Drought represents a major threat to food security. Mechanistic data describing plant responses to drought have been studied extensively and genes conferring drought resistance have been introduced into crop plants. However, plants with enhanced drought resistance usually display lower growth, highlighting the need for strategies to uncouple drought resistance from growth. Here, we show that overexpression of BRL3, a vascular-enriched member of the brassinosteroid receptor family, can confer drought stress tolerance in Arabidopsis. Whereas loss-of-function mutations in the ubiquitously expressed BRI1 receptor leads to drought resistance at the expense of growth, overexpression of BRL3 receptor confers drought tolerance without penalizing overall growth. Systematic analyses reveal that upon drought stress, increased BRL3 triggers the accumulation of osmoprotectant metabolites including proline and sugars.Transcriptomic analysis suggests that this results from differential expression of genes in the vascular tissues. Altogether, this data suggests that manipulating BRL3 expression could be used to engineer drought tolerant crops.
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.
In many fruit trees, heavy fruit load in one year reduces flowering in the following year, creating a biennial fluctuation in yield termed alternate bearing (AB). In subtropical trees, where flowering induction is mostly governed by the accumulation of chilling hours, fruit load is thought to generate a signal (AB signal) that blocks the perception of cold induction. Fruit removal during a heavy-fruit-load year is effective at inducing flowering only if performed one to a few months before the onset of the flowering induction period. We previously showed that following fruit removal, the content of the auxin indoleacetic acid (IAA) in citrus buds is reduced, suggesting that the hormone plays a role in the AB signal. Here, we demonstrate that fruit presence generates relatively strong polar auxin transport in citrus and olive stems. Upon fruit removal, polar auxin transport is reduced and allows auxin release from the bud. Furthermore, using immunolocalization, hormone, and gene expression analyses, we show that in citrus, IAA level in the bud and specifically in the apical meristem is reduced upon fruit removal. Overall, our data provide support for the notion that fruit presence generates an auxin signal in the bud, which may affect flowering induction.
There is increasing interest in using novel rootstocks to confer resistance to abiotic stresses in horticultural species, and to understand the physiological mechanism(s) conferring these responses. The same scion (Solanum lycopersicum cv. Boludo F1, Monsanto) was grafted onto 144 different rootstocks: six accessions from S. lycopersicum (cv cerasiforme) and S. pimpinellifolium, selected for drought tolerance (sourced from AVRDC); nine introgression lines from S. lycopersicum x S. pennellii and x S. habrochaites, selected for high root/shoot ratio, salinity and drought tolerances (sourced from TGRC); and a population of 129recombinant inbred lines (RILs) derived from a salt sensitive genotype of S. lycopersicum var.Cerasiforme and a salt tolerant line from S. pimpinellifolium L. (sourced from IVIA). Plants were grown in greenhouses in hydroponics (salinity stress) or soil (soil drying or high soil mechanical impedance) for 2-5 weeks (during the vegetative stage), and shoot fresh weight (SFW) was recorded at the end of each experiment. Although rootstock effects on SFW were significantly correlated in the soil drying and impedance assays, no significant correlations were detected between SFW under salinity and SFW under the other stresses. Indeed, the best rootstocks for drought stress were different to those that were the best for salinity and high soil impedance. For each abiotic stress, some graft combinations had higher SFW (up to 90% more) than the self-grafted commercial cultivar Boludo F1. The search for genetic factors contributing to this variation will be the objective of a future study.
Tomato production is influenced by shoot branching, which is controlled by different hormones. Here we produced tomato plants overexpressing the cytokinin-deactivating gene CYTOKININ OXYDASE 2 (CKX2). CKX2-overexpressing (CKX2-OE) plants showed an excessive growth of axillary shoots, the opposite phenotype expected for plants with reduced cytokinin content, as evidenced by LC-MS analysis and ARR5-GUS staining. The TCP transcription factor SlBRC1b was downregulated in the axillary buds of CKX2-OE and its excessive branching was dependent on a functional version of the GRAS-family gene LATERAL SUPPRESSOR (LS). Grafting experiments indicated that increased branching in CKX2-OE plants is unlikely to be mediated by root-derived signals. Crossing CKX2-OE plants with transgenic antisense plants for the strigolactone biosynthesis gene CAROTENOID CLEAVAGE DIOXYGENASE (CCD7-AS) produced an additive phenotype, indicating independent effects of cytokinin and strigolactones on increased branching. On the other hand, CKX2-OE plants showed reduced polar auxin transport and their bud outgrowth was reduced when combined with auxin mutants. Accordingly, CKX2-OE basal buds did not respond to auxin applied in the decapitated apex. Our results suggest that tomato shoot branching depends on a fine-tuning of different hormonal balances and that perturbations in the auxin status could compensate for the reduced cytokinin levels in CKX2-OE plants.
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