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      Different Modes of Hydrogen Peroxide Action During Seed Germination

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          Abstract

          Hydrogen peroxide was initially recognized as a toxic molecule that causes damage at different levels of cell organization and thus losses in cell viability. From the 1990s, the role of hydrogen peroxide as a signaling molecule in plants has also been discussed. The beneficial role of H 2O 2 as a central hub integrating signaling network in response to biotic and abiotic stress and during developmental processes is now well established. Seed germination is the most pivotal phase of the plant life cycle, affecting plant growth and productivity. The function of hydrogen peroxide in seed germination and seed aging has been illustrated in numerous studies; however, the exact role of this molecule remains unknown. This review evaluates evidence that shows that H 2O 2 functions as a signaling molecule in seed physiology in accordance with the known biology and biochemistry of H 2O 2. The importance of crosstalk between hydrogen peroxide and a number of signaling molecules, including plant phytohormones such as abscisic acid, gibberellins, and ethylene, and reactive molecules such as nitric oxide and hydrogen sulfide acting on cell communication and signaling during seed germination, is highlighted. The current study also focuses on the detrimental effects of H 2O 2 on seed biology, i.e., seed aging that leads to a loss of germination efficiency. The dual nature of hydrogen peroxide as a toxic molecule on one hand and as a signal molecule on the other is made possible through the precise spatial and temporal control of its production and degradation. Levels of hydrogen peroxide in germinating seeds and young seedlings can be modulated via pre-sowing seed priming/conditioning. This rather simple method is shown to be a valuable tool for improving seed quality and for enhancing seed stress tolerance during post-priming germination. In this review, we outline how seed priming/conditioning affects the integrative role of hydrogen peroxide in seed germination and aging.

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          Seed dormancy and the control of germination.

          Seed dormancy is an innate seed property that defines the environmental conditions in which the seed is able to germinate. It is determined by genetics with a substantial environmental influence which is mediated, at least in part, by the plant hormones abscisic acid and gibberellins. Not only is the dormancy status influenced by the seed maturation environment, it is also continuously changing with time following shedding in a manner determined by the ambient environment. As dormancy is present throughout the higher plants in all major climatic regions, adaptation has resulted in divergent responses to the environment. Through this adaptation, germination is timed to avoid unfavourable weather for subsequent plant establishment and reproductive growth. In this review, we present an integrated view of the evolution, molecular genetics, physiology, biochemistry, ecology and modelling of seed dormancy mechanisms and their control of germination. We argue that adaptation has taken place on a theme rather than via fundamentally different paths and identify similarities underlying the extensive diversity in the dormancy response to the environment that controls germination.
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            Oxidative modifications to cellular components in plants.

            Reactive oxygen species (ROS) and reactive nitrogen species (RNS) are produced in many places in living cells and at an increased rate during biotic or abiotic stress. ROS and RNS participate in signal transduction, but also modify cellular components and cause damage. We first look at the most common ROS and their properties. We then consider the ways in which the cell can regulate their production and removal. We critically assess current knowledge about modifications of polyunsaturated fatty acids (PUFAs), DNA, carbohydrates, and proteins and illustrate this knowledge with case stories wherever possible. Some oxidative breakdown products, e.g., from PUFA, can cause secondary damage. Other oxidation products are secondary signaling molecules. We consider the fate of the modified components, the energetic costs to the cell of replacing such components, as well as strategies to minimize transfer of oxidatively damaged components to the next generation.
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              Epigenetic regulation of stress responses in plants.

              Gene expression driven by developmental and stress cues often depends on nucleosome histone post-translational modifications and sometimes on DNA methylation. A number of studies have shown that these DNA and histone modifications play a key role in gene expression and plant development under stress. Most of these stress-induced modifications are reset to the basal level once the stress is relieved, while some of the modifications may be stable, that is, may be carried forward as 'stress memory' and may be inherited across mitotic or even meiotic cell divisions. Epigenetic stress memory may help plants more effectively cope with subsequent stresses. Comparative studies on stress-responsive epigenomes and transcriptomes will enhance our understanding of stress adaptation of plants.
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                Author and article information

                Contributors
                Journal
                Front Plant Sci
                Front Plant Sci
                Front. Plant Sci.
                Frontiers in Plant Science
                Frontiers Media S.A.
                1664-462X
                04 February 2016
                2016
                : 7
                : 66
                Affiliations
                [1]Department of Plant Physiology, Institute of Experimental Biology, Adam Mickiewicz University in Poznan Poznan, Poland
                Author notes

                Edited by: Naser A. Anjum, University of Aveiro, Portugal

                Reviewed by: Lam-Son Tran, RIKEN Center for Sustainable Resource Science, Japan; Abdul Wahid, University of Agriculture Faisalabad, Pakistan

                *Correspondence: Łukasz Wojtyla, lukasz.wojtyla@ 123456amu.edu.pl

                Present address: Szymon Kubala, Department of Protein Biosynthesis, Institute of Biochemistry and Biophysics, Polish Academy of Sciences, Warsaw, Poland

                This article was submitted to Plant Physiology, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2016.00066
                4740362
                26870076
                47fbb093-fc77-4597-8619-6de8efa514ca
                Copyright © 2016 Wojtyla, Lechowska, Kubala and Garnczarska.

                This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) or licensor are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.

                History
                : 05 November 2015
                : 14 January 2016
                Page count
                Figures: 3, Tables: 0, Equations: 0, References: 190, Pages: 16, Words: 0
                Funding
                Funded by: Narodowe Centrum Nauki 10.13039/501100004281
                Award ID: 2011/03/B/NZ/00068
                Funded by: Ministerstwo Nauki i Szkolnictwa Wyzszego 10.13039/501100004569
                Award ID: ZFD.74.53.2014.2.AS
                Categories
                Plant Science
                Review

                Plant science & Botany
                dormancy,germination,hydrogen peroxide,phytohormone,priming,reactive oxygen species,seed,signaling molecule

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