Climate change is increasing the frequency of extreme heat events that aggravate its negative impact on plant development and agricultural yield. Most experiments designed to study plant adaption to heat stress apply homogeneous high temperatures to both shoot and root. However, this treatment does not mimic the conditions in natural fields, where roots grow in a dark environment with a descending temperature gradient. Excessively high temperatures severely decrease cell division in the root meristem, compromising root growth, while increasing the division of quiescent center cells, likely in an attempt to maintain the stem cell niche under such harsh conditions. Here, we engineered the TGRooZ, a device that generates a temper-ature gradient for in vitro or greenhouse growth assays. The root systems of plants exposed to high shoot temperatures but cultivated in the TGRooZ grow efficiently and maintain their functionality to sustain proper shoot growth and development. Furthermore, gene expression and rhizosphere or root microbiome compo-sition are significantly less affected in TGRooZ-grown roots than in high-temperature-grown roots, corre-lating with higher root functionality. Our data indicate that use of the TGRooZ in heat-stress studies can improve our knowledge of plant response to high temperatures, demonstrating its applicability from laboratory studies to the field.

Ethylene signaling plays a major role in the regulation of plant growth, but its cross-talk with other phytohormones is still poorly understood. Here, we investigated whether or not a defect in ethylene signaling, particularly in the ETHYLENE INSENSITIVE3 (EIN3) transcription factor, alters plant growth and influences the contents of other phytohormones. With this aim, a hormonal profiling approach using ultrahigh performance liquid chromatography coupled to tandem mass spectrometry (UHPLC-MS/MS) was used to unravel organ-specific responses (in roots, leaves and fruits) in the ein3-1 mutant and wild-type A. thaliana plants exposed to contrasting phosphate (Pi) availability. A defect in ethylene signaling in the ein3-1 mutant increased the biomass of roots, leaves and fruits, both at 0.5 mM and 1 mM Pi, thus indicating the growth-inhibitory role of ethylene in all tested organs. The hormonal profiling in roots revealed a cross-talk between ethylene signaling and other phytohormones, as indicated by increases in the contents of auxin, gibberellins and the stress-related hormones, abscisic acid, salicylic acid and jasmonic acid. The ein3-1 mutant also showed increased cytokinin contents in leaves. Reduced Pi availability (from 1 mM to 0.5 mM Pi) affected fruit growth, but not root and leaf growth, thus indicating mild Pi deficiency. It is concluded that ethylene signaling plays a major role in the modulation of plant growth in A. thaliana and that the ein3-1 mutant is not only altered in ethylene signaling but in the contents of several phytohormones in an organ-specific manner, thus indicating a hormonal cross-talk.

The transition phase in plants, which is marked by the onset of flowering and seed production, is very sensitive to the physical environment. Moreover, dimorphic plant species can show important sex-related differences in the response to nutrient availability. How availability of inorganic phosphate (Pi) during the transition phase influences sex ratios and whether or not male and female plants respond differently to nutrient stress is still however very poorly understood. Here, we examined whether contrasting Pi availability during the transition phase may influence sexual differentiation in stinging nettle (U. dioica L.) plants. We also evaluated to what extent males and females respond differently to varying Pi concentrations in terms of nutrient accumulation, C and N isotopic composition, photo-oxidative stress and hormone profiling. Results showed that reduced Pi availability delays plant maturation in males, causing as well an increase in the sex ratio towards females. Females showed enhanced lipid peroxidation (as indicated by higher lipid hydroperoxide contents) compared to males under Pi starvation, but sexes did not differ in photosystem II efficiency. Furthermore, reduced Pi availability led to enhanced 12-oxo-phytodienoic acid contents, the immediate precursor of jasmonic acid, particularly in females. This increase was, however, not accompanied by increases in other jasmonates, neither free nor conjugated jasmonic acid, thus suggesting a specific role for 12-oxo-phytodienoic acid in chemical defense in females under Pi starvation. Sex biased ratios in favour of females over males and enhanced 12-oxo-phytodienoic acid contents in the former under Pi starvation illustrate the great adaptive capacity of dimorphic species to reduced nutrient availability.

Vitamin E inhibits the propagation of lipid peroxidation and helps protecting photosystem II from photoinhibition, but little is known about its possible role in plant response to Pi availability. Here, we aimed at examining the effect of vitamin E deficiency in Arabidopsis thaliana vte mutants on phytohormone contents and the expression of transcription factors in plants exposed to contrasting Pi availability. Plants were subjected to two doses of Pi, either unprimed (controls) or previously exposed to low Pi (primed). In the wild type, α-tocopherol contents increased significantly in response to repeated periods of low Pi, which was paralleled by increased growth, indicative of a priming effect. This growth-stimulating effect was, however, abolished in vte mutants. Hormonal profiling revealed significant effects of Pi availability, priming and genotype on the contents of jasmonates and salicylates; remarkably, vte mutants showed enhanced accumulation of both hormones under low Pi. Furthermore, expression profiling of 1,880 transcription factors by qRT-PCR revealed a pronounced effect of priming on the transcript levels of 45 transcription factors mainly associated with growth and stress in wild-type plants in response to low Pi availability; while distinct differences in the transcriptional response were detected in vte mutants. We conclude that α-tocopherol plays a major role in the response of plants to Pi availability not only by protecting plants from photo-oxidative stress, but also by exerting a control over growth- and defense-related transcriptional reprogramming and hormonal modulation.

Dimorphic plant species can show distinct nutrient needs due to sex-related differences in nutrient allocation to reproductive structures, which can potentially affect their sensitivity to photoinhibition and photo-oxidative stress. Here, we investigated sex-related differences in the extent of photo-oxidative stress in male and female individuals of U. dioica exposed to a combination of severe drought and nutrient starvation. Male and female individuals of U. dioica subject to severe drought stress were exposed to various levels of nutrient availability. First, a set of plants grown under field conditions and exposed to summer drought was used to test the effects of nutrient supply (given as NPK fertilizer). Secondly, the effects of various phosphate concentrations in the nutrient solution were tested in drought-stressed potted plants. The Fv/Fm ratio (maximum efficiency of PSII photochemistry), photoprotection capacity (levels of carotenoids, including the xanthophyll cycle, and vitamins C and E), and the extent of lipid peroxidation (hydroperoxide levels) were measured. Results showed that an application of the NPK fertilizer to the soil had a positive effect on drought-stressed plants, reducing the extent of lipid peroxidation in both males and females. P deficiency led to residual photoinhibition, as indicated by significant reductions in the Fv/Fm ratio, and enhanced lipid peroxidation in females, but not in males. We conclude that (i) increased nutrient availability in the soil can alleviate photo-oxidative stress in drought-stressed U. dioica plants, and (ii) U. dioica plants show sexual secondary dimorphism in terms of photoinhibition and photo-oxidative stress, but this is only apparent when stress infringed on plants is very severe.

BACKGROUND AND AIMS:Vitamin E helps to control the cellular redox state by reacting with singlet oxygen and preventing the propagation of lipid peroxidation in thylakoid membranes. Both plant ageing and phosphorus deficiency can trigger accumulation of reactive oxygen species, leading to damage to the photosynthetic apparatus. This study investigates how phosphorus availability and vitamin E interact in the control of plant longevity in the short-lived annual Arabidopsis thaliana.METHODS:The responses of tocopherol cyclase (VTE1)- and γ-tocopherol methyltransferase (VTE4)-null mutants to various levels of phosphorus availability was compared with that of wild-type plants. Longevity (time from germination to rosette death) and the time taken to pass through different developmental stages were determined, and measurements were taken of photosynthetic efficiency, pigment concentration, lipid peroxidation, vitamin E content and jasmonate content.KEY RESULTS:The vte1 mutant showed accelerated senescence under control conditions, excess phosphorus and mild phosphorus deficiency, suggesting a delaying, protective effect of α-tocopherol during plant senescence. However, under severe phosphorus deficiency the lack of α-tocopherol paradoxically increased longevity in the vte1 mutant, while senescence was accelerated in wild-type plants. Reduced photoprotection in vitamin E-deficient mutants led to increased levels of defence chemicals (as indicated by jasmonate levels) under severe phosphorus starvation in the vte4 mutant and under excess phosphorus and mild phosphorus starvation in the vte1 mutant, indicating a trade-off between the capacity for photoprotection and the activation of chemical defences (jasmonate accumulation).CONCLUSIONS:Vitamin E increases plant longevity under control conditions and mild phosphorus starvation, but accelerates senescence under severe phosphorus limitation. Complex interactions are revealed between phosphorus availability, vitamin E and the potential to synthesize jasmonates, suggesting a trade-off between photoprotection and the activation of chemical defences in the plants.