Rootstock: Scion combinations and nutrient uptake in grapevines

C H A P T E R

21 Rootstock: Scion combinations and nutrient uptake in grapevines Antonio Ibacache, Nicola´s Verdugo-Va´squez, Andr
es Zurita-Silva* Instituto de Investigaciones Agropecuarias (INIA), Centro Regional de Investigacio´n Intihuasi, La Serena, Chile *Corresponding author. E-mail: [email protected]

O U T L I N E 1 Introduction

297

2 Characteristics of rootstocks associated with the uptake of water and nutrients

298

3 Rootstocks differences on the uptake of macro and micronutrients 3.1 Macronutrients

298 301

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation

302

4.1 Chloride toxicity 4.2 Sodium toxicity 4.3 Boron toxicity

304 307 309

5 Carbohydrates and nitrogen reserves

310

6 Rootstock strategies to cope with salinity

311

Acknowledgments

314

References

314

1 Introduction The use of rootstocks in viticulture was initially intended to overcome phylloxera infections that infected European vineyards in the late 19th century (Corso and Bonghi, 2004). However, the recognition of the benefits of rootstocks has since expanded to include nematode control, nutrient absorption, water uptake, vine vigor, yield, and fruit quality (Satisha et al., 2010; Walker et al., 2000; Keller, 2001; Tambe and Gawade, 2004; Ibacache and Sierra, 2009; Ibacache et al., 2016). Also, in some arid or semiarid areas, rootstocks are used to replace old or unproductive vineyards (Ibacache and Sierra, 2009; Satisha et al., 2010). Most of the world viticulture is based on grafted grapevines, where the scion is a cultivar of Vitis vinifera and the rootstock is either an American Vitis species or an interspecific Vitis hybrid. The different effects of the rootstocks on scions take place in a more or less indirect manner and are consequences of interactions between environmental factors and the physiology of both scion and rootstock cultivars employed. Grapevine cultivars are known to exhibit wide differences in mineral nutrient status. Likewise, the use of rootstocks with the resistance and tolerance to phylloxera and/or nematodes can also have a major influence on the mineral nutrient status of scion cultivar (Garcia et al., 2001; Bavaresco et al., 2003; Fisarakis et al., 2004; Ibacache and Sierra, 2009). This influence has important implications for decisions involving soil adaptability, grapevine fertilizer requirements, canopy management, yield, and fruit quality. Rootstocks are also known to affect the uptake of mineral nutrients that can be damaging to grapevines, including sodium, chloride, and boron (Stevens and Harvey, 1995; Walker et al., 2004). This should be considered when choosing a rootstock for soils, which have potentially damaging levels of salts. Although information about the interactions among the cultivar, the rootstock, and nutrient content in grapevines is reduced, several studies have shown that rootstocks differ in their effect on the nutrient levels in the grafted cultivar

A.K. Srivastava, Chengxiao Hu (eds.) Fruit Crops: Diagnosis and Management of Nutrient Constraints https://doi.org/10.1016/B978-0-12-818732-6.00021-6

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© 2020 Elsevier Inc. All rights reserved.

298

21. Rootstock: Scion combinations and nutrient uptake in grapevines

(Grant and Matthews, 1996; Nikolaou et al., 2000; Garcia et al., 2001; Bavaresco et al., 2003; Fisarakis et al., 2004; Robinson, 2005). To select the type of rootstock to be used in a specific edaphoclimatic condition, it is important to carry out local long-term studies, given the interactions that exist among the rootstock, the cultivar, and the environment. This implies that the results obtained with a particular cultivar-rootstock combination in a particular environment cannot be extrapolated to other situations (Keller et al., 2001).

2 Characteristics of rootstocks associated with the uptake of water and nutrients Root system is the interface between the grapevine and the soil. It provides anchorage for grapevine in the soil and is responsible for water uptake and nutrient mining, therefore enabling combination of genotypes and cultivated scion to express the productive potential and attributes of the target cultivar. The roots also serve as storage organs for carbohydrates and other nutrients, which support the initial growth of scion and roots in spring, and for water. In addition, they are a source of plant hormones (cytokinins and ABA), which modify shoot physiology. Commonly used rootstocks are either individual Vitis species or crosses of two or more species, which harbors traits of agronomic relevance in grapevine rootstocks for grapevine production (Table 21.1, adapted after Keller, 2015). The practice of rootstocks on grapevines is a technological tool widely used, where about 10 rootstocks occupy 90% of grafted grapevines in the world (Keller, 2015). Despite this limited genetic diversity, new rootstock breeding programs have been developed in recent years with the aim of not only expanding frontiers of grapevine production, using marginal soils, dealing with biotic constrains (root parasites, such as phylloxera and nematodes), and higher efficiency in the use of water and nutrients (Tortosa et al., 2016; Bianchi et al., 2018; Romero et al., 2018), but also for their ability to influence crop maturity or their tolerance of adverse abiotic soil conditions such as waterlogging, lime, acid or saline soils, and excess of toxic mineral elements (Walker et al., 2018). Indeed, a major goal is to develop rootstocks that can influence scion growth and productivity under drought, particularly those that can increase water conservation through reducing the need for irrigation while ameliorating negative impacts on yields (Zhang et al., 2016). Therefore, grapevine is grown in a great diversity of geographical areas in the world, with a wide variety of cultivars and productive purposes (table grapes, wine, spirits, raisins, etc.). Notwithstanding grafting does not directly affect the quality-relevant traits of the grapevines produced by the scion cultivar, because the rootstock to which a scion cultivar is grafted may alter water and nutrient uptake and distribution, plant growth, and yield formation, it seems logical to expect the rootstock to influence fruit composition too (Keller, 2015). Thus, an indirect effect on fruit composition, especially on acidity, is possible due to the potential influence of the rootstock on scion vigor, canopy configuration, and yield components (Keller et al., 2001; Ruhl and Alleweldt, 1990). Nevertheless, the rootstock may alter amino acids, especially arginine, in the berries of its grafting partner. Reports had determined that 140 Ruggeri and 101-14 Mgt may sometimes lead to considerably lower amino acid concentrations in fruits of their grafting partner than some other rootstocks and own-rooted vines (Treeby et al., 1998).

3 Rootstocks differences on the uptake of macro and micronutrients The vineyard fertilization is a key management within the productive cycle, determining the yield and quality of the production. To carry out an adequate fertilization of macro- and micronutrients, information about the soil and climatic conditions, quality of the irrigation water, and the combination of cultivar rootstock should be considered. As already mentioned, rootstocks differ in the capacity to capture water and nutrients, and this can be affected by the cultivar used and environmental conditions (Ibacache and Sierra, 2009; Pachnowska and Ochmian, 2018). Although there are rootstock comparative tables in the literature and their effect on the absorption of few nutrients (Table 21.1; Keller, 2015), the aim is to present an update of the recent studies, which have been carried out in new geographical areas and the incorporation of new rootstocks and cultivars. This information can serve as a guide to establish guidelines for fertilization, considering the rootstock employed, increasing efficiency in the fertilizer use, reducing costs, and avoiding environmental contamination.

TABLE 21.1 Traits of agronomic relevance in grapevine rootstocks. Resistance Rootstock

Parent species

Phyll

Nem

Riparia Gloire

V. riparia

E

P

Rupestris St. George

V. rupestris

H

P

Rupestris du Lot

V. rupestris

420A Millardet et de Grasset

V. berlandieri
V. riparia

H

P

5BB Kober

V. berlandieri
V. riparia

H

P

SO4

V. berlandieri
V. riparia

H

P

8B

V. berlandieri
V. riparia

H

5C Teleki

V. berlandieri
V. riparia

161-49 Couderc

CGa

Tolerance Phyt

Lime

Salty

P

L

L

P

P

L

M

P

L

P

M

L

M

P

L

P

L

P

P

P

H

H

L

P

P

M

P

H

V. berlandieri
V. riparia

E

P

L

M

L

P

P

M

99 Richter

V. berlandieri
V. rupestris

H

P

L

M

P

L

L

M

P

Y

Y

H

110 Richter

V. berlandieri
V. rupestris

H

P

L

L

E

P

P

M

M

M

Y

Y

1103 Paulsen

V. berlandieri
V. rupestris

H

P

P

P

M

P

H

P

M

M

N

140 Ruggeri

V. berlandieri
V. rupestris

H

L

L

P

H

M

H

H

M

H

N

M L

P

L

P

M

ClayS

SandS

Mg-Def

L

P

N

L

L

P

L

M

P

P

L

P

L

K-Def

Influenced traits

Droug

L

AcidS

Susceptibility SciV

Mat

B-Graft

RootA

L

A

P

M

H

D

H

D

E

Y

P

D

P

L

Y

Y

P

D

P

P

Y

N

M

P

P

M

L

N

A

M

P

P

L

L

D

E

P

M

D

H

L

Y

M

D

M

M

Y

H

D

M

P Continued

TABLE 21.1 Traits of agronomic relevance in grapevine rootstocks.—cont’d Resistance Rootstock

Parent species

Phyll

Nem

CGa

44-53 Malègue

V. riparia
V. rupestris

H

M

3309 Couderc

V. riparia
V. rupestris

H

L

H

101-14 Millardet et de Grasset

V. riparia
V. rupestris

H

P

H

Schwarzmann

V. riparia
V. rupestris

H

Gravesac

V. berl.
V. rip.
V. rupestris

1616 Couderc

Tolerance Droug

Lime

H

L

L

P

L

L

P

P

L

M

M

P

P

M

H

L

M

P

V. solonis
V. riparia

M

L

L

M

L

Salt Creek (Ramsey)

V. champini

P

H

H

P

H

DogRidge

V. champini

P

H

H

P

Harmony

V. champini
V. sol.
V. riparia

P

H

H

Freedom

V. champini
V. sol.
V. riparia

P

H

H

L

Phyt

Salty

Susceptibility

Influenced traits

AcidS

ClayS

SandS

Mg-Def

K-Def

SciV

Mat

B-Graft

RootA

M

M

L

Y

N

M

A

E

H

P

P

N

P

A

P

M

P

L

M

A

P

M

M

H

M

P

M

M

L

M

M

H M

L

P

P

L

H

H

L

L

H

L

H

H

L

L

P

H

L

P

P

M

H

P

H

L

M

P

M

M

L

A

Scale: A, advanced; D, delayed; E, excellent; H, high; L, low; M, intermediate; N, absent; P, poor; Y, present. AcidS, acid soil; B-Graft, bench grafting; CGa, crown gall; ClayS, clay soil; Droug, drought; K-Def, K deficiency; Lime, lime; Mat, maturation; Mg-Def, Mg deficiency; Nem, nematode; Phyll, phylloxera; Phyt, phytophthora; RootA, rooting ability; Salty, salinity; SandS, sandy soil; SciV, scion vigor. Modified from Keller, M., 2015. The Science of Grapevines, second ed. Elsevier. 491 p.

3 Rootstocks differences on the uptake of macro and micronutrients

301

3.1 Macronutrients Nitrogen is the most abundant soil-derived macronutrient in grapevine, influencing plant growth and playing a major role in most of the biological functions and processes of both grapevine and fermentative microorganisms (Bell and Henschke, 2005). On the other hand, the nutritional status of the grapevine with respect to nitrogen is associated with the quality of wine or must and physiological disorders, mainly due to problems of vegetative balance. Viticultural practices aimed at reaching nitrogen requirements are of special relevance, because interactions between rootstock and vineyard supply strongly influence scion mineral nutrient status and shoot vigor and, via those processes, fruit composition (Holzapfel and Treeby, 2007). In this regard, it is important to carry out an adequate nitrogen fertilization to each productive system (table grapes, wine, etc.), considering from the vineyard design the rootstock to stablish. Table 21.2 shows rootstock-scion combinations that have displayed a higher absorption of nitrogen and phosphorus, compared with other rootstocks in each cited study. In addition, the rootstock parentals and type of study and tissue analyzed are mentioned. There are nine rootstocks that presented a higher nitrogen absorption compared with own-rooted plants and other rootstocks (statistically significant), according to field studies that have been conducted in the last 18 years. Frequently, more vigorous rootstocks result in higher N levels on the grafted variety. Thus, when a cultivar is grafted onto rootstocks that exhibit high nitrogen uptake, fertilization programs for this element should be adjusted. Excessive applications of nitrogen can have negative effects on the environment (contaminating ground water with nitrates) and increasing pest and diseases damage, physiological disorders, and reducing fruit quality (Keller et al., 2001). Phosphorus (P) is considered as a major limiting factor of crop production. Phosphorus deficiency is currently mitigated by P fertilizer application. However, P fertilizers are mainly produced from nonrenewable phosphate rock, and concerns have been expressed that this natural resource will be exhausted in the near future (Gautier et al., 2018). In Table 21.2, seven rootstocks that showed a higher absorption of phosphorus compared with own-rooted plants and other rootstocks are shown (statistically significant); thus, most of studies were carried out in the field. It is important to underline that some authors showed that own-rooted plants have a higher absorption of this element with respect to rootstocks (Vijaya and Rao, 2015). Information regarding the macronutrients, potassium (K), magnesium (Mg), and calcium (Ca), is presented in Table 21.3. Potassium corresponds to major cation in grape juice (Mpelasoka et al., 2003). A high concentration of K in grape juice can lead to high juice pH (e.g., >3.8) and, in turn, lower quality wines, for example, reduced color stability and poor taste (Kodur, 2011). Potential vineyard management options to manipulate berry K accumulation include selective use of rootstock/scion combination, canopy management, and irrigation strategies. With respect to rootstocks, the use of low K accumulating rootstocks can be an alternative to avoid quality problems in wine, associated with a high K concentration. Sixteen rootstocks that have shown a higher K absorption compared with own-rooted plants and other rootstocks are shown (Table 21.3). It is important to remark that potassium-magnesium and potassium-calcium cations may present antagonism in their uptake, and therefore, rootstocks with a high capacity to absorb potassium may have a low absorption of Mg-Ca or vice versa. This aspect should be considered since a normal fertilization, without differentiating by established rootstock can increase deficiencies of magnesium and/or calcium. For example, SO4 rootstock presents a high absorption of K, but low Mg and Ca (Table 21.3). In this fashion, it is important to consider the balance between these cations to avoid deficiencies. According to Delas and Pouget (1979), an adequate range for K/Mg in grape petioles at veraison is between 3 and 7. Values below 1 indicate K deficiency and above 10, Mg deficiency. Leaf symptoms of cv. Muscat of Alexandria grafted on 110 Richter with magnesium deficiency are shown (Fig. 21.1). The leaves with symptoms had 45% less petiole magnesium content than normal leaves. A yellowing of the basal leaves is observed, keeping the green vein (intervening chlorosis) together with burns in the leaf margin (Fig. 21.1). On the other hand, magnesium deficiency is related to the development of the physiological disorder called “bunch stem necrosis” that may develop during ripening (Keller, 2015). Information regarding micronutrients is presented (Table 21.4). Micronutrients such as zinc, boron, and molybdenum and the macronutrient calcium are essential for the process of pollination and fertilization in grapevines. Kidman et al. (2014) found that rootstocks affect the sequestration of nutrients, which affected reproductive performance on the cv. Syrah. Specifically, Kidman et al. (2014) showed that 1103 Paulsen rootstock had a significantly higher amount of boron and a lower number of seedless berries and a lower millerandage index. On the other hand, Zn deficiency was observed for 110 Richter and 140 Ruggeri. Finally, it is important to remark that tables presented should be used with caution, since site-cultivar-specific responses can be generated in each study. Therefore, the results may not be strictly extrapolated to other edaphic-climatic conditions or cultivars.

302

21. Rootstock: Scion combinations and nutrient uptake in grapevines

TABLE 21.2

Rootstock–scion combination that has presented a higher absorption of nitrogen and phosphorus, compared with other rootstocks in each mentioned study.

Nutrient

Rootstock

Nitrogen (N)

Phosphorus (P)

a b

Parentage

Cultivars

Reference

Type of study/analysis

- Salt creek

- V. champini

- Flame Seedless - Thompson Seedless - Superior Seedless - Red Globe

Ibacache and Sierra (2009)

Field/petioles at flowering

- 420A - 110 Richterb - 1103 Paulsenb - 99 Richterb

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

Nikolaou et al. (2000)

Field/blades at veraison

- 161-49 C - Sori

- V. riparia
V. berlandieri - V. solonis
V. riparia

- Regent

Pachnowska and Ochmian (2018)

Field/leaves and berries at fruit set

- 140 Ruggeri

- V. berlandieri
V. rupestris

- Chardonnay - Pinot Noir

Wooldridge et al. (2010)

Field/petioles at fruit set

- IAC 572

-V. caribaea
101-14

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

- Salt Creek

- V. champini

- Flame Seedless - Thompson Seedless - Superior Seedless - Red Globe

Ibacache and Sierra (2009)

Field/petioles at flowering

- 125 A Kober - B€ orner

- V. berlandieri
V. riparia - V. riparia 183 GM
V. cinerea

- Regent

Pachnowska and Ochmian (2018)

Field/leaves at fruit set

- Harmony

- Couderc 1613
V. champini

- Pinot Noir

Candolfi and Castagnoli (1997)

Field/petioles at flowering

- Thompson Seedless

Morales et al. (2014)

Glasshouse/petioles at veraison

a

- R99

- V. berlandieri
V. rupestris

- Niagara Rosada - Concord

Dalbó et al. (2011)

Field/petioles at veraison

- 140 Ruggeri

- V. berlandieri
V. rupestris

- Chardonnay - Pinot Noir

Wooldridge et al. (2010)

Field/petioles at fruit set

- IAC 572

- V. caribaea
10114

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

Syn. Ramsey. Corresponding to V. berlandieri
V. rupestris.

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation High yields and grapevine quality are best achieved with soils and irrigation water that have optimal levels of salinity and the correct composition of salts. In regions with lower rainfall, where leaching of soluble salts is often incomplete, soil salinity can be a serious constraint to grape production. Field problems often associated with salinity include decreased soil-water availability, and accumulation of specific elements like chloride, sodium and boron that lead to toxic levels in plant tissues. Grapevines are classified as moderately sensitive to salinity (Walker, 1994) and sensitive to boron excess (Christensen et al., 1978). Depending on concentration, salt accumulation in the medium may lead to a poor growth and yield performance of the vines (Walker et al., 1997, 2014; Zhang et al., 2002), and high concentrations may even cause death of plants

303

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation

TABLE 21.3 Rootstock-scion combination that has presented a higher absorption of potassium, magnesium, and calcium, compared with other rootstocks in each mentioned study. Nutrient

Rootstock

Parentage

Cultivars

Reference

Type of study/analysis

Potassium (K)

- Harmony - 1613C

- Couderc 1613
V. champini - (V. solonis
V. vinifera)
(V. labrusca
V. riparia)

- Flame Seedless - Thompson Seedless - Red Globe

Ibacache and Sierra (2009)

Field/petioles at flowering

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

Nikolaou et al. (2000)

Field/petioles at veraison

- Dogridge

- V. champini

- Thompson Seedless

Vijaya and Rao (2015)

Field/petioles at flowering

- SO4b

- V. berlandieri
V. riparia

- N
egrette

Garcia et al. (2001)

Greenhouse/blades at veraison

- Chardonnay - Pinot Noir

Wooldridge et al. (2010)

Field/petioles at fruit set

-

Magnesium (Mg)

- Sori

- V. solonis
V. riparia

- Regent

Pachnowska and Ochmian (2018)

Field/leaves and berries at fruit set

- 44-53c

- Riparia grand glabre
144M

- Pinot Noir

Candolfi and Castagnoli (1997)

Field/petioles at flowering

- 140 Ruggeri - 101-14 - Saint George

- V. berlandieri
V. rupestris - V. riparia
V. rupestris - V. rupestris

- Chardonnay - Shiraz

Walker and Blackmore (2012)

Field/petioles at flowering

- 043-43c

- V. vinifera
V. rotundifolia

- Niagara Rosada - Concord

Dalbó et al. (2011)

Field/petioles at veraison

- IAC 766

- V. caribaea
Traviú (106-8)

- Niagara Rosada

Dalbó et al. (2011)

Field/petioles at veraison

- IAC 572

- V. caribaea
101-14

Tecchio et al. (2014)

Field/branches at pruning

- 3309 C

- V. riparia
V. rupestris

- N
egrette

Garcia et al. (2001)

Greenhouse/blades at veraison

- Regent

Pachnowska and Ochmian (2018)

Field/leaves at fruit set

-

Sori 125A Kober B€ orner 5BB Kober

-

V. solonis
V. riparia V. berlandieri
V. riparia V. riparia 183 GM
V. cinerea V. riparia
V. berlandieri

- Harmony

- Couderc 1613
V. champini

- Pinot Noir

Candolfi and Castagnoli (1997)

Field/petioles at flowering

- 140 Ruggeri

- V. berlandieri
V. rupestris

- Chardonnay - Pinot Noir

Wooldridge et al. (2010)

Field/petioles at fruit set

- -IAC 572

- V. caribaea
101-14

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

- Isabellad

- V. labrusca

- Niagara Rosada

Dalbó et al. (2011)

Field/petioles at veraison

- IAC766 - 420-Ad

- V. caribaea
Traviú (106-8) - V. berlandieri
V. riparia

- Concord

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

Nikolaou et al. (2000)

Field/petioles at veraison

- 3309 C

- Riparia tomenteuse
Rupestris Martin

- N
egrette

Garcia et al. (2001)

Greenhouse/blades at veraison

- Sori

- V. solonis
V. riparia

- Regent

Calcium (Ca)

420A 110 Richtera 1103 Paulsena 99 Richtera

-420A 110 Richtera 1103 Paulsena 99 Richtera

Field/leaves at fruit set Continued

304

21. Rootstock: Scion combinations and nutrient uptake in grapevines

TABLE 21.3 Rootstock-scion combination that has presented a higher absorption of potassium, magnesium, and calcium, compared with other rootstocks in each mentioned study.—cont’d Nutrient

Rootstock -

b c d

-

Cultivars

V. berlandieri
V. riparia V. riparia 183 GM
V. cinerea V. riparia
V. berlandieri V. riparia
V. berlandieri

Reference

Type of study/analysis

Pachnowska and Ochmian (2018)

- 140 Ruggeri

- V. berlandieri
V. rupestris

- Chardonnay - Pinot Noir

Wooldridge et al. (2010)

Field/petioles at fruit set

- IAC 572

- V. caribaea
101-14

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

-Nikolaou et al. (2000)

Field/blades at veraison

a

125A Kober B€ orner 5BB Kober 161-49 C

Parentage

420A 110 Richtera 1103 Paulsena 99 Richtera

Corresponding to V. berlandieri
V. rupestris. Less magnesium and calcium absorption were recorded. Less magnesium absorption was recorded. Less potassium absorption was recorded.

(Troncoso et al., 1999). Physiological effects to salinity exposure in grapevine include reduced stomatal conductance and photosynthesis systemic disturbances, which can lead to reductions in growth, biomass accumulation, and yield (Downton, 1977a; Ben-Asher et al., 2006; Walker et al., 2004). The inhibition of grapevine growth and CO2 assimilation by high salinity is mainly due to changes in stomatal conductance, electron transport rate, leaf water potential, chlorophyll, fluorescence, osmotic potential, and leaf ion concentration (Stevens and Harvey, 1995; Cramer et al., 2007). Among the strategies adopted for sustaining growth and productivity of vine cultivars under salinity, the use of tolerant rootstocks is widely accepted (Walker, 1994). Moreover, in areas affected by salinity constrains one of the more important uses of rootstocks is for modifying fruit composition, because rootstocks can exclude much of the salt dissolved in the soil solution from root uptake and xylem transport, the scions grafted to them accumulate less Na+ and Cl in the fruit (Walker et al., 2000, 2010). Susceptibility or rootstock tolerance to high salinity is a coordinated action of multiple factors. Salinity tolerance mediated by rootstocks is attributed to root system restricting the movement and/or limiting absorption and accumulation of toxic ions from saline soils (Hepaksoy et al., 2006; Walker et al., 2002). Fisarakis et al. (2001) showed that there is a great variability in the uptake and accumulation of Na+ and Cl exclusion among rootstocks. Specifically, they demonstrated that Vitis berlandieri species had a great ability for Cl and/or Na+ exclusion, although this ability is reduced in hybrids having V. vinifera as parent. On the other hand, Walker et al. (2004) working with field-grown vines concluded that a high innate vigor of a rootstock combined with moderate to high chloride and sodium exclusion ability represents the best combination for salt tolerance in Sultana grapevines as measured by yield at moderate to high salinity.

4.1 Chloride toxicity The visible symptoms that develop on vines growing under saline conditions are, in most cases, due to accumulation of chloride toxic concentrations in leaves. The symptoms appear first as marginal chlorosis on leaves, followed by necrosis developing inward from leaf margins. This effect is frequently referred to as “leaf burn” (Walker, 1994). Petiole Cl concentrations that exceed 1.0% in spring are considered excessive (Robinson, 2005). Petiole analysis has been shown to provide a good tool to assess Cl concentration in Cl-stressed vines (Christensen et al., 1978). In a long-term field study carried out under semiarid conditions of northern Chile, we determined the Cl, Na, and B petiole concentration in two cultivars, Flame Seedless (table grape) and Muscat of Alexandria (syn. Moscatel de Alejandría and Muscat Gordo Blanco), a cultivar that is used to produce a distilled spirit called Pisco in Chile. Both varieties were grafted onto 10 different rootstocks, 1613 Couderc, Freedom, Harmony, 1103 Paulsen, 110 Richter, 99 Richter, 140 Ruggeri, SO4, Salt Creek (syn. Ramsey), and Saint George, which were compared with own-rooted vines. The study was carried out in a slightly saline soil (electrical conductivity 1.0 dS/m in saturated paste) located at the Vicuña Experimental Center (30°020 S, 70°440 W) of the Instituto de Investigaciones Agropecuarias (INIA).

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation

305

FIG. 21.1 Leaf of Muscat of Alexandria grafted on 110 Richter with magnesium deficiency.

The rootstock effect on chloride levels in Flame Seedless and Muscat of Alexandria cultivars are shown (Figs. 21.2 and 21.3), respectively. Own-rooted vines accumulated the highest amount of petiole chloride compared with grafted vines in both cultivars. Among the rootstocks, Harmony accumulated more chloride than the others rootstock studied. The higher chloride concentration in petioles at full flowering stage of cv. Flame Seedless and cv. Muscat of Alexandria on own roots, relative to cultivars on the various rootstocks, reflects the poor capacity of V. vinifera vines for chloride exclusion (Downton, 1977). A number of studies have been undertaken comparing chloride exclusion ability of grapevine species and varieties (Walker, 1994). Downton (1977) ranked them as follows: Vitis rupestris < V. berlandieri, Vitis riparia < Vitis champini < V. vinifera. Also, Fisarakis et al. (2001) demonstrated that V. berlandieri species had a greater ability for chloride exclusion than hybrids having V. vinifera as parent. In our study, V. berlandieri
V. rupestris rootstocks, 99 Richter, 110 Richter, 140 Ruggeri, and 1103 Paulsen were all comparable with V. rupestris Saint George rootstock performance. According to Walker et al. (2010), 1103 Paulsen and 140 Ruggeri rootstocks were among the lowest petiole chloride concentration in cv. Shiraz at full flowering. 140 Ruggeri is also mentioned as a good chloride excluder (Walker et al., 2018). The greater chloride exclusion capacity of this rootstock appears to be associated with restricted entry of Cl to xylem and lower root-to-shoot Cl transport (Tregeagle et al., 2010). While there is uncertainty as to whether high chloride accumulation by certain cultivars is linked to growth reductions, there is a general agreement that grapevine chloride content is an important factor for vine health under saline conditions (Walker et al., 2004). Chloride exclusion has subsequently been regarded as an important rootstock trait and therefore commonly used as a screening test for salt tolerance in grapevine rootstock breeding programs (Antcliff et al., 1983; Newman and Antcliff, 1984; Sykes, 1985; Walker, 1994). Studies on grapevine salt tolerance have shown that high uptake and root-to-shoot transport of chloride, resulting in excessive accumulation in leaf tissues, is a major factor in impaired leaf function and damage (Walker et al., 1997). Chloride-excluding rootstocks have similarly been thought to protect against excessive chloride accumulation and therefore contribute to salt tolerance (Walker et al., 2002).

306

21. Rootstock: Scion combinations and nutrient uptake in grapevines

TABLE 21.4 Rootstock-scion combination that has presented a higher absorption of micronutrients, compared with other rootstocks in each mentioned study. Nutrient

Rootstock

Parentage

Cultivars

Reference

Type of study/analysis

Iron (Fe)

- 161-49 C

- V. riparia
V. berlandieri

- Regent

Pachnowska and Ochmian (2018)

Field/leaves at fruit set

- 41B - 8B Teleki

- V. vinifera
V. berlandieri - V. riparia
V. berlandieri

- Thompson Seedless

Nikolaou et al. (2000)

Field/petioles at veraison

- 1613 C - Salt Creek

- (V. solonis
V. vinifera)
(V. labrusca
V. riparia) - V. champini

Vijaya and Rao (2015)

Field/petioles at flowering

- 420A - 110 Richtera - 1103 Paulsena - 99 Richtera

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

Nikolaou et al. (2000)

Field/blades and petioles at veraison

- Sori - B€ orner - 5BB Kober

- V. solonis
V. riparia - V. riparia 183 GM
V. cinerea - V. riparia
V. berlandieri

- Regent

Pachnowska and Ochmian (2018)

Field/leaves and berries at fruit set

- IAC 572

- V. caribaea
101-14

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

- 420A - 110 Richtera - 1103 Paulsena - 99 Richtera

- V. riparia
V. berlandieri - V. berlandieri
V. rupestris

- Thompson Seedless

Nikolaou et al. (2000)

Field/petioles at veraison

- 125AA Kober - 5BB Kober

- V. berlandieri
V. riparia - V. riparia
V. berlandieri

- Regent

Pachnowska and Ochmian (2018)

Field/leaves at fruit set

- 44-53 - 3309 C

- Riparia grand glabre
144M - V. riparia
V. rupestris

- Pinot Noir

Candolfi and Castagnoli (1997)

Field/petioles at flowering

- 1103 Paulsen - 110 Richter

- V. berlandieri
V. rupestris

- Shiraz

Kidman et al. (2014)

Field/petioles at flowering

- IAC 572

- V. caribaea
101-14

- Niagara Rosada

Tecchio et al. (2014)

Field/branches at pruning

- 41B - 8B Teleki

- V. vinifera
V. berlandieri - V. riparia
V. berlandieri

- Thompson Seedless

Nikolaou et al. (2000)

Field/blades and petioles at veraison

- 1613 C

- (V. solonis
V. vinifera)
(V. labrusca
V. riparia)

Vijaya and Rao (2015)

Field/petioles at flowering

- 5BB Kober

- V. riparia
V. berlandieri

Pachnowska and Ochmian (2018)

Field/leaves at fruit set

Zinc (Zn)

Copper (Cu)

Boron (B)

Manganese (Mn)

a

Corresponding to V. berlandieri
V. rupestris.

- Regent

307

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation

0.7

c

Chloride (%)

0.6 0.5

b

0.4 0.3

a

a

a

0.2

a

a

a a

a

a

0.1 0 1613 Couderc

Own roots

Freedom Harmony 1103

110 Paulsen Richter

99 140 Richter Ruggeri

SO4

Salt Creek

Saint George

Rootstocks

FIG. 21.2 Chloride concentration in petioles from cv. Flame Seedless on various rootstocks. Each bar represents the average of 4 years (2006–09) standard error. Different letters denote significant differences (P < .05).

0.7

d

Chloride (%)

0.6 0.5 0.4 0.3

bc

c

abc a

0.2

a

a

a

ab

ab

a

0.1 0 1613 Own Couderc roots

Freedom Harmony 1103 110 99 140 Paulsen Richter Richter Ruggeri

SO4

Salt Creek

Saint George

Rootstocks

FIG. 21.3 Chloride concentration in petioles from cv. Muscat of Alexandria on various rootstocks. Each bar represents the average of 4 years (2006–09) standard error. Different letters denote significant differences (P < .05).

4.2 Sodium toxicity The principal effects of sodium (Na) excess in grapevines are driven by soil physical and permeability difficulties. However, grapevines can also accumulate fairly high levels of Na from highly sodic soils. The direct effects of Na excess in plant tissue are not always clear, because excess Na is most commonly associated with excess chloride uptake as well (Christensen et al., 1978). Dag et al. (2015) showed that more sodium than chloride was accumulated in woody tissue of 5-year-old Cabernet Sauvignon grapevines, evaluated under three irrigation salinity levels. As a result, mortality rates as high as 17.5% were found for poor salt-excluding rootstocks irrigated with the highest salty water. The apparent collapse of tolerance mechanisms, leading to salt damage and vine mortality, might be due to sodium reaching critical levels in woody tissues. This contradicts earlier findings of Ehlig (1960), who concluded that chloride was the dominant cause of salt toxicity in grapevines. The prevalence of sodium as the governing factor in salinity damage has been suggested previously for Fisarakis et al. (2001) and Stevens et al. (2011), who similarly found stronger long-term accumulation of sodium than chloride in Colombard vines grafted onto the chloride-excluding Ramsey rootstock. It is well known that an excess of Na causes a decrease of other cations due to an ionic antagonism (Troncoso et al., 1999). Downton (1985) showed a Na-Ca antagonism in grapevines. Garcia and Charbaji (1993) related that Cabernet Sauvignon vines grown hydroponically and using a standard nutrient solution with various NaCl doses brought about an increase in the NaCl content in all plant vegetative organs. Thus, Na-K antagonism was shown by the decrease in K content, even at low NaCl doses. The Ca and Mg contents of the different plant organs were also decreased along with NaCl content that was increased in the nutrient solution. Also, Upreti and Murti (2010)

308

21. Rootstock: Scion combinations and nutrient uptake in grapevines

concluded that grape rootstocks exhibited considerable variations in salinity tolerance as evidence from changes in Na and K contents and their corresponding ratios. In our study carried out in northern Chile, sodium concentration in petioles of cv. Flame Seedless (Fig. 21.4) and cv. Muscat of Alexandria (Fig. 21.5) were comparatively higher in own-rooted vines compared with grafted vines on various rootstocks. There were nonsignificant differences in Na concentration among the rootstocks examined. These results are in agreement with Walker et al. (2004, 2010), who showed that own-rooted vines contained higher petiole Na concentrations than grafted vines. Rootstocks may also affect berry sodium concentrations. Berries from own-rooted Shiraz vines contain two- to threefold Na concentrations higher than grafted vines, but the difference was much less for Cabernet Sauvignon (Downton, 1977). Concentrations in petioles at flowering greater than 0.5% are regarded as excessive (Robinson, 2005). Regarding Cl-Na accumulation ratio in grapevines, Kuiper (1968) reported an inverse relation between chloride and sodium transport capacities by grape root lipids, suggesting that transport chloride readily should restrict sodium transport and vice versa. Walker et al. (2004) found that chloride concentrations in petioles of Sultana vines were higher than sodium concentrations by 1.3–22.1-fold at flowering stage. In the same way, chloride concentrations in laminae were higher than sodium concentrations by 1.6–25.2-fold at harvest stage. The same trend was found by Dag et al. (2015) who showed that Na concentrations in leaves of 5-year-old cv. Cabernet Sauvignon were much lower than Cl concentrations. There is often a negative correlation between Na and Cl concentration in the leaves. Walker et al. (2004) say that for any given treatment combination, Cl-ion concentration in plant parts are generally higher than

0.16 0.14

Sodium (%)

0.12 0.1 0.08

b

0.06 0.04

a

a

a

a

a

a

a

a

a

SO4

Salt Creek

a

0.02 0

1613 Own Couderc roots

Freedom Harmony 1103 110 99 140 Paulsen Richter Richter Ruggeri

Saint George

Rootstocks FIG. 21.4

Sodium concentration in petioles of cv. Flame Seedless on various rootstocks. Each bar represents the average of 4 years (2006–09) standard error. Different letters denote significant differences (P < .05).

0.16

b

0.14

Sodium (%)

0.12 0.1 0.08 0.06 0.04

a

a

a

a

a

a

a

a

a

SO4

Salt Creek

a

0.02 0 1613 Own Couderc roots

Freedom Harmony 1103 110 99 140 Paulsen Richter Richter Ruggeri

Saint George

Rootstocks

FIG. 21.5

Sodium concentration in petioles from cv. Muscat of Alexandria on various rootstocks. Each bar represents the average of 4 years (2006–09) standard error. Different letters denote significant differences (P < .05).

4 Rootstocks differences in chloride (Cl), sodium (Na), and boron (B) accumulation

309

the corresponding Na-ion concentrations. In making those comparisons, note the ordinate scale differences between Cl and Na in (Figs. 21.2–21.5).

4.3 Boron toxicity Boron toxicity is an important nutritional disorder in arid and semiarid environments. Boron excess can be avoided simply by not planting in affected areas. Grapevines should never be planted where the irrigation water source contains 1 ppm B and above. Even with supply of good water quality, vineyard establishment should be delayed until soil B levels are lowered by leaching to near 1 ppm or below (Christensen et al., 1978). Symptoms of B toxicity are characterized as chlorosis and necrosis of leaves beginning at their margins, reduced leaf size, and reduced internodal distance between adjacent leaves (Yermiyahu and Ben-Gal, 2006). Boron tends to accumulate along the leaf margins until the concentration becomes sufficiently high to be toxic to leaf tissues (Christensen et al., 1978) (Fig. 21.6). Boron toxicity symptoms are commonly observed during the vegetative growing periods of grapevines, especially at the end of the seasons (Yermiyahu and Ben-Gal, 2006). Christensen et al. (1978) and Robinson (2005) mention that values of boron greater than 150 mg/kg in blades taken at flowering are indicative of B toxicity. There is little information about the influence of rootstocks on B uptake in grapevines. Vines of cv. Sugraone grafted on Ramsey and Ruggeri rootstocks and irrigated at two salinity levels did not show a significant difference in the accumulation of boron in leaves (Yermiyahu et al., 2007). Table grapes are commonly grafted onto rootstocks developed for their hardiness or tolerance to environmental conditions. Both Ramsey (V. champini) and 140 Ruggeri (V. berlandieri
V. rupestris) are rootstocks that have shown relatively high tolerance to conditions of root-zone salinity (Walker et al., 2002; Zhang et al., 2002) and are found in commercial vineyards. Data comparison for two rootstocks, 41B (V. vinifera
V. berlandieri) and 1103 Paulsen (V. berlandieri
V. rupestris), showed no differences in boron content in leaves (Soylemezoglu et al., 2009). The influence of rootstocks on B accumulation in the leaf laminae in cv. Flame Seedless and cv. Muscat of Alexandria grown in the arid zone of northern Chile is shown (Figs. 21.7 and 21.8, respectively). The leaf laminae are used to test B concentration in vines because B accumulates more in the blades (Christensen et al., 1978; Pech et al., 2013). Boron is considered highly immobile in most plants in that it is restricted to transpiration flow and accumulates in leaves (Nable et al., 1997), especially in leaf edges where necrotic specks may also develop along the leaf margins (Christensen et al., 1978). Own-rooted vines of cv. Flame Seedless accumulated less B than 1103 Paulsen, 110 Richter, and Salt Creek rootstocks (Fig. 21.7), and B content in vines of Muscat of Alexandria grafted on Salt Creek was significantly lower than vines grafted onto SO4 rootstock (Fig. 21.8). In general, B concentration in leaves of cv. Flame Seedless variety was higher than in Muscat of Alexandria vines, independent of the rootstock (Figs. 21.7 and 21.8) effect that was also demonstrated by Walker et al. (2014) for chloride and sodium accumulation in trunk wood and grape juice of cv. Chardonnay and cv. Shiraz over own roots and also over a range of rootstocks. FIG. 21.6 Boron toxicity symptoms along the leaf margins (at harvest stage, cv. Thompson Seedless, own rooted).

310

21. Rootstock: Scion combinations and nutrient uptake in grapevines

250

b

Boron (ppm)

200

ab

ab 150

b b

ab

ab

ab

ab

ab

a

100 50 0 1613 Own Couderc roots

Freedom Harmony 1103 110 99 140 Paulsen Richter Richter Ruggeri

SO4

Salt Creek

Saint George

Rootstocks

FIG. 21.7 Boron concentration in petioles from cv. Flame Seedless on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05). 250

Boron (ppm)

200 150

ab ab

100

ab

ab

ab ab

ab

ab

b

ab a

50 0 1613 Couderc

Own Freedom Harmony 1103 110 99 140 roots Paulsen Richter Richter Ruggeri

SO4 SaltCreek Saint George

Rootstocks

FIG. 21.8 Boron concentration in petioles from cv. Muscat of Alexandria on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05).

To determine the tolerance to boron, Pech et al. (2013) evaluated various Vitis species, including V. acerifolia, V. berlandieri, V. caribaea, V. champini, V. labrusca, V. rupestris, V. vinifera, and hybrids. They determined that concentration of B in leaves from boron treatments was equivalent within comparisons. However, genotypes V. acerifolia
(V. vinifera
[V. riparia
V. labrusca]) (1613 Couderc), V. acerifolia, and V. vinifera
V. rupestris (1202 Couderc) showed tolerance to excess B in terms of absolute and/or relative growth.

5 Carbohydrates and nitrogen reserves Both reserve carbohydrates and nitrogen are essential for the initial growth and development of grapevine in spring, as they provide energy and building blocks for the new growth before any net carbon assimilation and significant root uptake of nitrogen takes place (Cheng and Xia, 2004). The majority of the stored assimilates that are needed in the subsequent year for successful growth in the vineyard is accumulated in the roots (Zapata et al., 2004; Vrsic et al., 2009). According to Zapata et al. (2004), roots contained more than 90% of the carbohydrates and 75% of the nitrogen stored in the dormant Pinot noir vines. In the same fashion, Bates et al. (2002) found that roots were the major storage organ for carbohydrates and nutrients, accounting for 84% of starch and 75% of nitrogen stored in the vines at the season start. Ruhl and Alleweldt (1990) mention that in late summer, starch is major carbohydrate in roots and those concentrations of sugars other than glucose are low at that stage. In their study, root starch

311

6 Rootstock strategies to cope with salinity

e

40 35

Starch (%)

30

de de

cde

abcde abcd

de bcde

ab a

25

abc

20 15 10 5 0 1613 Freedom Harmony Couderc

Own roots

1103 110 Paulsen Richter

99 Richter

140 Ruggeri

SO4

Saint George

Salt Creek

Rootstocks

FIG. 21.9 Starch concentration in roots from cv. Flame Seedless on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05).

concentrations ranged from 11.0% to 26.3%. Zapata et al. (2004) found that root starch concentration at dormancy was 29% dry weight, four times higher than in other perennial tissues. Along with carbohydrates, nitrogen reserves play a crucial role in supporting early season growth in woody plants (Conradie, 1980). In grapevines, nitrogen reserves are made mainly by amino acids, mostly arginine (Kliewer, 1991). Total nitrogen in the form of proteins are free amino acids typically accounts for less than 2% of dry matter of a dormant vine (Xia and Cheng, 2004). Compared with starch storage, nitrogen was far less represented at dormancy (33% vs 1.6% of root dry weight, respectively), in cv. Pinot noir grafted onto SO4 rootstock (Zapata et al., 2004). In our long-term study conducted in arid zone of northern Chile, we determined the levels of reserve carbohydrates and nitrogen accumulated in roots of cv. Flame Seedless and cv. Muscat of Alexandria grafted onto various rootstocks (Fig. 21.9–21.12). The results of our study show that in cv. Flame Seedless 1103 Paulsen, 110 Richter, 99 Richter, and SO4 rootstocks accumulated more starch in roots than own-rooted vines and Harmony rootstock. However, there was no significant effect of rootstocks in starch root accumulation in cv. Muscat of Alexandria. Significant differences in arginine root accumulation were found in both grapevine cultivars. Salt Creek rootstock had almost three times more arginine in roots than 110 Richter in cv. Flame Seedless (3.23% and 1.11%, respectively). In this cultivar, neither of the rootstocks accumulated more arginine than the own-rooted vines. On the other hand, in cv. Muscat of Alexandria SO4, Salt Creek and Harmony rootstocks accumulated arginine 1.7-fold more, in average, than own-rooted vines and Saint George rootstock. Because there are scarce available data about the rootstock influences in reserves accumulation under field conditions, it was not possible to compare our information. In general, high yield and low leaf area per vine decreased the nitrogen concentration in roots. The leaf-fruit ratio, expressed as the “light-exposed leaf area per kg fruit,” substantially influenced the nitrogen and starch concentration in the roots (Zufferey et al., 2015).

6 Rootstock strategies to cope with salinity In semiarid and arid regions, the competition for scarce water resources inevitably reduces the supplies of fresh water for irrigation; thus, agriculture is forced to utilize low-quality water for irrigation, increasing the risks of soil salinization where about 6% of the world’s land is already affected by salinity and 20% of irrigated land (Ollat et al., 2016). Considering the context of global change, precipitation patterns are also likely to change with a reduction of the balance between precipitation and evapotranspiration, leading to an acceleration of salinization in dry (and drying) regions (Keller, 2010). As moderately sensitive to salinity, grapevine responses depend on several factors, such as the rootstock-scion combination, irrigation system, soil type, and climate (Ollat et al., 2016). It is well known that variations in salt exclusion exist between grapevine species and cultivars, and rootstocks are considered as one important means for improving grapevine salt tolerance (Sykes, 1985; Walker et al., 2010; Ollat et al., 2016). Physiological effects of exposure to salinity in grapevines include reduced stomatal conductance and photosynthesis (Ben-Asher et al., 2006; Downton, 1977b); systemic disturbances can lead to reduced growth, vegetative biomass

312

21. Rootstock: Scion combinations and nutrient uptake in grapevines

35

ns

30

Starch (%)

25 20 15 10 5 0 1613 Freedom Harmony Couderc

Own roots

1103 110 Paulsen Richter

99 Richter

140 Ruggeri

SO4

Saint George

Salt Creek

Rootstocks

FIG. 21.10

Starch concentration in roots from cv. Muscat of Alexandria on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05).

c

4 bc

3.5 Arginine (%)

2.5

bc

ab

3

abc ab

ab

1.5

ab

ab

ab

2

a

1 0.5 0 1613 Freedom Harmony Couderc

Own roots

1103 110 Paulsen Richter

99 Richter

140 Ruggeri

SO4

Saint George

Salt Creek

Rootstocks Arginine concentration in roots from cv. Flame Seedless on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05).

FIG. 21.11

3

b

Arginine (%)

2.5

ab

ab

b b

ab

ab

ab

ab

2

a

a

1.5 1 0.5 0 1613 Freedom Harmony Couderc

Own roots

1103 110 Paulsen Richter

99 Richter

140 Ruggeri

SO4

Saint George

Salt Creek

Rootstocks

FIG. 21.12 Arginine concentration in roots from cv. Muscat of Alexandria on various rootstocks. Each bar represents the average of 4 years (2006–09)  standard error. Different letters denote significant differences (P < .05).

6 Rootstock strategies to cope with salinity

313

(Shani and Ben-Gal, 2005), and yield (Walker et al., 2010; Zhang et al., 2002). Similar to other plants, grapevines have osmotic and toxicity-related responses to salinity, and tolerance mechanisms involve exclusion of salt uptake, accumulation and storage of ions in cellular compartments, and restriction of toxic ions in leaves (Shani and Ben-Gal, 2005). The differences in nutrient uptake and distribution may be explained in several ways. First, rootstocks may have different absorption capability or affinity for some specific nutrients and minerals. In this concern, Bavaresco et al. (1991) pointed out that rootstocks with lime tolerance have a strategy to overcome chlorosis with high root iron uptake and reducing capacity. Grant and Matthews (1996) observed that different rootstocks might have different abilities to absorb phosphorus. Ruhl (2000) also found a high potassium acquisition mechanism on some rootstocks, which would affect pH of fruit and wines. Second, translocation and nutrient distribution may differ among rootstocks. Giorgessi et al. (1997) found differences in number and size of the xylem vessels between rootstocks and own-rooted grapevines. Third, some nutrients might be assimilated mostly by roots, thus reducing the amount translocated to the shoots. Ruhl (1993) points out that rootstocks delivering low amounts of potassium to the cultivar accumulate majority of absorbed cations in the vacuole of root cells, and Keller et al. (2001) discovered that over 85% of nitrogen was assimilated by means of vine root metabolism. In all cases, further studies are required to understand the influence of the root system in mineral absorption (Nikolaou et al., 2000). Salt exclusion by roots is considered a main mechanism to contribute to tolerance, since previous studies have shown that high uptake and root-to-shoot transport of chloride resulted in its excessive accumulation in leaf tissues, causing impaired leaf function and damage (Walker et al., 2002). Chloride exclusion by roots may prevent chloride accumulation in leaves contributing to salt tolerance (Walker et al., 1997, 2000, 2002; Stevens et al., 2011; Fisarakis et al., 2001; Zhang et al., 2002). Indeed, a much lower chloride concentration was found in xylem sap and shoot tissue of potted grape vines when grafted on to a salt tolerant (140 Ru) than on to a salt-sensitive rootstock (K51-40). The ability of rootstock-scion combinations to inhibit salt accumulation in leaf tissue is used to categorize sensitivity to salinity. One management strategy for vineyards with saline irrigation water is to select rootstocks based on their ability to prevent salt uptake and accumulation. There is significant variation in scion Cl content among rootstocks (Walker et al., 2004, 2007). Dag et al. (2015) determined that apparent breakdown of tolerance mechanisms, leading to salt damage and vine mortality, might be due to Na-reaching critical levels in woody tissues. The ability to exclude Na and Cl from shoots and fruit was found to (a) increase wine quality by reducing concentrations of salt ions in must and wine and (b) reduce mortality rates that result from long-term exposure to salt, since Vitis spp. rootstocks can mediate salt exclusion from grafted V. vinifera scions enabling higher grapevine yields and production of superior wines with lower salt content. Recently, by using a cross between two Vitis interspecific hybrid rootstocks, Henderson et al. (2017) mapped a dominant quantitative trait locus (QTL) associated with leaf Na + exclusion (NaE) under salinity stress. The NaE locus encodes six high-affinity potassium transporters (HKT). Transcript profiling and functional characterization in heterologous systems identified VisHKT1;1 as the best candidate gene for controlling leaf Na + exclusion. The origin of the recessive VisHKT1;1 alleles was traced to V. champini and V. rupestris and therefore is possible to assist breeding Na+tolerant grapevine rootstocks through the genetic and functional data generated (Henderson et al., 2017). In the field, tolerance to salt may be defined as the ability to maintain shoot growth and yield under high-salinity conditions. Over time, it appears that vine vigor and intrinsic ability for salt tolerance are both required to maintain yield performance in saline environments (Walker et al., 2010). Moreover, as grapevine rootstocks differ widely in their ability to exclude chloride from the shoot and in their salinity tolerance, field experiments showed that Ramsey was one of the best performing salt-tolerant rootstocks combined with various scion varieties, soils, and climate conditions (summarized by Ollat et al., 2016). 140Ru was also considered as highly salinity tolerant, and differences among extreme genotypes are maintained in the field in a range of environments and salinities (Walker et al., 2010); nevertheless, large genotype–environment interactions have been reported (Gong et al., 2011), suggesting that chloride exclusion ability of certain rootstocks can vary with long-term exposure to salinity (Tregeagle et al., 2010). These variations could be related to the volume of irrigation, the salt concentration in the soil, the leaching conditions, and the evapotranspiration levels in each site. Finally, Sivritepe et al. (2010) observed a strong effect of scion varieties on the salt tolerance of grafted plants, reinforcing the notion of a terroir based cultivar/rootstock responses. Rootstocks have a key role in the grapevine response to the environment and in grape berry composition as they represent a physical and biological link between the soil and the aboveground part of the plant. Nevertheless, as they are the hidden half of the grapevine, our knowledge about their functioning is still very scarce (Ollat et al., 2016). The features involved in the influence of grapevine rootstocks on scion growth and nutrient uptake and the interactions between rootstock and scion in a grafted system are gathering renewed interest from scientific community. Since grafting is required in the cultivation of grapevine in most areas in the world, rootstocks have a wide range of impacts on scion behavior (summarized from Keller, 2015; Ollat et al., 2016). The study of rootstock-scion interaction is incredibly complex integrating structural changes at the graft interface, hydraulic integration, hormonal communication,

314

21. Rootstock: Scion combinations and nutrient uptake in grapevines

and even exchange of genetic materials (Zhang et al., 2016). It has been described that the effect of the interaction between the two genotypes is, in general, larger than the rootstock effect. By analogy with genotype-environment studies, the concept of plasticity was introduced to characterize the phenotypic variations produced by a genotype in response to grafting partners. There is a good amount of data describing the variability among rootstocks, but the mechanisms underlying their responses to the environment are still unknown. Also, further research is still needed to elucidate the genetic determinism of these traits and how rootstocks can be used to mitigate plant stress in the context of the ongoing climate changes. Indeed, Tandonnet et al. (2018) have identified key QTLs for traits assessed on field-grown grafted grapevines, where root number and section had the largest phenotypic variance explained. Genetic control of root and aerial traits was independent, when analyzing genetic architecture of root traits in a segregating progeny from an interspecific cross between V. vinifera cv. Cabernet Sauvignon
V. riparia cv. Gloire de Montpellier. Identified QTLs for aerialto-root biomass ratio suggested that aerial and root traits are controlled independently, opening new venues for breeding rootstocks with improved root development capacities. Moreover, studies concerning the influences of rootstocks on scion growth, fruit composition, or wine quality do not always produce consistent results, possibly due to experimental conditions (e.g., potted vs field and young vines vs old vines), soil type and/or climatic conditions, scion variety, etc. Metaanalysis studies could be useful to better understand and integrate the studies that have already been carried out. Further studies aimed at understanding the physiology and traits responsible for rootstock control over scion behavior might benefit by integrating different approaches (genetic, transcriptomic, metabolic, hydraulic, etc.) in the experimental approach. There is still a lot to be gained from investigations in ungrafted material creating a foundation of understanding regarding the differences between the rootstock genotypes themselves (Zhang et al., 2016). Finally, future research demands the application of new technologies in the vineyard and the integration of multidisciplinary approaches. One of the most common goals pursued by grapevine researchers is the development of new rootstock varieties that meet growers’ demands, especially in the context of climate change (Zhang et al., 2016). The complexity of the responses, putative mechanisms, and interactions with environment present significant challenges, but the rootstock breeding is critical for the development of new sustainable approaches to adapt and mitigate climate change effects in viticulture.

Acknowledgments The authors are grateful to their technical staff who perform a lot of experimental work related to rootstock studies, including María Isabel Rojas, Elizabeth Past
en, Carmen Jopia, and Cristián González for their valuable contribution in maintaining the experimental trials and technical support. They also thank the financial support of Instituto de Investigaciones Agropecuarias (INIA) (long-term rootstocks studies), FONDECYT Regular Grant no. 1140039 (to AZS and AI), and Postdoctoral FONDECYT 2018 N°3180252 (to NVV).

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