Jesús Cuartero

Tomato plant-water uptake and plant-water relationships under saline growth conditions

Growth and water uptake both decreases when tomato plants are irrigated with saline water. To determine the relative contribution of physiological traits to these decreases plant fresh and dry weight, leaf area, leaf water (Psi(w)) and osmotic (Psi(Pi)) potentials, gas exchange parameters, stomatal density, leaf chlorophyll and Na content were investigated in the tomato (Lycopersicon esculentum) cultivars, Daniela and Moneymaker. Plants were grown in greenhouse, in sand culture, and irrigated with a complete nutrient solution supplied with 0 (control), 35 and 70 mM NaCl over a period of 2 months. Salinity reduced plant dry weight, height and number of leaves even at 35 mM NaCl. Leaf Psi(w) and Psi(Pi) decreased with salinity but leaf turgor pressures were significantly higher in salinised than in control plants which suggests that bulk tissue turgor did not limit growth under the saline conditions tested. Increasing salinity in the irrigation solution led to both morphological changes [(reduction of plant leaf area and stomatal density) and physiological changes [reduction of stomatal conductance, transpiration, and net CO(2) assimilation (A(CO(2)))] Plant water uptake, measured as the difference between volume of nutrient solution supplied and drainage collected, was closely related to transpiration, stomatal conductance, and stomatal density. Chlorophyll content per unit of leaf area increased with salinity. Reduction of net A(CO(2)) with salinity was explained in higher degree by stomatal conductance and stomatal density than by Na accumulation in the leaves. Although plant water uptake was similar for the two cultivars, Daniela transported, per unit of water uptake, more Na to the leaves than did Moneymaker. However, Daniela reduced leaf area less than did Moneymaker. Water use efficiency, calculated either as the ratio between total plant dry matter and total plant water uptake, or as the ratio between net A(CO(2)) and transpiration, did not change under our saline growth conditions. The contribution of the observed salt-responses to reduction in shoot water loss, plant water uptake and salt loading, while keeping water use efficiency, is discussed in relation to salt tolerance. Because some of these salt-responses take a long time to develop, growing seedlings in seedbeds with saline media could be of interest to better tolerate further salty conditions in the field or greenhouse.

Rootstock-mediated changes in xylem ionic and hormonal status are correlated with delayed leaf senescence, and increased leaf area and crop productivity in salinized tomato

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 x 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 (F(v)/F(m)) 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.

An overview on plant cuticle biomechanics

Plant biomechanics combines the principles of physics, chemistry and engineering to answer questions about plant growth, development and interaction with the environment. The epidermal-growth-control theory, postulated in 1867 and verified in 2007, states that epidermal cells determine the rate of organ elongation since they are under tension, while inner tissues are under compression. The lipid cuticle layer is deposited on the surface of outer epidermal cell walls and modifies the chemical and mechanical nature of these cell walls. Thus, the plant cuticle plays a key role in plant interaction with the environment and in controlling organ expansion. Rheological analyses indicate that the cuticle is a mostly viscoelastic and strain-hardening material that stiffens the comparatively more elastic epidermal cell walls. Cuticle stiffness can be attributed to polysaccharides and flavonoids present in the cuticle whereas a cutin matrix is mainly responsible for its extensibility. Environmental conditions such as temperature and relative humidity have a plasticizing effect on the mechanical properties of cuticle since they lower cuticle stiffness and strength. The external appearance of agricultural commodities, especially fruits, is of great economic value. Mechanical properties of the cuticle can have a positive or negative effect on disorders like fruit cracking, fungal pathogen penetration and pest infestation. Cuticle rheology has significant variability within a species and thus can be subjected to selection in order to breed cultivars resistant to pests, infestation and disorders.

Identification of Two Loci in Tomato Reveals Distinct Mechanisms for Salt Tolerance

Salt stress is one of the most serious environmental factors limiting the productivity of crop plants. To understand the molecular basis for salt responses, we used mutagenesis to identify plant genes required for salt tolerance in tomato. As a result, three tomato salt-hypersensitive (tss) mutants were isolated. These mutants defined two loci and were caused by single recessive nuclear mutations. The tss1 mutant is specifically hypersensitive to growth inhibition by Na+ or Li+ and is not hypersensitive to general osmotic stress. The tss2 mutant is hypersensitive to growth inhibition by Na+ or Li+ but, in contrast to tss1, is also hypersensitive to general osmotic stress. The TSS1 locus is necessary for K+ nutrition because tss1 mutants are unable to grow on a culture medium containing low concentrations of K+. Increased Ca2+ in the culture medium suppresses the growth defect of tss1 on low K+. Measurements of membrane potential in apical root cells were made with an intracellular microelectrode to assess the permeability of the membrane to K+ and Na+. K+-dependent membrane potential measurements indicate impaired K+ uptake in tss1 but not tss2, whereas no differences in Na+ uptake were found. The TSS2 locus may be a negative regulator of abscisic acid signaling, because tss2 is hypersensitive to growth inhibition by abscisic acid. Our results demonstrate that the TSS1 locus is essential for K+ nutrition and NaCl tolerance in tomato. Significantly, the isolation of the tss2 mutant demonstrates that abscisic acid signaling is also important for salt and osmotic tolerance in glycophytic plants.

Relative humidity and temperature modify the mechanical properties of isolated tomato fruit cuticles

The mechanical properties of enzymatically isolated cuticular membrane (CM) from ripe tomato fruits were investigated at 10 to 45°C and relative humidity (RH) of 40 to wet. CM samples were stressed by uniaxial tension loads to determine their tensile modulus, E, breaking stress (strength), σ(max), and maximum elongation, ε(max). The CM stress-strain curves revealed a biphasic behavior when tested at RH values below wet conditions. In the first phase, CM responded to the loads by instantaneous extension with no further extension recorded until a further load was added: defined as pure elastic strain (E(e)). In the second phase, CM responded by instantaneous extension and by some additional time-dependent extension, defined as viscoelastic strain (E(v)). When CMs were submerged in aqueous solution (wet), the stress-strain curves were monophasic, with both elastic and viscoelastic strain. E(e) depended on RH and was higher than E(v), which was independent of RH. Temperature decreased E(e) and σ(max) of tomato fruit CM. Temperature response was not linear but consisted of two temperature-independent phases separated by a transition temperature. This transition zone has been related previously to the presence of a secondary phase transition in the cutin matrix of the tomato fruit CM.

Salinity tolerance of normal-fruited and cherry tomato cultivars

The salinity tolerances (NaCl) of 8 normal-fruited tomato cultivars (Lycopersicon esculentum Mill.) and 4 cherry tomato cultivars (L. esculentum var.cerasiforme) were determined by yield-substrate EC response curves, according to the Mass-Hoffman model, modified by van Genuchten and Hoffman (1984). The same model was used to determine the response curves of leaf dry-weight, stem dry-weight, and plant height against substrate EC and also between yield and leaf concentrations of Cl- and Na ions.According to the salinity-threshold (maximum EC-value without yield reduction) and slope (yield decrease per unit EC increase) parameters, determined from the yield-EC response curves, the cherry tomato cultivars were more salt-tolerant than the normal-fruited ones. However, on the basis of vegetative growth characters-EC response curves, cherry tomato cultivars and normal-fruited ones were similarly affected by NaCl.The ranking of the cultivars by their salinity tolerance, determined from the plots of yield vs. leaf concentrations of Cl- and Na ions, was the same as that evaluated from the yield vs. substrate EC plots.

Genetic analysis of Na + and K + concentrations in leaf and stem as physiological components of salt tolerance in Tomato

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 F8 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.

The high fruit soluble sugar content in wild Lycopersicon species and their hybrids with cultivars depends on sucrose import during ripening rather than on sucrose metabolism

Soluble sugar content has been studied in relation to sucrose metabolism in the hexose-accumulating cultivated tomato Lycopersicon esculentum Mill, the wild relative species Lycopersicon cheesmanii Riley, in the sucrose-accumulating wild relative species Lycopersicon chmielewskii Rick, Kesicky, Fobes & Holle. and in two hexose-accumulating interspecific F1 hybrids (L. esculentum × L. cheesmanii; L. esculentum × L. chmielewskii), cultivated under two irrigation regimes (control: EC = 2.1 and saline: EC = 8.4 dS m-1). Under control conditions the total soluble sugar content (as hexose equivalents) in the ripe fruits of L. cheesmanii was 3-fold higher than in L. esculentum, while L. chmielewskii and both F1 hybrids contained twice as much as the cultivar. With the exception of L. esculentum × L. cheesmanii, salinity increased the sugar content by 1.3 (wild species) and 1.7 times (cultivar and L. esculentum × L. chmielewskii) with respect to control fruits. Wild germplasm or salinity provided two different mechanisms for the increases in fruit sugar content. The hexoses accumulated in ripe fruits were strongly influenced by those accumulated at the start of ripening, but the hydrolysed starch before start of ripening only partially explained the final hexose levels and especially the increase under salinity. The early cell wall acid invertase and the late neutral invertase activities appeared to be related to the amount of hexoses accumulated in ripe fruits. However, no metabolic parameter was positively related to the amount of sugar accumulated (including sucrose). The major differences between genotypes appeared in ripe fruits, in which up to 50% of the total amount of sugars accumulated in the wild species (mainly in L. cheesmanii) and hybrids cannot be explained by the sugars accumulated and the starch hydrolysed before the start of ripening stage. As a consequence, the higher fruit quality of the wild species compared with L. esculentum may depend more on the continuation of sucrose import during ripening than on osmotic or metabolic particularities such as the hexose / sucrose-accumulator character or specific enzyme activities.

Phase transitions in the biopolyester cutin isolated from tomato fruit cuticles

The specific heat of isolated tomato fruit cuticles and their corresponding cutins have been measured by first time for the physiological temperature in the range of 0–55 ◦ C. Variation of specific heat of the different isolates during fruit growth have been also measured. Isolated cuticles and cutin from young tomato fruits presented a clear glass transition temperature around 23 ◦ C. Water sorption on cutin samples shifted the glass transition temperature to 16.3 ◦ C indicating a clear plasticization of the biopolymer. The presence of these second-order transitions in these lipophilic plant material that act as a molecular barrier between the atmosphere and the plant cell, determine the mechanical and rheological properties of this biological barrier modulating the mass transfer between the environment ant the plant cell.

Relationship between tomato fruit growth and fruit osmotic potential under salinity.

To investigate the relationship between fruit growth and fruit osmotic potential (Psi(s)) in salty conditions, a sensitive tomato cultivar (Lycopersicon esculentum Mill.) and a tolerant accession of the wild species Lycopersicon pimpinellifolium Mill. were grown in a greenhouse with 0 and 70 mM NaCl, and the growth of the fruit studied from 15 to 70 days after anthesis (DAA). L. pimpinellifolium did not reduce significantly fruit weight in salty conditions throughout the growth period, whereas L. esculentum fruit weights decreased significantly with salinity from 45 DAA. L. esculentum fruit fresh weight reductions resulted from both less dry matter and water accumulation, although the fruit water content was affected by salinity before the fruit weight. In both species, fruit osmotic potential (Psi(s)) decreased significantly with salinity during the rapid fruit growth phase, although the changes were different. Thus, fruits from L. pimpinellifolium salt treated plants showed a Psi(s) reduction at the beginning (15 DAA) twice as high as that found in L. esculentum. As the advanced growth stage (from 15 to 55 DAA), the Psi(s) reduction percentages induced by salinity were quite similar in L. pimpinellifolium fruits, while increased in L. esculentum. Under saline conditions, the solutes contributing to reduce the fruit Psi(s) during the first 55 DAA were the inorganic solutes in both species, while in the ripe fruits they were hexoses. L. esculentum fruits accumulated K(+) as the main osmoticum in salty conditions, while L. pimpinellifolium fruits were able to use not only K(+) but also the Na(+) provided by the salt.