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Grafting Improves Tomato Salinity Tolerance Through Sodium

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Grafting Improves Tomato Salinity Tolerance Through Sodium HORTSCIENCE 48(7):855–862. 2013. several physiological and biochemical changes that inhibit plant growth and production and affect the fruit quality (Cuartero and Fernandez- Grafting Improves Tomato Salinity Mun˜oz, 1999). Graftinghasbeenshowntobeavalid Tolerance through Sodium Partitioning technique to enhance tomato plant growth and productivity (Colla et al., 2010; Estan˜ within the Shoot et al., 2005; Ferna´ndez-Garcı´a et al., 2004). Unlike traditional breeding, grafting enables Francesco Di Gioia and Angelo Signore1 the combination of desired high-yielding Dipartimento di Scienze agro-ambientali e territoriali, Universita` degli Studi and/or high-quality tomato genotypes, often di Bari ‘‘Aldo Moro,’’ via Amendola 165/A, 70126 Bari, Italy susceptible to SS, with rootstocks character- ized by higher vigor and, potentially, capable Francesco Serio of improving the scion tolerance to SS con- Istituto di Scienze delle Produzioni Alimentari, Consiglio Nazionale delle ditions (Bolarin et al., 1991; Estan˜ et al., 2009; Ghanem et al., 2011). Ricerche, via Amendola 122/O, 70126 Bari, Italy Enhanced tolerance to SS in grafted veg- etable crops has mainly been attributed to the Pietro Santamaria + ` Na exclusion (Edelstein et al., 2011; Estan˜ Dipartimento di Scienze agro-ambientali e territoriali, Universita degli Studi et al., 2005) and/or inclusion capacity in the di Bari ‘‘Aldo Moro,’’ via Amendola 165/A, 70126 Bari, Italy shoot and/or root vacuoles (Albacete et al., Additional index words. rootstock, sodium compartmentation, ionic homeostasis, salt stress, 2009; Edelstein et al., 2011). Instead, grafting and/or rootstocks seem to be less effective fruit quality in restricting and/or regulating Cl– transport Abstract. Two greenhouse experiments were carried out to analyze the shoot sodium and/or concentration in the shoot (Colla et al., (Na+) partitioning, yield, and fruit quality of ‘Cuore di Bue’, a salt-sensitive heirloom 2006; Edelstein et al., 2011; Savvas et al., tomato (Solanum lycopersicum L.), ungrafted or grafted onto interspecific tomato hybrid 2011). Nevertheless, the mechanism of Na+ rootstocks (S. lycopersicum 3 S. habrochaites) ‘Maxifort’ and ‘Arnold’ in 2009, ‘Arnold’ exclusion and/or inclusion in the shoot/root and ‘Armstrong’ in 2010, grown at different salinity stress (SS) levels (0, 20, and 40 mM of of grafted plants and the role played by the NaCl in 2009; 0 and 20 mM of NaCl in 2010). In both experiments, an interaction was scion and rootstock genotypes remain still observed between grafting combinations and SS levels in terms of fruit yield, and fruit unclear. juice Na+ content. Under no SS conditions, plant grafted onto ‘Maxifort’ and ‘Arm- In a recent study conducted on melon and strong’ provided the highest yield in 2009 and 2010 experiments, respectively. In the pumpkin with reciprocal grafting, Edelstein + presence of 20 mM of NaCl, plants grafted onto ‘Arnold’ provided a marketable yield et al. (2011) attributed the lower Na concen- 23.5% (on average) higher than plants grafted onto ‘Maxifort’ or ungrafted in 2009 and tration observed in plants grafted on pumpkin 33% (on average) higher than plants grafted onto ‘Armstrong’ or ungrafted in 2010. The rootstock mainly to the Na+ exclusion and further increase of SS to 40 mM of NaCl considerably reduced the productivity of all retention by the rootstock, rejecting the hy- + grafting combinations. At 20 mM of NaCl, plants grafted onto ‘Arnold’ showed also pothesis that 1) Na transport to the scion can a higher capacity to modulate shoot Na+ partitioning with respect to ungrafted plants by be restricted by grafting itself; and 2) the scion increasing Na+ accumulation in older leaves (52%) and reducing Na+ content in younger may play a role in Na+ partitioning within the and most active leaves (24%), thus enabling the maintenance of higher K+/Na+,Ca2+/Na+, plant. However, physiological studies suggest and Mg2+/Na+ ratios compared with ungrafted plants. Fruit total soluble solids content, that, in tomato and other solanaceous species, titratable acidity, and dry matter were unaffected by grafting at any SS level, whereas salinity tolerance (ST) is mainly related to under SS, the fruit juice Na+ content of grafted plants was consistently lower (from 19% aNa+ inclusion mechanism (Bolarin et al., up to 68%) than that of ungrafted plants. Under moderate SS conditions (20 mM of NaCl), 1991; Chen et al., 2003; Pe´rez-Alfocea et al., the use of rootstock genotypes such as ‘Arnold’ having a particular ability to reduce Na+ 1993) and is closely correlated to the ability accumulation in younger and most active leaves may increase tomato yield and enhance of the plant to regulate Na+ concentration in tomato nutritional value by reducing the fruit juice Na+ content. the leaf tissue (Cuartero and Fernandez-Mun˜oz, 1999; Sacher et al., 1983; Savvas et al., 2011) and to maintain a high K+/Na+ ratio. Al- As the use of tomato grafting is gaining specific abiotic stress conditions (King et al., though maintenance of higher K+/Na+ ratios popularity worldwide (Kubota et al., 2008; 2010; Savvas et al., 2010; Schwarz et al., in grafted plants grown under SS conditions Lee et al., 2010), breeders and seed compa- 2010). has often been associated with a higher K+ nies are eager to select and develop new Salinity is one of the most critical envi- uptake capacity of the rootstock (Albacete rootstocks characterized by high yield and ronmental stresses that limit agriculture world- et al., 2009; Leonardi and Giuffrida, 2006; fruit quality performance, multidisease resis- wide. Over 20% of irrigated land is affected Santa-Cruz et al., 2002), Na+ partitioning in tance (Barrett et al., 2012; Guan et al., 2012; by salinity (Rozema and Flowers, 2008), and young and mature leaves represents a key as- King et al., 2008), and possibly tolerance to although arid and semiarid zones are more pect of such regulation (Maggio et al., 2007; subject to this phenomenon, salinization is Shannon, 1997; Shannon et al., 1987). increasingly affecting also less extreme en- In the present study we hypothesized that vironments being strictly associated with the grafting onto vigorous interspecific hybrids Received for publication 6 Feb. 2013. Accepted for irrigation practice itself (Tanji and Kielen, (S. lycopersicum · S. habrochaites) might publication 17 May 2013. 2002). raise the ST of tomato also through the The research was funded by MIUR-PRIN 2007 and The detrimental effects of salinity on crops regulation of Na+ partitioning in young and 2009 (Ministry of Education, University and Re- are mainly related to 1) osmotic stress, result- mature leaves. In this regard, a complete search, Italy, Projects ‘‘Vegetable grafting: Bio- ing from the accumulation of high solute analysis of the Na+ partitioning within the physiological basis, effects on crop and product concentrations in the rooting zone; 2) ion- shoot portions of grafted plants is still miss- quality’’ and ‘‘Physiological response, growth, + yield, and quality of grafted tomato under com- specific toxicity, mainly caused by Na and ing in the literature and is required for a better – bined excess boron and salinity stress.’’ Cl ; and 3) ion imbalance in cells, especially comprehension of the rootstock–scion inter- 1 + 2+ 2– To whom reprint requests should be addressed; lower concentration of K ,Ca ,SO4 , and action under SS conditions and to also clarify – + e-mail [email protected]. NO3 (Giuffrida et al., 2009), leading to possible effects on fruit Na content and HORTSCIENCE VOL. 48(7) JULY 2013 855 quality (Davis et al., 2008; Rouphael et al., in the greenhouse was on average 21.0 °C, management approach was used to control all 2010). Although SS may improve the organ- and daily minimum and maximum air tem- major diseases and pests. oleptic quality of tomato fruits, it may also perature ranged from 5.0 to 26.0 °C and from Nutrient solution management. In both increase the fruit Na+ content, causing a loss 19.0 to 38.0 °C, respectively (Fig. 1A). In the experiments, the control nutrient solution of nutritional value given the notorious Na+ 2010 experiment, daily air temperature in (0 mM of NaCl) was prepared with deionized negative effects on human health (Karppanen the greenhouse was on average 19.5 °C, and water and contained: nitrogen (10 mM), phos- and Marvaala, 2006). daily minimum and maximum air tempera- phorus (2 mM), potassium (6 mM), magne- Therefore, the objectives of this study ture ranged from 4.5 to 22.5 °C and from 17.6 sium (2 mM), calcium (3 mM), iron (20 mM), were to analyze the shoot Na+ partitioning to 34.8 °C, respectively (Fig. 1B). Daily solar manganese (5 mM), zinc (2 mM), boron of grafted and ungrafted plants of ‘Cuore radiation was on average 356 W·m–2 in 2009 (25 mM), copper (0.5 mM), and molybdenum di Bue’, an ‘‘oxheart’’ heirloom tomato type and 353 W·m–2 in the 2010 experiment. Daily (0.1 mM), resulting in an electrical conduc- that is characterized by big-sized fruits and relative humidity was on average 77% and tivity (EC) of 1.8 dS·m–1. The saline nutrient considered to be particularly sensitive to SS, ranged between a minimum of 51% and a solutions had the same composition plus 20 and to evaluate the interaction between sa- maximum of 90%. and 40 mM of NaCl, resulting in an EC of 3.7 linity levels and different commercial tomato Plants were trained vertically to one stem and 5.6 dS·m–1, respectively. When needed rootstocks (claimed to be particularly vigor- around each plastic string using the V-shaped the nutrient solution pH was adjusted to 5.5 ous and tolerant to SS conditions) in terms of method and were topped at the sixth floral using 2 M H2SO4. In both experiments, plant Na+ content, yield, and fruit quality.
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