Contents
 	'Summary'	32	
I.	'Introduction'	32	
II.	'Effects of stress combination on growth, yield and physiological traits in plants and crops'	34	
III.	'The complexity of stress response signaling during stress combination'	38	
IV.	'Conclusions'	39	
 	'Acknowledgements'	41	
 	References	41	


Summary
Environmental stress conditions such as drought, heat, salinity, cold, or pathogen infection can have a devastating impact on plant growth and yield under field conditions. Nevertheless, the effects of these stresses on plants are typically being studied under controlled growth conditions in the laboratory. The field environment is very different from the controlled conditions used in laboratory studies, and often involves the simultaneous exposure of plants to more than one abiotic and/or biotic stress condition, such as a combination of drought and heat, drought and cold, salinity and heat, or any of the major abiotic stresses combined with pathogen infection. Recent studies have revealed that the response of plants to combinations of two or more stress conditions is unique and cannot be directly extrapolated from the response of plants to each of the different stresses applied individually. Moreover, the simultaneous occurrence of different stresses results in a high degree of complexity in plant responses, as the responses to the combined stresses are largely controlled by different, and sometimes opposing, signaling pathways that may interact and inhibit each other. In this review, we will provide an update on recent studies focusing on the response of plants to a combination of different stresses. In particular, we will address how different stress responses are integrated and how they impact plant growth and physiological traits.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.12797

Abiotic and biotic stress combinations

Under field conditions crops are routinely subjected to a number of different abiotic stress factors simultaneously. Recent studies revealed that the response of plants to a combination of different abiotic stresses is unique and cannot be directly extrapolated from simply studying each of the different stresses applied individually. These studies have also identified specific regulatory transcripts, combinations of metabolites and proteins, and physiological responses that are unique to specific stress combinations, highlighting the importance of studying abiotic stress combination in plants. Here we describe the interactions between drought and other abiotic stresses with emphasis on drought and heat stress. We compile new data about the different molecular, physiological and metabolic adaptations of different plants and crops to this stress combination and we highlight the importance of reactive oxygen species (ROS) metabolism and stomatal responses for plant acclimation to drought and heat stress combination. We further emphasize the need for developing crops with enhanced tolerance to drought and heat stress combination in order to mitigate the negative impacts of predicted global climatic changes on agricultural production worldwide.

Plant adaptations to the combination of drought and high temperatures

Global warming leads to the concurrence of a number of abiotic and biotic stresses, thus affecting agricultural productivity. Occurrence of abiotic stresses can alter plant-pest interactions by enhancing host plant susceptibility to pathogenic organisms, insects, and by reducing competitive ability with weeds. On the contrary, some pests may alter plant response to abiotic stress factors. Therefore, systematic studies are pivotal to understand the effect of concurrent abiotic and biotic stress conditions on crop productivity. However, to date, a collective database on the occurrence of various stress combinations in agriculturally-prominent areas is not available. This review attempts to assemble published information on this topic, with a particular focus on the impact of combined drought and pathogen stresses on crop productivity. In doing so, this review highlights some agriculturally important morpho-physiological traits that can be utilized to identify genotypes with combined stress tolerance. In addition, this review outlines potential role of recent genomic tools in deciphering combined stress tolerance in plants. This review will, therefore, be helpful for agronomists and field pathologists in assessing the impact of the interactions between drought and plant-pathogens on crop performance. Further, the review will be helpful for physiologists and molecular biologists to design agronomically relevant strategies for the development of broad spectrum stress tolerant crops.

/pdf/impact-of-combined-abiotic-and-biotic-stresses-on-plant-1loq95whio.pdf

Impact of Combined Abiotic and Biotic Stresses on Plant Growth and Avenues for Crop Improvement by Exploiting Physio-morphological Traits

Drought is a major environmental stress threatening wheat productivity worldwide. Global climate models predict changed precipitation patterns with frequent episodes of drought. Although drought impedes wheat performance at all growth stages, it is more critical during the flowering and grain-filling phases (terminal drought) and results in substantial yield losses. The severity and duration of the stress determine the extent of the yield loss. The principal reasons for these losses are reduced rates of net photosynthesis owing to metabolic limitations—oxidative damage to chloroplasts and stomatal closure—and poor grain set and development. A comprehensive understanding of the impact of terminal drought is critical for improving drought resistance in wheat, with marker-assisted selection being increasingly employed in breeding for this resistance. The limited success of molecular breeding and physiological strategies suggests a more holistic approach, including interaction of drought with other stresses a...

Drought Stress in Wheat during Flowering and Grain-filling Periods

In field conditions, plants are often simultaneously exposed to multiple biotic and abiotic stresses resulting in substantial yield loss. Plants have evolved various physiological and molecular adaptations to protect themselves under stress combinations. Emerging evidences suggest that plant responses to a combination of stresses are unique from individual stress responses. In addition, plants exhibit shared responses which are common to individual stresses and stress combination. In this review, we provide an update on the current understanding of both unique and shared responses. Specific focus of this review is on heat-drought stress as a major abiotic stress combination and, drought-pathogen and heat-pathogen as examples of abiotic-biotic stress combinations. We also comprehend the current understanding of molecular mechanisms of cross talk in relation to shared and unique molecular responses for plant survival under stress combinations. Thus, the knowledge of shared responses of plants from individual stress studies and stress combinations can be utilized to develop varieties with broad spectrum stress tolerance.

/pdf/shared-and-unique-responses-of-plants-to-multiple-individual-x6e06t304d.pdf

Shared and unique responses of plants to multiple individual stresses and stress combinations: physiological and molecular mechanisms.

High temperature and drought stress are among the two most important environmental factors influencing crop growth, development and yield processes. These two stresses commonly occur in combination. Objectives of this research were to investigate the independent and combined effects of high temperature and drought stress during grain filling on physiological, vegetative and yield traits and expression of a chloroplast protein synthesis elongation factor (EF-Tu) of wheat (Triticum aestivum L.). Two spring wheat cultivars (Pavon-76 and Seri-82) were grown at control temperatures (CT; day/night, 24/14 � C; 16/ 8 h photo/dark period) from sowing to heading. Thereafter, one half of the plants were exposed to high temperature stress (HT; 31/18 � C in Exp. 1 and 34/ 22 � C in Exp. 2), drought stress (withholding water), or a combination of both HT and drought stress. There were significant influences of HT and/or drought stress on physiological, growth and yield traits. There was no cultivar or cultivar by temperature or cultivar by drought interaction effects on most traits. The decreases in leaf photosynthesis were greater at HT compared with drought alone throughout the stress period, and the combination of HT and drought had the lowest leaf photosynthetic rates. Overall, HT or drought had similar effects (about 48‐56 % decrease) on spikelet fertility, grain numbers and grain yield. High temperature decreased grain numbers (by 56 % averaged across both experiments) and individual grain weight (by 25 %), while, respective decreases due to drought were 48 % and 35 %. This suggests that the grain numbers were more sensitive to HT and grain weights to drought for the range of temperatures tested in this research. The interaction between HT and drought stress was significant for total dry weights, harvest index and spikelet fertility, particularly when HT stress was severe (34/22 � C). The combined effects of HT and drought were greater than additive effects of HT or drought alone for leaf chlorophyll content, grain numbers and harvest index. High temperature stress and the combination of HT and drought stress but not drought stress alone resulted in the overexpression of EF-Tu in both spring wheat cultivars.

Independent and Combined Effects of High Temperature and Drought Stress During Grain Filling on Plant Yield and Chloroplast EF‐Tu Expression in Spring Wheat

S. R. Pisipati

Papers

Independent and Combined Effects of High Temperature and Drought Stress During Grain Filling on Plant Yield and Chloroplast EF‐Tu Expression in Spring Wheat