Drought and heat stress are among the two most important environmental factors influencing crop growth, development, and yield processes. A comprehensive understanding of the impact of drought and heat stress will be critical in evaluating the impact of climate change and climate variabil ity on crop production. Both drought and heat stress influence an array of processes including physiological, growth, developmental, yield, and quality of crop. The objective of this review is to provide an overview of the influences of these two stresses on the above processes independently and in combination. Our review suggests a clear need of information on interactive effects of stresses particularly of drought and heat stress which mostly occur in combination. Both short- and long-term stresses can signifi cantly influence growth and yield processes when stress occurs at sensitive stages. Crops are generally more sensitive to drought and/or heat stress during reproductive stages of development, which mainly influences seed numbers. Some of the important traits associated with drought- and/or heat-stress tolerance are indicated and discussed. The impacts of drought and heat stress are often different, and tolerance mechanisms may also be different. There is a wide range of crop modeling approaches (simple empirical models and more mechanistic models) that try to quantify the impact of stresses on growth, development, and yield and yield quality traits. These crop models should have the capability to quantify the impact of both short- and long-term stress events on growth, development, and yield processes. Modeling growth, development, sink-source relation, grain yield, and grain quality of crops can improve understanding of physiological and genetic nature of tolerance which can lead to increased grain yield and quality of crops. Improved models can enhance our capacity to predict crop performance in future climates and also to identify traits that can potentially be improved or exploited to obtain higher and more stable crop yields under stressed environments.

Impacts of Drought and/or Heat Stress on Physiological, Developmental, Growth, and Yield Processes of Crop Plants

The heat wave of summer 2003 was the largest and the most persistent ever experienced in Central Europe and has fuelled concern about the effects of climate change on European ecosystems. Since forests constitute the most important European ecosystems, in this review article we assess current knowledge on the effects of heat and drought on key metabolic processes for growth and productivity of forest trees. In particular, the general consequences of heat and drought on (1) photosynthesis and respiration at the cellular and community level, and (2) on nutrient uptake, partitioning and competition for nutrients are summarized. The latter are a major sink for photosynthetic energy and, therefore, are indirectly but strongly connected to the performance of photosynthesis. In addition, the interaction of heat and drought with stress compensation mechanisms and emission of biogenic volatile organic compounds (BVOC) are discussed, since these processes are directly connected to carbon metabolism. Effects on the emission of BVOC are also included because they constitute an important feedback mechanism on ozone formation and, thus, on atmospheric pollution. As far as available, data collected during the 2003 heat wave are included and discussed.

Physiological responses of forest trees to heat and drought.

The effects of plants on the biosphere, atmosphere and geosphere are key determinants of terrestrial ecosystem functioning. However, despite substantial progress made regarding plant belowground components, we are still only beginning to explore the complex relationships between root traits and functions. Drawing on the literature in plant physiology, ecophysiology, ecology, agronomy and soil science, we reviewed 24 aspects of plant and ecosystem functioning and their relationships with a number of root system traits, including aspects of architecture, physiology, morphology, anatomy, chemistry, biomechanics and biotic interactions. Based on this assessment, we critically evaluated the current strengths and gaps in our knowledge, and identify future research challenges in the field of root ecology. Most importantly, we found that belowground traits with the broadest importance in plant and ecosystem functioning are not those most commonly measured. Also, the estimation of trait relative importance for functioning requires us to consider a more comprehensive range of functionally relevant traits from a diverse range of species, across environments and over time series. We also advocate that establishing causal hierarchical links among root traits will provide a hypothesis-based framework to identify the most parsimonious sets of traits with the strongest links on functions, and to link genotypes to plant and ecosystem functioning.

https://hal.archives-ouvertes.fr/hal-03007086/document

Root traits as drivers of plant and ecosystem functioning: current understanding, pitfalls and future research needs

In the context of a recent massive increase in research on plant root functions and their impact on the environment, root ecologists currently face many important challenges to keep on generating cutting-edge, meaningful and integrated knowledge. Consideration of the below-ground components in plant and ecosystem studies has been consistently called for in recent decades, but methodology is disparate and sometimes inappropriate. This handbook, based on the collective effort of a large team of experts, will improve trait comparisons across studies and integration of information across databases by providing standardised methods and controlled vocabularies. It is meant to be used not only as starting point by students and scientists who desire working on below-ground ecosystems, but also by experts for consolidating and broadening their views on multiple aspects of root ecology. Beyond the classical compilation of measurement protocols, we have synthesised recommendations from the literature to provide key background knowledge useful for: (1) defining below-ground plant entities and giving keys for their meaningful dissection, classification and naming beyond the classical fine-root vs coarse-root approach; (2) considering the specificity of root research to produce sound laboratory and field data; (3) describing typical, but overlooked steps for studying roots (e.g. root handling, cleaning and storage); and (4) gathering metadata necessary for the interpretation of results and their reuse. Most importantly, all root traits have been introduced with some degree of ecological context that will be a foundation for understanding their ecological meaning, their typical use and uncertainties, and some methodological and conceptual perspectives for future research. Considering all of this, we urge readers not to solely extract protocol recommendations for trait measurements from this work, but to take a moment to read and reflect on the extensive information contained in this broader guide to root ecology, including sections I-VII and the many introductions to each section and root trait description. Finally, it is critical to understand that a major aim of this guide is to help break down barriers between the many subdisciplines of root ecology and ecophysiology, broaden researchers' views on the multiple aspects of root study and create favourable conditions for the inception of comprehensive experiments on the role of roots in plant and ecosystem functioning.

https://hal.archives-ouvertes.fr/hal-02918834/document

A starting guide to root ecology: strengthening ecological concepts and standardising root classification, sampling, processing and trait measurements

Contents
 	'Summary'	513	
I.	'Introduction'	513	
II.	'Integrating nutrients into the hydraulic framework predicting mechanisms of drought survival, mortality, and recovery'	514	
III.	'Integration'	517	
 	'Acknowledgements'	518	
 	References	518	


Summary
Global forests are experiencing rising temperatures and more severe droughts, with consistently dire forecasts for negative future impacts. Current research on the physiological mechanisms underlying drought impacts is focused on the water- and carbon-associated mechanisms. The role of nutrients is notably missing from this research agenda. Here, we investigate what role, if any, forest nutrition plays for survival and recovery of forests during and after drought. High nutrient availability may play a detrimental role in drought survival due to preferential biomass allocation aboveground that (1) predispose plants to hydraulic constraints limiting photosynthesis and promoting hydraulic failure, (2) increases carbon costs during periods of carbon starvation, and (3) promote biotic attack due to low tissue carbon: nitrogen (C : N). When nutrient uptake occurs during drought, high nutrient availability can increase water use efficiency thus minimizing negative feedbacks between carbon and nutrient balance. Nutrients are released after drought ceases, which might promote faster recovery but the temporal dynamics of microbial immobilization and nutrient leaching have a significant impact on nutrient availability. We provide a framework for understanding nutrient impacts on drought survival that allows a more complete analysis of forest ecosystem responses.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1111/nph.14340

The role of nutrients in drought-induced tree mortality and recovery

The effects of local climate and silvicultural treatment on the inorganic N availability, net N uptake capacity of mycorrhizal beech roots and microbial N conversion were assessed in order to characterise changes in the partitioning of inorganic N between adult beech and soil microorganisms. Fine root dynamics, inorganic N in the soil solution and in soil extracts, nitrate and ammonium uptake kinetics of beech as well as gross ammonification, nitrification and denitrification rates were determined in a beech stand consisting of paired sites that mainly differed in aspect (SW vs. NE) and stand density (controls and thinning treatments). Nitrate was the only inorganic N form detectable in the soil water. Its concentration was high in control plots of the NE aspect, but only in canopy gaps and not influenced by thinning. Neither thinning nor aspect affected the abundance of root tips in the soil. Maximum nitrate net uptake by mycorrhizal fine roots of beech, however, differed with aspect, showing significantly lower values at the SW aspect with warm–dry local climate. There were no clear-cut significant effects of local climate or thinning on microbial N conversion, but a tendency towards higher ammonification and nitrification and lower denitrification rates on the untreated controls of the SW as compared to the NE aspect. Apparently, the observed sensitivity of beech towards reduced soil water availability is at least partially due to impaired N acquisition. This seems to be mainly a consequence of reduced N uptake capacity rather than of limited microbial re-supply of inorganic N or of changed patterns of inorganic N partitioning between soil bacteria and roots.

Climate and forest management influence nitrogen balance of European beech forests: microbial N transformations and inorganic N net uptake capacity of mycorrhizal roots

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Climate and forest management influence nitrogen balance of European beech forests: microbial N transformations and inorganic N net uptake capacity of mycorrhizal roots