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Aims: Since producing more with less is required for increasing agricultural sustainability and reducing its environmental impact, breeding varieties with increased yield stability under reduced fertilizer application is an important goal, particularly in high valued horticultural crops such as tomato (Solanum lycopersicum L.). However, because of the difficulties to conciliate yield and fertilizer use efficiency through breeding, the graft-compatible genetic biodiversity existing in horticultural species offers the possibility to directly approach this objective in high-yielding elite varieties through improving nutrient capture and promoting ecosystem services such as insect pollination. We hypothesized that rootstocks affect pollinator foraging decisions through the nutritional status that impacts yield. Methods: Fifteen genetically diverse experimental rootstocks were grafted to a scion tomato variety and cultivated under optimal and reduced (25% of optimal) P and NPK fertilization in the presence of managed bumblebee pollinators (Bombus terrestris). Results: Up to twofold yield variability between rootstocks was associated with leaf nutrition and photosynthesis of the scion. Interestingly, fertilization regime and the rootstock genotype influenced the pollinator foraging decisions since bumblebees showed feeding preference for plants cultivated under low P, and for the most yielding and nutritious graft combinations under reduced but not under optimal fertilization. Bumblebees can sense plant nutritional status through source-sink relations, as supported by the consistent relationship between pollinator preferences and leaf carbon concentration. Conclusions: This study opens new perspectives for using pollinators as "phenotypers" to select the most resilient plants under suboptimal conditions and/or genotypes that synergistically increase crop productivity by promoting the ecosystem service provided by the insects. [ABSTRACT FROM AUTHOR]

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−1), 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 GA3 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. ABA overproducing tomato rootstocks increase shoot vigour and fruit yield under saline conditions by altering stress‐related responses, improving photosynthesis and optimizing source‐sink relationships. [ABSTRACT FROM AUTHOR]

Overexpression of a cell wall invertase gene promotes dehydration avoidance and WUE, and delays senescence in tomato through limiting stomatal conductance and inducing sink metabolism and photosynthesis in source leaves.Drought stress conditions modify source–sink relations, thereby influencing plant growth, adaptive responses, and consequently crop yield. Invertases are key metabolic enzymes regulating sink activity through the hydrolytic cleavage of sucrose into hexose monomers, thus playing a crucial role in plant growth and development. However, the physiological role of invertases during adaptation to abiotic stress conditions is not yet fully understood. Here it is shown that plant adaptation to drought stress can be markedly improved in tomato (Solanum lycopersicum L.) by overexpression of the cell wall invertase (cwInv) gene CIN1 from Chenopodium rubrum. CIN1 overexpression limited stomatal conductance under normal watering regimes, leading to reduced water consumption during the drought period, while photosynthetic activity was maintained. This caused a strong increase in water use efficiency (up to 50%), markedly improving water stress adaptation through an efficient physiological strategy of dehydration avoidance. Drought stress strongly reduced cwInv activity and induced its proteinaceous inhibitor in the leaves of the wild-type plants. However, the CIN1-overexpressing plants registered 3- to 6-fold higher cwInv activity in all analysed conditions. Surprisingly, the enhanced invertase activity did not result in increased hexose concentrations due to the activation of the metabolic carbohydrate fluxes, as reflected by the maintenance of the activity of key enzymes of primary metabolism and increased levels of sugar-phosphate intermediates under water deprivation. The induced sink metabolism in the leaves explained the maintenance of photosynthetic activity, delayed senescence, and increased source activity under drought stress. Moreover, CIN1 plants also presented a better control of production of reactive oxygen species and sustained membrane protection. Those metabolic changes conferred by CIN1 overexpression were accompanied by increases in the concentrations of the senescence-delaying hormone trans-zeatin and decreases in the senescence-inducing ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC) in the leaves. Thus, cwInv critically functions at the integration point of metabolic, hormonal, and stress signals, providing a novel strategy to overcome drought-induced limitations to crop yield, without negatively affecting plant fitness under optimal growth conditions. [ABSTRACT FROM AUTHOR]

Mixed nitrate/ammonium fertilization can partially alleviate the negative effects of salinity on growth of some plant species compared to all-nitrate or all-ammonium fertilization. To gain insights about the mechanisms involved, tomato ( Solanum lycopersicum L. cv Moneymaker) plants were grown hydroponically for 3 weeks with two NO/NH fertilization regimes (6/0.5 and 5/1.5; N = 6.5 mM) in the absence (control) or presence of salt stress (100 mM NaCl). Ammonium enrichment had no effect on growth and other parameters under control conditions. Under salinity, however, ammonium enrichment improved shoot and root biomass by 20% and maintained leaf PSII efficiency close to control levels. These changes were related to higher leaf K, NO, and NH concentrations and activities of the N-assimilatory enzymes glutamate synthase (GOGAT) and glutamine synthase (GS) in the leaves. Ammonium enrichment also attenuated the salt-induced increase in leaf abscisic acid (ABA) concentration and decrease in leaf concentrations of indole 3-acetic acid (IAA) and the cytokinins trans-zeatin ( tZ) and trans-zeatin riboside ( tZR). Enhanced cytokinin status was probably due to maintenance of root-to-shoot cytokinin transport and decreased leaf induction of the cytokinin-degrading enzyme cytokinin oxidase/dehydrogenase (CKX) under ammonium-enriched conditions. It is concluded that nitrogen form modifies salinity-induced physiological responses and that these modifications are associated with changes in plant hormone status. [ABSTRACT FROM AUTHOR]

Salinity limits crop productivity, in part by decreasing shoot concentrations of the growth-promoting and senescence-delaying hormones cytokinins. Since constitutive cytokinin overproduction may have pleiotropic effects on plant development, two approaches assessed whether specific root-localized transgenic IPT (a key enzyme for cytokinin biosynthesis) gene expression could substantially improve tomato plant growth and yield under salinity: transient root IPT induction (HSP70::IPT) and grafting wild-type (WT) shoots onto a constitutive IPT-expressing rootstock (WT/35S::IPT). Transient root IPT induction increased root, xylem sap, and leaf bioactive cytokinin concentrations 2- to 3-fold without shoot IPT gene expression. Although IPT induction reduced root biomass (by 15%) in control (non-salinized) plants, in salinized plants (100 mM NaCl for 22 d), increased cytokinin concentrations delayed stomatal closure and leaf senescence and almost doubled shoot growth (compared with WT plants), with concomitant increases in the essential nutrient K+ (20%) and decreases in the toxic ion Na+ (by 30%) and abscisic acid (by 20–40%) concentrations in transpiring mature leaves. Similarly, WT/35S::IPT plants (scion/rootstock) grown with 75 mM NaCl for 90 d had higher fruit trans-zeatin concentrations (1.5- to 2-fold) and yielded 30% more than WT/non-transformed plants. Enhancing root cytokinin synthesis modified both shoot hormonal and ionic status, thus ameliorating salinity-induced decreases in growth and yield. [ABSTRACT FROM PUBLISHER]

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. [ABSTRACT FROM AUTHOR]

The effect of short-term treatments (10 days) by a high salt level (150mmNaCl) on vegetative and reproductive development was investigated in tomato plants (Solanum lycopersicumL. cv. Ailsa Craig) at two developmental stages. Salinity applied during flowering transition reduced shoot biomass and delayed the appearance of the first inflorescence. Both shoot and root biomasses were reduced when salt was applied during the development of the first inflorescence. At both stages, areas of young leaves decreased and time to first anthesis increased, while total number of flowers in the first inflorescence was not affected. Flower abortion, reduction of pollen number and viability were higher when salinity was applied during inflorescence development. Na+accumulated in all organs while K+decreased. Laser ablation inductively coupled plasma mass spectrometry microanalysis revealed that Na+accumulated in style, ovaries and anther intermediate layers but not in the tapetum nor in the pollen grains when salinity was applied during inflorescence development. K+was not significantly affected in these structures. Soluble carbohydrates dramatically increased in leaves and decreased in the inflorescence under salt stress conditions. The failure of inflorescence to develop normally under salt stress can be better explained in terms of altered source–sink relationships rather than accumulation of toxic ions. [ABSTRACT FROM AUTHOR]

Plants have a remarkable regenerative capacity, which allows them to survive tissue damage after biotic and abiotic stresses. In this study, we use Solanum lycopersicum 'Micro-Tom' explants as a model to investigate wound-induced de novo organ formation, as these explants can regenerate the missing structures without the exogenous application of plant hormones. Here, we performed simultaneous targeted profiling of 22 phytohormone-related metabolites during de novo organ formation and found that endogenous hormone levels dynamically changed after root and shoot excision, according to region-specific patterns. Our results indicate that a defined temporal window of high auxin-to-cytokinin accumulation in the basal region of the explants was required for adventitious root formation and that was dependent on a concerted regulation of polar auxin transport through the hypocotyl, of local induction of auxin biosynthesis, and of local inhibition of auxin degradation. In the apical region, though, a minimum of auxin-to-cytokinin ratio is established shortly after wounding both by decreasing active auxin levels and by draining auxin via its basipetal transport and internalization. Cross-validation with transcriptomic data highlighted the main hormonal gradients involved in wound-induced de novo organ formation in tomato hypocotyl explants. [ABSTRACT FROM AUTHOR]

Following exposure to salinity, the root/shoot ratio is increased (an important adaptive response) due to the rapid inhibition of shoot growth (which limits plant productivity) while root growth is maintained. Both processes may be regulated by changes in plant hormone concentrations. Tomato plants (Solanum lycopersicum L. cv Moneymaker) were cultivated hydroponically for 3 weeks under high salinity (100 mM NaCl) and five major plant hormones (abscisic acid, ABA; the cytokinins zeatin, Z, and zeatin-riboside, ZR; the auxin indole-3-acetic acid, IAA; and the ethylene precursor 1-aminocyclopropane-1-carboxylic acid, ACC) were determined weekly in roots, xylem sap, and leaves. Salinity reduced shoot biomass by 50–60% and photosynthetic area by 20–25% both by decreasing leaf expansion and delaying leaf appearance, while root growth was less affected, thus increasing the root/shoot ratio. ABA and ACC concentrations strongly increased in roots, xylem sap, and leaves after 1 d (ABA) and 15 d (ACC) of salinization. By contrast, cytokinins and IAA were differentially affected in roots and shoots. Salinity dramatically decreased the Z+ZR content of the plant, and induced the conversion of ZR into Z, especially in the roots, which accounted for the relative increase of cytokinins in the roots compared to the leaf. IAA concentration was also strongly decreased in the leaves while it accumulated in the roots. Decreased cytokinin content and its transport from the root to the shoot were probably induced by the basipetal transport of auxin from the shoot to the root. The auxin/cytokinin ratio in the leaves and roots may explain both the salinity-induced decrease in shoot vigour (leaf growth and leaf number) and the shift in biomass allocation to the roots, in agreement with changes in the activity of the sink-related enzyme cell wall invertase. [ABSTRACT FROM PUBLISHER]

Leaf senescence is one of the most limiting factors to plant productivity under salinity. Both the accumulation of specific toxic ions (e.g. Na+) and changes in leaf hormone relations are involved in the regulation of this process. Tomato plants (Solanum lycopersicum L. cv Moneymaker) were cultivated for 3 weeks under high salinity (100 mM NaCl) and leaf senescence-related parameters were studied during leaf development in relation to Na+ and K+ contents and changes in abscisic acid (ABA), cytokinins, the ethylene precursor 1-aminocyclopropane-1-carboxylic acid (ACC), and the auxin indole-3-acetic acid (IAA). Na+ accumulated to a similar extent in both leaves 4 and 5 (numbering from the base of the plant) and more quickly during the third week, while concurrently K+ contents sharply decreased. However, photosystem II efficiency, measured as the Fv/Fm ratio, decreased from the second week of salinization in leaf 4 but only at the end of the third week in the younger leaf 5. In the prematurely senescent leaf 4, ABA content increased linearly while IAA strongly decreased with salinization time. Although zeatin (Z) levels were scarcely affected by salinity, zeatin-riboside (ZR) and the total cytokinin content (Z+ZR) progressively decreased by 50% from the imposition of the stress. ACC was the only hormonal compound that increased in leaf tissue coincident with the onset of oxidative damage and the decline in chlorophyll fluorescence, and prior to massive Na+ accumulation. Indeed, (Z+ZR) and ACC contents and their ratio (Z+ZR/ACC) were the hormonal parameters best correlated with the onset and progression of leaf senescence. The influence of different hormonal changes on salt-induced leaf senescence is discussed. [ABSTRACT FROM PUBLISHER]

Developing drought-tolerant crops is an important strategy to mitigate climate change impacts. Modulating root system function provides opportunities to improve crop yield under biotic and abiotic stresses. With this aim, a commercial hybrid tomato variety was grafted on a genotyped population of 123 recombinant inbred lines (RILs) derived from Solanum pimpinellifolium, and compared with self- and non-grafted controls, under contrasting watering treatments (100% vs. 70% of crop evapotranspiration). Drought tolerance was genetically analyzed for vegetative and flowering traits, and root xylem sap phytohormone and nutrient composition. Under water deficit, around 25% of RILs conferred larger total shoot dry weight than controls. Reproductive and vegetative traits under water deficit were highly and positively correlated to the shoot water content. This association was genetically supported by linkage of quantitative trait loci (QTL) controlling these traits within four genomic regions. From a total of 83 significant QTLs, most were irrigation-regime specific. The gene contents of 8 out of 12 genomic regions containing 46 QTLs were found significantly enriched at certain GO terms and some candidate genes from diverse gene families were identified. Thus, grafting commercial varieties onto selected rootstocks derived from S. pimpinellifolium provides a viable strategy to enhance drought tolerance in tomato. [ABSTRACT FROM AUTHOR]

• Constitutive ABA overproduction reduces shoot and root growth and close stomata, under optimal conditions. • Constitutive ABA overproduction reduces the percentage loss in shoot and root growth, under salinity conditions. • Constitutive ABA overproduction increases the total root length, under salinity conditions. • Constitutive ABA overproduction limited plant growth under optimal conditions by altering growth regulatory gene networks. • Salinity differentially altered the growth regulatory gene networks in ABA overproducing plants, thereby sustaining growth. To better understand abscisic acid (ABA)'s role in the salinity response of tomato (Solanum lycopersicum L.), two independent transgenic lines, sp5 and sp12, constitutively overexpressing the LeNCED1 gene (encoding 9- cis -epoxycarotenoid dioxygenase, a key enzyme in ABA biosynthesis) and the wild type (WT) cv. Ailsa Craig, were cultivated hydroponically with or without the addition of 100 mM NaCl. Independent of salinity, LeNCED1 overexpression (OE) increased ABA concentration in leaves and xylem sap, and salinity interacted with the LeNCED1 transgene to enhance ABA accumulation in xylem sap and roots. Under control conditions, LeNCED1 OE limited root and shoot biomass accumulation, which was correlated with decreased leaf gas exchange. In salinized plants, LeNCED1 OE reduced the percentage loss in shoot and root biomass accumulation, leading to a greater total root length than WT. Root qPCR analysis of the sp12 line under control conditions revealed upregulated genes related to ABA, jasmonic acid and ethylene synthesis and signalling, gibberellin and auxin homeostasis and osmoregulation processes. Under salinity, LeNCED1 OE prevented the induction of genes involved in ABA metabolism and GA and auxin deactivation that occurred in WT, but the induction of ABA signalling and stress-adaptive genes was maintained. Thus, complex changes in phytohormone and stress-related gene expression are associated with constitutive upregulation of a single ABA biosynthesis gene, alleviating salinity-dependent growth limitation. [ABSTRACT FROM AUTHOR]

Water availability is the most important factor limiting food production, thus developing new scientific strategies to allow crops to more efficiently use water could be crucial in a world with a growing population. Tomato is a highly water consuming crop and improving its water use efficiency (WUE) implies positive economic and environmental effects. This work aimed to study and exploit root-derived hormonal traits to improve WUE in tomato by grafting on selected rootstocks. Firstly, root-related hormonal parameters associated to WUE were identified in a population of recombinant inbred lines (RILs) derived from the wild tomato species Solanum pimpinellifolium . A principal component analysis (PCA) revealed that some hormonal traits were associated with productivity (plant biomass and photosynthesis) and WUE in the RIL population. Leaf ABA concentration was associated to the first component (PC1) of the PCA, which explained a 60% of the variance in WUE, while the ethylene precursor ACC and the ratio ACC/ABA were also associated to PC1 but in the opposite direction. Secondly, we selected RILs according to their extreme biomass (high, B, low, b) and water use (high, W, low, w), and studied the differential effect of shoot and root on WUE by reciprocal grafting. In absence of any imposed stress, there were no rootstock effects on vegetative shoot growth and water relations. Finally, we exploited the previously identified root-related hormonal traits by grafting a commercial tomato variety onto the selected RILs to improve WUE. Interestingly, rootstocks that induced low biomass and water use, ‘bw’, improved fruit yield and WUE (defined as fruit yield/water use) by up to 40% compared to self-grafted plants. Although other hormonal factors appear implicated in this response, xylem ACC concentration seems an important root-derived trait that inhibits leaf growth but does not limit fruit yield. Thus tomato WUE can be improved exploiting rootstock-derived hormonal signals which control leaf growth. [ABSTRACT FROM AUTHOR]