Heat Stress Tolerance In Plants Research Articles

Melatonin Reverses High-Temperature-Stress-Inhibited Photosynthesis in the Presence of Excess Sulfur by Modulating Ethylene Sensitivity in Mustard.

Melatonin is a pleiotropic, nontoxic, regulatory biomolecule with various functions in abiotic stress tolerance. It reverses the adverse effect of heat stress on photosynthesis in plants and helps with sulfur (S) assimilation. Our research objective aimed to find the influence of melatonin, along with excess sulfur (2 mM SO42-), in reversing heat stress's impacts on the photosynthetic ability of the mustard (Brassica juncea L.) cultivar SS2, a cultivar with low ATP-sulfurylase activity and a low sulfate transport index (STI). Further, we aimed to substantiate that the effect was a result of ethylene modulation. Melatonin in the presence of excess-S (S) increased S-assimilation and the STI by increasing the ATP-sulfurylase (ATP-S) and serine acetyltransferase (SAT) activity of SS2, and it enhanced the content of cysteine (Cys) and methionine (Met). Under heat stress, melatonin increased S-assimilation and diverted Cys towards the synthesis of more reduced glutathione (GSH), utilizing excess-S at the expense of less methionine and ethylene and resulting in plants' reduced sensitivity to stress ethylene. The treatment with melatonin plus excess-S increased antioxidant enzyme activity, photosynthetic-S use efficiency (p-SUE), Rubisco activity, photosynthesis, and growth under heat stress. Further, plants receiving melatonin and excess-S in the presence of norbornadiene (NBD; an ethylene action inhibitor) under heat stress showed an inhibited STI and lower photosynthesis and growth. This suggested that ethylene was involved in the melatonin-mediated heat stress reversal effects on photosynthesis in plants. The interaction mechanism between melatonin and ethylene is still elusive. This study provides avenues to explore the melatonin-ethylene-S interaction for heat stress tolerance in plants.

Exogenous Methyl Jasmonate Mediated MiRNA-mRNA Network Improves Heat Tolerance of Perennial Ryegrass

Heat stress can hinder the growth of perennial ryegrass (Lolium perenne L.). Methyl jasmonate (MeJA) applied exogenously can increase heat stress tolerance in plants; however, the regulatory mechanisms involved in heat tolerance mediated by MeJA are poorly understood in perennial ryegrass. Here, the microRNA (miRNA) expression profiles of perennial ryegrass were assessed to elucidate the regulatory pathways associated with heat tolerance induced by MeJA. Plants were subjected to four treatments, namely, control (CK), MeJA pre-treatment (T), heat stress treatment (H), and MeJA pre-treatment + heat stress (TH). According to the results, 102 miRNAs were up-regulated in all treatments, with 20, 27, and 33 miRNAs being up-regulated in the T, H, and TH treatment groups, respectively. The co-expression network analysis between the deferentially expressed miRNAs and their corresponding target genes showed that 20 miRNAs modulated 51 potential target genes. Notably, the miRNAs that targeted genes related to with regards to heat tolerance were driven by MeJA, and they were involved in four pathways: novel-m0258-5p mediated signal transduction, novel-m0350-5p mediated protein homeostasis, miR397-z, miR5658-z, and novel-m0008-5p involved in cell wall component, and miR1144-z and miR5185-z dominated chlorophyll degradation. Overall, the findings of this research paved the way for more research into the heat tolerance mechanism in perennial ryegrass and provided a theoretical foundation for developing cultivars with enhanced heat tolerance.

Insights into morphological and physio-biochemical adaptive responses in mungbean (Vigna radiata L.) under heat stress.

Mungbean (Vigna radiata L. Wilczek) is an important food legume crop which contributes significantly to nutritional and food security of South and Southeast Asia. The crop thrives in hot and humid weather conditions, with an optimal temperature range of 28°-35°C, and is mainly cultivated under rainfed environments. However, the rising global temperature has posed a serious threat to mungbean cultivation. Optimal temperature is a vital factor in cellular processes, and every crop species has evolved with its specific temperature tolerance ability. Moreover, variation within a crop species is inevitable, given the diverse environmental conditions under which it has evolved. For instance, various mungbean germplasm can grow and produce seeds in extreme ambient temperatures as low as 20°C or as high as 45°C. This range of variation in mungbean germplasm for heat tolerance plays a crucial role in developing heat tolerant and high yielding mungbean cultivars. However, heat tolerance is a complex mechanism which is extensively discussed in this manuscript; and at the same time individual genotypes have evolved with various ways of heat stress tolerance. Therefore, to enhance understanding towards such variability in mungbean germplasm, we studied morphological, anatomical, physiological, and biochemical traits which are responsive to heat stress in plants with more relevance to mungbean. Understanding heat stress tolerance attributing traits will help in identification of corresponding regulatory networks and associated genes, which will further help in devising suitable strategies to enhance heat tolerance in mungbean. The major pathways responsible for heat stress tolerance in plants are also discussed.

Thermopriming mitigates the effects of heat stress by modulating the expression of Heat shock factors in Brassica juncea (Indian mustard)

Brassica juncea (also known as Indian mustard) is an important vegetable oil seed crop whose growth and productivity is severely affected by heat stress. Interestingly, pre-exposure to sublethal heat stress, referred as thermopriming can remarkably enhance the plant heat stress tolerance. However, little is known about the impact of thermopriming on heat stress response of the important oil seed crop, B. juncea. In this study, we investigated the basal and thermopriming-induced heat stress response of 16 different cultivated, agronomically important varieties of Brassica juncea. Based on their basal heat stress response, we classify the varieties as heat sensitive, moderately tolerant or tolerant. Notably, almost all the varieties displayed enhanced heat stress tolerance upon thermopriming (acquired thermotolerance), albeit to varying magnitudes. Strikingly, the high oil-yielding, drought tolerant, heat sensitive variety Pusa Bold showed remarkable acquired thermotolerance upon thermopriming. Investigations in Pusa Bold indicate that the thermopriming-induced acquired thermotolerance predominantly correlates with the rapid activation of ROS scavenging mechanisms and differential expression of Heat Shock Family of transcription factors (BjHSFs). Taken together, our study reveals the positive impact of thermopriming in alleviating the harmful effects of heat stress on Brassica juncea seedlings and paves the path to enhance heat stress tolerance for improved productivity in the important oilseed crop Indian mustard.

Expression of ZmNAGK in tobacco enhances heat stress tolerance via activation of antioxidant-associated defense

Heat stress severely inhibits plant growth and limits crop yields. Thus, it is crucial to identify genes that are associated with plant heat stress responses. Here, we report a maize (Zea mays L.) gene, N-acetylglutamate kinase (ZmNAGK), that positively enhances plant heat stress tolerance. The ZmNAGK expression level was significantly up-regulated by heat stress in maize plants, and ZmNAGK was found to be localized in maize chloroplasts. Phenotypic analysis showed that overexpressing of ZmNAGK enhanced the tolerance of tobacco to heat stress both in the seed germination and seedling growth stages. Further physiological analysis showed that ZmNAGK overexpression in tobacco could alleviate oxidative damages that occurred during heat stress via activation of antioxidant defense signaling. Transcriptome analysis revealed that ZmNAGK could modulate the expression of antioxidant-enzyme encoding genes, such as ascorbate peroxidase 2 (APX2) and superoxide dismutase C (SODC), and heat shock network genes. Taken together, we have identified a maize gene that can provide plants with heat tolerance through the induction of antioxidant-associated defense signaling.

Role of Salicylic Acid in Combating Heat Stress in Plants: Insights into Modulation of Vital Processes.

In the present era of climate change and global warming, high temperatures have increased considerably, posing a threat to plant life. Heat stress affects the biochemistry, physiology and molecular makeup of the plant by altering the key processes, i.e., photosynthesis, respiration and reproduction which reduces its growth and development. There is a dire need to manage this problem sustainably for plant conservation as well as the food security of the human population. Use of phytohormones to induce thermotolerance in plants can be a sustainable way to fight the adversities of heat stress. Phytohormone-induced thermotolerance proves to be a compelling approach to sustainably relieve the damaging effects of heat stress on plants. Salicylic acid (SA) is an essential molecule in biotic and abiotic defense response signal transduction pathways. When supplied externally, it imparts heat stress tolerance to the plants by different means, viz., increased Heat Shock Proteins (HSP) production, Reactive oxygen species (ROS) scavenging, protection of the reproductive system and enhancing photosynthetic efficiency. The effect of SA on plants is highly dependent on the concentration applied, plant species, plant age, type of tissues treated, and duration of the treatment. The present review paper summarizes the mechanism of thermotolerance induced by salicylic acid in plants under heat stress conditions. It includes the regulatory effects of SA on heat shock proteins, antioxidant metabolism, and maintenance of Ca2+ homeostasis under heat stress. This review combines the studies conducted to elucidate the role of SA in the modulation of different mechanisms which lead to heat stress tolerance in plants. It discusses the mechanism of SA in protecting the photosynthetic machinery and reproductive system during high-temperature stress.

ZmNAC074, a maize stress-responsive NAC transcription factor, confers heat stress tolerance in transgenic Arabidopsis.

The harsh environment such as high temperature greatly limits the growth, development and production of crops worldwide. NAC (NAM, ATAF1/2, and CUC2) transcription factors (TFs) play key regulatory roles in abiotic stress responses of plants. However, the functional roles of NAC TFs in heat stress response of maize remain elusive. In our present study, we identified and isolated a stress-responsive NAC transcription factor gene in maize, designated as ZmNAC074 and orthologous with rice OsNTL3. Further studies revealed that ZmNAC074 may encode a membrane-bound transcription factor (MTF) of NAC family in maize, which is comprised of 517 amino acid residues with a transmembrane domain at the C-terminus. Moreover, ZmNAC074 was highly expressed and induced by various abiotic stresses in maize seedlings, especially in leaf tissues under heat stress. Through generating ZmNAC074 transgenic plants, phenotypic and physiological analyses further displayed that overexpression of ZmNAC074 in transgenic Arabidopsis confers enhanced heat stress tolerance significantly through modulating the accumulation of a variety of stress metabolites, including reactive oxygen species (ROS), antioxidants, malondialdehyde (MDA), proline, soluble protein, chlorophyll and carotenoid. Further, quantitative real-time PCR analysis showed that the expression levels of most ROS scavenging and HSR- and UPR-associated genes in transgenic Arabidopsis were significantly up-regulated under heat stress treatments, suggesting that ZmNAC074 may encode a positive regulator that activates the expression of ROS-scavenging genes and HSR- and UPR-associated genes to enhance plant thermotolerance under heat stress conditions. Overall, our present study suggests that ZmNAC074 may play a crucial role in conferring heat stress tolerance in plants, providing a key candidate regulatory gene for heat stress tolerance regulation and genetic improvement in maize as well as in other crops.

A Review on the Role of Endophytes and Plant Growth Promoting Rhizobacteria in Mitigating Heat Stress in Plants.

Among abiotic stresses, heat stress is described as one of the major limiting factors of crop growth worldwide, as high temperatures elicit a series of physiological, molecular, and biochemical cascade events that ultimately result in reduced crop yield. There is growing interest among researchers in the use of beneficial microorganisms. Intricate and highly complex interactions between plants and microbes result in the alleviation of heat stress. Plant–microbe interactions are mediated by the production of phytohormones, siderophores, gene expression, osmolytes, and volatile compounds in plants. Their interaction improves antioxidant activity and accumulation of compatible osmolytes such as proline, glycine betaine, soluble sugar, and trehalose, and enriches the nutrient status of stressed plants. Therefore, this review aims to discuss the heat response of plants and to understand the mechanisms of microbe-mediated stress alleviation on a physio-molecular basis. This review indicates that microbes have a great potential to enhance the protection of plants from heat stress and enhance plant growth and yield. Owing to the metabolic diversity of microorganisms, they can be useful in mitigating heat stress in crop plants. In this regard, microorganisms do not present new threats to ecological systems. Overall, it is expected that continued research on microbe-mediated heat stress tolerance in plants will enable this technology to be used as an ecofriendly tool for sustainable agronomy.

Reproductive-Stage Heat Stress in Cereals: Impact, Plant Responses and Strategies for Tolerance Improvement.

Reproductive-stage heat stress (RSHS) poses a major constraint to cereal crop production by damaging main plant reproductive structures and hampering reproductive processes, including pollen and stigma viability, pollination, fertilization, grain setting and grain filling. Despite this well-recognized fact, research on crop heat stress (HS) is relatively recent compared to other abiotic stresses, such as drought and salinity, and in particular, RSHS studies in cereals are considerably few in comparison with seedling-stage and vegetative-stage-centered studies. Meanwhile, climate change-exacerbated HS, independently or synergistically with drought, will have huge implications on crop performance and future global food security. Fortunately, due to their sedentary nature, crop plants have evolved complex and diverse transient and long-term mechanisms to perceive, transduce, respond and adapt to HS at the molecular, cell, physiological and whole plant levels. Therefore, uncovering the molecular and physiological mechanisms governing plant response and tolerance to RSHS facilitates the designing of effective strategies to improve HS tolerance in cereal crops. In this review, we update our understanding of several aspects of RSHS in cereals, particularly impacts on physiological processes and yield; HS signal perception and transduction; and transcriptional regulation by heat shock factors and heat stress-responsive genes. We also discuss the epigenetic, post-translational modification and HS memory mechanisms modulating plant HS tolerance. Moreover, we offer a critical set of strategies (encompassing genomics and plant breeding, transgenesis, omics and agronomy) that could accelerate the development of RSHS-resilient cereal crop cultivars. We underline that a judicious combination of all of these strategies offers the best foot forward in RSHS tolerance improvement in cereals. Further, we highlight critical shortcomings to RSHS tolerance investigations in cereals and propositions for their circumvention, as well as some knowledge gaps, which should guide future research priorities. Overall, our review furthers our understanding of HS tolerance in plants and supports the rational designing of RSHS-tolerant cereal crop cultivars for the warming climate.

What Did We Learn From Current Progress in Heat Stress Tolerance in Plants? Can Microbes Be a Solution?

Temperature is a significant parameter in agriculture since it controls seed germination and plant growth. Global warming has resulted in an irregular rise in temperature posing a serious threat to the agricultural production around the world. A slight increase in temperature acts as stress and exert an overall negative impact on different developmental stages including plant phenology, development, cellular activities, gene expression, anatomical features, the functional and structural orientation of leaves, twigs, roots, and shoots. These impacts ultimately decrease the biomass, affect reproductive process, decrease flowering and fruiting and significant yield losses. Plants have inherent mechanisms to cope with different stressors including heat which may vary depending upon the type of plant species, duration and degree of the heat stress. Plants initially adapt avoidance and then tolerance strategies to combat heat stress. The tolerance pathway involves ion transporter, osmoprotectants, antioxidants, heat shock protein which help the plants to survive under heat stress. To develop heat-tolerant plants using above-mentioned strategies requires a lot of time, expertise, and resources. On contrary, plant growth-promoting rhizobacteria (PGPRs) is a cost-effective, time-saving, and user-friendly approach to support and enhance agricultural production under a range of environmental conditions including stresses. PGPR produce and regulate various phytohormones, enzymes, and metabolites that help plant to maintain growth under heat stress. They form biofilm, decrease abscisic acid, stimulate root development, enhance heat shock proteins, deamination of ACC enzyme, and nutrient availability especially nitrogen and phosphorous. Despite extensive work done on plant heat stress tolerance in general, very few comprehensive reviews are available on the subject especially the role of microbes for plant heat tolerance. This article reviews the current studies on the retaliation, adaptation, and tolerance to heat stress at the cellular, organellar, and whole plant levels, explains different approaches, and sheds light on how microbes can help to induce heat stress tolerance in plants.

The Functional Interplay between Ethylene, Hydrogen Sulfide, and Sulfur in Plant Heat Stress Tolerance.

Plants encounter several abiotic stresses, among which heat stress is gaining paramount attention because of the changing climatic conditions. Severe heat stress conspicuously reduces crop productivity through changes in metabolic processes and in growth and development. Ethylene and hydrogen sulfide (H2S) are signaling molecules involved in defense against heat stress through modulation of biomolecule synthesis, the antioxidant system, and post-translational modifications. Other compounds containing the essential mineral nutrient sulfur (S) also play pivotal roles in these defense mechanisms. As biosynthesis of ethylene and H2S is connected to the S-assimilation pathway, it is logical to consider the existence of a functional interplay between ethylene, H2S, and S in relation to heat stress tolerance. The present review focuses on the crosstalk between ethylene, H2S, and S to highlight their joint involvement in heat stress tolerance.

Melatonin: A Vital Pro-Tectant for Crops against Heat Stress: Mechanisms and Prospects

Heat stress (HS) is a serious environmental stress that negatively affects crop growth and productivity across the globe. The recent increase in atmospheric temperature caused by global warming has increased its intensity, which is a serious challenge that needs to be addressed. Plant growth and development involves a series of physiological, metabolic, and biochemical processes that are negatively affected by heat-induced oxidative stress, disorganization of cellular membranes and disturbed plant water relations, nutrient uptake, photosynthetic efficiency, and antioxidant activities. Plant tolerance to abiotic stresses can be substantially increased by the application of bio-stimulants, without posing a threat to the ecosystem. Melatonin (MT) is a multi-functional signaling molecule that has the potential to protect plants from the adverse impacts of HS. MT protects the cellular membranes, maintains the leaf water content, and improves the water use efficiency (WUE) and nutrient homeostasis; thereby, improving plant growth and development under HS. Moreover, MT also improves gene expression, crosstalk of hormones, and osmolytes, and reduces the accumulation of reactive oxygen species (ROS) by triggering the antioxidant defense system, which provides better resistance to HS. High endogenous MT increases genes expression and antioxidant activities to confer HS tolerance. Thus, it is important to understand the detailed mechanisms of both exogenous and endogenous MT, to induce HS tolerance in plants. This review highlights the versatile functions of MT in various plant responses, to improve HS tolerance. Moreover, we also discussed the MT crosstalk with other hormones, antioxidant potential of MT, and success stories of engineering MT to improve HS tolerance in plants. Additionally, we also identified various research gaps that need to be filled in future research using this important signaling molecule. Thus, this review will help the readers to learn more about MT under changing climatic conditions and will provide knowledge to develop heat tolerance in crops.

Impact of heat stress of varying severity on papaya (Carica papaya) leaves: Major changes in stress volatile signatures, but surprisingly small enhancements of total emissions

Tropical plants are considered to be highly heat stress resistant, but heat waves of different frequency and duration induced by global climate change can exceed the plant heat stress tolerance in the tropics. To gain insight into the overall heat resistance, leaves of papaya were exposed to mild (37 and 41 °C), moderate (46 °C) and severe (49 °C) heat stress (applied for 5 min), and plant recovery was monitored for 48 h upon return to pre-stress conditions. Leaf photosynthetic characteristics were reduced immediately after stress in all cases, but photosynthesis rate almost fully recovered after mild heat treatment, and partly recovered in 48 h after moderate stress, while no recovery was observed under severe stress. Constitutive BVOC emissions dominated by long-chained aldehydes, and glucosinolate breakdown products, attained the emission maxima at 46 °C. Acute heat stress led only to a minor induction of BVOC emissions, and the greatest share of induced emissions was observed in leaves exposed to 46 °C. Induced emissions were dominated by oxygenated monoterpenes, indicating an increase in leaf oxidative status. Our results demonstrate that papaya could withstand moderately severe heat stress of up to 46 °C, but this level of stress will have negative impacts on plant carbon gain due to delayed photosynthetic recovery after heat stress. Although volatile profiles sensitively responded to heat stress, the magnitude and degree of elicitation were surprisingly small throughout the recovery, suggesting that papaya is relatively heat resistant and possess a suite of heat-protective traits. • Papaya photosynthetic recovery was rapid under mild heat stress, delayed under moderate stress and absent under severe stress. • Heat stress of different severity had only moderate effects on volatile emissions, but the composition of emitted volatiles was strongly altered. • Volatile emission blend rather than the absolute amount characterizes the impact of heat stress on leaves of papaya.

Analyzing the regulatory role of heat shock transcription factors in plant heat stress tolerance: a brief appraisal.

An increase in ambient temperature throughout the twenty-first century has been described as a "worldwide threat" for crop production. Due to their sessile lifestyles, plants have evolved highly sophisticated and complex heat stress response (HSR) mechanisms to respond to higher temperatures. The HSR allows plants to minimize the damages caused by heat stress (HS), thus enabling cellular protection. HSR is crucial for their lifecycle and yield, particularly for plants grown in the field. At the cellular level, HSR involves the production of heat shock proteins (HSPs) and other stress-responsive proteins to counter the negative effects of HS. The expression of most HSPs is transcriptionally regulated by heat shock transcription factors (HSFs). HSFs are a group of evolutionary conserved regulatory proteins present in all eukaryotes and regulate various stress responses and biological processes in plants. In recent years, significant progress has been made in deciphering the complex regulatory network of HSFs, and several HSFs not only from model plants but also from major crops have been functionally characterized. Therefore, this review explores the progress made in this fascinating research area and debates the further potential to breed thermotolerant crop cultivars through the modulation of HSF networks. Furthermore, we discussed the role of HSFs in plant HS tolerance in a class-specific manner and shed light on their functional diversity, which is evident from their mode of action. Additionally, some research gaps have been highlighted concerning class-specific manners.

De novo analysis reveals transcriptomic responses to heat stress in loquat leaves

AbstractTemperature has a vital impact on plant growth and development, and heat stress severely threatens agricultural productivity and food security. High temperatures can cause annual yield reductions and disease in loquat [Eriobotrya japonica (Thunb.) Lindl.]. However, the molecular mechanisms underlying the heat stress response in loquat are not known. In this study, the transcriptome of loquat leaves in response to heat stress was sequenced and assembled de novo, and differentially expressed genes (DEGs) were identified. A total of 16,120 and 11,668 DEGs were identified at 1 and 2 h posttreatment, respectively. Gene ontology (GO) analysis indicated that the DEGs were located in the membrane and endoplasmic reticulum and were related to signal transduction and protein modification. Kyoto Encyclopedia of Genes and Genomes pathway analysis revealed that the heat‐induced DEGs were mainly related to the activation of protein processing in the endoplasmic reticulum and transcription. Ninety‐six heat shock proteins (HSPs) were also found in the protein processing pathway in the endoplasmic reticulum, and most HSPs were induced by heat stress. Heat shock transcription factors regulating the expression of downstream HSPs were induced by heat together with HSPs. This research provides insights into the mechanism of loquat plant heat stress tolerance and paves the way for heat‐tolerant loquat breeding.

Unfolding molecular switches in plant heat stress resistance: Acomprehensive review.

Plant heat stress response is a multi-factorial trait that is precisely regulated by the complex web of transcription factors from various families that modulate heat stress responsive gene expression. Global warming due to climate change affects plant growth and development throughout its life cycle. Adds to this, the frequent occurrence of heat waves is drastically reducing the global crop yield. Molecular plant scientists can help crop breeders by providing genetic markers associated with stress resistance. Plant heat stress response (HSR), however, is a multi-factorial trait and using a single stress resistance trait might not be ideal to develop thermotolerant crops. Transcription factors participate in regulation of plant biological processes and environmental stress responses. Recent studies have revealed that plant HSR is precisely regulated by the complex web of transcription factors from various families. These transcription factors enhance plant heat stress tolerance by regulating the expression level of several stress-responsive genes independently or in cross talk with different other transcription factors. This review explores how signaling pathways triggered by heat stress are regulated by multiple transcription factor families. To our knowledge, we for the first time analyze the role of major transcription factor families in plant HSR along with their regulatory mechanisms. In the end, we will also discuss the potential of emerging technologies to improve thermotolerance in plants.

Mitochondrial ATP synthase subunit d, a component of the peripheral stalk, is essential for growth and heat stress tolerance in Arabidopsis thaliana.

As rapid changes in climate threaten global crop yields, an understanding of plant heat stress tolerance is increasingly relevant. Heat stress tolerance involves the coordinated action of many cellular processes and is particularly energy demanding. We acquired a knockout mutant and generated knockdown lines in Arabidopsis thaliana of the d subunit of mitochondrial ATP synthase (gene name: ATPQ, AT3G52300, referred to hereafter as ATPd), a subunit of the peripheral stalk, and used these to investigate the phenotypic significance of this subunit in normal growth and heat stress tolerance. Homozygous knockout mutants for ATPd could not be obtained due to gametophytic defects, while heterozygotes possess no visible phenotype. Therefore, we used RNA interference to create knockdown plant lines for further studies. Proteomic analysis and blue native gels revealed that ATPd downregulation impairs only subunits of the mitochondrial ATP synthase (complex V). Knockdown plants were more sensitive to heat stress, had abnormal leaf morphology, and were severely slow growing compared to wild type. These results indicate that ATPd plays a crucial role in proper function of the mitochondrial ATP synthase holoenzyme, which, when reduced, leads to wide-ranging defects in energy-demanding cellular processes. In knockdown plants, more hydrogen peroxide accumulated and mitochondrial dysfunction stimulon (MDS) genes were activated. These data establish the essential structural role of ATPd and support the importance of complex V in normal plant growth, and provide new information about its requirement for heat stress tolerance.

Role of Phytohormones in Regulating Heat Stress Acclimation in Agricultural Crops

Heat stress (HS) seriously affects crop growth, causing significant crop yield losses worldwide. The regulatory mechanisms controlling HS tolerance in plants are not well understood. Phytohormones are important molecules for coordinating myriad of phenomena related to plant growth and development. They are also essential endogenous signaling molecules that actively mediate numerous physiological responses under abiotic stress by triggering stress-responsive regulatory genes involved in plant growth. This review updates the central role of various phytohormones—indole acetic acid, gibberellic acid, abscisic acid, cytokinins, ethylene, salicylic acid, brassinosteroids, strigolactone, and jasmonic acid—in regulating the HS response so that plants can adapt to increasing temperature stress. We also reveal how these stress-responsive phytohormones switch on various regulatory gene(s) and genes encoding antioxidants and heat shock proteins (HSPs) to combat HS in various plant species.

Plant heat stress: Concepts directing future research.

Predicted increases in future global temperatures require us to better understand the dimensions of heat stress experienced by plants. Here we highlight four key areas for improving our approach towards understanding plant heat stress responses. First, although the term 'heat stress' is broadly used, that term encompasses heat shock, heat wave and warming experiments, which vary in the duration and magnitude of temperature increase imposed. A greater integration of results and tools across these approaches is needed to better understand how heat stress associated with global warming will affect plants. Secondly, there is a growing need to associate plant responses to tissue temperatures. We review how plant energy budgets determine tissue temperature and discuss the implications of using leaf versus air temperature for heat stress studies. Third, we need to better understand how heat stress affects reproduction, particularly understudied stages such as floral meristem initiation and development. Fourth, we emphasise the need to integrate heat stress recovery into breeding programs to complement recent progress in improving plant heat stress tolerance. Taken together, we provide insights into key research gaps in plant heat stress and provide suggestions on addressing these gaps to enhance heat stress resilience in plants.

Genome-wide association study of multiple traits linked to heat tolerance in emmer-derived hexaploid wheat genotypes.

The online version contains supplementary material available at 10.1007/s11032-021-01222-3.