Plant metabolic response to stress in an arid ecosystem is mediated by the presence of neighbors.

Plant neighbors in arid environments can ameliorate abiotic stress by reducing insolation, but they also attract herbivores and pathogens, especially when neighbors are close relatives that share similar antagonists. Plants' metabolic profiles provide a chemical fingerprint of the physiological processes behind plant responses to different environmental stresses. For example, abscisic acid and proline, mainly involved in stomatal closure and osmotic adjustment, can induce plant responses to abiotic stress, while jasmonic acid and salicylic acid primarily regulate plant defense to herbivory or pathogens. Neighbor plants can generate contrasting ecological contexts, modulating plant responses to abiotic and biotic stresses. We hypothesize that plant metabolic profile is modulated by its neighbors in a vegetation patch, expecting a higher investment in metabolites related to biotic-stress tolerance (i.e., herbivory or pathogens) when growing associated with other plants, especially to phylogenetically close relatives, compared to plants growing alone. We show that plants from five species growing with neighbors invest more in biotic-stress tolerance while their conspecifics, growing alone, invest more in abiotic-stress tolerance. This tendency in plants' metabolic profiles was not affected by the phylogenetic diversity of their neighborhood. Linking physiological snapshots with community processes can contribute to elucidating metabolic profiles derived from plant-plant interactions.

The role of salicylic acid on glutathione metabolism under endoplasmic reticulum stress in tomato.

The endoplasmic reticulum (ER) stress and the unfolded protein response (UPR) are highly dependent on phytohormones such as salicylic acid (SA). In this study, the effect of SA supplementation and the lack of endogenous SA on glutathione metabolism were investigated under ER stress in wild-type (WT) and transgenic SA-deficient NahG tomato (Solanum lycopersicum L.) plants. The expression of the UPR marker gene SlBiP was dependent on SA levels and remained lower in NahG plants. Exogenous application of the chemical chaperone 4-phenylbutyrate (PBA) also reduced tunicamycin (Tm)-induced SlBiP transcript accumulation. At the same time, Tm-induced superoxide and hydrogen peroxide production were independent of SA, whereas the accumulation of reduced form of glutathione (GSH) and the oxidised glutathione (GSSG) was regulated by SA. Tm increased the activity of glutathione reductase (GR; EC 1.6.4.2) independently of SA, but the activities of dehydroascorbate reductase (DHAR; EC 1.8.5.1) and glutathione S-transferases (GSTs; EC 2.5.1.18) were increased by Tm in a SA-dependent manner. SlGR2, SlGGT and SlGSTT2 expression was activated in a SA-dependent way upon Tm. Although expression of SlGSH1, SlGSTF2, SlGSTU5 and SlGTT3 did not change upon Tm treatment in leaves, SlGR1 and SlDHAR2 transcription decreased. PBA significantly increased the expression of SlGR1, SlGR2, SlGSTT2, and SlGSTT3, which contributed to the amelioration of Tm-induced ER stress based on the changes in lipid peroxidation and cell viability. Malondialdehyde accumulation and electrolyte leakage were significantly higher in WT as compared to NahG tomato leaves under ER stress, further confirming the key role of SA in this process.

Morphological, physiological, and molecular scion traits are determinant for salt-stress tolerance of grafted citrus plants.

Introduction: Citrus productivity has been decreasing in the last decade in the Mediterranean basin as a consequence of climate change and the high levels of salinity found in the aquifers. Citrus varieties are cultivated grafted onto a rootstock, which has been reported as responsible for plant tolerance to adverse situations. However, other important factors for stress tolerance relying in the scion have been less studied. The aim of this study was to evaluate the effect of the grafted scion on citrus tolerance to salt stress. Methods: Four different citrus rootstock/scion combinations were subjected to salt stress for 30 days, using Carrizo citrange (CC) or Citrus macrophylla (CM) as rootstocks, and Navelina orange (NA) or Oronules mandarin (OR) as scions. CM-OR was the most tolerant combination, whereas CC-NA was the most sensitive one. Results and discussion: Our results support the idea that the rootstock plays an important role in salt stress tolerance, but scion is also crucial. Thus, photosynthesis and transpiration, processes regulated by abscisic acid and jasmonic acid, are determinant of plant performance. These photosynthetic parameters were not affected in plants of the salt-tolerant combination CM-OR, probably due to the lower intoxication with Cl- ions, allowing a better performance of the photosynthetic machinery under stress conditions. The different stomatal density of the two citrus scions used in this work (higher in the sensitive NA in comparison to the tolerant OR) also contributes to the different tolerance of the grafted plants to this adverse condition. Additionally, CsDTX35.1 and CsDTX35.2, genes codifying for Cl- tonoplast transporters, were exclusively overexpressed in plants of the salt-tolerant combination CM-OR, suggesting that these transporters involved in Cl- compartmentalization could be crucial for salt stress tolerance. It is concluded that to improve citrus tolerance to high salinity, it is important that scions have a versatile photosynthetic system, an adequate stomatal density, and a proper modulation of genes coding for Cl- transporters in the tonoplast.

Exogenous spermine alleviates the negative effects of combined salinity and paraquat in tomato plants by decreasing stress-induced oxidative damage.

Plants are frequently exposed to different combinations of soil constraints including salinity and different herbicides. These abiotic conditions negatively affect photosynthesis, growth and plant development resulting in limitations in agriculture production. To respond to these conditions, plants accumulate different metabolites that restore cellular homeostasis and are key for stress acclimation processes. In this work, we analyzed the role of exogenous spermine (Spm), a polyamine involved in plant tolerance to abiotic stress, in tomato responses to the combination of salinity (S) and the herbicide paraquat (PQ). Our findings showed that application of Spm reduced leaf damage and enhanced survival, growth, photosystem II function and photosynthetic rate of tomato plants subjected to the combination of S and PQ. In addition, we revealed that exogenous Spm reduced H2O2 and malondialdehyde (MDA) accumulation in plants subjected to S+PQ, suggesting that the role of exogenous Spm in alleviating the negative effects of this stress combination could be attributed to a decrease in stress-induced oxidative damage in tomato plants. Taken together, our results identify a key role for Spm in improving plant tolerance to combined stress.

Climate change-associated multifactorial stress combination: A present challenge for our ecosystems.

Humans negatively influence Earth ecosystems and biodiversity causing global warming, climate change as well as man-made pollution. Recently, the number of different stress factors have increased, and when impacting simultaneously, the multiple stress conditions cause dramatic declines in plant and ecosystem health. Although much is known about how plants and ecosystems are affected by each individual stress, recent research efforts have diverted into how these biological systems respond to several of these stress conditions applied together. Studies of such "multifactorial stress combination" concept have reported a severe decrease in plant survival and microbiome biodiversity along the increasing number of factors in a consistent directional trend. In addition, these results are in concert with studies about how ecosystems and microbiota are affected by natural conditions imposed by climate change. Therefore, all this evidence should serve as an important warning in order to decrease pollutants, create strategies to deal with global warming, and increase the tolerance of plants to multiple stressful factors in combination. Here we review recent studies focused on the impact of abiotic stresses on plants, agrosystems and different ecosystems including forests and microecosystems. In addition, different strategies to mitigate the impact of climate change in ecosystems are discussed.

Particle film technology modulates xanthophyll cycle and photochemical dynamics of grapevines grown in the Douro Valley.

Field-grown grapevines are often exposed to multiple environmental stresses, which challenges wine-growers to develop sustainable measures to sustain vine growth, yield, and quality. Under field conditions this task is demanding, due to differences in the magnitudes of stresses and associated plant responses. In this study we explored the hypothesis that kaolin-particle film application improves grapevine photoprotection through the regulation of xanthophyll cycle genes, limiting the thermal dissipation of excess energy under harsh environmental conditions. Hence, we selected two grapevine varieties, Touriga-Nacional (TN) and Touriga-Franca (TF), grown in the Douro Demarcated Region, and evaluated changes in light dissipation mechanisms, xanthophyll cycle components, and the expression of xanthophyll cycle genes during the 2017 summer season. The results showed that, from veraison to ripening, kaolin triggered the up-regulation of violaxanthin de-epoxidase (VvVDE1) and zeaxanthin epoxidase (VvZEP1) genes, indicating optimised regulation of the xanthophyll cycle. Kaolin treatment also decreased chlorophyll (Chla, Chlb, Chl(a+b)) and carotenoid (Car) accumulation under increasing summer stress conditions in both varieties and lowered the non-photochemical quenching (NPQ) of grapevines on ripening, suggesting a long-term response to summer stress. In addition, kaolin-treated grapevines showed increased Chla/Chlb and lower Chl(a+b)/Car ratios, displaying some features of high light adapted leaves. Overall, this study suggests that kaolin application enabled grapevines to benefit from fluctuating periods of summer stress by managing chlorophyll and carotenoid content and limiting down-regulation of both photochemistry and photoinhibition processes. Under Mediterranean field conditions, kaolin application can be considered an efficient method of minimising summer stress impact on grapevines.

Water Stress Responses of Tomato Mutants Impaired in Hormone Biosynthesis Reveal Abscisic Acid, Jasmonic Acid and Salicylic Acid Interactions.

To investigate the putative crosstalk between JA and ABA in Solanum lycopersicum plants in response to drought, suppressor of prosystemin-mediated responses2 (spr2, JA-deficient) and flacca (flc, ABA-deficient) mutants together with the naphthalene/salicylate hydroxylase (NahG) transgenic (SA-deficient) line were used. Hormone profiling and gene expression of key enzymes in ABA, JA and SA biosynthesis were analyzed during early stages of drought. ABA accumulation was comparable in spr2 and wild type (WT) plants whereas expression of 9-cis-epoxycarotenoid dioxygenase 1 (NCED1) and NCED2 was different, implying a compensation mechanism between NCED genes and an organ-specific regulation of NCED1 expression. JA levels and 12-oxo-phytodienoic acid reductase 3 (OPR3) expression in flc plants suggest that ABA regulates the induction of the OPR3 gene in roots. By contrast, ABA treatment to flc plants leads to a reduction of JA and SA contents. Furthermore, different pattern of SA accumulation (and expression of isochorismate synthase and phenylalanine ammonia lyase 1) was observed between WT seedlings and mutants, suggesting that SA plays an important role on the early response of tomato plants to drought and also that JA and ABA modulate its biosynthesis. Finally, hormone profiling in spr2 and NahG plants indicate a crosstalk between JA and SA that could enhance tolerance of tomato to water stress.

Hormonal and hydroxycinnamic acids profiles in banana leaves in response to various periods of water stress.

The pattern of change in the endogenous levels of several plant hormones and hydroxycinnamic acids in addition to growth and photosynthetic performance was investigated in banana plants (Musa acuminata cv. "Grand Nain") subjected to various cycles of drought. Water stress was imposed by withholding irrigation for six periods with subsequent rehydration. Data showed an increase in abscisic acid (ABA) and indole-3-acetic acid (IAA) levels, a transient increase in salicylic acid (SA) concentration, and no changes in jasmonic acid (JA) after each period of drought. Moreover, the levels of ferulic (FA) and cinnamic acids (CA) were increased, and plant growth and leaf gas exchange parameters were decreased by drought conditions. Overall, data suggest an involvement of hormones and hydroxycinnamic acids in plant avoidance of tissue dehydration. The increase in IAA concentration might alleviate the senescence of survival leaves and maintained cell elongation, and the accumulation of FA and CA could play a key role as a mechanism of photoprotection through leaf folding, contributing to the effect of ABA on inducing stomatal closure. Data also suggest that the role of SA similarly to JA might be limited to a transient and rapid increase at the onset of the first period of stress.

Biotechnological approaches to study plant responses to stress.

Multiple biotic and abiotic environmental stress factors affect negatively various aspects of plant growth, development, and crop productivity. Plants, as sessile organisms, have developed, in the course of their evolution, efficient strategies of response to avoid, tolerate, or adapt to different types of stress situations. The diverse stress factors that plants have to face often activate similar cell signaling pathways and cellular responses, such as the production of stress proteins, upregulation of the antioxidant machinery, and accumulation of compatible solutes. Over the last few decades advances in plant physiology, genetics, and molecular biology have greatly improved our understanding of plant responses to abiotic stress conditions. In this paper, recent progresses on systematic analyses of plant responses to stress including genomics, proteomics, metabolomics, and transgenic-based approaches are summarized.

Modulation of the antioxidant system in Citrus under waterlogging and subsequent drainage.

Soil flooding induces an impairment of the photosynthetic system that often leads to an accumulation of reactive oxygen species (ROS) in plant tissues. Moreover, flooding release by drainage can cause a sudden oxygen burst that exacerbates oxidative damage. To examine the influence of different anoxic and post-anoxic periods on citrus physiology, citrumelo CPB4475, a moderate flood-tolerant genotype, was subjected to three different periods of soil flooding followed by drainage. Plant performance in terms of visible damage, photosynthetic activity, malondialdehyde (MDA) and hydrogen peroxide accumulation was examined together with the plant antioxidant response. The results indicated that coordinated antioxidant activity, involving increased activities of superoxide dismutase (SOD) (EC 1.15.1.1) and catalase (CAT) (EC 1.11.1.6), together with a modulation of the ascorbate-glutathione cycle, allowed plants to cope with flooding-induced oxidative stress up to a certain point. Elevated ascorbate peroxidase (APX) (EC 1.11.1.11) activity or discrete increases in AsA or glutathione concentrations seemed inefficient in maintaining low levels of oxidative damage. Waterlogging stress release by soil drainage did not improve plant performance but, on the contrary, enhanced oxidative stress and even accelerated plant injury. This appears to be the result of sudden oxygen burst soon after release of water.

Membrane transporters and carbon metabolism implicated in chloride homeostasis differentiate salt stress responses in tolerant and sensitive Citrus rootstocks.

Salinity tolerance in Citrus is strongly related to leaf chloride accumulation. Both chloride homeostasis and specific genetic responses to Cl(-) toxicity are issues scarcely investigated in plants. To discriminate the transcriptomic network related to Cl(-) toxicity and salinity tolerance, we have used two Cl(-) salt treatments (NaCl and KCl) to perform a comparative microarray approach on two Citrus genotypes, the salt-sensitive Carrizo citrange, a poor Cl(-) excluder, and the tolerant Cleopatra mandarin, an efficient Cl(-) excluder. The data indicated that Cl(-) toxicity, rather than Na(+) toxicity and/or the concomitant osmotic perturbation, is the primary factor involved in the molecular responses of citrus plant leaves to salinity. A number of uncharacterized membrane transporter genes, like NRT1-2, were differentially regulated in the tolerant and the sensitive genotypes, suggesting its potential implication in Cl(-) homeostasis. Analyses of enriched functional categories showed that the tolerant rootstock induced wider stress responses in gene expression while repressing central metabolic processes such as photosynthesis and carbon utilization. These features were in agreement with phenotypic changes in the patterns of photosynthesis, transpiration, and stomatal conductance and support the concept that regulation of transpiration and its associated metabolic adjustments configure an adaptive response to salinity that reduces Cl(-) accumulation in the tolerant genotype.

Antioxidant enzymatic activity is linked to waterlogging stress tolerance in citrus.

Soil flooding constitutes a seasonal factor that negatively affects plant performance and crop yields. In this work, the relationship between oxidative damage and flooding sensitivity was addressed in three citrus genotypes with different abilities to tolerate waterlogging. We examined leaf visible damage, oxidative damage in terms of malondialdehyde (MDA) concentration, leaf proline concentration, leaf and root ascorbate and glutathione contents and the antioxidant enzyme activities superoxide dismutase (EC 1.15.1.1), ascorbate peroxidase (EC 1.11.1.11), catalase (EC 1.11.1.6) and glutathione reductase (EC 1.8.1.7). No differences in the extent of oxidative damage relative to controls were found among genotypes. However, a different ability to delay the apparition of oxidative damage was associated to a higher tolerance to waterlogging. This ability was linked to an enhanced activated oxygen species' scavenging capacity in terms of an increased antioxidant enzyme activity and higher content in polar antioxidant compounds. Therefore, the existence of a direct relationship between stress sensitivity and the early accumulation of MDA is proposed. In addition, data indicate that the protective role of proline has to be considered minimal as its accumulation was inversely correlated with tolerance to the stress. The positive antioxidant response in Carrizo citrange (Poncirus trifoliata L. Raf. x Citrus sinensis L. Osb.) and Citrumelo CPB 4475 (Poncirus trifoliata L. Raf. x Citrus paradisi L. Macf.) might be responsible for a higher tolerance to flooding stress, whereas in Cleopatra mandarin (Citrus reshni Hort. Ex Tan.), the early accumulation of MDA seems to be associated to an impaired ability for H2O2 scavenging.