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      Stone Fruits: Growth and Nitrogen and Organic Acid Metabolism in the Fruits and Seeds—A Review

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          Abstract

          Stone fruits of the Rosaceae family consist of several distinct parts, and these include the flesh, woody endocarp, and seed. To understand the metabolism of these fruits, it is necessary to have knowledge of both their structure and growth characteristics. The nitrogen metabolism of the different tissues of stone fruits is interlinked. For example, there is an import and storage of nitrogenous compounds in the endocarp that are then exported to the seed. Moreover, there are links between the metabolism of nitrogen and that of malic/citric acids. In this article, the structure and growth characteristics, together with the import/export, contents, metabolism, and functions of nitrogenous compounds and organic acids in the different parts of stone fruits and their seeds are reviewed.

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          An Overview of Sucrose Synthases in Plants

          Sucrose is the end product of photosynthesis and the primary sugar transported in the phloem of most plants. Sucrose synthase (SuSy) is a glycosyl transferase enzyme that plays a key role in sugar metabolism, primarily in sink tissues. SuSy catalyzes the reversible cleavage of sucrose into fructose and either uridine diphosphate glucose (UDP-G) or adenosine diphosphate glucose (ADP-G). The products of sucrose cleavage by SuSy are available for many metabolic pathways, such as energy production, primary-metabolite production, and the synthesis of complex carbohydrates. SuSy proteins are usually homotetramers with an average monomeric molecular weight of about 90 kD (about 800 amino acids long). Plant SuSy isozymes are mainly located in the cytosol or adjacent to plasma membrane, but some SuSy proteins are found in the cell wall, vacuoles, and mitochondria. Plant SUS gene families are usually small, containing between four to seven genes, with distinct exon-intron structures. Plant SUS genes are divided into three separate clades, which are present in both monocots and dicots. A comprehensive phylogenetic analysis indicates that a first SUS duplication event may have occurred before the divergence of the gymnosperms and angiosperms and a second duplication event probably occurred in a common angiosperm ancestor, leading to the existence of all three clades in both monocots and dicots. Plants with reduced SuSy activity have been shown to have reduced growth, reduced starch, cellulose or callose synthesis, reduced tolerance to anaerobic-stress conditions and altered shoot apical meristem function and leaf morphology. Plants overexpressing SUS have shown increased growth, increased xylem area and xylem cell-wall width, and increased cellulose and starch contents, making SUS high-potential candidate genes for the improvement of agricultural traits in crop plants. This review summarizes the current knowledge regarding plant SuSy, including newly discovered possible developmental roles for SuSy in meristem functioning that involve sugar and hormonal signaling.
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            The sucrose-cleaving enzymes of plants are crucial for development, growth and carbon partitioning.

            Sink organs of most plant species are supplied with carbon and energy in the form of sucrose. The channeling of sucrose into sink metabolism requires its cleavage by several isoforms of invertase and sucrose synthase, which are localized in different subcellular compartments. These activities regulate the entry of sucrose into distinct biochemical pathways, such as respiration or biosynthesis of cell wall polysaccharides and storage reserves. Other vital roles for the sucrose-cleaving enzymes include invertase activity at the site of phloem unloading and vacuolar invertase and sucrose synthase in sink organs, which drives the long-distance transport of sucrose. In addition, invertases have been implicated in the defense response and in turgor-driven cell expansion, and sucrose synthase expression is associated with low temperature and anaerobiosis responses. Finally, because sugars also regulate gene expression, the sucrose-cleaving enzymes play a fundamental role in controlling cell differentiation and development.
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              Asparagine in 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
                25 September 2020
                2020
                : 11
                : 572601
                Affiliations
                [1] 1Dipartimento di Scienze Agrarie, Alimentari e Ambientali, Università degli Studi di Perugia , Perugia, Italy
                [2] 2Department of Agronomy, Food, Natural Resources, Animals and Environment, University of Padova Agripolis , Legnaro, Italy
                [3] 3College of Life Science, University of Dundee , Dundee, United Kingdom
                [4] 4Facultad de Ciencias Bioquímicas y Farmacéuticas, Centro de Estudios Fotosintéticos y Bioquímicos, Consejo Nacional de Investigaciones Científicas y Técnicas, Universidad Nacional de Rosario , Rosario, Argentina
                [5] 5Istituto di Ricerca sugli Ecosistemi Terrestri, Consiglio Nazionale delle Ricerche , Porano (TR), Italy
                [6] 6CREA Centro di ricerca Olivicoltura, Frutticoltura e Agrumicoltura , Spoleto (PG), Italy
                [7] 7Department of Agricultural, Food, Environmental, and Animal Sciences, University of Udine , Udine, Italy
                Author notes

                Edited by: Flavia Guzzo, University of Verona, Italy

                Reviewed by: Maria Elena Maldonado, University of Antioquia, Colombia; Yuepeng Han, Chinese Academy of Sciences, China

                *Correspondence: Franco Famiani, franco.famiani@ 123456unipg.it ; Robert P. Walker, rob.walker@ 123456talktalk.net

                This article was submitted to Plant Metabolism and Chemodiversity, a section of the journal Frontiers in Plant Science

                Article
                10.3389/fpls.2020.572601
                7546786
                33101339
                707afec1-55ab-4fc0-89cd-fc0e3f3fd67a
                Copyright © 2020 Famiani, Bonghi, Chen, Drincovich, Farinelli, Lara, Proietti, Rosati, Vizzotto and Walker

                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) and the copyright owner(s) 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
                : 15 June 2020
                : 31 August 2020
                Page count
                Figures: 4, Tables: 1, Equations: 0, References: 118, Pages: 16, Words: 11016
                Categories
                Plant Science
                Review

                Plant science & Botany
                endocarp metabolism,fruit growth,nitrogen compounds,organic acids,seed metabolism

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