In this paper, a number of strategies to overcome the deleterious effects of salinity on plants will be reviewed; these strategies include using molecular markers and genetic transformation as tools to develop salinity-tolerant genotypes, and some cultural techniques. For more than 12 years, QTL analysis has been attempted in order to understand the genetics of salt tolerance and to deal with component traits in breeding programmes. Despite innovations like better marker systems and improved genetic mapping strategies, the success of marker-assisted selection has been very limited because, in part, of inadequate experimental design. Since salinity is variable in time and space, experimental design must allow the study of genotype x environment interaction. Genetic transformation could become a powerful tool in plant breeding, but the growing knowledge from plant physiology must be integrated with molecular breeding techniques. It has been shown that the expression of several transgenes promotes a higher level of salt tolerance in some species. Despite this promising result, the development of a salt-tolerant cultivar by way of transgenesis has still not been achieved. Future directions in order to overcome the present limitations are proposed. Three cultural techniques have proved useful in tomato to overcome, in part, the effects of salinity: treatment of seedlings with drought or NaCl ameliorates the adaptation of adult plants to salinity; mist applied to tomato plants grown in Mediterranean conditions improves vegetative growth and yield in saline conditions; and grafting tomato cultivars onto appropriate rootstocks could reduce the effects of salinity.

The location of major quantitative trait loci (QTL) contributing to stem and leaf [Na

Tomato crop productivity under salinity can be improved by grafting cultivars onto salt-tolerant wild relatives, thus mediating the supply of root-derived ionic and hormonal factors that regulate leaf area and senescence.A tomato cultivar was grafted onto rootstocks from a population of recombinant inbred lines (RILs) derived from a Solanum lycopersicum ¥ Solanum cheesmaniae cross and cultivated under moderate salinity (75 mM NaCl). Concentrations of Na + , K + and several phytohormones [abscisic acid (ABA); the cytokinins (CKs) zeatin, Z; zeatin riboside, ZR; and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC)] were analysed in leaf xylem sap in graft combinations of contrasting vigour. Scion leaf area correlated with photosystem II (PSII) efficiency (Fv/Fm) and determined fruit productivity. Xylem K + (but not Na + ), K + /Na + , the active CK Z, the ratio with its storage form Z/ZR and especially the ratio between CKs and ACC (Z/ACC and Z + ZR/ACC) were positively loaded into the first principal component (PC) determining both leaf growth and PSII efficiency. In contrast, the ratio ACC/ABA was negatively correlated with leaf biomass. Although the underlying physiological mechanisms by which rootstocks mediate leaf area or chlorophyll fluorescence (and thus influence tomato salt tolerance) seem complex, a putative potassium-CK interaction involved in regulating both processes merits further attention.

The joint analysis of genotype marker segregation and phenotypic values of individuals or lines enables the detection and location of loci affecting quantitative traits (QTL). The availability of DNA markers and powerful biometric methods has led to considerable progress in QTL mapping in plants.The most obvious applications of QTL analysis seem to be marker-assisted selection (MAS) in breeding and pre-breeding and QTL cloning. However, other areas are envisaged where QTL analysis can contribute decisively. These are: the understanding of complex traits such as plant-pathogen interaction; plant genomics, connecting proteins and regulatory elements of known functions to QTL by candidate gene analysis; and germplasm enhancement through a characterization that allows its efficient utilization. The success in all these applications depends primarily on the reliability and accuracy of the QTL analysis itself. Therefore, the discussion of its limitations will constitute an important part of this review.

The sodium and potassium concentrations in leaf and stem have been genetically studied as physiological components of the vegetative and reproductive development in two populations of F(8) lines, derived from a salt sensitive genotype of Solanum lycopersicum cv. Cerasiforme, as female parent, and two salt tolerant lines, as male parents, from S. pimpinellifolium, the P population (142 lines), and S. cheesmaniae, the C population (116 lines). Genetic parameters of ten traits under salinity and five of them under control conditions were studied by ANOVA, correlation, principal component and QTL analysis to understand the global response of the plant. Two linkage maps including some tomato flowering time and salt tolerance candidate genes encoding for SlSOS1, SlSOS2, SlSOS3, LeNHX1, LeNHX3, were used for the QTL detection. Thirteen and 20 QTLs were detected under salinity in the P and C populations, respectively, and four under control conditions. Highly significant and contributing QTLs (over 40%) for the concentrations of Na(+) and K(+) in stems and leaves have been detected on chromosome 7 in both the populations. This is the only genomic position where the concentration QTLs for both the cations locate together. The proportion of QTLs significantly affected by salinity was larger in the P population (64.3%, including all QTLs detected under control) than in the C population (21.4%), where the estimated genetic component of variance was larger for most traits. A highly significant association between the leaf area and fruit yield under salinity was found only in the C population, which is supported by the location of QTLs for these traits in a common region of chromososome C1. As far as breeding for salt tolerance is concerned, only two sodium QTLs (lnc1.1 and lnc8.1) map in genomic regions of C1 and C8 where fruit yield QTLs are also located but in both the cases the profitable allele corresponds to the salt sensitive, cultivated species. One of those QTLs, lnc1.1 might involve LeNHX3.