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Abstract Background Plant hormones play a pivotal role in several physiological processes during a plant's life cycle, from germination to senescence, and the determination of endogenous concentrations of hormones is essential to elucidate the role of a particular hormone in any physiological process. Availability of a sensitive and rapid method to quantify multiple classes of hormones simultaneously will greatly facilitate the investigation of signaling networks in controlling specific developmental pathways and physiological responses. Due to the presence of hormones at very low concentrations in plant tissues (10-9 M to 10-6 M) and their different chemistries, the development of a high-throughput and comprehensive method for the determination of hormones is challenging. Results The present work reports a rapid, specific and sensitive method using ultrahigh-performance liquid chromatography coupled to electrospray ionization tandem spectrometry (UPLC/ESI-MS/MS) to analyze quantitatively the major hormones found in plant tissues within six minutes, including auxins, cytokinins, gibberellins, abscisic acid, 1-amino-cyclopropane-1-carboxyic acid (the ethylene precursor), jasmonic acid and salicylic acid. Sample preparation, extraction procedures and UPLC-MS/MS conditions were optimized for the determination of all plant hormones and are summarized in a schematic extraction diagram for the analysis of small amounts of plant material without time-consuming additional steps such as purification, sample drying or re-suspension. Conclusions This new method is applicable to the analysis of dynamic changes in endogenous concentrations of hormones to study plant developmental processes or plant responses to biotic and abiotic stresses in complex tissues. An example is shown in which a hormone profiling is obtained from leaves of plants exposed to salt stress in the aromatic plant, Rosmarinus officinalis.

The advanced molecular tools provide critical inputs in uncovering the regulatory mechanisms underlying plants' adaptation to abiotic stress. Presented holistic studies were done on the barley crown tissue being essential for plant performance under various environmental stimuli. To investigate the effect of brassinosteroids (BRs), the known players in stress management, on molecular response of this tissue to drought, the genotypes with different BRs signal transduction efficiency were employed. Large-scale transcriptomic and proteomic profiling confirmed the specific re-modeling of behavior of the BRs-insensitive barley uzu1.a mutant under drought. On the other hand, a set of genes expressed independently of the genotype was identified, including dehydrin encoding genes. This study also uncovered the candidate genes to be linkers of phytohormones crosstalk. Importantly, we detected the converging upregulation of several proteins and encoding genes under drought, including late embryogenesis abundant proteins and chaperones; they represent a promising target for cereals' improvement. Moreover, the greatest variation between genotypes in accumulation of BRs in the crown tissue exposed to drought was observed for castasterone. Presented multi-omics, high-throughput results enhanced the understanding of molecular response to drought in crown tissue. The new insight was provided into the relationships between gene expression, protein and phytohormone content in barley plants of different BRs signaling., Competing Interests: The authors have declared that no competing interests exist., (Copyright: © 2025 Kuczyńska et al. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.)

Background: Flower load in peach is an important determinant of final fruit quality and is subjected to cost-effective agronomical practices, such as the thinning, to finely balance the sink-source relationships within the tree and drive the optimal amount of assimilates to the fruits. Floral transition in peach buds occurs as a result of the integration of specific environmental signals, such as light and temperature, into the endogenous pathways that induce the meristem to pass from vegetative to reproductive growth. The cross talk and integration of the different players, such as the genes and the hormones, are still partially unknown. In the present research, transcriptomics and hormone profiling were applied on bud samples at different developmental stages. A gibberellin treatment was used as a tool to identify the different phases of floral transition and characterize the bud sensitivity to gibberellins in terms of inhibition of floral transition., Results: Treatments with gibberellins showed different efficacies and pointed out a timeframe of maximum inhibition of floral transition in peach buds. Contextually, APETALA1 gene expression was shown to be a reliable marker of gibberellin efficacy in controlling this process. RNA-Seq transcriptomic analyses allowed to identify specific genes dealing with ROS, cell cycle, T6P, floral induction control and other processes, which are correlated with the bud sensitivity to gibberellins and possibly involved in bud development during its transition to the reproductive stage. Transcriptomic data integrated with the quantification of the main bioactive hormones in the bud allowed to identify the main hormonal regulators of floral transition in peach, with a pivotal role played by endogenous gibberellins and cytokinins., Conclusions: The peach bud undergoes different levels of receptivity to gibberellin inhibition. The stage with maximum responsiveness corresponded to a transcriptional and hormonal crossroad, involving both flowering inhibitors and inductors. Endogenous gibberellin levels increased only at the latest developmental stage, when floral transition was already partially achieved, and the bud was less sensitive to exogenous treatments. A physiological model summarizes the main findings and suggests new research ideas to improve our knowledge about floral transition in peach., (© 2024. The Author(s).)

Desiccation tolerance in vegetative tissues enables resurrection plants to remain quiescent under severe drought and rapidly recover full metabolism once water becomes available. Barbacenia graminifolia is a resurrection plant that occurs at high altitudes, typically growing on rock slits, exposed to high irradiance and limited water availability. We analyzed the levels of reactive oxygen species (ROS) and antioxidants, carotenoids and its cleavage products, and stress-related phytohormones in fully hydrated, dehydrated, and rehydrated leaves of B. graminifolia . This species exhibited a precise adjustment of its antioxidant metabolism to desiccation. Our results indicate that this adjustment is associated with enhanced carotenoid and apocarotenoids, α-tocopherol and compounds of ascorbate-glutathione cycle. While α-carotene and lutein increased in dried-leaves suggesting effective protection of the light-harvesting complexes, the decrease in β-carotene was accompanied of 10.2-fold increase in the content of β-cyclocitral, an apocarotenoid implicated in the regulation of abiotic stresses, compared to hydrated plants. The principal component analysis showed that dehydrated plants at 30 days formed a separate cluster from both hydrated and dehydrated plants for up to 15 days. This regulation might be part of the protective metabolic strategies employed by this resurrection plant to survive water scarcity in its inhospitable habitat., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2024 Vieira, Gaspar, Caldeira, Munné-Bosch and Braga.)

Mature forests and their extremely old trees are rare and threatened ancient vestiges in remote European high-mountain regions. Here, we analyze the role that extremely long-living trees have in mature forests biodiversity in relation to their singular traits underlying longevity. Tree size and age determine relative growth rates, bud abortion, and the water status of long-living trees. The oldest trees suffer indefectible age-related constraints but possess singular evolutionary traits defined by fitness adaptation, modular autonomy, and a resilient metabolism that allow them to have irreplaceable roles in the ecosystem as biodiversity anchors of vulnerable lichen species like Letharia vulpina . We suggest that the role of ancient trees as unique biodiversity reservoirs is linked to their singular physiological traits associated with longevity. The set of evolutionarily plastic tools that can only be provided by centuries or millennia of longevity helps the oldest trees of mature forests drive singular ecological relationships that are irreplaceable and necessary for ecosystem dynamics., Competing Interests: Competing interests statement:The authors declare no competing interest.

Mediterranean-type ecosystems provide a unique opportunity to study parasitic plant-host interactions, such as the relationship between the dominant shrub Cistus albidus L. and the root holoparasitic plant Cytinus hypocistis L. We examined this interaction (i) locally, by measuring the hormonal profiling of the interaction zone between the holoparasitic plant and the host, and (ii) systemically, by examining the hormonal profiling and physiological status of leaves from infested and uninfested plants. Furthermore, we explored how temporal variation (seasonal effects) and geographical location influenced the systemic hormonal and physiological response of leaves. Results shed light on tissue-related variations in hormones, suggesting the parasite exerted a sink effect, mainly influenced by cytokinins. Jasmonates triggered a defense response in leaves, far from the infestation point, and both jasmonates and abscisic acid (ABA) appeared to be involved in the tolerance to holoparasitism when plants were simultaneously challenged with summer drought. Parasitism did not have any major negative impact on the host, as indicated by physiological stress markers in leaves, thus indicating a high tolerance of the shrub C. albidus to the root holoparasitic plant C. hypocistis. Rather, parasitism seemed to exert a priming-like effect and some compensatory effects were observed (increased chlorophyll contents) in the host under mild climatic conditions. We conclude that (i) cytokinins, jasmonates and ABA play a role at the local and systemic levels in the response of C. albidus to the biotic stress caused by C. hypocistis, and that (ii) seasonal changes in environmental conditions and geographical location may impact holoparasitic plant-host interactions in the field, modulating the physiological response., (© The Author(s) 2023. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Increasing drought phenomena pose a serious threat to agricultural productivity. Although plants have multiple ways to respond to the complexity of drought stress, the underlying mechanisms of stress sensing and signaling remain unclear. The role of the vasculature, in particular the phloem, in facilitating inter-organ communication is critical and poorly understood. Combining genetic, proteomic and physiological approaches, we investigated the role of AtMC3, a phloem-specific member of the metacaspase family, in osmotic stress responses in Arabidopsis thaliana. Analyses of the proteome in plants with altered AtMC3 levels revealed differential abundance of proteins related to osmotic stress pointing into a role of the protein in water-stress-related responses. Overexpression of AtMC3 conferred drought tolerance by enhancing the differentiation of specific vascular tissues and maintaining higher levels of vascular-mediated transportation, while plants lacking the protein showed an impaired response to drought and inability to respond effectively to the hormone abscisic acid. Overall, our data highlight the importance of AtMC3 and vascular plasticity in fine-tuning early drought responses at the whole plant level without affecting growth or yield., (© 2023 The Authors. New Phytologist © 2023 New Phytologist Foundation.)

Phytohormones are naturally occurring small organic molecules found at low concentrations in plants. They perform essential functions in growth and developmental processes, from organ initiation to senescence, including fruit ripening. These regulatory molecules are studied using different experimental approaches, such as performing exogenous applications, evaluating endogenous levels, and/or obtaining genetically modified lines. Here, we discuss the advantages and limitations of current experimental approaches used to study active biomolecules modulating fruit ripening, focusing on melatonin. Although melatonin has been implicated in fruit ripening in several model fruit crops, current knowledge is affected by the different experimental approaches used, which have given different and sometimes even contradictory results. The methods of application and the doses used have produced different results in studies based on exogenous applications, while different measurement methods and ways of expressing results explain most of the variability in studies using correlative analyses. Furthermore, studies on genetically modified crops have focused on tomato (Solanum lycopersicum L.) plants only. However, TILLING and CRISPR methodologies are becoming essential tools to complement the results from the experimental approaches described above. This will not only help the scientific community better understand the role of melatonin in modulating fruit ripening, but it will also help develop technological advances to improve fruit yield and quality in major crops. The combination of various experimental approaches will undoubtedly lead to a complete understanding of the function of melatonin in fruit ripening in the near future, so that this knowledge can be effectively transferred to the field., Competing Interests: Conflict of interest statement. None declared., (© American Society of Plant Biologists 2023. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

The raspberry ( Rubus idaeus L.) fruit is characterized by its richness in functional molecules and high nutritional value, but the high rate of fruit softening limits its quality during postharvest. Raspberry drupelets have a particular ripening regulation, depending partially on the effect of ethylene produced from the receptacle. However, the possible role of abscisic acid (ABA) in the modulation of quality parameters during the ripening of raspberry is unclear. This study characterized the fruit quality-associated parameters and hormonal contents during fruit development in two seasons. The quality parameters showed typical changes during ripening: a drastic loss of firmness, increase in soluble solids content, loss of acidity, and turning to a red color from the large green stage to fully ripe fruit in both seasons. A significant increase in the ABA content was observed during the ripening of drupelets and receptacles, with the higher content in the receptacle of ripe and overripe stages compared to the large green stage. Moreover, identification of ABA biosynthesis-(9-cis-epoxycarotenoid dioxygenase/NCED) and ABA receptor-related genes (PYRs-like receptors) showed three genes encoding RiNCEDs and nine genes for RiPYLs. The expression level of these genes increased from the large green stage to the full-ripe stage, specifically characterized by a higher expression of RiNCED1 in the receptacle tissue. This study reports a consistent concomitant increase in the ABA content and the expression of RiNCED1, RiPYL1, and RiPYL8 during the ripening of the raspberry fruit, thus supporting the role for ABA signaling in drupelets.

Ancient trees are life history longevity winners that mostly persist in remote and environmentally harsh mountainous areas. Here, we performed a multifeature analysis in a protected mature mountain pine (Pinus uncinata) forest to identify the morphological and physiological traits that make these trees unique. We compared the physiology of meristematic and somatic tissues (apical buds and needles, respectively) from juvenile, mature young, mature old, and mature ancient trees under cold stress and nonstress conditions. We successfully identified key morphological features of extreme longevity at the organism level, as well as various growth, vigor, stress, and dormancy markers underlying extreme longevity in old and ancient trees. Results indicated that evolution has exerted selective pressure on specific physiological traits that make trees become longevity winners (<0.1% of the tree population were ancient trees, with an average trunk diameter >100 cm and an estimated age of 700 years). Traits entailing longevity not only included apical dominance loss, epicormic growth, and modular senescence, but also an extreme plasticity in both meristematic and somatic tissues (buds and needles, respectively), as shown by various physiological markers. In conclusion, ancient trees are oddities that not only possess a unique ecological value but also show divergent physiological behaviors selected during their evolution to allow them to cope with adversities and attain long life., Competing Interests: Conflict of interest statement. All authors declare no conflict of interest., (© American Society of Plant Biologists 2022. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Grapevine cultivation, such as the whole horticulture, is currently challenged by several factors, among which the extreme weather events occurring under the climate change scenario are the most relevant. Within this context, the present study aims at characterizing at the berry level the physiological response of Vitis vinifera cv. Sauvignon Blanc to sequential stresses simulated under a semi-controlled environment: flooding at bud-break followed by multiple summer stress (drought plus heatwave) occurring at pre-vèraison. Transcriptomic and metabolomic assessments were performed through RNASeq and NMR, respectively. A comprehensive hormone profiling was also carried out. Results pointed out a different response to the heatwave in the two situations. Flooding caused a developmental advance, determining a different physiological background in the berry, thus affecting its response to the summer stress at both transcriptional levels, with the upregulation of genes involved in oxidative stress responses, and metabolic level, with the increase in osmoprotectants, such as proline and other amino acids. In conclusion, sequential stress, including a flooding event at bud-break followed by a summer heatwave, may impact phenological development and berry ripening, with possible consequences on berry and wine quality. A berry physiological model is presented that may support the development of sustainable vineyard management solutions to improve the water use efficiency and adaptation capacity of actual viticultural systems to future scenarios.

Hairy roots are made after the integration of a small set of genes from Agrobacterium rhizogenes in the plant genome. Little is known about how this small set is linked to their hormone profile, which determines development, morphology, and levels of secondary metabolite production. We used C. asiatica hairy root line cultures to determine the putative links between the rol and aux gene expressions with morphological traits, a hormone profile, and centelloside production. The results obtained after 14 and 28 days of culture were processed via multivariate analysis and machine-learning processes such as random forest, supported vector machines, linear discriminant analysis, and neural networks. This allowed us to obtain models capable of discriminating highly productive root lines from their levels of genetic expression ( rol and aux genes) or from their hormone profile. In total, 12 hormones were evaluated, resulting in 10 being satisfactorily detected. Within this set of hormones, abscisic acid (ABA) and cytokinin isopentenyl adenosine (IPA) were found to be critical in defining the morphological traits and centelloside content. The results showed that IPA brings more benefits to the biotechnological platform. Additionally, we determined the degree of influence of each of the evaluated genes on the individual hormone profile, finding that aux1 has a significant influence on the IPA profile, while the rol genes are closely linked to the ABA profile. Finally, we effectively verified the gene influence on these two specific hormones through feeding experiments that aimed to reverse the effect on root morphology and centelloside content., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Alcalde, Müller, Munné-Bosch, Landín, Gallego, Bonfill, Palazon and Hidalgo-Martinez.)

Improved nutrient use efficiency together with the use of biostimulants have been little explored thus far to improve fruit yield and quality in economically relevant crops. The aim of this study was to determine the additive or synergistic effects, if any, of the application of an enzyme hydrolyzed animal protein biostimulant (Pepton) combined with priming with low nitrogen (N) in the production and quality of greenhouse tomatoes. Biostimulant treatment (Pepton at a dose equivalent of 4 kg/ha) was applied by ferti-irrigation for 2 months during the vegetative phase both in controls (watered with nutrient solution) and nutrient efficient crop (NEC), in which plants were primed with low N by exposing them to a 30% N deficiency for 2 months, and then recovered for 1 month before fruit production. Foliar water and N contents, pigments, maximum PSII efficiency (Fv/Fm ratio), and phytohormones [including abscisic acid (ABA), salicylic acid (SA), jasmonic acid (JA), and cytokinins] were measured prior and at 4 and 8 weeks after the first application. Fruit production and quality [as indicated by total soluble sugars (TSS) and acidity (TA), and the contents of lycopene, vitamin E, and vitamin C] were measured 1 month later at harvest. Priming with low N availability (NEC plants) doubled ( p  < 0.001) fruit production (due to an increase in the number of fruits), tended to increase ( p  = 0.057) by 20% the amount of TSS and increased ( p  < 0.05) the contents of lycopene (by 90%) and vitamin E (by 40%). Pepton displayed a tendency, almost significant, to improve ( p  = 0.054) total fruit production both in control and NEC plants, thus showing an additive effect to low N priming in boosting fruit production. Pepton maintained fruit quality in terms of sugar accumulation, total acidity and the contents of carotenoids, vitamins C and E. Pepton-related improvement in fruit production seemed to be related, at least partially, to an increased accumulation of cytokinins and photosynthetic pigments in leaves, which might favor vegetative vigor and ultimately fruit yield. In conclusion, Pepton application was effective in improving the yield of greenhouse tomatoes showing additive effect with low N priming, without negatively affecting fruit quality., Competing Interests: JP is employed by APC Europe S.L. The remaining authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2022 Mesa, Polo, Casadesús, Gómez and Munné-Bosch.)

Leaf senescence, the last stage of the developmental program of leaves, can be induced by both internal and external signals. Cold stress-induced leaf senescence is an efficient strategy to overcome winter temperatures. In this work, we studied leaf senescence in yellow flag ( Iris pseudacorus L.) individuals growing in a natural wetland, not only considering its relationship with external and internal cues, but also the plant developmental program, and the biological significance of rhizomes, storage organs that remain viable through winter. Total chlorophyll contents and the maximum efficiency of PSII ( F v / F m ratio) decreased in senescing leaves, which was associated with a sharp increase in abscisic acid (ABA) contents. Furthermore, total cytokinin and 2-isopentenyladenine contents decreased in December compared to November, as plants became more stressed due to a decline in air temperatures. ABA increases in senescing leaves increased in parallel to reductions in violaxanthin. Rhizomes also accumulated large amounts of ABA during winter, while roots did not, and neither roots nor rhizomes accumulated 9- cis -epoxycarotenoids, thus suggesting ABA, which might play a role in conferring cold tolerance to this subterranean organ, may result from phloem transport from senescing leaves. It is concluded that (i) leaf senescence is a highly regulated physiological process in yellow flag playing a key role in the modulation of the entire plant developmental program, and (ii) ABA plays a major role not only in the regulation of leaf senescence but also in the establishment of cold tolerance in rhizomes, two processes that appear to be intimately interconnected., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Caselles, Casadesús and Munné-Bosch.)

Drought can lead to important shifts in population dynamics if it occurs during seedling establishment. With the aim of elucidating the underlying mechanisms of drought tolerance and resilience, here we monitored the survival of seedlings of the Mediterranean shrub Cistus albidus L. throughout a year growing in the natural Park of the Montserrat Mountains (Spain) and, additionally, we studied the response to severe drought and subsequent recovery after rewatering of seedlings grown in growth chambers. To find possible mechanisms explaining how seedlings respond to drought, growth and survival together with physiological-related parameters such as chlorophyll contents, vitamin E and stress-related phytohormones were measured. We found that survival decreased by 30% at the end of summer and that the main proxy of seedling survival was total chlorophyll. This proxy was further confirmed in the growth chambers, where we found that seedlings that recovered from drought had higher levels of total chlorophyll compared with the seedlings that did not recover. Furthermore, modulation of vitamin E and jasmonates contents appeared to be crucial in the drought response of C. albidus seedlings. We propose a prediction model of survival that includes total chlorophyll height, leaf mass area and maximum photosystem II efficiency with chlorophyll contents being a good long-term predictor of C. albidus seedling survival under severe stress, which, in turn, could help to better foresee population fluctuations in the field., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

Premise: Biological invasions and climate change are major threats to biodiversity. It is therefore important to anticipate how the climate changes projected for Southern Europe would affect the ecophysiological performance of the invasive South African plant, Carpobrotus edulis (ice plant or sour fig), and its capacity to undergo rapid adaptive evolution., Methods: We manipulated the climate conditions in a field plot located on the island of Sálvora (northwest of the Iberian Peninsula) to establish a full factorial experiment with C. edulis plants transplanted from four native (southern African) and four invasive (northwestern Iberian Peninsula) populations. Throughout 14 months we measured growth and functional traits of this species under two temperatures (control vs. increased), and two rainfall levels (control vs. reduced)., Results: Temperature increased photochemical efficiency and relative growth rate of C. edulis. Rainfall modulated some of the effects of temperature on C and N isotopic composition, and pigment contents. Invasive populations showed lower root mass allocation and higher survival rates, as well as increased water use efficiency, lipid peroxidation, chlorophyll, and xanthophyll cycle pigment contents than native populations., Conclusions: The increased growth and physiological performances observed under our experimental conditions suggest that the expected climate changes would further promote the invasion of C. edulis. Differences between native and invasive genotypes in survival and functional traits revealed that populations have diverged during the process of invasion, what gives support to the invasiveness hypothesis. Our findings highlight the importance of analyzing intraspecific variability in functional responses to better predict how invasive species will respond to environmental changes., (© 2021 The Authors. American Journal of Botany published by Wiley Periodicals LLC on behalf of Botanical Society of America.)

Mastic trees (Pistacia lentiscus L.) are dioecious perennial plants that are highly adapted to Mediterranean climates but display a high sensitivity to winter periods. In order to understand how sex, leaf phenology and ecological context could condition sensitivity to winter and associated mechanisms to acclimate to these conditions, photoinhibition and photo-oxidative stress markers were examined in mastic trees (P. lentiscus) from a natural population growing in the Garraf Natural Park for a consecutive 12-month period (seasonal study), as well as in three populations naturally growing in the Montseny Natural Park, including the highest altitudes described for this species, during winter (altitudinal study). Results from these studies indicate that both the winter period and higher elevation influenced the degree of photoinhibition, but this was not conditioned by sex. In fact, winter photoinhibition occurred transiently even though it was accompanied by chlorophyll loss and malondialdehyde contents. Stress acclimation was achieved through biochemical adjustments in chloroplasts, characterized by anthocyanin shielding, increased de-epoxidation state of the xanthophyll cycle as well as tocopherol accumulation, and phenological adaptations, the latter allowing a complete resetting of the physiological performance of leaves. Moreover, although females showed higher lipid peroxidation than males during the coldest winter months, at the highest elevation and during flowering in spring, this oxidative stress was mild and transient with no negative consequences for the physiology of plants. It is concluded that evergreen mastic trees acclimate to winter conditions and higher elevations by activation of antioxidant defenses together with phenological adjustments, altogether playing a crucial role in plant survival. Sexual dimorphism in mastic trees appears as a relevant factor when considering sensitivity to photo-oxidative stress in winter and altitudinal conditions., (© The Author(s) 2021. Published by Oxford University Press. All rights reserved. For permissions, please e-mail: journals.permission@oup.com.)

Photosynthesis is not only essential for plants, but it also sustains life on Earth. Phytohormones play crucial roles in developmental processes, from organ initiation to senescence, due to their role as growth and developmental regulators, as well as their central role in the regulation of photosynthesis. Furthermore, phytohormones play a major role in photoprotection of the photosynthetic apparatus under stress conditions. Here, in addition to discussing our current knowledge on the role of the phytohormones auxin, cytokinins, gibberellins, and strigolactones in promoting photosynthesis, we will also highlight the role of abscisic acid beyond stomatal closure in modulating photosynthesis and photoprotection under various stress conditions through crosstalk with ethylene, salicylates, jasmonates, and brassinosteroids. Furthermore, the role of phytohormones in controlling the production and scavenging of photosynthesis-derived reactive oxygen species, the duration and extent of photo-oxidative stress and redox signaling under stress conditions will be discussed in detail. Hormones have a significant impact on the regulation of photosynthetic processes in plants under both optimal and stress conditions, with hormonal interactions, complementation, and crosstalk being important in the spatiotemporal and integrative regulation of photosynthetic processes during organ development at the whole-plant level., (© The Author(s) 2021. Published by Oxford University Press on behalf of American Society of Plant Biologists.)

Although photosynthesis is essential to sustain life on Earth, not all plants use sunlight to synthesize nutrients from carbon dioxide and water. Holoparasitic plants, which are important in agricultural and natural ecosystems, are dependent on other plants for nutrients. Phytohormones are crucial in holoparasitic plant-host interactions, from seed germination to senescence, not only because they act as growth and developmental regulators, but also because of their central role in the regulation of host photosynthesis and source-sink relations between the host and the holoparasitic plant. Here, we compile and discuss current knowledge on the impact and ecophysiology of holoparasitic plants (such as the broomrapes Orobanche sp. and Phelipanche sp.) that infest economically important dicotyledonous crops in Mediterranean agroecosystems (legumes [Fabaceae], sunflowers [Helianthus sp.], or tomato [Solanum lycopersicum] plants). We also highlight the role of holoparasitic plant-host interactions (such as those between Cytinus hypocistis and various shrubs of the genus Cistus) in shaping natural Mediterranean ecosystems. The roles of phytohormones in controlling plant-host interactions, abiotic factors in parasitism, and the biological significance of natural seed banks and how dormancy and germination are regulated, will all be discussed. Holoparasitic plants are unique organisms; improving our understanding of their interaction with hosts as study models will help us to better manage parasitic plants, both in agricultural and natural ecosystems., (© American Society of Plant Biologists 2021. All rights reserved. For permissions, please email: journals.permissions@oup.com.)

Sweet cherry is an important non-climacteric fruit with a high commercial interest, but exploitation of sweet cherry trees ( Prunus avium L.) in orchards is usually subject to important economic losses due to fruit decay by pathogenic fungi and other microorganisms. Sweet cherries development and ripening are characterized by profound physiological changes in the fruit, among which the phytohormone abscisic acid (ABA) plays a pivotal role. In addition, sweet cherries are usually affected by fruit decay pathogens, and the role of other stress-related hormones such as jasmonic acid (JA) and salicylic acid (SA) may also be of paramount importance, not only from a developmental point of view, but also from a fruit-microbe interaction perspective. Here, a tissue-specific hormone quantification by LC-MS/MS, including the contents of JA, SA, and ABA, in the fruit exocarp and mesocarp of sweet cherries during fruit development from trees growing in a commercial orchard was carried out. Additionally, this study was complemented with the characterization of the culturable epiphytic and endophytic microbial communities of sweet cherries at various stages of fruit development and during cracking lesion formation. Our results revealed a completely differential behavior of phytohormones between both tissues (the exocarp and mesocarp), with a more dynamic exocarp in front of a more stable mesocarp, and with marked variations during fruit development. Microbial epiphytic community was mainly composed by yeasts, although rot-causing fungi like Alternaria spp. were always also present throughout fruit development. Endophytic colonization was poor, but it increased throughout fruit development. Furthermore, when the exocarp was naturally disrupted in sweet cherries suffering from cracking, the colonization by Alternaria spp. markedly increased. Altogether, results suggest that the fruit exocarp and mesocarp are very dynamic tissues in which endogenous phytohormones not only modulate fruit development and ripening but also fruit-microbe interactions., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Fresno and Munné-Bosch.)

Agricultural practices in grapevines management include water restrictions due to its positive effect on wine quality, especially when applied at fruit ripening. Although the effects of water stress in some groups of phytohormones have already been described in leaves and whole grapes, information regarding tissue-specific variations in hormones during ripening in grapes is scarce. Field-grown grapevines from the cv. "Merlot" were subjected to two differential water supplies, including only rainfed, non-irrigated vines (T0) and vines additionally irrigated with 25Lweek -1 vine -1 (T1). Tissue-specific variations in the hormonal profiling of grapes [including changes in the contents of abscisic acid (ABA), jasmonic acid (JA), salicylic acid (SA), the ethylene precursor 1-amino-cyclopropane-1-carboxylic acid (ACC), the auxin indole-3-acetic acid, gibberellins 1, 3, 4, and 7 (GA 1 , GA 3 , GA 4 , and GA 7 ), the cytokinins trans -zeatin, and 2-isopentenyl adenine, including as well their respective ribosylated forms] were periodically evaluated from veraison to harvest. The hormonal profiling in leaves was also measured at the beginning and end of the season for comparison. Results showed that grape growth dynamics were transiently affected by the differences in water regimes, the increased water supply leading to an accelerated growth, slightly reduced accumulation of sugars, and transiently lowered pH, although grape quality did not differ between treatments at harvest. Hormonal profiling of whole berries did not reveal any difference in the endogenous contents of phytohormones between treatments, except for a transient decrease in GA 4 contents in T1 compared to T0 vines, which was not confirmed at the tissular level. Hormonal profiling at the tissue level highlighted a differential accumulation of phytohormones during ripening in berry tissues, with pulps being particularly poor in ABA, JA, and SA contents, seeds particularly accumulating ACC, gibberellins, and zeatin-type cytokinins, and the skin being particularly rich in auxin and active cytokinins. Changes in water supply led to very small and transient changes in the endogenous contents of phytohormones in the seeds, pulp, and skin of berries, the most remarkable variations being observed in cytokinin contents, which increased earlier [between 5 and 12days after veraison (DAV)] but later kept more constant in the skin from T1 compared to T0 vines and were also 3-fold higher at 40 DAV in seeds of T1 compared to T0 vines. It is concluded that small changes in water supply can trigger hormonal-driven physiological adjustments at the tissular level affecting the evolution of fruit growth and quality throughout grape berry ripening., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2021 Ribalta-Pizarro, Muñoz and Munné-Bosch.)

Aging, stress, and senescence in plants are interconnected processes that determine longevity. We focus here on compiling and discussing our current knowledge on the mechanisms of development that long-lived perennial plants have evolved to prevent and delay senescence. Clonal and nonclonal perennial herbs of various life forms and longevities will be particularly considered to illustrate what biological diversity can teach us about aging as a universal phenomenon. Source-sink relations and redox signaling will also be discussed as examples of regulatory mechanisms of senescence at the organ level. Whether or not effective mechanisms that biological diversity has evolved to completely prevent the wear and tear of aging will be applicable to human aging in the near future ultimately depends on ethical aspects.

The identification of drought-tolerant olive tree genotypes has become an urgent requirement to develop sustainable agriculture in dry lands. However, physiological markers linking drought tolerance with mechanistic effects operating at the cellular level are still lacking, in particular under severe stress, despite the urgent need to develop these tools in the current frame of global change. In this context, 1-year-old olive plants growing in the greenhouse and with a high intra-specific variability (using various genotypes obtained either from cuttings or seeds) were evaluated for drought tolerance under severe stress. Growth, plant water status, net photosynthesis rates, chlorophyll contents and the extent of photo- and antioxidant defenses (including the de-epoxidation state of the xanthophyll cycle, and the contents of carotenoids and vitamin E) were evaluated under well-watered conditions and severe stress (by withholding water for 60 days). Plants were able to continue photosynthesizing under severe stress, even at very low leaf water potential of -4 to -6 MPa. This ability was achieved, at least in part, by the activation of photo- and antioxidant mechanisms, including not only increased xanthophyll cycle de-epoxidation, but also enhanced α-tocopherol contents. "Zarrazi" (obtained from seeds) and "Chemlali" (obtained from cuttings) showed better performance under severe water stress compared to the other genotypes, which was associated to their ability to trigger a higher antioxidant protection. It is concluded that (i) drought tolerance among the various genotypes tested is associated with antioxidant protection in olive trees, (ii) the extent of xanthophyll cycle de-epoxidation is strongly inversely related to photosynthetic rates, and (iii) vitamin E accumulation is sharply induced upon severe chlorophyll degradation., Competing Interests: The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest., (Copyright © 2020 Baccari, Elloumi, Chaari-Rkhis, Fenollosa, Morales, Drira, Ben Abdallah, Fki and Munné-Bosch.)

Oxylipins (compounds derived from the oxidation of polyunsaturated fatty acids) are essential in retrograde signaling emanating from plastids to the nucleus during plant developmental and stress responses. In this graphical review, we provide an overview of the chemical structure, biosynthesis and role of oxylipins, as both redox and hormonal signals, in controlling plant development and stress responses. We also briefly summarize current gaps in the understanding of the involvement of oxylipins in plastidial retrograde signaling to highlight future avenues for research., (Copyright © 2020. Published by Elsevier B.V.)

A complex network of cellular receptors, RNA targeting pathways, and small-molecule signaling provides robust plant immunity and tolerance to viruses. To maximize their fitness, viruses must evolve control mechanisms to balance host immune evasion and plant-damaging effects. The genus Potyvirus comprises plant viruses characterized by RNA genomes that encode large polyproteins led by the P1 protease. A P1 autoinhibitory domain controls polyprotein processing, the release of a downstream functional RNA-silencing suppressor, and viral replication. Here, we show that P1Pro, a plum pox virus clone that lacks the P1 autoinhibitory domain, triggers complex reprogramming of the host transcriptome and high levels of abscisic acid (ABA) accumulation. A meta-analysis highlighted ABA connections with host pathways known to control RNA stability, turnover, maturation, and translation. Transcriptomic changes triggered by P1Pro infection or ABA showed similarities in host RNA abundance and diversity. Genetic and hormone treatment assays showed that ABA promotes plant resistance to potyviral infection. Finally, quantitative mathematical modeling of viral replication in the presence of defense pathways supported self-control of polyprotein processing kinetics as a viral mechanism that attenuates the magnitude of the host antiviral response. Overall, our findings indicate that ABA is an active player in plant antiviral immunity, which is nonetheless evaded by a self-controlled RNA virus.

Biostimulants may be particularly interesting for application in agricultural and horticultural crops since they can exert a growth-promoting effect on roots. This may be important for promoting longitudinal and lateral root growth and therefore increasing belowground vegetative growth, which may in turn lead to improved aboveground vegetative growth and increased yields. Here, we examined the effects and mechanism of action of an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) on the root growth of tomato plants, with an emphasis on its possible role on chorismate-derived hormones (auxin, salicylic acid, and melatonin). Tomato plants growing in hydroponic systems were exposed to either nutrient stress conditions (experiment 1) or suboptimal temperatures (experiment 2) in a greenhouse, and the concentration of auxin, salicylic acid, and melatonin in roots were measured just prior and after the application of the biostimulant. Results showed that the application of Pepton exerted a growth-promoting effect on roots in plants growing under suboptimal conditions, which might be associated with enhanced salicylic acid levels in roots. The extent of effects of this enzymatically hydrolyzed animal protein-based biostimulant might strongly depend on the growth conditions and stage of root system development. It is concluded that an enzymatically hydrolyzed animal protein-based biostimulant (Pepton) may exert a positive effect enhancing primary and lateral root growth of tomato plants growing under suboptimal conditions, by stimulating the biosynthesis of specific hormonal pathways, such as salicylic acid under stress., (Copyright © 2020 Casadesús, Pérez-Llorca, Munné-Bosch and Polo.)

Background: Seed longevity and vigor assessment is crucial for efficient ex situ biodiversity conservation in genebanks but may also have potential applications for the understanding of ecological processes and in situ biodiversity conservation. In fact, one of the factors determining the persistence of invasive species, a main threat to global biodiversity, is the generation of soil seed banks where seeds may remain viable for several years. Artificial seed aging tests using high temperatures and high relative humidity have been described for seed longevity estimation but have been mainly optimized for species with commercial interest. Thus, the aim of the study is to define a rapid and sensitive method to assess seed longevity and vigor through accelerated aging in the worldwide distributed invasive species Carpobrotus edulis to provide tools to biodiversity managers to evaluate invasive potential and develop effective post-eradication plans., Results: Slow seed deterioration rate was obtained when C. edulis seeds were subjected to common accelerated aging temperatures (43-45 °C). This contrasts with the rapid viability decay between 24-72 h when seeds were subjected to temperatures superior to 55 °C, a strong inflection point for this species' thermosensitivity. Relative humidity also played a role in defining seed survival curves, but only at high temperatures, speeding up the deterioration process. The selected aging conditions, 55 °C at 87% relative humidity were tested over two C. edulis populations and three measures were proposed to parametrize the differential sigmoidal seed survival curves, defining the seed resistance to deterioration (L 5 , aging time where 95% of seeds maintain their viability), medium longevity (L 50 , 50% of seeds lose their viability) and lethal aging time (L 95 , 95% of viability loss)., Conclusions: An accelerated aging test at 55 °C and 87% relative humidity constitutes a rapid and sensitive method that can be performed within a working week, allowing managers to easily test seed vigor and longevity. This test may contribute to assess invasive potential, design effective monitoring programs and soil seed bank eradication treatments., Competing Interests: Competing interestsThe authors declare that they have no competing interests., (© The Author(s) 2020.)

Background: Jasmonates play an important role in plant stress and defence responses and are also involved in the regulation of anthocyanin synthesis in response to sucrose availability. Here we explore the signalling interactions between sucrose and jasmonates in response to cold stress in Arabidopsis., Results: Sucrose and cold treatments increased anthocyanin content additively. Comprehensive profiling of phytohormone contents demonstrated that jasmonates, salicylic acid and abscisic acid contents increased in response to sucrose treatment in plants grown on agar, but remained considerably lower than in plants grown in compost. The gibberellin GA 3 accumulated in response to sucrose treatment but only at warm temperature. The role of jasmonate signalling was explored using the jasmonate response mutants jar1-1 and coi1-16. While the jar1-1 mutant lacked jasmonate-isoleucine and jasmonate-leucine, it accumulated 12-oxo-phytodienoic acid at low temperature on agar medium. Altered patterns of abscisic acid accumulation and higher sugar contents were found in the coi1-16 mutant when grown in compost. Both mutants were able to accumulate anthocyanin and to cold acclimate, but the jar-1-1 mutant showed a larger initial drop in whole-rosette photosystem II efficiency upon transfer to low temperature., Conclusions: Hormone contents are determined by interactions between temperature and sucrose supply. Some of these effects may be caused indirectly through senescence initiation in response to sucrose availability. During cold stress, the adjustments of hormone contents may compensate for impaired jasmonate signalling, enabling cold acclimation and anthocyanin accumulation in Arabidopsis jasmonate response mutants, e.g. through antagonistic interactions between gibberellin and jasmonate signalling.

Background and Aims: Monocarpic plants are those that flower, produce seeds and then die. Although most monocarpic plants are annual or biennial, some of them are perennial. However, relatively little is known regarding the biology of monocarpic perennials. Pyrenean saxifrage (Saxifraga longifolia) is a monocarpic perennial that is well adapted to high-mountain ecosystems. Here, we evaluated altitudinal changes in clonality in various populations growing in their natural habitat with particular emphasis on the physiological costs of clonal growth., Methods: We assessed the percentage of clonal plants in nine populations growing in their natural habitat, as well as the plant stress response of clonal and non-clonal plants, in terms of photoprotection and accumulation of stress-related phytohormones, in a 3-year study at Las Blancas (2100 m a.s.l.). We also evaluated the influence of plant size on the activation of defensive responses to biotic and abiotic stresses., Key Results: We found that 12 % of Pyrenean saxifrage plants growing at the highest altitudes (2100 m a.s.l.) produced lateral rosettes which survived the flowering of the main rosette and shared the same axonomorphic root, thus escaping monocarpic senescence. This clonal growth did not worsen the physiological performance of plants growing at this altitude. Furthermore, increased plant size did not negatively affect the physiology of plants, despite adjustments in endogenous stress-related phytohormones. In contrast, maturity led to rapid physiological deterioration of the rosette, which was associated with monocarpic senescence., Conclusions: This study shows that the evolution of clonality has allowed Pyrenean saxifrage to survive harsh environmental conditions and it provides evidence that harsh environments push plant species to their limits in terms of life form and longevity., (© The Author(s) 2019. Published by Oxford University Press on behalf of the Annals of Botany Company. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com.)

When plants establish outside their native range, their ability to adapt to the new environment is influenced by both demography and dispersal. However, the relative importance of these two factors is poorly understood. To quantify the influence of demography and dispersal on patterns of genetic diversity underlying adaptation, we used data from a globally distributed demographic research network comprising 35 native and 18 nonnative populations of Plantago lanceolata Species-specific simulation experiments showed that dispersal would dilute demographic influences on genetic diversity at local scales. Populations in the native European range had strong spatial genetic structure associated with geographic distance and precipitation seasonality. In contrast, nonnative populations had weaker spatial genetic structure that was not associated with environmental gradients but with higher within-population genetic diversity. Our findings show that dispersal caused by repeated, long-distance, human-mediated introductions has allowed invasive plant populations to overcome environmental constraints on genetic diversity, even without strong demographic changes. The impact of invasive plants may, therefore, increase with repeated introductions, highlighting the need to constrain future introductions of species even if they already exist in an area., Competing Interests: The authors declare no competing interest., (Copyright © 2020 the Author(s). Published by PNAS.)

Invasive plants are expanding their geographical distribution across new regions. Expansion modeling is crucial for geographic prioritization in management policies. However, the assumption of niche conservatism and the lack of information of the species physiological response to the environmental factors determining species presence may hinder predictions. In this study, we aimed to understand the expansion of the widely distributed plant Carpobrotus edulis in Europe. We contrasted introduced and native C. edulis ecological niches and explored the experimental response to temperature, a major determining factor for species distribution, of native and invasive individuals in terms of different biochemical markers. Niche analysis revealed an expansion of the introduced niche to occupy colder climates. Introduced and native individuals showed differential mechanisms facing low temperatures. Individuals from the native range showed an increased sensitivity to chilling, as reflected by photosynthetic pigment degradation, increased de-epoxidation of xanthophylls and the accumulation of the lipophilic antioxidant alpha-tocopherol. The found physiological differentiation towards an increased invasive chilling tolerance of invasive C. edulis individuals together with a high propagule pressure may explain the introduced climatic niche shift to colder climates observed, allowing the extensive expansion of this species in Europe., (© The Author(s) 2019. Published by Oxford University Press and the Society for Experimental Biology.)

Alfalfa ( Medicago sativa L.) is frequently constrained by environmental conditions such as drought. Within this context, it is crucial to identify the physiological and metabolic traits conferring a better performance under stressful conditions. In the current study, two alfalfa cultivars (San Isidro and Zhong Mu) with different physiological strategies were selected and subjected to water limitation conditions. Together with the physiological analyses, we proceeded to characterize the isotopic, hormone, and metabolic profiles of the different plants. According to physiological and isotopic data, Zhong Mu has a water-saver strategy, reducing water lost by closing its stomata but fixing less carbon by photosynthesis, and therefore limiting its growth under water-stressed conditions. In contrast, San Isidro has enhanced root growth to replace the water lost through transpiration due to its more open stomata, thus maintaining its biomass. Zhong Mu nodules were less able to maintain nodule N 2 fixing activity (matching plant nitrogen (N) demand). Our data suggest that this cultivar-specific performance is linked to Asn accumulation and its consequent N-feedback nitrogenase inhibition. Additionally, we observed a hormonal reorchestration in both cultivars under drought. Therefore, our results showed an intra-specific response to drought at physiological and metabolic levels in the two alfalfa cultivars studied.

Mediterranean ecosystems harbor a very important part of Earth's biodiversity, and they are a conservation priority due to the effects of global change. Here, we examined the performance of the semi-deciduous shrub Cistus albidus under Mediterranean conditions during winter, including changes in leaf angle governed by diurnal, seasonal, and positional effects and their relationship with winter photoinhibition and photoprotection. We found marked diurnal variations in leaf angle during the day in autumn, which disappeared as the photoperiod shortened during winter due to a progressive decrease in the predawn leaf angle from November to January. During this period, auxin contents decreased, while those of melatonin increased, and the F v / F m ratio, chlorophyll, and tocopherol contents kept unaltered, thus indicating the absence of photoinhibitory damage. A marked decrease in the leaf angle toward the shoot apex occurred during winter, which was associated with slightly higher F v / F m ratios. An analysis of the inter-individual variability and sun orientation effects on leaf movements in a population growing in the Montserrat mountains revealed a very marked variability of 46.8% in the leaf angle, while F v / F m ratio showed a variation of 7.5% only. West orientation, which was associated with reduced leaf temperatures, but with no differences in the photosynthetic photon flux density, led to enhanced tocopherol contents, while leaf angle, F v / F m ratio, chlorophyll, auxin, and melatonin contents kept unaltered. It is concluded that (1) C. albidus has very effective and fine-regulated photoprotection mechanisms, including an adequate orientation of decussate leaves as part of the developmental program, (2) leaf angle is modulated on a diurnal and seasonal basis, thus contributing to prevent photoinhibition as a first line of defense, and (3) enhanced tocopherol contents help withstand combined high light with low temperature stress in C. albidus growing at high elevation.

Biostimulants may promote growth or alleviate the negative effects of abiotic stress on plant growth eventually resulting in enhanced yields. We examined the mechanism of action of an enzymatically hydrolyzed animal protein-based biostimulant (Pepton), which has previously been shown to benefit growth and yield in several horticultural crops, particularly under stressful conditions. Tomato plants were exposed to well-watered and water-stressed conditions in a greenhouse and the hormonal profiling of leaves was measured during and after the application of Pepton. Results showed that the Pepton application benefited antioxidant protection and exerted a major hormonal effect in leaves of water-stressed tomatoes by increasing the endogenous content of indole-3-acetic acid (auxin), trans -zeatin (cytokinin), and jasmonic acid. The enhanced jasmonic acid content may have contributed to an increased production of tocochromanols because plastochromanol-8 concentration per unit of chlorophyll was higher in Pepton-treated plants compared to controls. In conclusion, the tested Pepton application may exert a positive effect on hormonal balance and the antioxidant system of plants under water stress in an economically important crop, such as tomato plants.

Climacteric and non-climacteric fruits are differentiated by the ripening process, in particular by the involvement of ethylene, high respiration rates and the nature of the process, being autocatalytic or not, respectively. Here, we focus on the biosynthesis, metabolism and function of three compounds (auxin, salicylic acid and melatonin) sharing not only a common precursor (chorismate), but also regulatory functions in plants, and therefore in fruits. Aside from describing their biosynthesis in plants, with a particular emphasis on common precursors and points of metabolic diversion, we will discuss recent advances on their role in fruit ripening and the regulation of bioactive compounds accumulation, both in climacteric and non-climacteric fruits.

Vitamin E is involved in heat stress acclimation. In this issue of Developmental Cell, Fang et al. (2018) uncover a new link between Vitamin E and plant retrograde signaling and show that vitamin E is required for the accumulation of 3'-phosphoadenosine 5'-phosphate, an inhibitor of exoribonucleases, to protect microRNAs from degradation., (Copyright © 2019 Elsevier Inc. All rights reserved.)

Vitamin E, a potent antioxidant either presents in the form of tocopherols and/or tocotrienols depending on the plant species, tissue and developmental stage, plays a major role in protecting lipids from oxidation in seeds. Unlike tocopherols, which have a more universal distribution, the occurrence of tocotrienols is limited primarily to monocot seeds. Dwarf fan palm (Chamaerops humilis var. humilis) seeds accumulate tocotrienols in quiescent and dormant seeds, while tocopherols are de novo synthesized during germination. Here, we aimed to elucidate whether tocopherol biosynthesis is regulated at the transcriptional level during germination in this species. We identified and quantified the expression levels of five genes involved in vitamin E biosynthesis, including TYROSINE AMINOTRANSFERASE (ChTAT), HOMOGENTISATE PHYTYLTRANSFERASE (ChHPT), HOMOGENTISATE GERANYLGERANYL TRANSFERASE (ChHGGT), TOCOPHEROL CYCLASE (ChTC) and TOCOPHEROL γ-METHYLTRANSFERASE (Chγ-TMT). Furthermore, we evaluated to what extent variations in the endogenous contents of hormones and hydrogen peroxide (H2O2) correlated with transcriptional regulation. Results showed an increase of ChTAT and ChHPT levels during seed germination, which correlated with an increase of jasmonic acid (JA), gibberellin4 (GA4), and H2O2 contents, while ChHGGT and Chγ-TMT expression levels decreased, thus clearly indicating vitamin E biosynthesis is diverted to tocopherols rather than to tocotrienols. Exogenous application of jasmonic acid increased tocopherol, but not tocotrienol content, thus confirming its regulatory role in vitamin E biosynthesis during seed germination. It is concluded that the biosynthesis of vitamin E is regulated at the transcriptional level during germination in dwarf fan palm seeds, with ChHPT playing a key role in the diversion of the vitamin E pathway towards tocopherols instead of tocotrienols.

Background: While the role of ethylene in fruit ripening has been widely studied, the contributions of additional plant hormones are less clear. Here we examined the interactions between the transcription factor MaMADS2-box which plays a major role in banana fruit ripening and hormonal regulation. Specifically, we used MaMADS2 repressed lines in transcriptome and hormonal analyses throughout ripening and assessed hormone and gene expression perturbations as compared to wild-type (WT) control fruit., Results: Our analyses revealed major differences in hormones levels and in expression of hormone synthesis and signaling genes mediated by MaMADS2 especially in preclimacteric pulp. Genes encoding ethylene biosynthesis enzymes had lower expression in the pulp of the repressed lines, consistent with reduced ethylene production. Generally, the expression of other hormone (auxin, gibberellins, abscisic acid, jasmonic acid and salicylic acid) response pathway genes were down regulated in the WT pulp prior to ripening, but remained high in MaMADS2 repressed lines. Hormone levels of abscisic acid were also higher, however, active gibberellin levels were lower and auxin levels were similar with MaMADS2 repression as compared to WT. Although abscisic level was higher in MaMADS2 repression, exogenous abscisic acid shortened the time to ethylene production and increased MaMADS2 mRNA accumulation in WT. Exogenous ethylene did not influence abscisic acid level. CRE - a cytokinin receptor, increased its expression during maturation in WT and was lower especially at prebreaker in the repressed line and zeatin level was lower at mature green of the repressed line in comparison to WT., Conclusions: In addition to previously reported effects of MaMADS2 on ethylene, this transcription factor also influences other plant hormones, particularly at the pre-climacteric stage. The cytokinin pathway may play a previously unanticipated role via MaMADS2 in banana ripening. Finally, abscisic acid enhances MaMADS2 expression to promote ripening, but the transcription factor in turn auto inhibits ABA synthesis and signaling. Together, these results demonstrate a complex interaction of plant hormones and banana fruit ripening mediated by MaMADS2.