Molecular chaperones, including the heat-shock proteins (Hsps), are a ubiquitous feature of cells in which these proteins cope with stress-induced denaturation of other proteins. Hsps have received the most attention in model organisms undergoing experimental stress in the laboratory, and the function of Hsps at the molecular and cellular level is becoming well understood in this context. A complementary focus is now emerging on the Hsps of both model and nonmodel organisms undergoing stress in nature, on the roles of Hsps in the stress physiology of whole multicellular eukaryotes and the tissues and organs they comprise, and on the ecological and evolutionary correlates of variation in Hsps and the genes that encode them. This focus discloses that (a) expression of Hsps can occur in nature, (b) all species have hsp genes but they vary in the patterns of their expression, (c) Hsp expression can be correlated with resistance to stress, and (d) species' thresholds for Hsp expression are correlated with levels of stress that they naturally undergo. These conclusions are now well established and may require little additional confirmation; many significant questions remain unanswered concerning both the mechanisms of Hsp-mediated stress tolerance at the organismal level and the evolutionary mechanisms that have diversified the hsp genes.

/pdf/heat-shock-proteins-molecular-chaperones-and-the-stress-p3ghklf1z7.pdf

Heat-shock proteins, molecular chaperones, and the stress response: evolutionary and ecological physiology

Heat tolerance in plants: An overview

Stressful environments such as salinity, drought, and high temperature (heat) cause alterations in a wide range of physiological, biochemical, and molecular processes in plants. Photosynthesis, the most fundamental and intricate physiological process in all green plants, is also severely affected in all its phases by such stresses. Since the mechanism of photosynthesis involves various components, including photosynthetic pigments and photosystems, the electron transport system, and CO2 reduction pathways, any damage at any level caused by a stress may reduce the overall photosynthetic capacity of a green plant. Details of the stress-induced damage and adverse effects on different types of pigments, photosystems, components of electron transport system, alterations in the activities of enzymes involved in the mechanism of photosynthesis, and changes in various gas exchange characteristics, particularly of agricultural plants, are considered in this review. In addition, we discussed also progress made during the last two decades in producing transgenic lines of different C3 crops with enhanced photosynthetic performance, which was reached by either the overexpression of C3 enzymes or transcription factors or the incorporation of genes encoding C4 enzymes into C3 plants. We also discussed critically a current, worldwide effort to identify signaling components, such as transcription factors and protein kinases, particularly mitogen-activated protein kinases (MAPKs) involved in stress adaptation in agricultural plants.

Photosynthesis under stressful environments: An overview

Temperature is a primary factor affecting the rate of plant development. Warmer temperatures expected with climate change and the potential for more extreme temperature events will impact plant productivity. Pollination is one of the most sensitive phenological stages to temperature extremes across all species and during this developmental stage temperature extremes would greatly affect production. Few adaptation strategies are available to cope with temperature extremes at this developmental stage other than to select for plants which shed pollen during the cooler periods of the day or are indeterminate so flowering occurs over a longer period of the growing season. In controlled environment studies, warm temperatures increased the rate of phenological development; however, there was no effect on leaf area or vegetative biomass compared to normal temperatures. The major impact of warmer temperatures was during the reproductive stage of development and in all cases grain yield in maize was significantly reduced by as much as 80−90% from a normal temperature regime. Temperature effects are increased by water deficits and excess soil water demonstrating that understanding the interaction of temperature and water will be needed to develop more effective adaptation strategies to offset the impacts of greater temperature extreme events associated with a changing climate.

Temperature extremes: Effect on plant growth and development

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

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

Climate models predict greater increases in nighttime temperature in the future. The impacts of high nighttime temperature on wheat (Triticum aestivum L.) are not well understood. Objectives of this research were to quantify the impact of high nighttime temperatures during reproductive development on phenology, physiological, vegetative, and yield traits of wheat. Two spring wheat cultivars (Pavon-76 and Seri-82) were grown at optimum temperatures (day/night, 24/14°C; 16/8 h light/dark photoperiod) from sowing to booting. Thereafter, plants were exposed to four different nighttime temperatures (14, 17, 20, 23°C) until maturity. The daytime temperature was 24°C across all treatments. There were signifi cant infl uences of high nighttime temperatures on physiological, growth, and yield traits, but no cultivar or cultivar by temperature interactions were observed. High nighttime temperatures (>14°C) decreased photosynthesis after 14 d of stress. Grain yields linearly decreased with increasing nighttime temperatures, leading to lower harvest indices at 20 and 23°C. High nighttime temperature (≥20°C) decreased spikelet fertility, grains per spike, and grain size. Compared to the control (14°C), grain fi lling duration was decreased by 3 and 7 d at night temperatures of 20 and 23°C, respectively. High nighttime temperature increased the expression of chloroplast protein synthesis elongation factor in both cultivars suggesting possible involvement of this protein in plant response to stress.

Impact of Nighttime Temperature on Physiology and Growth of Spring Wheat

Determining mechanisms associated with heat tolerance and identifying screening methods are vital for improvement of heat tolerance in plants. The objectives of this study were to investigate the relationship between the heat stability of thylakoids and loss of chlorophyll in winter wheat (Triticum aestivum L.) under heat stress, and to examine whether chlorophyll loss can be used as an indicator of heat tolerance in wheat. We assessed heat tolerance and measured chlorophyll content in 12 cultivars of winter wheat at fl owering stage during exposure to 16-d-long heat stress. Heat tolerance was assessed using fl uorescence to determine the heat stability of thylakoids, and chlorophyll content was measured with a chlorophyll meter. Experiments were conducted under controlled conditions. Heat stress caused damage to thylakoids in all cultivars as indicated by the increase in the ratio of constant fl uorescence (O) and the peak of variable fl uorescence (P). Heat stress also caused a decline in chlorophyll content in most cultivars. A strong negative correlation between heat-induced increases in O/P and chlorophyll content was seen. The results suggest that heat-induced damage to thylakoids and chlorophyll loss are closely associated in winter wheat. Measurements of chlorophyll content with a chlorophyll meter could be useful for high throughput screening for heat tolerance in wheat.

Zoran Ristic

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

Impact of Nighttime Temperature on Physiology and Growth of Spring Wheat

Correlation between Heat Stability of Thylakoid Membranes and Loss of Chlorophyll in Winter Wheat under Heat Stress

Leaf cuticle and water loss in maize lines differing in dehydration avoidance

Heat-induced accumulation of chloroplast protein synthesis elongation factor, EF-Tu, in winter wheat