Abiotic stresses are one of the major constraints to crop production and food security worldwide. The situation has aggravated due to the drastic and rapid changes in global climate. Heat and drought stress are undoubtedly the two most important stresses having huge impact on growth and productivity of the crops. It is very important to understand the physiological, biochemical and ecological interventions related to these stresses for better management. A wide range of plant responses to these stresses could be generalized into morphological, physiological and biochemical responses. Interestingly, this review provides a detailed account of plant responses to heat and drought stresses with special focus on highlighting the commonalities and differences. Crop growth and yields are negatively affected by sub-optimal water supply and abnormal temperatures due to physical damages, physiological disruptions and biochemical changes. Both these stresses have multi-lateral impacts and therefore, complex in mechanistic action. A better understanding of plant responses to these stresses has pragmatic implication for remedies and management. A comprehensive account of conventional as well as modern approaches to deal with heat and drought stresses have also been presented here. A side-by-side critical discussion on salient responses and management strategies for these two important abiotic stresses provides a unique insight into the phenomena. A holistic approach taking into account the different management options to deal with heat and drought stress simultaneously could be a win-win approach in future.

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Crop Production under Drought and Heat Stress: Plant Responses and Management Options

Plant biostimulants are diverse substances and microorganisms used to enhance plant growth. The global market for biostimulants is projected to increase 12 % per year and reach over $2,200 million by 2018. Despite the growing use of biostimulants in agriculture, many in the scientific community consider biostimulants to be lacking peer-reviewed scientific evaluation. This article describes the emerging definitions of biostimulants and reviews the literature on five categories of biostimulants: i. microbial inoculants, ii. humic acids, iii. fulvic acids, iv. protein hydrolysates and amino acids, and v. seaweed extracts. The large number of publications cited for each category of biostimulants demonstrates that there is growing scientific evidence supporting the use of biostimulants as agricultural inputs on diverse plant species. The cited literature also reveals some commonalities in plant responses to different biostimulants, such as increased root growth, enhanced nutrient uptake, and stress tolerance.

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Agricultural uses of plant biostimulants

Plants in nature are continuously exposed to several biotic and abiotic stresses. Among these stresses, drought stress is one of the most adverse factors of plant growth and productivity and considered a severe threat for sustainable crop production in the conditions on changing climate. Drought triggers a wide variety of plant responses, ranging from cellular metabolism to changes in growth rates and crop yields. Understanding the biochemical and molecular responses to drought is essential for a holistic perception of plant resistance mechanisms to water-limited conditions. This review describes some aspects of drought induced changes in morphological, physiological and biochemical changes in plants. Drought stress progressively decreases CO2 assimilation rates due to reduced stomatal conductance. It reduces leaf size, stems extension and root proliferation, disturbs plant water relations and reduces water-use efficiency. It disrupts photosynthetic pigments and reduces the gas exchange leading to a reduction in plant growth and productivity. The critical roles of osmolyte accumulation under drought stress conditions have been actively researched to understand the tolerance of plants to dehydration. In addition, drought stress-induced generation of active oxygen species is well recognized at the cellular level and is tightly controlled at both the production and consumption levels, through increased antioxidative systems. This review focuses on the ability and strategies of higher plants to respond and adapt to drought stress.

	 

	Key words: Drought stress, growth, yield, gas exchange, photosynthetic pigments, antioxidative system.

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Morphological, physiological and biochemical responses of plants to drought stress

Brassinosteroids (BRs) are a group of polyhydroxylated plant steroid hormones crucial for many aspects of plant life. BRs were originally characterized for their function in cell elongation, but it is becoming clear that they play major roles in plant growth, development and responses to several stresses such as temperature and drought. A BR signaling pathway from cell surface receptors to central transcription factors has been well characterized. Here we summarize recent progress towards understanding the BR pathway including BR perception and the molecular mechanisms of BR signaling. Next, we discuss the roles of BRs in development and stress responses. Finally, we show how knowledge of the BR pathway is being applied to manipulate the growth and stress responses in crops. These studies highlight the complex regulation of BR signaling, multiple points of crosstalk between BRs and other hormones or stress responses, and finely tuned spatiotemporal regulation of BR signaling.

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Brassinosteroids: Multidimensional Regulators of Plant Growth, Development, and Stress Responses.

Use of plant growth promoting rhizobacteria (PGPRs) with multiple plant growth promoting traits in stress agriculture: Action mechanisms and future prospects

Brassinolides (BRs) are naturally occurring substances, which modulate plant growth and development events and have been known to improve the crop tolerance to abiotic stresses. In this study, possible role of exogenously applied brassinolide (BR) in alleviating the detrimental effects of drought in maize was evaluated in a rain-protected wire-house. Maize was subjected to drought at the start of tasseling for 6 days by withholding water application followed by foliar spray of BR (0.1 mg l−1) to assess the changes in growth, gas exchange, chlorophyll contents, protein, relative leaf water contents (RLWC), proline, malonialdehyde (MDA) and enzymatic antioxidants. Drought substantially reduced the maize growth in terms of plant height, leaf area and plant biomass. Moreover, substantial decrease in gas exchange attributes (net photosynthetic rate (A), transpiration rate (E), stomatal conductance (gs), water use efficiency (WUE), instantaneous water use efficiency (WUEi) and intercellular CO2 (Ci) was also recorded. However, exogenous application of BR remarkably improved the gas exchange attributes, plant height, leaf area, cobs per plant, seedling dry weight both under drought and well-watered conditions. BR-induced promotion in growth and physiological and metabolic activities were mediated through increased protein synthesis enabling maintenance of tissue water potential and activities of antioxidant enzymes lowering the lipid peroxidation under drought.

Brassinolide Application Improves the Drought Tolerance in Maize Through Modulation of Enzymatic Antioxidants and Leaf Gas Exchange

Methyl jasmonate (MeJA), a plant-signalling molecule, is involved in an array of plant development and the defence responses. This study was conducted to explore the role of exogenous MeJA application in alleviating the adversities of drought stress in soybean (Glycine max L. Merrill.). Soybean plants were grown under normal conditions until blooming and were then subjected to drought by withholding irrigation followed by foliar application of (50 lm) MeJA. Drought stress substantially suppressed the yield and yield-related traits, whereas it accelerated the membrane lipid peroxidation. Nonetheless, substantial increase in activities of enzymatic antioxidants (superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT)), proline, relative water contents (RWC) with simultaneous decrease in membrane lipid peroxidation was observed in MeJA-treated plants under drought. These beneficial effects led to improvement in biological and grain yield, and harvest index under drought. Interestingly, MeJA application was also useful under well-watered conditions. These results suggest the involvement of MeJA in improving the drought tolerance of soybean by modulating the membrane lipid peroxidation and antioxidant activities.

Methyl Jasmonate‐Induced Alteration in Lipid Peroxidation, Antioxidative Defence System and Yield in Soybean Under Drought

Water deficit is perhaps the most severe threat to sustainable crop production in the conditions of changing climate. Researchers are striving hard to develop resistance against water deficit in crop plants to ensure food security for the coming generations. This study was conducted to establish the role of fulvic acid (FA) application in improving the performance of hybrid maize (Zea mays L.) under drought. Maize plants were grown under normal conditions till tasselling and were then subjected to drought by cessation of water followed by foliar application of FA (1.5âmgâlâ1). Drought stress disrupted the photosynthetic pigments and reduced the gas exchange leading to reduction in plant growth and productivity. Nonetheless, exogenous FA application substantially ameliorated the adversities of drought by sustaining the chlorophyll contents and gas exchange possibly by enhanced levels of antioxidant enzyme (superoxide dismutase (SOD), peroxidase (POD), and catalase (CAT)) activities and proline. These beneficial effects yielded in terms of plant growth and allometry, and grain yield. It is interesting to note that FA application also improved the crop performance under wellâwatered conditions. Hence, FA may be applied to improve the crop performance under drought and wellâwatered conditions.

Fulvic Acid Application Improves the Maize Performance under Well‐watered and Drought Conditions

Glycinebetaine acts as an osmoprotectant and is closely related with drought resistance. In the present study, glycinebetaine (GB) was exogenously-applied to two contrasting maize cultivars, Dongdan-60 and ND-95, to see whether GB improves drought resistance. Maize cultivars were grown with normal water supply till the heading stage and then exposed to two levels of soil moisture, well-watered control and drought-stressed, and then GB solution of 100 mmol was foliar applied five days after moisture treatments were imposed. The gas exchange and chlorophyll concentration were substantially declined in both maize cultivars under water stressed conditions. However, this reduction was less in Dongdan-60 than ND-95. Nonetheless, GB-treated plants considerably maintained higher gas exchange rate and chlorophyll concentration during drought stress than non-GB treated plants. The GB-induced improvement in gas exchange and chlorophyll synthesis under water stress ultimately resulted in improved growth and yield in both maize cultivars. Furthermore, the positive responses to exogenous GB application were more pronounced in Dongdan-60 as compared to ND-95 in all traits examined under water-deficit con ditions. In conclusion, exogenously applied GB to maize crops could improve gas exchange, chlorophyll synthesis, growth and yield of maize.

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L. L. Xue

Papers

Brassinolide Application Improves the Drought Tolerance in Maize Through Modulation of Enzymatic Antioxidants and Leaf Gas Exchange

Methyl Jasmonate‐Induced Alteration in Lipid Peroxidation, Antioxidative Defence System and Yield in Soybean Under Drought

Fulvic Acid Application Improves the Maize Performance under Well‐watered and Drought Conditions

Gas exchange and chlorophyll synthesis of maize cultivars are enhanced by exogenously-applied glycinebetaine under drought conditions.