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      Comparative Analysis of the Combined Effects of Different Water and Phosphate Levels on Growth and Biological Nitrogen Fixation of Nine Cowpea Varieties

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          Abstract

          Water deficit and phosphate (Pi) deficiency adversely affect growth and biological nitrogen fixation (BNF) of legume crops. In this study, we examined the impact of interaction between soil water conditions and available soil-Pi levels on growth, nodule development and BNF potential of nine cowpea varieties grown on dry savanna soils. In our experimental design, soils with different available soil-Pi levels, i.e., low, moderate, and high soil-Pi levels, collected from various farming fields were used to grow nine cowpea varieties under well-watered and water-deficit conditions. Significant and severe water deficit-damaging effects on BNF, nodulation, growth, levels of plant-nitrogen (N) and -phosphorus (P), as well as shoot relative water content and chlorophyll content of cowpea plants were observed. Under well-watered and high available soil-Pi conditions, cowpea varieties IT07K-304-9 and Dan'Ila exhibited significantly higher BNF potential and dry biomass, as well as plant-N and -P contents compared with other tested ones. Significant genotypic variations among the cowpeas were recorded under low available soil-Pi and water-deficit conditions in terms of the BNF potential. Principal component (PC) analysis revealed that varieties IT04K-339-1, IT07K-188-49, IT07K-304-9, and IT04K-405-5 were associated with PC1, which was better explained by performance for nodulation, plant biomass, plant-N, plant-P, and BNF potential under the combined stress of water deficit and Pi deficiency, thereby offering prospects for development of varieties with high growth and BNF traits that are adaptive to such stress conditions in the region. On another hand, variety Dan'Ila was significantly related to PC2 that was highly explained by the plant shoot/root ratio and chlorophyll content, suggesting the existence of physiological and morphological adjustments to cope with water deficit and Pi deficiency for this particular variety. Additionally, increases in soil-Pi availability led to significant reductions of water-deficit damage on dry biomass, plant-N and -P contents, and BNF potential of cowpea varieties. This finding suggests that integrated nutrient management strategies that allow farmers to access to Pi-based fertilizers may help reduce the damage of adverse water deficit and Pi deficiency caused to cowpea crop in the regions, where soils are predominantly Pi-deficient and drought-prone.

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

          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. 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.
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            How do plants respond to nutrient shortage by biomass allocation?

            Plants constantly sense the changes in their environment; when mineral elements are scarce, they often allocate a greater proportion of their biomass to the root system. This acclimatory response is a consequence of metabolic changes in the shoot and an adjustment of carbohydrate transport to the root. It has long been known that deficiencies of essential macronutrients (nitrogen, phosphorus, potassium and magnesium) result in an accumulation of carbohydrates in leaves and roots, and modify the shoot-to-root biomass ratio. Here, we present an update on the effects of mineral deficiencies on the expression of genes involved in primary metabolism in the shoot, the evidence for increased carbohydrate concentrations and altered biomass allocation between shoot and root, and the consequences of these changes on the growth and morphology of the plant root system.
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              Plant and mycorrhizal regulation of rhizodeposition

              The loss of carbon from roots (rhizodeposition) and the consequent proliferation of microorganisms in the surrounding soil, coupled with the physical presence of a root and processes associated with nutrient uptake, gives rise to a unique zone of soil called the rhizosphere. In this review, we bring together evidence to show that roots can directly regulate most aspects of rhizosphere C flow either by regulating the exudation process itself or by directly regulating the recapture of exudates from soil. Root exudates have been hypothesized to be involved in the enhanced mobilization and acquisition of many nutrients from soil or the external detoxification of metals. With few exceptions, there is little mechanistic evidence from soil-based systems to support these propositions. We conclude that much more integrated work in realistic systems is required to quantify the functional significance of these processes in the field. We need to further unravel the complexities of the rhizosphere in order to fully engage with key scientific ideas such as the development of sustainable agricultural systems and the response of ecosystems to climate change. Contents I. Introduction 460 II. What is rhizodeposition? 460 III. Regulation of rhizodeposition 460 IV. How large is the root exudation C flux? 463 V. How responsive is the root exudation C flux? 463 VI. How responsive is the microbial community to root exudation? 464 VII. The role of root exudates in nutrient acquisition 464 VIII. Mycorrhizal fungi and rhizodeposition 471 IX. Future thoughts 474 Acknowledgements 474 References 474.
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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
                19 December 2017
                2017
                : 8
                : 2111
                Affiliations
                [1] 1AgroBiosciences Division, Mohammed VI Polytechnic University (UM6P) , Ben Guerir, Morocco
                [2] 2Office Chérifien des Phosphates (OCP)-Africa , Casablanca, Morocco
                [3] 3Department of Agronomy, Faculty of Agriculture, University of Khartoum , Shambat, Sudan
                [4] 4Department of Agronomy, University of Ibadan , Ibadan, Nigeria
                [5] 5Botany and Microbiology Department, College of Science, King Saud University , Riyadh, Saudi Arabia
                [6] 6Mycology and Plant Disease Survey Department, Plant Pathology Research Institute, ARC , Giza, Egypt
                [7] 7Plant Production Department, College of Food and Agricultural Sciences, King Saud University , Riyadh, Saudi Arabia
                [8] 8Institute of Research and Development, Duy Tan University , Da Nang, Vietnam
                [9] 9Signalling Pathway Research Unit, RIKEN Center for Sustainable Resource Science , Yokohama, Japan
                Author notes

                Edited by: Marcello Mastrorilli, Consiglio per la Ricerca in Agricoltura e l'Analisi dell'Economia Agraria (CREA), Italy

                Reviewed by: Alvina Gul, National University of Sciences and Technology, Pakistan; Jin Xu, Xishuangbanna Tropical Botanical Garden (CAS), China

                *Correspondence: Martin Jemo m.jemo@ 123456ocpafrica.com
                Lam-Son Phan Tran son.tran@ 123456riken.jp

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

                †Elsayed Fathi Abd_Allah orcid.org/0000-0002-8509-8953

                Article
                10.3389/fpls.2017.02111
                5742256
                29312379
                aa1e5af1-4753-4519-a0f3-7a5ab21761ab
                Copyright © 2017 Jemo, Sulieman, Bekkaoui, Olomide, Hashem, Abd_Allah, Alqarawi and Tran.

                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
                : 06 July 2017
                : 27 November 2017
                Page count
                Figures: 4, Tables: 3, Equations: 8, References: 87, Pages: 16, Words: 13595
                Categories
                Plant Science
                Original Research

                Plant science & Botany
                biological nitrogen fixation potential,cowpea varieties,water-deficit damage,dry savanna of nigeria,phosphate deficiency

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