While much recent science has focused on understanding and exploiting root traits as new opportunities for crop improvement, the use of rootstocks has enhanced productivity of woody perennial crops for centuries. Grafting of vegetable crops has developed very quickly in the last 50 years, mainly to induce shoot vigour and to overcome soil-borne diseases in solanaceous and cucurbitaceous crops. In most cases, such progress has largely been due to empirical interactions between farmers, gardeners, and botanists, with limited insights into the underlying physiological mechanisms. Only during the last 20 years has science realized the potential of this old activity and studied the physiological and molecular mechanisms involved in rootstock×scion interactions, thereby not only explaining old phenomena but also developing new tools for crop improvement. Rootstocks can contribute to food security by: (i) increasing the yield potential of elite varieties; (ii) closing the yield gap under suboptimal growing conditions; (iii) decreasing the amount of chemical (pesticides and fertilizers) contaminants in the soil; (iv) increasing the efficiency of use of natural (water and soil) resources; (v) generating new useful genotypic variability (via epigenetics); and (vi) creating new products with improved quality. The potential of grafting is as broad as the genetic variability able to cross a potential incompatibility barrier between the rootstock and the scion. Therefore, understanding the mechanisms underlying the phenotypic variability resulting from rootstock×scion×environment interactions will certainly contribute to developing and exploiting rootstocks for food security.

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To determine whether root-supplied ABA alleviates saline stress, tomato (Solanum lycopersicum L. cv. Sugar Drop) was grafted onto two independent lines (NCED OE) overexpressing the SlNCED1 gene (9-cis-epoxycarotenoid dioxygenase) and wild type rootstocks. After 200 days of saline irrigation (EC = 3.5 dS m À1 ), plants with NCED OE rootstocks had 30% higher fruit yield, but decreased root biomass and lateral root development. Although NCED OE rootstocks upregulated ABA-signalling (AREB, ATHB12), ethylene-related (ACCs, ERFs), aquaporin (PIPs) and stress-related (TAS14, KIN, LEA) genes, downregulation of PYL ABA receptors and signalling components (WRKYs), ethylene synthesis (ACOs) and auxin-responsive factors occurred. Elevated SlNCED1 expression enhanced ABA levels in reproductive tissue while ABA catabolites accumulated in leaf and xylem sap suggesting homeostatic mechanisms. NCED OE also reduced xylem cytokinin transport to the shoot and stimulated foliar 2-isopentenyl adenine (iP) accumulation and phloem transport. Moreover, increased xylem GA 3 levels in growing fruit trusses were associated with enhanced reproductive growth. Improved photosynthesis without changes in stomatal conductance was consistent with reduced stress sensitivity and hormone-mediated alteration of leaf growth and mesophyll structure. Combined with increases in leaf nutrients and flavonoids, systemic changes in hormone balance could explain enhanced vigour, reproductive growth and yield under saline stress.

Selection and breeding of rootstocks that can tolerate low K supply may increase crop productivity in low fertility soils and reduce fertilizer application. However, the underlying physiological traits are still largely unknown. In this study, 16 contrasting recombinant inbred lines (RILs) derived from a cross between domestic and wild tomato species (Solanum lycopersicum × Solanum pimpinellifolium) have been used to analyse traits related to the rootstock-mediated induction of low (L, low shoot fresh weight) or high (H, high shoot fresh weight) vigor to a commercial F1 hybrid grown under control (6 mM, c) and low-K (1 mM, k). Based on hormonal and ionomic composition in the root xylem sap and the leaf nutritional status after long-term (7 weeks) exposure low-K supply, a model can be proposed to explain the rootstocks effects on shoot performance with the ethylene precursor aminocyclopropane-1-carboxylic acid (ACC) playing a pivotal negative role. The concentration of this hormone was higher in the low-vigor Lc and Lk rootstocks under both conditions, increased in the sensitive HcLk plants under low-K while it was reduced in the high-vigor Hk ones. Low ACC levels would promote the transport of K vs. Na in the vigorous Hk grafted plants. Along with K, Ca, and S, micronutrient uptake and transport were also activated in the tolerant Hk combinations under low-K. Additionally, an interconversion of trans-zeatin into trans-zeatin riboside would contribute to decrease ACC in the tolerant LcHk plants. The high vigor induced by the Hk plants can also be explained by an interaction of ACC with other hormones (cytokinins and salicylic, abscisic and jasmonic acids). Therefore, Hk rootstocks convert an elite tomato F1 cultivar into a (micro) nutrient-efficient phenotype, improving growth under reduced K fertilization.

To determine whether root-supplied ABA alleviates saline stress, tomato (Solanum lycopersicum L. cv. Sugar Drop) was grafted onto two independent lines overexpressing the SlNCED1 (9-cis-epoxycarotenoid dioxygenase) gene (NCED OE) and wild type rootstocks. After 200 days of salinity irrigation (EC = 3.5 dS m-1), plants with NCED OE rootstocks had 30% higher fruit yield, but root biomass and lateral root development was reduced. Although NCED OE rootstocks upregulated ABA-signalling (AREB, ATHB12), ethylene-related (ACCs, ERFs), aquaporin (PIPs) and stress-related (TAS14, KIN, LEA) genes, downregulation of PYL ABA receptors and signalling components (WRKYs), ethylene synthesis (ACOs) and auxin responsive factors occurred. Elevated SlNCED1 expression enhanced ABA levels in reproductive tissue while ABA catabolites accumulated in leaf and xylem sap suggesting homeostatic mechanisms. NCED OE also reduced xylem cytokinin transport to the shoot and stimulated foliar 2-isopentenyl adenine (iP) accumulation and phloem transport. Moreover, increased xylem gibberellin GA3 levels in growing fruit trusses was associated with enhanced reproductive growth. Improved photosynthesis without changes in stomatal conductance was consistent with hormone-mediated alteration of leaf growth and mesophyll structure, which combined with lower assimilate requirement in the roots and systemic changes in hormone balances could explain enhanced vigour, reproductive growth and yield under saline stress.