural Water Management 177 (2016) 410–418
Marsal, J., Mata, M., Del Campo, J., Arbones, A., Vallverdú, X., Girona, J., Olivo, N., 2008. Evaluation of partial root-zone drying for potential field use as a deficit irrigation technique in commercial vineyards according to two different pipeline layouts. Irrig. Sci. 26 (4), 347–356. Medrano, H., Escalona, J.M., Bota, J., Gulías, J., Flexas, J., 2002. Regulation of photosynthesis of C3 plants in response to progressive drought: stomatal conductance as a reference parameter. Ann. Bot. 89, 895–905, http://dx.doi. org/10.1093/aob/mcf079. Myburgh, P.A., 2012. Comparing irrigation systems and strategies for table grapes in the weathered granite-gneiss soils of the Lower Orange River Region. S. Afr. J. Enol. Vitic. 33, 184–197. Padgett-Johnson, M., Williams, L.E., Walker, M.A., 2003. Vine water relations, gas exchange, and vegetative growth of seventeen Vitis species grown under irrigated and non-irrigated conditions in California. J. Am. Soc. Hortic. Sci. 128, 269–276. Pou, A., Medrano, H., Tomás, M., Martorell, S., Ribas-Carbó, M., Flexas, J., 2012. An anisohydric grapevine variety performs better under moderate water stress and recovery than isohydric varieties. Plant Soil 359, 335–349, http://dx.doi. org/10.1007/s11104-012-1206-7. R Core Team, 2016. R: A language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria. URL http://www.R-project.org/. Rogiers, S.Y., Greer, D.H., Hatfield, J.M., Hutton, R.J., Clarke, S.J., Hutchinson, P.A., Somers, A., 2012. Stomatal response of an anisohydric grapevine cultivar to evaporative demand, available soil moisture and abscisic acid. Tree Physiol. 32 (3), 249–261. Romero, P., Fernández-Fernández, J.I., Cutillas, A., 2010. Physiological thresholds for efficient regulated deficit-irrigation management in winegrapes grown under semiarid conditions. Am. J. Enol. Vitic. 61, 300–312. Salleo, S., Lo Gullo, M.A., 1989. Xylem cavitation in nodes and internodes of Vitis vinifera L. plants subjected to water stress. Limits of restoration of water conduction in cavitated xylem conduits. In: Kreeb, K.H., Richter, H., Hinckley, T.M. (Eds.), Structural and Functional Responses to Environmental Stresses. SPB Academic Publishing, The Hague,The Netherlands, pp. 33–42. Santesteban, L.G., Royo, J.B., 2006. Water status, leaf area and fruit load influence on berry weight and sugar accumulation of cv.‘Tempranillo’ under semiarid conditions. Sci. Hortic. 109 (1), 60–65. Santesteban, L.G., Miranda, C., Royo, J.B., 2009. Effect of water deficit and rewatering on leaf gas exchange and transpiration decline of excised leaves of four grapevine (Vitis vinifera L.) cultivars. Sci. Hortic. 121 (4), 434–439. Schultz, H.R., 2003. Differences in hydraulic architecture account for near isohydric and anisohydric behaviours of two field-grown Vitis vinifera L. cultivars during drought. Plant Cell Environ. 26, 1393–1405, http://dx.doi.org/10.1046/j.13653040.2003.01064.x. Sebastian, B., Baeza, P., Santesteban, L.G., Sanchez de Miguel, P., De La Fuente, M., Lissarrague, J.R., 2015. Response of grapevine cv Syrah to irrigation frequency
and water distribution pattern in a clay soil. Agric. Water Manage. 148, 269–279. Selles, G., Ferreyra, R., Contreras, G., Ahumada, R., Valenzuela, J., Bravo, R., 2004. Effect of three irrigation frequencies applied by drip irrigation over table grapes (Vitis vinifera L. cv Thompson Seedless) located in the Aconcagua Valley (Chile). Acta Hortic. 646, 175–181. Smasjtrla, A.G., Harrison, D.S., Clark, G.A., 1985. Trickle Irrigation Scheduling I. Duration of Water Application IFAS Bulletin 204. University of Florida, Gainesville. Soar, C.J., Dry, P.R., Loveys, B.R., 2006. Scion photosynthesis and leaf gas exchange in Vitis vinifera L. cv. Shiraz: mediation of rootstock effect via xylem sap ABA. Aust. J. Grape Wine Res. 12, 82–96. Soil Survey Staff, 2013. Simplified Guide to Soil Taxonomy. USDA, Natural Resources Conservation Service. National Soil Survey Center, Lincoln, NE http://www.nrcs.usda.gov/wps/portal/nrcs/detail/soils/survey/class/ ?cid=nrcs142p2 053580. Tardieu, F., Simonneau, T., 1998. Variability among species of stomatal control under fluctuating soil water status and evaporative demand: modelling isohydric and anisohydric behaviours. J. Exp. Bot. 49 (Secial Issue), 419–432. Tardieu, F., 1989. Root system responses to soil structural properties: micro-and macro-Scale. In: Mechanics and Related Processes in Structured Agricultural Soils. Springer, Netherlands, pp. 153–172. Tomas, M., Medrano, H., Brugnoli, E., Escalona, J.M., Martorell, S., Pou, A., Ribas-Carbó, M., Flexas, J., 2014. Variability of mesophyll conductance in grapevine cultivars under water stress conditions in relation to leaf anatomy and water use efficiency. Aust. J. Grape Wine Res. 20, 272–280. Tramontini, S., Van Leeuwen, C., Domec, J.-C., Destrac-Irvine, A., Basteau, C., Vitali, M., Mosbach-Schulz, O., Lovisolo, C., 2013. Impact of soil texture and water availability on the hydraulic control of plant and grape-berry development. Plant Soil 368, 215–230. Turner, N.C., Long, M.J., 1980. Errors arising from rapid water loss in the measurements of leaf water potential by the pressure chamber technique. Aust. J. Plant Physiol. 7, 527–537. Turner, N.C., 1988. Measurement of plant water status by the pressure chamber technique. Irrig. Sci. 9, 289–308. Van Leeuwen, C., Tregoat, O., Chone, X., Bois, B., Pernet, D., Gaudillere, J.P., 2009. Vine water status is a key factor in grape ripening and vintage quality for red Bordeaux wine. How can it be assessed for vineyard management purposes? J. Int. Sci. Vigne Vin 43, 121–134. Vandeleur, R.K., Mayo, G., Shelden, M.C., Gilliham, M., Kaiser, B.N., Tyerman, S.D., 2009. The role of plasma membrane intrinsic protein aquaporins in water transport through roots: diurnal and drought stress responses reveal different strategies between isohydric and anisohydric cultivars of grapevine. Plant Physiol. 149, 445–460. Wang, F.X., Kang, Y., Liu, S.P., 2006. Effects of drip irrigation frequency on soil wetting pattern and potato growth in North China Plain. Agric. Water Manage. 79 (3), 248–264.