Synergistic regulation at physiological, transcriptional and metabolic levels in tomato plants subjected to a combination of salt and heat stress.

With global warming and climate change, abiotic stresses often simultaneously occur. Combined salt and heat stress was a common phenomenon that was severe, particularly in arid/semi-arid lands. We aimed to reveal the systematic responsive mechanisms of tomato genotypes with different salt/heat susceptibilities to combined salt and heat stress. Morphological and physiological responses of salt-tolerant/sensitive and heat-tolerant/sensitive tomatoes at control, heat, salt and combined stress were investigated. Based on leaf Fv /Fm and H2 O2 content, samples from tolerant genotype at the four treatments for 36 h were taken for transcriptomics and metabolomics. We found that plant height, dry weight and net photosynthetic rate decreased while leaf Na+ concentration increased in all four genotypes under salt and combined stress than control. Changes in physiological indicators such as photosynthetic parameters and defence enzyme activities in tomato under combined stress were regulated by the expression of relevant genes and the accumulation of key metabolites. We screened five key pathways in tomato responding to a combination of salt and heat stress, such as oxidative phosphorylation (map00190). Synergistic regulation at morphological, physiological, transcriptional and metabolic levels in tomato plants was induced by combined stress. Heat stress was considered as a dominant stressor for tomato plants under the current combined stress. The oxidative phosphorylation pathway played a key role in tomato in response to combined stress, where tapped key genes (e.g. alternative oxidase, Aox1a) need further functional analysis. Our study will provide a valuable resource important for studying stress combination and improving tomato tolerance.

A synchronized, large-scale field experiment using Arabidopsis thaliana reveals the significance of the UV-B photoreceptor UVR8 under natural conditions.

This study determines the functional role of the plant ultraviolet-B radiation (UV-B) photoreceptor, UV RESISTANCE LOCUS 8 (UVR8) under natural conditions using a large-scale 'synchronized-genetic-perturbation-field-experiment'. Laboratory experiments have demonstrated a role for UVR8 in UV-B responses but do not reflect the complexity of outdoor conditions where 'genotype × environment' interactions can mask laboratory-observed responses. Arabidopsis thaliana knockout mutant, uvr8-7, and the corresponding Wassilewskija wild type, were sown outdoors on the same date at 21 locations across Europe, ranging from 39°N to 67°N latitude. Growth and climatic data were monitored until bolting. At the onset of bolting, rosette size, dry weight, and phenolics and glucosinolates were quantified. The uvr8-7 mutant developed a larger rosette and contained less kaempferol glycosides, quercetin glycosides and hydroxycinnamic acid derivatives than the wild type across all locations, demonstrating a role for UVR8 under field conditions. UV effects on rosette size and kaempferol glycoside content were UVR8 dependent, but independent of latitude. In contrast, differences between wild type and uvr8-7 in total quercetin glycosides, and the quercetin-to-kaempferol ratio decreased with increasing latitude, that is, a more variable UV response. Thus, the large-scale synchronized approach applied demonstrates a location-dependent functional role of UVR8 under natural conditions.

Supplementary UV-A and UV-B radiation differentially regulate morphology in Ocimum basilicum.

UV-A- or UV-B-enriched growth light was given to basil plants at non-stress-inducing intensities. UV-A-enriched growth light gave rise to a sharp rise in the expression of PAL and CHS genes in leaves, an effect that rapidly declined after 1-2 days of exposure. On the other hand, leaves of plants grown in UV-B-enriched light had a more stable and long-lasting increase in the expression of these genes and also showed a stronger increase in leaf epidermal flavonol content. UV supplementation of growth light also led to shorter more compact plants with a stronger UV effect the younger the tissue. The effect was more prominent in plants grown under UV-B-enriched light than in those grown under UV-A. Parameters particularly affected were internode lengths, petiole lengths and stem stiffness. In fact, the bending angle of the 2nd internode was found to increase as much as 67% and 162% for plants grown in the UV-A- and UV-B-enriched treatments, respectively. The decreased stem stiffness was probably caused by both an observed smaller internode diameter and a lower specific stem weight, as well as a possible decline in lignin biosynthesis due to competition for precursors by the increased flavonoid biosynthesis. Overall, at the intensities used, UV-B wavelengths are stronger regulators of morphology, gene expression and flavonoid biosynthesis than UV-A wavelengths.

The light spectrum differentially influences morphology, physiology and metabolism of Chrysanthemum × morifolium without affecting biomass accumulation.

The development of light emitting diodes (LED) gives new possibilities to use the light spectrum to manipulate plant morphology and physiology in plant production and research. Here, vegetative Chrysanthemum × morifolium were grown at a photosynthetic photon flux density of 230 µmol m-2 s-1 under monochromatic blue, cyan, green, and red, and polychromatic red:blue or white light with the objective to investigate the effect on plant morphology, gas exchange and metabolic profile. After 33 days of growth, branching and leaf number increased from blue to red light, while area per leaf, leaf weight fraction, flavonol index, and stomatal density and conductance decreased, while dry matter production was mostly unaffected. Plants grown under red light had decreased photosynthesis performance compared with blue or white light-grown plants. The primary and secondary metabolites, such as organic acids, amino acids and phenylpropanoids (measured by non-targeted metabolomics of polar metabolites), were regulated differently under the different light qualities. Specifically, the levels of reduced ascorbic acid and its oxidation products, and the total ascorbate pool, were significantly different between blue light-grown plants and plants grown under white or red:blue light, which imply photosynthesis-driven alterations in oxidative pressure under different light regimens. The overall differences in plant phenotype, inflicted by blue, red:blue or red light, are probably due to a shift in balance between regulatory pathways controlled by blue light receptors and/or phytochrome. Although morphology, physiology, and metabolism differed substantially between plants grown under different qualities of light, these changes had limited effects on biomass accumulation.

Downsizing in plants-UV light induces pronounced morphological changes in the absence of stress.

Ultraviolet (UV) light induces a stocky phenotype in many plant species. In this study, we investigate this effect with regard to specific UV wavebands (UV-A or UV-B) and the cause for this dwarfing. UV-A- or UV-B-enrichment of growth light both resulted in a smaller cucumber (Cucumis sativus L.) phenotype, exhibiting decreased stem and petiole lengths and leaf area (LA). Effects were larger in plants grown in UV-B- than in UV-A-enriched light. In plants grown in UV-A-enriched light, decreases in stem and petiole lengths were similar independent of tissue age. In the presence of UV-B radiation, stems and petioles were progressively shorter the younger the tissue. Also, plants grown under UV-A-enriched light significantly reallocated photosynthates from shoot to root and also had thicker leaves with decreased specific LA. Our data therefore imply different morphological plant regulatory mechanisms under UV-A and UV-B radiation. There was no evidence of stress in the UV-exposed plants, neither in photosynthetic parameters, total chlorophyll content, or in accumulation of damaged DNA (cyclobutane pyrimidine dimers). The abscisic acid content of the plants also was consistent with non-stress conditions. Parameters such as total leaf antioxidant activity, leaf adaxial epidermal flavonol content and foliar total UV-absorbing pigment levels revealed successful UV acclimation of the plants. Thus, the UV-induced dwarfing, which displayed different phenotypes depending on UV wavelengths, occurred in healthy cucumber plants, implying a regulatory adjustment as part of the UV acclimation processes involving UV-A and/or UV-B photoreceptors.

Salinity, waterlogging, and elevated [CO2] interact to induce complex responses in cultivated and wild tomato.

The effects of individual climatic factors on crops are well documented, whereas the interaction of such factors in combination has received less attention. The frequency of salinity and waterlogging stress is increasing with climate change, accompanied by elevated CO2 concentration (e[CO2]). This study explored how these three variables interacted and affected two tomato genotypes. Cultivated and wild tomato (Solanum lycopersicum and Solanum pimpinellifolium) were grown at ambient [CO2] and e[CO2], and subjected to salinity, waterlogging, and combined stress. Leaf photosynthesis, chlorophyll fluorescence, quenching analysis, pigment, and plant growth were analyzed. The response of tomatoes depended on both genotype and stress type. In cultivated tomato, photosynthesis was inhibited by salinity and combined stress, whereas in wild tomato, both salinity and waterlogging stress, alone and in combination, decreased photosynthesis. e[CO2] increased photosynthesis and biomass of cultivated tomato under salinity and combined stress compared with ambient [CO2]. Differences between tomato genotypes in response to individual and combined stress were observed in key photosynthetic and growth parameters. Hierarchical clustering and principal component analysis revealed genetic variations of tomatoes responding to the three climatic factors. Understanding the interacting effects of salinity and waterlogging with e[CO2] in tomato will facilitate improvement of crop resilience to climate change.

Dominant and Priming Role of Waterlogging in Tomato at e[CO2] by Multivariate Analysis.

The frequency of waterlogging episodes has increased due to unpredictable and intense rainfalls. However, less is known about waterlogging memory and its interaction with other climate change events, such as elevated CO2 concentration (e[CO2]). This study investigated the combined effects of e[CO2] and two rounds of waterlogging stress on the growth of cultivated tomato (Solanum lycopersicum) and wild tomato (S. pimpinellifolium). The aim is to elucidate the interaction between genotypes and environmental factors and thereby to improve crop resilience to climate change. We found that two rounds of treatments appeared to induce different acclimation strategies of the two tomato genotypes. S. pimpinellifolium responded more negatively to the first-time waterlogging than S. lycopersicum, as indicated by decreased photosynthesis and biomass loss. Nevertheless, the two genotypes respond similarly when waterlogging stress recurred, showing that they could maintain a higher leaf photosynthesis compared to single stress, especially for the wild genotype. This showed that waterlogging priming played a positive role in stress memory in both tomato genotypes. Multivariate analysis showed that waterlogging played a dominant role when combined with [CO2] for both the cultivated and wild tomato genotypes. This work will benefit agricultural production strategies by pinpointing the positive effects of e[CO2] and waterlogging memory.

Effects of UV radiation on transcript and metabolite accumulation are dependent on monochromatic light background in cucumber.

During recent years, we have advanced our understanding of plant molecular responses to ultraviolet radiation (UV, 280-400 nm); however, how plants respond to UV radiation under different spectral light qualities is poorly understood. In this study, cucumber plants (Cucumis sativus "Lausanna RZ F1") were grown under monochromatic blue, green, red, and broadband white light in combination with UV radiation. The effects of light quality and UV radiation on acclimatory responses were assessed by measuring transcript accumulation of ELONGATED HYPOCOTYL 5 (HY5), CHALCONE SYNTHASE 2 (CHS2), and LIGHT HARVESTING COMPLEX II (LHCII), and the accumulation of flavonoids and hydroxycinnamic acids in the leaves. The growth light backgrounds differentially regulated gene expression and metabolite accumulation. While HY5 and CHS2 transcripts were induced by blue and white light, LHCII was induced by white and red light. Furthermore, UV radiation antagonized the effects of blue, red, green, and white light on transcript accumulation in a gene-dependent manner. Plants grown under blue light with supplementary UV radiation increased phenylalanine, flavonol disaccharide I and caffeic acid contents compared to those exposed only to blue light. UV radiation also induced the accumulation of flavonol disaccharide I and II, ferulic acid hexose and coumaric acid hexose in plants grown under green light. Our findings provide a further understanding of plant responses to UV radiation in combination with different light spectra and contribute to the design of light recipes for horticultural practices that aim to modify plant metabolism and ultimately improve crop quality.

Interactive effects of elevated CO2 concentration and combined heat and drought stress on tomato photosynthesis.

BACKGROUND: Extreme weather events are predicted to increase, such as combined heat and drought. The CO2 concentration ([CO2]) is predicted to approximately double by 2100. We aim to explore how tomato physiology, especially photosynthesis, is affected by combined heat and drought under elevated [CO2] (e [CO2]). RESULTS: Two genotypes, 'OuBei' ('OB', Solanum lycopersicum) and 'LA2093' (S. pimpinellifolium) were grown at a [CO2] (atmospheric [CO2], 400 ppm) and e [CO2] (800 ppm), respectively. The 27-days-old seedlings were treated at 1) a [CO2], 2) a [CO2] + combined stress, 3) e [CO2] and 4) e [CO2] + combined stress, followed by recovery. The PN (net photosynthetic rate) increased at e [CO2] as compared with a [CO2] and combined stress inhibited the PN. Combined stress decreased the Fv/Fm (maximum quantum efficiency of photosystem II) of 'OB' at e [CO2] and that of 'LA2093' in regardless of [CO2]. Genotypic difference was observed in the e [CO2] effect on the gas exchange, carbohydrate accumulation, pigment content and dry matter accumulation. CONCLUSIONS: Short-term combined stress caused reversible damage on tomato while the e [CO2] alleviated the damage on photosynthesis. However, the e [CO2] cannot be always assumed have positive effects on plant growth during stress due to increased water consumption. This study provided insights into the physiological effects of e [CO2] on tomato growth under combined stress and contributed to tomato breeding and management under climate change.

The Alleviation of Photosynthetic Damage in Tomato under Drought and Cold Stress by High CO2 and Melatonin.

The atmospheric CO2 concentration (a[CO2]) is increasing at an unprecedented pace. Exogenous melatonin plays positive roles in the response of plants to abiotic stresses, including drought and cold. The effect of elevated CO2 concentration (e[CO2]) accompanied by exogenous melatonin on plants under drought and cold stresses remains unknown. Here, tomato plants were grown under a[CO2] and e[CO2], with half of the plants pre-treated with melatonin. The plants were subsequently treated with drought stress followed by cold stress. The results showed that a decreased net photosynthetic rate (PN) was aggravated by a prolonged water deficit. The PN was partially restored after recovery from drought but stayed low under a successive cold stress. Starch content was downregulated by drought but upregulated by cold. The e[CO2] enhanced PN of the plants under non-stressed conditions, and moderate drought and recovery but not severe drought. Stomatal conductance (gs) and the transpiration rate (E) was less inhibited by drought under e[CO2] than under a[CO2]. Tomato grown under e[CO2] had better leaf cooling than under a[CO2] when subjected to drought. Moreover, melatonin enhanced PN during recovery from drought and cold stress, and enhanced biomass accumulation in tomato under e[CO2]. The chlorophyll a content in plants treated with melatonin was higher than in non-treated plants under e[CO2] during cold stress. Our findings will improve the knowledge on plant responses to abiotic stresses in a future [CO2]-rich environment accompanied by exogenous melatonin.

Allopolyploidization in Cucumis contributes to delayed leaf maturation with repression of redundant homoeologous genes.

The important role of polyploidy in plant evolution is widely recognized. However, many questions remain to be explored to address how polyploidy affects the phenotype of the plant. To shed light on the phenotypic and molecular impacts of allopolyploidy, we investigated the leaf development of a synthesized allotetraploid (Cucumis × hytivus), with an emphasis on chlorophyll development. Delayed leaf maturation was identified in C. × hytivus, based on delayed leaf expansion, initial chlorophyll deficiency in the leaves and disordered sink-source transition. Anatomical observations also revealed disturbed chloroplast development in C. ×hytivus. The determination of chlorophyll biosynthesis intermediates suggested that the chlorophyll biosynthesis pathway of C. × hytivus is blocked at the site at which uroporphyrinogen III is catalysed to coproporphyrinogen III. Three chlorophyll biosynthesis-related genes, HEMA1, HEME2 and POR, were significantly repressed in C. × hytivus. Sequence alignment showed both synonymous and non-synonymous substitutions in the HEMA1, HEME2 and POR genes of the parents. Cloning of the chlorophyll biosynthetic genes suggested the retention of homoeologs. In addition, a chimeric clone of the HEMA1 gene that consisted of homologous genes from the parents was identified in C. × hytivus. Overall, our results showed that allopolyploidization in Cucumis has resulted in disturbed chloroplast development and reduced chlorophyll biosynthesis caused by the repressed expression of duplicated homologous genes, which further led to delayed leaf maturation in the allotetraploid, C. × hytivus. The preferential retention/loss of certain types of genes and non-reciprocal homoeologous recombination were also supported in the present study, which provides new insights into the impact of allopolyploidy.

Physiological analysis and transcriptome sequencing reveal the effects of combined cold and drought on tomato leaf.

BACKGROUND: Co-occurrence of cold and drought stress can alter the response of plants at morphological, physiological and molecular levels, which finally affect crop production, more than individual stress. Understanding the responses of crop to combined stress is necessary to improve tolerance and maintain crop production especially in the field where combined stress frequently occurs. We aimed to clarify the underlying leaf physiological and molecular mechanisms of tomato by imposing combining cold and drought on one popular tomato cultivar 'Jinlingmeiyu' as an example. RESULTS: The physiological and genetic responses were identified in tomatoes after 42 h exposure to control, cold, drought and combined treatments. As compared with control, water loss rate at the three stresses including cold, drought and combined stress significantly decreased until 40 min after taking samples from the plants. The content of H2O2, zeatin riboside (ZR) and melatonin in all stress treatments were significantly higher than the control. Drought stress alone and combined stress induced the accumulation of abscisic acid (ABA) and auxin (IAA) as compared with control. The individual cold and combined stress significantly decreased the maximum quantum efficiency of PSII (Fv/Fm), quantum yield of PSII (Fq'/Fm') and electron transport rate (ETR). In total, 7141, 1850 and 7841 genes were involved in the stress response to cold, drought and their combination. Functional analysis of the stress-inducible genes provided more insights concerning the complex regulatory mechanisms that were involved in combined stress. The expression level of 12 genes were validated by quantitative real-time PCR (qRT-PCR). CONCLUSIONS: We found that the expression of stress-specific genes changed with physiological variation, indicating the close crosstalk between physiological and genetic response especially under combined stress. This study provides new knowledge on the complex regulatory mechanism genes in tomato ('Jinlingmeiyu') leaf to abiotic stresses.

UV regulates the expression of phenylpropanoid biosynthesis genes in cucumber (Cucumis sativus L.) in an organ and spectrum dependent manner.

Expression of cucumber (Cucumis sativus) genes encoding the phenylpropanoid and flavonoid biosynthetic enzymes phenylalanine ammonia lyase (PAL), cinnamic acid 4-hydroxylase (C4H), and chalcone synthase (CHS), was studied under control light conditions (photosynthetically active radiation, PAR) in root, stem, and leaf. Furthermore, the expression was quantified in leaves illuminated with PAR and supplemental ultraviolet-A (315-400 nm) or ultraviolet-B (280-315 nm) radiation. The expression patterns of all twelve CsPAL, three CsC4H, and three CsCHS genes were established. Among the genes regulated by UV two general expression patterns emerge. One pattern applies to genes primarily regulated by enriched UV-A illumination (pattern 1). Another pattern (pattern 2) was found for the genes regulated by enriched UV-B. Three of the pattern 2 genes (CsPAL4, CsPAL10, and CsCHS2) displayed a particular sub-pattern (pattern 2b) with transcription enriched by at least 30-fold. In contrast to the other genes studied, the promoters of the genes regulated according to pattern 2b contained a combination of a number of cis-acting regulatory elements (MREs, ACEs, and G-boxes) that may be of importance for the particularly high enhancement of expression under UV-B-containing light. The regulation of phenylpropanoid and flavonoid biosynthesis genes in cucumber resembles that of a number of other plants. However, cucumber, due to its greater size, is an attractive species for combining more detailed studies of the morphology, physiological parameters and fine regulation of spatial and temporal expression of key genes. This, in turn, can facilitate the quantitative investigation of the relationships among different promoter motifs, the expression levels of each of these three genes, and metabolite accumulation profiles.

Physiological response of tomatoes at drought, heat and their combination followed by recovery.

In nature, crops encounter a combination of abiotic stresses that severely limit yield. Our aim was to dynamically expose the changes of tomatoes' physiological parameters to drought, heat and their combination and thereby clarify the relationship between the responses to single and combined stress. We studied the effect of single and combined drought and heat stresses on the shoot and root of two tomato cultivars (Sufen No.14 as CV1; Jinlingmeiyu as CV2). After being exposed to combined stress for 6 days, the dry weight of shoot and root significantly decreased. The Fq '/Fm ' (quantum yield of photosystem II) was significantly lower in CV1 upon drought and combined stress and in CV2 subjected to combined stress (between days 4 and 6) compared to control. The relative water content during combined stress was significantly lower than control from day 4 to recovery day 2. On days 3 and 6, the water loss rate significantly increased under heat stress and decreased at drought and combined stress, respectively. The combined stress caused severe damages on photosystem II and chloroplast ultrastructure. The root activity after stress recovered even though drought significantly increased the activity from day 2 to day 6. Combined stress result in complex responses during tomato growth. The CV1 was more heat tolerant than CV2, but there was no varietal difference at drought and combined stress. This study contributes to the understanding of the underlying physiological response mechanism of plant to combined stress and crop improvement by providing valuable information for abiotic stress-tolerant tomato breeding.

Drought stress had a predominant effect over heat stress on three tomato cultivars subjected to combined stress.

BACKGROUND: Abiotic stresses due to environmental factors could adversely affect the growth and development of crops. Among the abiotic stresses, drought and heat stress are two critical threats to crop growth and sustainable agriculture worldwide. Considering global climate change, incidence of combined drought and heat stress is likely to increase. The aim of this study was to shed light on plant growth performance and leaf physiology of three tomatoes cultivars ('Arvento', 'LA1994' and 'LA2093') under control, drought, heat and combined stress. RESULTS: Shoot fresh and dry weight, leaf area and relative water content of all cultivars significantly decreased under drought and combined stress as compared to control. The net photosynthesis and starch content were significantly lower under drought and combined stress than control in the three cultivars. Stomata and pore length of the three cultivars significantly decreased under drought and combined stress as compared to control. The tomato 'Arvento' was more affected by heat stress than 'LA1994' and 'LA2093' due to significant decreases in shoot dry weight, chlorophyll a and carotenoid content, starch content and NPQ (non-photochemical quenching) only in 'Arvento' under heat treatment. By comparison, the two heat-tolerant tomatoes were more affected by drought stress compared to 'Arvento' as shown by small stomatal and pore area, decreased sucrose content, ΦPSII (quantum yield of photosystem II), ETR (electron transport rate) and qL (fraction of open PSII centers) in 'LA1994' and 'LA2093'. The three cultivars showed similar response when subjected to the combination of drought and heat stress as shown by most physiological parameters, even though only 'LA1994' and 'LA2093' showed decreased Fv/Fm (maximum potential quantum efficiency of photosystem II), ΦPSII, ETR and qL under combined stress. CONCLUSIONS: The cultivars differing in heat sensitivity did not show difference in the combined stress sensitivity, indicating that selection for tomatoes with combined stress tolerance might not be correlated with the single stress tolerance. In this study, drought stress had a predominant effect on tomato over heat stress, which explained why simultaneous application of heat and drought revealed similar physiological responses to the drought stress. These results will uncover the difference and linkage between the physiological response of tomatoes to drought, heat and combined stress and be important for the selection and breeding of tolerant tomato cultivars under single and combine stress.

Spectral effects of LEDs on chlorophyll fluorescence and pigmentation in Phalaenopsis 'Vivien' and 'Purple Star'.

We examined the effect of light emitting diode (LED) lighting in greenhouse facilities on growth, chlorophyll fluorescence and pigmentation in Phalaenopsis 'Vivien' and 'Purple Star' under purpose-built LED arrays yielding c. 200 µmol m(-2) s(-1) at plant height for 14 h per day and 24/18°C day/night temperature, respectively, from January to April 2013. The light treatments were (1) 40% blue in 60% red (40% B/R), (2) 0% blue in 100% red (0% B/R) and (3) white LEDs with 32% blue in white (32% B/W, control), with background daylight under shade screens. The plants were harvested twice for leaf growth and pigmentation. There was no clear pattern in the spectral effect on growth since the order of leaf size differed between harvests in March and April. Fv /Fm was in the range of 0.52-0.72, but overall slightly higher in the control, which indicated a permanent downregulation of PSII in the colored treatments. The fluorescence quenching showed no acclimation to color in 'Purple Star', while 'Vivien' had lower ETR and higher NPQ in the 40% B/R, resembling low light acclimation. The pigmentation showed corresponding spectral response with increasing concentration of lutein while increasing the fraction of blue light, which increased the light absorption in the green/yellow part of the spectrum. The permanent downregulation of PSII moved a substantial part of the thermal dissipation from the light regulated NPQ to non-regulated energy losses estimated by ΦNPQ and ΦNO and the difference found in the balance between ΦPSII and ΦNPQ in 'Vivien' disappeared when ΦNO was included in the thermal dissipation.

Wheat cultivars selected for high Fv /Fm under heat stress maintain high photosynthesis, total chlorophyll, stomatal conductance, transpiration and dry matter.

The chlorophyll fluorescence parameter Fv /Fm reflects the maximum quantum efficiency of photosystem II (PSII) photochemistry and has been widely used for early stress detection in plants. Previously, we have used a three-tiered approach of phenotyping by Fv /Fm to identify naturally existing genetic variation for tolerance to severe heat stress (3 days at 40°C in controlled conditions) in wheat (Triticum aestivum L.). Here we investigated the performance of the previously selected cultivars (high and low group based on Fv /Fm value) in terms of growth and photosynthetic traits under moderate heat stress (1 week at 36/30°C day/night temperature in greenhouse) closer to natural heat waves in North-Western Europe. Dry matter accumulation after 7 days of heat stress was positively correlated to Fv /Fm . The high Fv /Fm group maintained significantly higher total chlorophyll and net photosynthetic rate (PN ) than the low group, accompanied by higher stomatal conductance (gs ), transpiration rate (E) and evaporative cooling of the leaf (ΔT). The difference in PN between the groups was not caused by differences in PSII capacity or gs as the variation in Fv /Fm and intracellular CO2 (Ci ) was non-significant under the given heat stress. This study validated that our three-tiered approach of phenotyping by Fv /Fm performed under increasing severity of heat was successful in identifying wheat cultivars differing in photosynthesis under moderate and agronomically more relevant heat stress. The identified cultivars may serve as a valuable resource for further studies to understand the physiological mechanisms underlying the genetic variability in heat sensitivity of photosynthesis.

Plant responses to co-occurring heat and water deficit stress: A comparative study of tolerance mechanisms in old and modern wheat genotypes.

Global climate change increases the likelihood of co-occurrence of hot and dry spells with increased intensity, frequency, and duration. Studying the impact of the two stresses provide a better understanding of tolerance mechanisms in wheat, and our study was focused on revealing plant stress responses to different severities of combined stress at two phenophases in old and modern wheat genotypes. During the stem elongation and anthesis stages, plants were exposed to four treatments: control, deficit irrigation, combined heat, and deficit irrigation at 31 °C (HD31) and 37 °C (HD37). The modern genotypes were less affected by deficit irrigation at stem elongation as they maintained higher photosynthesis, stomatal conductance, and leaf cooling than old genotypes. When the HD37 stress was imposed during anthesis, the modern genotypes exhibited superior performance compared to the old, which was due to their higher photosynthetic rates resulting from improved biochemical regulation and a higher chlorophyll content. The plant responses varied during two phenophases under the combined stress exposure. Genotypes subjected to HD37 stress during stem elongation, photosynthesis was mainly controlled by stomatal regulation, whereas at anthesis it was predominated by biochemical regulation. These findings contribute to a deeper comprehension of plant tolerance mechanisms in response to different intensities of co-occurring hot and dry weather conditions.