Influence of rootstock, irrigation level and recycled water on growth and yield of Soultanina grapevines

Environmental and Experimental Botany 52 (2004) 185–198

Influence of rootstock, irrigation level and recycled water on waterrelations and leaf gas exchange of Soultanina grapevines

Nikos V. Paranychianakisa,∗, Kostas S. Chartzoulakisb, Andreas N. Angelakisa

a NAGREF, Institute of Iraklio, P.O. Box 2229, 71307 Iraklio, Greeceb NAGREF, Institute of Olive Trees and Subtropical Plants, 73100 Chania, Greece

Accepted 10 February 2004

Abstract

The effects of rootstock variety (41B, 1103P and 110R), irrigation level (0.50, 0.75 and 1.00 of the evapotranspiration) and waterquality (recycled versus freshwater) on water relations and gas exchange of potted Soultanina grapevines were investigated duringtwo growing seasons. An early reduction of predawn leaf water potential (Ψpd) was detected for vines irrigated with recycledwater in both the seasons. However, assimilation rate (A) and stomatal conductance (gs) were reduced only late in the first year athigher irrigation levels. This was consistent with the higher reduction ofΨpd at these treatments indicating that the developmentof a water deficit due to salt accumulation reduced gas exchange. At midday during 1998, leaves on vines grafted on 1103P and110R had lower leaf water potentials and highergs andA. An opposite effect was however observed early in June the followingseason, although later in that season no differences were detected among rootstocks. That differentiation of rootstock effectbetween seasons may have been induced by alterations in source/sink relations due to differences in yield among rootstocksand its effects on leaf area. Irrigation level strongly affected water relations and gas exchange. BothA andgs were significantlycorrelated with soil moisture. A differentiation in these relations was observed among rootstocks late in 1998 which probablyresulted from the change in leaf area development among rootstocks during that period.© 2004 Elsevier B.V. All rights reserved.

Keywords: Grapevines; Rootstock; Recycled water; Irrigation level; Water relations; Gas exchange; Salts; Source/sink relations

1. Introduction

Water relations and leaf gas exchange parametersare affected by both the water availability and quality.Photosynthesis is adversely affected by low soil watersupply (Williams et al., 1994) and salt accumulation(Prior et al., 1992), mainly due to stomatal closure. The

∗ Corresponding author. Tel.:+30-2810-242870; mobile:30-6977396137; fax:+30-2810-245873.

E-mail address: [email protected] (N.V. Paranychianakis).

resulting reduction in carbohydrate production may bean important constraint for growth and yield (Prioret al., 1992; Zhu, 2001). In addition, nutrient avail-ability (Wong et al., 1985) and source/sink relations(Candolfi-Vasconcelos and Koblet, 1991) have beenreported to affect water relations and gas exchange.

Rootstocks have been widely reported to affect gasexchange (During, 1994; Candolfi-Vasconcelos et al.,1994), but reasons for the effect remain obscure dueto the possibly complex interactions among morpho-logical factors (root distribution and density, root hy-

0098-8472/$ – see front matter © 2004 Elsevier B.V. All rights reserved.doi:10.1016/j.envexpbot.2004.02.002

186 N.V. Paranychianakis et al. / Environmental and Experimental Botany 52 (2004) 185–198

draulic conductivity and leaf area per root ratio), nutri-ent status, water absorption and source/sink relations.Reported differences in assimilation rate among un-grafted Riesling vines, grafted to Riesling, 5BB, andSO4 were attributed to stomatal conductance and/orcarboxylation efficiency (During, 1994). The lowerstomatal conductance of Cabernet Sauvignon vinesgrafted on AxR1 compared to those grafted on St.George and 5C was related to higher number of rootintercepts<2 mm (Williams and Smith, 1991). Fur-thermore, leaf area development which has been foundto differ among rootstocks (Sommer and Clingellefer,1997) may also affect transpiration rates (Gomez deCampo et al., 1999).

Rootstocks were also found to modify leaf gasexchange of the scion under non-irrigated condi-tions, even though vine water status was not altered(Padget-Johnson et al., 2000). However, little isknown about the interactive effects of soil moistureand rootstock. In an early study, differences in waterabsorption among the grapevine rootstocks, whichwere assessed by multiplying the total leaf area withstomatal conductance, were observed only under se-vere water deficits (Carbonneau, 1985). In addition,Muller Thurgau vines grafted on H1 and H8 hybridrootstocks did not show any reduction in assimilationrate after 14 days of water stress, in contrast to thoseon H26 rootstock which showed a reduction (Iakonoet al., 1998). However, possible factors which causethis differentiation remain obscure.

Furthermore, differences in vegetative growth pat-terns and yield attributable to rootstocks (Ezzahouaniand Williams, 1995; McCarthy et al., 1997) may alsoaffect gas exchange by altering source/sink relations.An understanding of the interrelationships among wa-ter availability, gas exchange, vegetation and rootingpatterns and yield may lead to development of newmethods for improving vine productivity and wateruse efficiency.

The use of recycled water for irrigation has becamea common practice throughout the world, especiallyin areas with deficient water resources (Klein et al.,2000; Mauer et al., 1995). Recycled water containsessential nutrients and salts at concentrations whichcan affect gas exchange and/or water relations of theirrigated plants. Increased content of salts in irriga-tion water was found to reduce leaf gas exchange ofvines and that reduction was highly correlated with

lamina Cl− (Prior et al., 1992). On the other hand, noeffect of saline irrigation on midday leaf water poten-tial was reported.Walker et al. (1997)reported sim-ilar results and concluded that salt accumulation ingrapevine tissues and ion imbalance rather than a wa-ter deficit were responsible for gas exchange reduc-tion. Vine response to salinity is affected by rootstockvariety (Walker et al., 1997). Studies of various plantspecies have shown that the concentrations of certainelements, such as K+ and Ca2+ may also alleviate theeffect of salinity on affected parameters (Chow et al.,1990; Zhu, 2001). However, little is known about thecombined effect of low N concentration and relativelyhigh concentrations of Na+ and Cl− on vine leaf gasexchange parameters and how it interacts with root-stock and irrigation level. Thus, the main objectivesof this study were to evaluate the effects of rootstock,irrigation rate and water quality and their interactionson vine water relations and leaf gas exchange.

2. Materials and methods

2.1. Growth conditions and treatments

In May 1996, 1-year-old ‘Soultanina’ grapevines(Vitis vinifera L.) on three rootstocks, 41B (V. vinifera× V. berlandieri), 1103P (V. berlandieri × V. ru-pestris), and 110R (V. berlandieri × V. rupestris),were planted in 30 l pots filled with sandy loam soil(7.4 pH and 0.46 dS m−1 EC). The pots were spaced1.2 m within and 1.5 m between rows in an open fieldlocated at the National Foundation for AgriculturalResearch, Iraklio, Greece. All vines were pruned toone or two-node spur in March 1997 and one shootwas allowed to grow vertically until late June. Shootswere then trained along a horizontal trellis wire, 40 cmfrom the pot surface, to develop the cordon. Vineswere pruned to one or eight-node cane in 1998 andthree shoots were trained vertically and tied to wireslocated at 20, 40, 60 and 100 cm from the horizontalpart of the trunk; all inflorescences were removedbefore bloom. Vines were trained in a similar mannerin 1999, except that four shoots were retained andinflorescences were kept on two of the shoots.

Water demand for the 1.00ET treatment was deter-mined as follows: 12 vines (two vines per rootstockper water quality) were irrigated until tensiometer

N.V. Paranychianakis et al. / Environmental and Experimental Botany 52 (2004) 185–198 187

readings approached the zero or until drainage wasevident, and the average water consumption wasconsidered to be the 1.00ET level independently ofrootstock and water quality. The other two irrigationlevels 0.50ET and 0.75ET were proportional. Anydrainage was collected and reapplied to the pots. Ten-siometers, located at 20 cm depth, provided a rapidand continuous monitoring of soil water status at1.00ET irrigation treatments. Soil moisture at that irri-gation treatment was never allowed to fall bellow the15 kPa so that vines irrigated with this treatment donot experience water stress. As a result, the frequencyof irrigation changed during the growing season withclimatic conditions and leaf area development fromone irrigation per 2 days early in the season (April toMay) to two irrigations per day (July to August). Af-ter 1 August 1998 and 15 June 1999, when vines wereirrigated with recycled water, showed an increasingsoil water content, due to inhibition of shoot growth,irrigation needs were determined separately for eachwater quality (six vines per water quality treatment).Based on this estimate, vines irrigated with recycledwater received 8% less water in 1998 and 33% lesswater in 1999 than vines irrigated with freshwater(Table 1). Irrigation treatments were applied from20 May to 20 September each year. During wintermonths, all pots were irrigated in excess with fresh-water to leach any accumulated salts and to preventthe build up of salts in pots from season to season.

The composition of recycled and freshwater duringthe period of the study is shown inTable 2. The onlynutrients that vines irrigated with recycled water re-ceived for their fertilization were those found in thewater itself, while vines irrigated with freshwater re-ceived Hoagland solution. During 1998 and 1999, the

Table 1Total consumption of water (l) during the years 1998 and 1999for vines irrigated at 1.00ET level

Year Water May June July August Septembera Total

1998 FWb 41 68 93 102 64 368RWc 41 68 93 89 48 340

1999 FW 49 86 121 132 99 489RW 49 70 83 79 46 325

a Water consumption until 19 September.b FW: freshwater.c RW: recycled water.

Table 2Chemical composition of recycled water, and freshwater

Characteristics Recycled water Freshwater

1998 1999 1997–1999

EC (dS m−1) 1.9 1.9 0.6pH 7.5 7.3 7.1NH4–N (mg l−1) 5.7 4.2NO3–N (mg l−1) 4.6 4.1Total P (mg l−1) 10.7 9.4K+ (mg l−1) 37.4 34.6 2.6Ca2+ (mg l−1) 39.1 51.2Mg2+ (mg l−1) 9.3 12.4Na+ (mg l−1) 247 264 72Cl− (mg l−1) 401 436 118B3+ (mg l−1) 0.48 0.67Cu2+ (mg l−1) 0.013Fe2+ (mg l−1) 0.21Mn2+ (mg l−1) 0.03

fertilization supply was adjusted to irrigation level toensure that availability of nutrients was more propor-tional to differential vigor resulted from the irrigationtreatments. It was considered as necessary, since 1997when the same amount of nutrient solution was addedin the irrigation treatments leaf N content was lower at1.00ET treatment (Paranychianakis et al., 2000). Moredetailed in both seasons the application of nutrientsolution began on 1 May, simultaneously with recy-cled water irrigation. On 1 May 1998, one-half litterof Hoagland solution was provided to each vine andthen again on 10 and 20 May. After that date until 15September, the nutrient solution was applied throughthe irrigation system at 1 l per week for vines irrigatedat 1.00ET, and proportionally lower for the 0.50 and0.75ET treatments. The fertilization schedule resultedin 18.5 l of Hoagland solution for 1.00ET vines 13.25 lfor 0.75ET vines and 10 l for 0.50ET vines. In 1999,the same fertilization schedule was followed but thequantity of nutrient solution was increased by 50%due to the fact that root growth was not yet restrictedby the pot size and therefore a further increase in vinegrowth was expected.

2.2. Statistical analysis

Eighteen treatments were laid out as a three-factorexperiment involving three rootstocks, three irrigationlevels, and two water quality levels arranged in a ran-


domized complete block design with four replications.Each replication consisted of four vines in a row. Datawere analyzed using analysis of variance and meanswere separated by Tukey’s significant difference test.

2.3. Parameters measured

2.3.1. Soil water contentTo determine soil water content (SWC), soil sam-

ples from entire pot depth were taken with a soil sam-pler of 0.8 cm diameter. Soil sampling was performedjust before irrigation applied and simultaneously withgas exchange measurements to remove any variabil-ity from SWC versus gas exchange curves. Soil coreswere immediately weighted and afterwards were ovendried at 65◦C to a constant weight. One vine per treat-ment per block (four vines per treatment) were se-lected for SWC measurements.

2.3.2. Leaf water potentialPredawn leaf water potential (Ψpd) was measured

weekly on fully-expanded healthy leaves using a pres-sure chamber according to the technique ofScholanderet al. (1965). Midday leaf water potential (Ψmd) wasalso assessed in both seasons. It was first measured on14 June 1998 and thereafter followed theΨpd mea-surements in both the seasons. Measurements ofΨpdwere performed on pots that were to be sampled forSWC.

2.3.3. Gas exchangeAssimilation rate (A), stomatal conductance (gs)

and intercellular CO2 concentration (Ci ) were deter-mined using an open gas exchange system (CIRAS1, PP Systems, UK). The gas exchange unit wasoperated at 4.2 ml s−1 flow rate and ambient CO2partial pressure of 35–37 Pa. Measurements weretaken between 11:30 and 14:00 h on fully expandedhealthy, sun-exposed (photosynthetic photon flux den-sity (PPFD) > 1200�mol m−2 s−1) leaves adjacentto those used forΨpd determination. The carboxyla-tion efficiency (CE) was calculated by theA/Ci ratio(During, 1994).

2.3.4. Leaf area developmentLeaf area (LA) was estimated at monthly intervals

during 1998 growing season, and in 1999, it was esti-

mated at fruit set, veraison and on 20 September, usingthe following equation:

LA = 0.3119(VL + VR) + 2.0679

(R2 = 0.976, n = 120) (1)

where VL and VR are the lengths (cm) of the left andright veins, respectively.

2.3.5. Root dry weightOn 20 September 1998 and 1999, whole plants

where destructively harvested and the root dry weightswere assessed. One vine per treatment per block wassampled for above-ground and root dry weights. Rootswere thoroughly washed and were oven-dried at 65◦Cto a constant weight.

2.3.6. Leaf N and Cl− contentTotal nitrogen (N) in grapevine tissues samples was

assessed by the Kjeldahl method. Chloride was as-sessed in dilute nitric acid extract using a chloride me-ter (JENWAY PCLM3).

3. Results

3.1. Recycled water

A significant increase in SWC of vines irrigatedwith recycled water was observed after 17 July 1998(Fig. 1a) when leaf area significantly reduced inthat treatment (Fig. 2). This effect increased withincreasing irrigation level (Fig. 1a) as a result ofthe greater inhibition of leaf area at these treatments(Paranychianakis, 2001) while irrigation rates weremaintained the same for both water qualities. Theadjustment of irrigation needs separately for eachquality after 15 August, eliminated the difference.Likewise in 1999, the lower leaf area of vines irri-gated with recycled water resulted in higher SWCfor these vines from the beginning of June (Fig. 1b).Thereafter the adjustment of irrigation needs as theprevious year reduced the differences.

An early (mid-June) reduction inΨpd was the firstphysiological response of vines to irrigation with recy-cled water versus freshwater in both growing seasons(Fig. 3). This reduction became more evident as the


Fig. 1. Soil water content (SWC) changes under different irrigationlevels and water qualities during the 1998 (a) and 1999 season(b). Vertical bars represent±S.E.

season progressed. In contrast,Ψmd was not affectedby water quality in both seasons (Fig. 4).

Reductions ings andA for vines irrigated with re-cycled water compared to those irrigated with fresh-

Fig. 2. Effect of water quality on leaf area development during 1998growing season. Sets of bars without letters imply non-significantdifferences.

Fig. 3. Predawn leaf water potential (Ψpd) as affected by irrigationlevel and water quality during the 1998 (a) and 1999 season (b).Vertical bars represent±S.E.

water were observed only late in the 1998 growingseason. The reduction ings was first detected on 18August (Fig. 5). Assimilation rate was first reducedon 28 August (data not shown). During the followingseason, water quality did not affect leaf gas exchangeuntil 22 July when the measurements were completed,

Fig. 4. Effect of water quality onΨmd in 1998 growing season.Sets of bars without letters imply non-significant differences.


Fig. 5. Stomatal conductance (gs) variation under different irriga-tion levels and water qualities during the 1998 season. Verticalbars represent±S.E.

although higher lamina Cl− contents were reachedcompared to those of 1998 (Table 3). In addition, inboth seasons vines irrigated with recycled water hada lower leaf N content (Table 3).

3.2. Irrigation level

Soil water content decreased in response to deficitirrigation treatments. The lowest values of SWC,in response to the 0.50ET were recorded from 17July–15 August and thereafter SWC increased slightly(Fig. 1a). The following season, milder water deficitswere reached in deficit irrigation treatments compared

Table 3The effect of water quality on leaf Cl− and N content during 1998 and 1999 growing seasons

Water quality Year

28 July 1998 20 September 1998 1 July 1999 5 August 1999 20 September 1999

Leaf (Cl−) (% d.w.)Recycled water 0.770 1.316 0.834 1.468 2.250Freshwater 0.495 0.836 0.393 0.774 1.695Significance ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗

Leaf (N) (% d.w.)Recycled water 1.825 1.865 1.808 1.756 1.820Freshwater 1.945 2.046 1.978 1.840 1.836Significance ∗∗ ∗∗∗ ∗∗ ∗∗ ns

ns: not significant.∗∗ P < 0.01.∗∗∗ P < 0.001.

to these of 1998 year (Fig. 1b). It was likely the re-sult of the lower growth rates of shoots (14 and 28%on 20 May) that observed during that year in vinesirrigated at 0.75 and 0.50ET treatments compared tothose of 1.00ET.

Compared with vines irrigated at 1.00ET, a signif-icant reduction ofΨpd was first detected in vines ir-rigated with 0.50ET on 14 June 1998. Subsequently,Ψpd in vines under deficit irrigation treatments de-creased until the first week of August for vines irri-gated with freshwater but then slightly increased at theend of season. However, in vines irrigated with recy-cled water, the reduction ofΨpd continued for all irri-gation treatments until the end of the season (Fig. 3a).An interaction between irrigation level and water qual-ity was detected in both seasons (P < 0.01, ANOVA),indicating thatΨpd increase with increasing irrigationlevel was less in vines irrigated with recycled watercompared to those irrigated with freshwater (Fig. 3). In1999, significant differences between irrigation treat-ments had been developed from the first measurementon 4 June (P < 0.001, ANOVA). Ψpd maintained rel-atively constant in freshwater irrigated treatments butas in the previous season declined in those irrigatedwith recycled water (Fig. 3b). In both years,Ψpd wascurvilinearly related to SWC, independently of waterquality, with a diminishing response to SWC at higherlevels (Fig. 6). Higher values ofΨpd were recorded in1999 compared to those of the previous season (Fig. 6).In terms ofΨmd, vines irrigated with 0.50ET level hadsignificant lower values compared to those irrigated at0.75 and 1.00ET level (data not shown).


Fig. 6. Relationships between soil water content (SWC) andpredawn leaf water potential (Ψpd) during the 1998 and 1999growing seasons. Each point represents a single measurement.

Reduced rates of leaf gas exchange were detectedon 26 June 1998 in vines irrigated at 0.50ET (Fig. 5),after a significant reduction in SWC andΨpd had al-ready been observed. Later, larger differences inAand gs were developed among irrigation levels andremained until the end of the season. Stomatal con-ductance showed a large variation independent of theirrigation level. Higher readings ofgs were recordedearly in June, while a severe decrease was observedfrom the beginning of July to mid-July, butgs in-creased for all treatments after 22 July until the sea-son end (Fig. 5). A similar pattern was observed forA. Both A andgs were affected by an interaction be-tween irrigation level and water quality on 18 and 28August (P < 0.01, ANOVA). The increase in bothAandgs with increasing irrigation level was less whenirrigated with recycled rather than freshwater (Fig. 5).During 1999, significant differences in gas exchangeamong irrigation levels had developed by 6 June (datanot given). Higher values ofgs were found comparedto those of the previous season, however they werenot accompanied by increasedA. There was a sig-nificant curvilinear relationship betweenA and gs inboth seasons. A differentiation in theA versusgs re-lationship was observed not only between seasons butalso within the same season. For example in 1999 therelationship differed between the 6 June and the fol-lowing samplings (Fig. 7). Furthermore, bothA andgs were significantly correlated withΨpd and this re-lationship did not change with water quality (Fig. 8).

Fig. 7. Relationships between stomatal conductance (gs) and as-similation rate (A) during 1999 growing season. Each point repre-sents a single measurement. Fitted lines were statistically signifi-cant atP < 0.05.

Stomatal conductance was also significantly correlatedwith SWC, however that relationship was affected bywater quality after 18 August 1998, whengs of vinesirrigated with recycled water responded less to in-creasing SWC (gs = 10.7(SWC) + 12.1R2 = 0.42)

Fig. 8. Relationships of predawn leaf water potential (Ψpd) vs.stomatal conductance (gs) and assimilation rate (A) for both waterqualities. Fitted lines were statistically significant atP < 0.05.


Fig. 9. Relationship between soil water content (SWC) and stom-atal conductance (gs) of vines grafted on 41B and 110R on 5July 1998 (a) and 22 July 1998 (b). Fitted lines were statisticallysignificant atP < 0.05.

than didgs of vines irrigated with freshwater (gs =21(SWC)−43.9R2 = 0.70). A similar trend was alsofound for the correlations betweenA and SWC (datanot shown). With regards to rootstocks, significant re-lations between SWC and gas exchange were estab-lished with vines grafted on 41B to show showed lowerrates of gas exchange at all irrigation levels (Fig. 9).However, on 18 and 28 August 1998, significant in-teractions were detected between rootstock and irri-gation level in terms of gas exchange (P < 0.05 andP < 0.01, respectively, ANOVA). At low SWC,A andgs for vines on all rootstocks were similar but as SWCincreased vines grafted on 1103P and 110R showedgreater increases inA andgs than did vines grafted on41B (Fig. 10).

3.3. Rootstock

Rootstock affected SWC in both years. In 1998,vines grafted on 41B showed a significant higherSWC on 5 July compared to other rootstocks. Al-thoughgs for vines grafted on 41B remained loweruntil the late in the season the differences in SWC

Fig. 10. Relationship between soil water content (SWC) and stom-atal conductance (gs) (a) and SWC and assimilation rate (A) and(b) of vines grafted on 41B and 110R on 28 August 1998. Fittedlines were statistically significant atP < 0.05.

among rootstocks did not increase but they disap-peared on 7 August (Table 4). In the following season,rootstock did not affect SWC until 31 July. Afterthat date vines grafted on 1103P showed lower SWCvalues until 15 August when the measurements werecompleted.

Vines grafted on 1103P and 110R had lowerΨmdduring 1998 compared to those on 41B (Fig. 11a). In1999, vines grafted on 41B were lowest inΨmd on 9July. No differences inΨmd were detected among root-stocks on 21 July, while on 17 August, vines graftedon 1103P were lowest inΨmd, showing a similar toprevious year behavior (Fig. 11b).

Vines grafted on 1103P and 110R rootstocks hadhigher gs in six out of eight samplings and higherA in seven out of eight sampling dates comparedto vines grafted on 41B during the 1998 growingseason (Fig. 12). Also vines grafted on 1103P and110R showed higher carboxylation efficiency (datanot shown). In contrast, rootstock effects on leaf gasexchange parameters changed over the 1999 season.On 6 June,A andgs were higher in vines grafted on41B than on vines grafted on 1103P (Table 5). In the


Table 4The effect of rootstock on soil water content (SWC) (%) during 1998 season

Rootstock 28/6 5/7 16/7 22/7 1/8 9/8 14/8 20/8

41B 11.5 11.9 a 9.7 a 11.2 a 11.3 a 11.9 a 10.5 121103P 10.9 10.1 b 9b 10.4 b 9.8 b 10.3 b 10 11.5110R 10.3 10.7 b 9.1 a 11 ab 10.4 ab 10.6 b 10.5 11.6

Significance ns ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗ ns ns

ns: not significant. Numbers with different letters differ significantly at the 0.05 level by Tukey’s significant difference.∗∗ P < 0.01.∗∗∗ P < 0.001.

Table 5Effect of rootstock on assimilation rate (A) stomatal conductance (gs) and carboxylation efficiency (CE) on 6 June 1999

Rootstock A (�mol m−2 s−1) gs (mmol m−2 s−1) CE (�mol m−2 s−1 ppm−1)

41B 12.2 a 103 a 0.0981103P 10.4 b 88 b 0.077110R 11.3 ab 100 ab 0.088

Significance ∗∗ ∗ ns

ns: not significant. Numbers with different letters differ significantly at the 0.05 level by Tukey’s significant difference.∗ P < 0.05.∗∗ P < 0.01.

Fig. 11. The effect of rootstock on midday leaf water potential(Ψmd) during the 1998 (a) and 1999 growing season (b). Sets ofbars without letters imply non-significant differences.

Fig. 12. The effect of rootstock on stomatal conductance (gs) (a)and assimilation rate (A) (b) in 1998. Sets of bars without lettersimply non-significant differences.


Fig. 13. The effect of rootstock on leaf area development during1998 (a) and 1999 growing season (b). Sets of bars without lettersimply non-significant differences.

later samplings, no effect of rootstock on leaf gas ex-change was observed.

Rootstock also significantly affected both leaf areadevelopment and root dry weight. Vines grafted on41B developed more leaf area than vines on 110Rand 1103P on 30 July 1998 and these differencesmaintained until the end of the season (Fig. 13a).The following season, vines grafted on 1103P hadgreater leaf area by fruit set than did vines grafted

Table 6Effect of rootstock on root dry weight, root/leaf dry weight ratioand leaf area/yield ratio during the 1998 and 1999 growing seasons

Rootstock Root dryweight

Root/leaf dryweight ratio

Leaf area/yield ratio

1998 1999 1998 1999 1999a

41B 93 b 138 c 1.48 b 2.29 b 42.4 b1103P 129 a 225 a 2.55 a 3.46 a 85.5 a110R 100 b 163 b 1.79 b 2.59 b 84.2 a

Significance ∗∗∗ ∗∗∗ ∗∗∗ ∗∗∗ ∗∗

Any two means within a row not followed by the same letterare significantly different atP < 0.05 with Tukey’s significantdifference, ns: not significant.

a Leaf area in LA/yield ratio has been assessed at veraison.∗∗ P < 0.01.∗∗∗ P < 0.001.

Table 7The effect of rootstock and irrigation level on root dry weight (g)a

Rootstock Irrigation level 1998 1999

41B 0.50ET 75 d 107 e0.75ET 88 cd 122 de1.00ET 118 bc 186 bc

1103P 0.50ET 90 c 148 cd0.75ET 133 b 200 ab1.00ET 166 a 236 a

110R 0.50ET 64 d 97 e0.75ET 98 c 180 bc1.00ET 136 b 217 ab

a Numbers with different letters differ significantly at the 5%level by Tukey’s significant difference.

on other rootstocks. The rootstock effect on leaf areadisappeared at veraison but by 20 September vinesgrafted on 41B reached more leaf area than thoseon 1103P (Fig. 12b). Furthermore, during that seasonvines grafted on 41B had a significantly lower leafarea to yield ratio (Table 6).

In terms of root dry weight, vines grafted on 1103Phad the greatest root dry weights and root to leaf arearatio (Table 6). Moreover a significant interaction (P <

0.01, ANOVA) was detected. Vines grafted on 41Bshowed a slighter reduction of root dry weight withirrigation level in both seasons (Table 7).

4. Discussion

4.1. Recycled water

Irrigation with recycled water reducedΨpd whichattributed to the osmotic effect arising from salt ac-cumulation in rootzone. The lower response ofΨpdto increasing irrigation level for vines received recy-cled water is consistent with greater salt accumulationin these treatments. In contrastΨmd was not affectedby recycled water in agreement with previous stud-ies (Prior et al., 1992; Walker et al., 1997; Gibberdet al., 2003). It may probably result from the isohydricbehavior of grapevines according which water use iscontrolled to maintain the minimum leaf water poten-tial at a constant value (Winkel and Rambal, 1993).

Although the induced water deficit reduced dra-matically vines’ growth, leaf gas exchange were af-


fected only late in 1998. The reduction of leaf area(Fig. 2) and the ability of grapevines for osmoregu-lation (During and Dry, 1995; Patakas and Noitsakis,1999) seem to counteract the impacts of salinity onvine water use per unit of leaf area arising from re-ductions on root hydraulic conductivity (Ramos andKaufmann, 1979; Storey and Walker, 1999) and waterdeficit development (Storey and Walker, 1999). Evenwhen a reduction of gas exchange was observed itcould be explained by the water deficit developmentas results from the samegs versusΨpd curves of bothwater qualities (Fig. 8). Walker et al. (1997)however,based on the lack of an effect of saline irrigation onΨmd of grapevines concluded that a water deficit de-velopment did not cause the gas exchange reduction.Likely, the use ofΨmd as indicator of vine water sta-tus in that study prevented the detection of the waterdeficit.

A negative correlation between lamina Cl− con-centration and leaf gas exchange has been establishedfor grapevines in earlier studies (Prior et al., 1992;Walker et al., 1981; Downton, 1977) implying thatnon-stomatal factors dominate in gas exchange reduc-tion under salinity stress. According to these relation-ships grapevines grown in both field and pots showedan approximately 15% depression ofA when laminaCl− increased from 100 to 200 mmol kg−1 (Prior et al.,1992; Downton, 1977). In the present study, althoughlamina Cl− exceeded the latter levels from the end ofJuly 1998 onwards (Table 3), gas exchange did notdecrease until late in August. Moreover, the similarrelationships betweenΨpd and photosynthesis in bothwater qualities (Fig. 8b) indicate that decreased pho-tosynthesis could be fully explained by the osmoticcomponent of salinity at least at moderate levels ofstress. The uncoupling of lamina Cl− and leaf gas ex-change reduction in this study became more evidentin 1999, when although lamina Cl− at the beginningof August was double in comparison to the previousseason (Table 3) gas exchange did not decline. A lackof correlation between leaf Cl− content and gas ex-change has been also observed and in other species(Bañuls and Primo-Millo, 1995). These findings indi-cate that there is no particular threshold of Cl− lam-ina concentration above which a reduction of leaf gasexchange occurs. Factors such as, the composition ofirrigation water, availability of nutrients (Chow et al.,1990; Zhu, 2001), cultural practices, and environmen-

tal conditions (Prior et al., 1992) may affect this thresh-old.

Finally, the lower leaf N content of vines irrigatedwith recycled water does not appear to affectA. Thelack of a reduction of A early in 1999 season when thegreatest differences were observed in leaf N contentand the samegs versusA and Ψmd versusA curves(Fig. 8b) support this hypothesis.

4.2. Irrigation level

The curvilinear relationship betweenΨpd and SWCshows thatΨpd is relatively insensitive to soil dryinguntil a SWC threshold value of approximately 11%(w/w) is reached. Severe inhibition of shoot growthwas observed at that level of SWC (Paranychianakis,2001). Moreover,Ψpd sharply decreased as SWC fellbelow the threshold value. These results show thatΨpdby itself, is not the only parameter that should be con-sidered while assessing vine water status and schedul-ing irrigation for grapevines. The generally higherΨpdfound during 1999 cannot be explained only in termsof the increased SWC because the SWC versusΨpdcurves for the 2 years differed (Fig. 4). More likelythe greater root to leaf area ratio which observed in1999 compared to that of the previous year (Table 6)resulted in the higherΨpd values.

Leaf gas exchange is known to decrease with de-creasing irrigation level. The results obtained in thisstudy confirm these of previous ones where bothAand gs were significantly correlated with SWC (vanZyl, 1987; Williams et al., 1994) andΨpd (van Zyl,1987). Williams et al. (1994)suggest that the relativelylow coefficients of correlation found in their study in-dicate that both internal and external factors may beinvolved in the regulation of these relations which isin agreement with the results obtained in this study.Rootstock effect on gas exchange did not change withwater deficit development until 18 August. In con-trast to earlier studies which reported a differentiationof rootstock response when a severe water stress wasdeveloped (Iakono et al., 1998; Carbonneau, 1985).An interaction between rootstocks and irrigation levelswas however, observed at two last samplings (Fig. 10)which was consistent with the root dry weight responseto irrigation level of different rootstocks (Table 7).However, differences in root growth do not appear asthe possible cause of this effect since in that case a


same response should have been prevailed throughoutthe 1998 growing season. It is more possible that thehigher rate of leaf area development of vines graftedon 41B during that period (Fig. 13) and particularly at0.75 and 1.00ET irrigation levels where growth wasless inhibited by water stress led to reduction of gas ex-change in these vines.Gomez de Campo et al. (1999)have also reported a significant dependence of tran-spiration rate from leaf area in grapevines.

4.3. Rootstock

Rootstocks used in this study did not differenti-ate total water consumption during the course of ex-perimental trial season despite the differences in gasexchange rates, leaf area development and rootingpatterns. The greater leaf area developed from vinesgrafted on 41B after 30 July 1998 counteracted thehigher gs of vines grafted on 1103P and 110R andbalanced total transpiration losses. Likewise the fol-lowing season the greatergs for vines grafted on 41Bearly in the season was balanced by the greater leafarea of vines grafted on 1103P.

Rootstock appeared to exert an influence ongas exchange in agreement with previous results(Candolfi-Vasconcelos et al., 1994; During, 1994;Sommer and Clingellefer, 1997; Williams and Smith,1991). The highergs of vines grafted on 1103P wasrelated to greater root dry weight but not in vinesgrafted on 110R. Implying that root size is not theonly factor determining gas exchange rates. Takingalso into account the lower root to leaf area ratio ofvines grafted on 41B and 110R (Table 6) comparedto 1103P and the similar whole-season transpirationlosses it can be concluded that these rootstocks arecharacterized by higher root hydraulic conductiv-ity. Differences in root hydraulic conductivity havebeen also reported in citrus rootstocks (Syvertsen andYelenosky, 1988; Zekri and Parsons, 1989). The grea-ter A in vines grafted on 110R and 1103P in the 1998growing season can be attributed to both highergs andCE in agreement with the findings ofDuring (1994).

Vines grafted on 110R and 1103P had lowerΨmdthan those on 41B during 1998.Ezzahouani andWilliams (1995) reported higherΨmd for vines ofcultivar Ruby seedless grafted on 1103P and 41Bcompared to vines grafted on 110R. The lowerΨmdobserved in this study for vines grafted on 110R and

1103P, was consistent with the highergs found indi-cating thatgs at least partly mediateΨmd. Such anassumption is also verified from the following seasonresults, where the altered effect of rootstock onΨmd(Fig. 6) was followed by corresponding changes ings.Likewise, values ofΨmd were the same for own-rootedSoultana grapevines and grafted on Ramsey rootstockwhich displayed the samegs (Gibberd et al., 2003).

Crop load and particularly the ratio of leaf area toyield may dramatically change source/sink relationsand have been related with changes toA in grapevines(Naor et al., 1997; Candolfi-Vasconcelos and Koblet,1991). Therefore, the observed leaf area to yield dif-ferences among rootstocks (Table 6) may have causedthe differentiation on leaf gas exchange during 1999.In that season, vines grafted on 41B had higherA com-pared to those on 1103P in early June. That periodcoincided with Stage I of berry development when thedemands for assimilates may be high. This hypothesisis supported by the similarA despite lowergs in earlyJune, compared with that found on later dates (Fig. 7).During that period vines grafted on 41B had lower leafarea and higher yield compared with vines on 1103Pwhich may have stimulatedA. IncreasedA, which as-cribed to a highergs, following a defoliation treatmenthas been reported in vines (Candolfi-Vasconcelos andKoblet, 1991). Naor et al. (1997)found that vines bear-ing two clusters per shoot had highergs and A thanvines bearing one cluster per shoot. Since in that studyleaf area did not differ among crop load treatments, itis suggested that the highergs may have been inducedby an increase in root permeability. Such an effectmay also have induced highergs and henceA in vinesgrafted on 41B in this study. In addition, biochemicalprocesses have also been found to contribute to theenhancement of A in defoliated vines (Iakono et al.,1995) but a significant of effect was not observed inthis study (Table 5). However, the increase in leaf Ncontent of vines grafted on 41B in 1999 to similarlevels to those of other rootstocks (Paranychianakis,2001) may imply that biochemical processes also con-tributed in enhancement ofA in the present study.

Acknowledgements

Financial support for this study was provided byEu-Interreg II programme. The authors are grateful to


Prof. K.A. Roubelakis-Angelakis for supervising thestudy and reviewing the manuscripts. Thanks are alsodue to Mr. Y. Spanakis from the Technological Insti-tute of Crete for permitting the use of gas exchangeequipment.

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Influence of rootstock, irrigation level and recycled water on growth and yield of Soultanina grapevines