Several reactive oxygen species (ROS) are continuously produced in plants as byproducts of aerobic metabolism. Depending on the nature of the ROS species, some are highly toxic and rapidly detoxified by various cellular enzymatic and nonenzymatic mechanisms. Whereas plants are surfeited with mechanisms to combat increased ROS levels during abiotic stress conditions, in other circumstances plants appear to purposefully generate ROS as signaling molecules to control various processes including pathogen defense, programmed cell death, and stomatal behavior. This review describes the mechanisms of ROS generation and removal in plants during development and under biotic and abiotic stress conditions. New insights into the complexity and roles that ROS play in plants have come from genetic analyses of ROS detoxifying and signaling mutants. Considering recent ROS-induced genome-wide expression analyses, the possible functions and mechanisms for ROS sensing and signaling in plants are compared with those in animals and yeast.

REACTIVE OXYGEN SPECIES: Metabolism, Oxidative Stress, and Signal Transduction

Reactive oxygen species and antioxidant machinery in abiotic stress tolerance in crop plants

1 Plant ecology at high elevations.- The concept of limitation.- A regional and historical account.- The challenge of alpine plant research.- 2 The alpine life zone.- Altitudinal boundaries.- Global alpine land area.- Alpine plant diversity.- Origin of alpine floras.- Alpine growth forms.- 3 Alpine climate.- Which alpine climate.- Common features of alpine climates.- Regional features of alpine climates.- 4 The climate plants experience.- Interactions of relief, wind and sun.- How alpine plants influence their climate.- The geographic variation of alpine climate.- 5 Life under snow: protection and limitation.- Temperatures under snow.- Solar radiation under snow.- Gas concentrations under snow.- Plant responses to snowpack.- 6 Alpine soils.- Physics of alpine soil formation.- The organic compound.- The interaction of organic and inorganic compounds.- 7 Alpine treelines.- About trees and lines.- Current altitudinal positions of climatic treelines.- Treeline-climate relationships.- Intrazonal variations and pantropical plateauing of alpine treelines.- Treelines in the past.- Attempts at a functional explanation of treelines.- A hypothesis for treeline formation.- Growth trends near treelines.- Evidence for sink limitation.- 8 Climatic stress.- Survival of low temperature extremes.- Avoidance and tolerance of low temperature extremes.- Heat stress in alpine plants.- Ultraviolet radiation - a stress factor.- 9 Water relations.- Ecosystem water balance.- Soil moisture at high altitudes.- Plant water relations - a brief review of principles.- Water relations of alpine plants.- Desiccation stress.- Water relations of special plant types.- 10 Mineral nutrition.- Soil nutrients.- The nutrient status of alpine plants.- Nutrient cycling and nutrient budgets.- Nitrogen fixation.- Mycorrhiza.- Responses of vegetation to variable nutrient supply.- 11 Uptake and loss of carbon.- Photosynthetic capacity of alpine plants.- Photosynthetic responses to the environment.- Daily carbon gain of leaves.- The seasonal carbon gain of leaves.- C4 and CAM photosynthesis at high altitudes.- Tissue respiration of alpine plants.- Ecosystem carbon balance.- 12 Carbon investments.- Non-structural carbohydrates.- Lipids and energy content.- Carbon costs of leaves and roots.- Whole plant carbon allocation.- 13 Growth dynamics and phenology.- Seasonal growth.- Diurnal leaf extension.- Rates of plant dry matter accumulation.- Functional duration of leaves and roots.- 14 Cell division and tissue formation.- Cell size and plant size.- Mitosis and the cell cycle.- From meristem activity to growth control.- 15 Plant biomass production.- The structure of alpine plant canopies.- Primary productivity of alpine vegetation.- Plant dry matter pools.- Biomass losses through herbivores.- 16 Plant reproduction.- Flowering and pollination.- Seed development and seed size.- Germination.- Alpine seed banks and natural recruitment.- Clonal propagation.- Alpine plant age.- Community processes.- 17 Global change at high elevation.- Alpine land use.- The impact of altered atmospheric chemistry.- Climatic change and alpine ecosystems.- References (with chapter annotation).- Taxonomic index (genera).- Geographical index.- Color plates.- Plant life forms.- The alpine life zone.- Environmental stress.- The human dimension.

Alpine plant life

Water deficit and salinity, especially under high light intensity or in combination with other stresses, disrupt photosynthesis and increase photorespiration, altering the normal homeostasis of cells and cause an increased production of reactive oxygen species (ROS). ROS play a dual role in the response of plants to abiotic stresses functioning as toxic by-products of stress metabolism, as well as important signal transduction molecules. In this review, we provide an overview of ROS homeostasis and signalling in response to drought and salt stresses and discuss the current understanding of ROS involvement in stress sensing, stress signalling and regulation of acclimation responses.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2009.02041.x

Reactive oxygen species homeostasis and signalling during drought and salinity stresses

Contents
I. Introduction  424
II. The phosphorus conundrum 424
III. Adaptations to low P 424
IV. Uptake of P  424
V. P deficiency alters root development and function 426
VI. P deficiency modifies carbon metabolism 431
VII. Acid phosphatase 436
VIII. Genetic regulation of P responsive genes 437
IX. Improving P acquisition 439
X. Synopsis  440


Summary
Phosphorus (P) is limiting for crop yield on > 30% of the world's arable land and, by some estimates, world resources of inexpensive P may be depleted by 2050. Improvement of P acquisition and use by plants is critical for economic, humanitarian and environmental reasons. Plants have evolved a diverse array of strategies to obtain adequate P under limiting conditions, including modifications to root architecture, carbon metabolism and membrane structure, exudation of low molecular weight organic acids, protons and enzymes, and enhanced expression of the numerous genes involved in low-P adaptation. These adaptations may be less pronounced in mycorrhizal-associated plants. The formation of cluster roots under P-stress by the nonmycorrhizal species white lupin (Lupinus albus), and the accompanying biochemical changes exemplify many of the plant adaptations that enhance P acquisition and use. Physiological, biochemical, and molecular studies of white lupin and other species response to P-deficiency have identified targets that may be useful for plant improvement. Genomic approaches involving identification of expressed sequence tags (ESTs) found under low-P stress may also yield target sites for plant improvement. Interdisciplinary studies uniting plant breeding, biochemistry, soil science, and genetics under the large umbrella of genomics are prerequisite for rapid progress in improving nutrient acquisition and use in plants.

/pdf/phosphorus-acquisition-and-use-critical-adaptations-by-56im8qzi8m.pdf

Phosphorus acquisition and use: critical adaptations by plants for securing a nonrenewable resource

A novel insight into Arabidopsis mitochondrial function was revealed from a large experimental proteome derived by liquid chromatography-tandem mass spectrometry. Within the experimental set of 416 identified proteins, a significant number of low-abundance proteins involved in DNA synthesis, transcriptional regulation, protein complex assembly, and cellular signaling were discovered. Nearly 20% of the experimentally identified proteins are of unknown function, suggesting a wealth of undiscovered mitochondrial functions in plants. Only approximately half of the experimental set is predicted to be mitochondrial by targeting prediction programs, allowing an assessment of the benefits and limitations of these programs in determining plant mitochondrial proteomes. Maps of putative orthology networks between yeast, human, and Arabidopsis mitochondrial proteomes and the Rickettsia prowazekii proteome provide detailed insights into the divergence of the plant mitochondrial proteome from those of other eukaryotes. These show a clear set of putative cross-species orthologs in the core metabolic functions of mitochondria, whereas considerable diversity exists in many signaling and regulatory functions.

Experimental Analysis of the Arabidopsis Mitochondrial Proteome Highlights Signaling and Regulatory Components, Provides Assessment of Targeting Prediction Programs, and Indicates Plant-Specific Mitochondrial Proteins

Treatment of Arabidopsis cell culture for 16 h with H2O2, menadione or antimycin A induced an oxidative stress decreasing growth rate and increasing DCF fluorescence and lipid peroxidation products. Treated cells remained viable and maintained significant respiratory rates. Mitochondrial integrity was maintained, but accumulation of alternative oxidase and decreased abundance of lipoic acid-containing components during several of the treatments indicated oxidative stress. Analysis of the treatments was undertaken by IEF/SDS-PAGE, comparison of protein spot abundances and tandem mass spectrometry. A set of 25 protein spots increased >3-fold in H2O2/menadione treatments, a subset of these increased in antimycin A-treated samples. A set of 10 protein spots decreased significantly during stress treatments. A specific set of mitochondrial proteins were degraded by stress treatments. These damaged components included subunits of ATP synthase, complex I, succinyl CoA ligase, aconitase, and pyruvate and 2-oxoglutarate dehydrogenase complexes. Nine increased proteins represented products of different genes not found in control mitochondria. One is directly involved in antioxidant defense, a mitochondrial thioredoxin-dependent peroxidase, while another, a thioredoxin reductase-dependent protein disulphide isomerase, is required for protein disulfide redox homeostasis. Several others are generally considered to be extramitochondrial but are clearly present in a highly purified mitochondrial fraction used in this study and are known to play roles in stress response. Using H2O2 as a model stress, further work revealed that this treatment induced a protease activity in isolated mitochondria, putatively responsible for the degradation of oxidatively damaged mitochondrial proteins and that O2 consumption by mitochondria was significantly decreased by H2O2 treatment.

https://onlinelibrary.wiley.com/doi/pdf/10.1046/j.1365-313X.2002.01474.x

The impact of oxidative stress on Arabidopsis mitochondria.

Mitochondrial respiration in plants provides energy for biosynthesis, and its balance with photosynthesis determines the rate of plant biomass accumulation. We describe recent advances in our understanding of the mitochondrial respiratory machinery of cells, including the presence of a classical oxidative phosphorylation system linked to the cytosol by transporters, discussed alongside nonphosphorylating (and, therefore, non-energy conserving) bypasses that alter the efficiency of ATP synthesis and play a role in oxidative stress responses in plants. We consider respiratory regulation in the context of the contrasting roles mitochondria play in different tissues, from photosynthetic leaves to nutrient-acquiring roots. We focus on the molecular nature of this regulation at transcriptional and post-transcriptional levels that allow the respiratory apparatus of plants to help shape organ development and the response of plants to environmental stress. We highlight the challenges for future research considerin...

Organization and regulation of mitochondrial respiration in plants.

Infection of legume roots or stems with soil bacteria of the Rhizobiaceae results in the formation of nodules that become symbiotic nitrogen-fixing organs. Within the infected cells of these nodules, bacteria are enveloped in a membrane of plant origin, called the peribacteroid membrane (PBM), and divide and differentiate to form nitrogen-fixing bacteroids. The organelle-like structure comprised of PBM and bacteroids is termed the symbiosome, and is the basic nitrogen-fixing unit of the nodule. The major exchange of nutrients between the symbiotic partners is reduced carbon from the plant, to fuel nitrogenase activity in the bacteroid, and fixed nitrogen from the bacteroid, which is assimilated in the plant cytoplasm. However, many other metabolites are also exchanged. The metabolic interaction between the plant and the bacteroids is regulated by a series of transporters and channels on the PBM and the bacteroid membrane, and these form the focus of this review.

David A. Day

Papers

Experimental Analysis of the Arabidopsis Mitochondrial Proteome Highlights Signaling and Regulatory Components, Provides Assessment of Targeting Prediction Programs, and Indicates Plant-Specific Mitochondrial Proteins

The impact of oxidative stress on Arabidopsis mitochondria.

Organization and regulation of mitochondrial respiration in plants.

Metabolite transport across symbiotic membranes of legume nodules

Metabolite transport across symbiotic membranes of legume nodules