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.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2004.01130.x

Plant and mycorrhizal regulation of rhizodeposition

Nitrogen uptake, assimilation and remobilization in plants: challenges for sustainable and productive agriculture

†Background Productive agriculture needs a large amount of expensive nitrogenous fertilizers. Improving nitrogen use efficiency (NUE) of crop plants is thus of key importance. NUE definitions differ depending on whether plants are cultivated to produce biomass or grain yields. However, for most plant species, NUE mainly depends on how plants extract inorganic nitrogen from the soil, assimilate nitrate and ammonium, and recycle organic nitrogen. Efforts have been made to study the genetic basis as well as the biochemical and enzymatic mechanisms involved in nitrogen uptake, assimilation, and remobilization in crops and model plants. The detection of the limiting factors that could be manipulated to increase NUE is the major goal of such research. †Scope An overall examination of the physiological, metabolic, and genetic aspects of nitrogen uptake, assimilation and remobilization is presented in this review. The enzymes and regulatory processes manipulated to improve NUE components are presented. Results obtained from natural variation and quantitative trait loci studies are also discussed. †Conclusions This review presents the complexity of NUE and supports the idea that the integration of the numerous data coming from transcriptome studies, functional genomics, quantitative genetics, ecophysiology and soil science into explanatory models of whole-plant behaviour will be promising.

REVIEW: PART OF A SPECIAL ISSUE ON PLANT NUTRITION Nitrogen uptake, assimilation and remobilization in plants: challenges for sustainable and productive agriculture

Root structure and functioning for efficient acquisition of phosphorus: Matching morphological and physiological traits.

Nitrogen (N) is the main mineral element in plant tissues and almost all of this nutrient is acquired from the soil by the roots. Nitrogen is available in many different forms in the soil, but the three most abundant forms are nitrate, ammonium and amino acids. The relative importance of these different soil N pools to a plant is difficult to measure and depends on many different environmental factors. Changes in the available amounts and imbalance in the supply of some N forms can even be toxic to plants and in extreme cases can lead to changes in the vegetation. However, the importance of this element for agriculture is reflected in the amounts of N-fertiliser applied to crops and this is a major cost (economic and environmental) for world agriculture. This review covers the molecular mechanisms that the plant uses for accessing these soil N pools and briefly includes consideration of the root N assimilatory pathways that exist in the plant. The soil forms of N that are used by plants depend on many factors, but a series of different transporter and assimilatory genes that can provide access to these pools have been identified. This information can now provide the molecular tools to identify the N sources accessed by a plant and the relative importance of these different pools.

Root Nitrogen Acquisition and Assimilation

https://febs.onlinelibrary.wiley.com/doi/pdf/10.1016/S0014-5793%2801%2902096-8

An arabidopsis T-DNA mutant affected in Nrt2 genes is impaired in nitrate uptake.

Coordination between the activity of ion transport systems in the root and photosynthesis in the shoot is a main feature of the integration of ion uptake in the whole plant. However, the mechanisms that ensure this coordination are largely unknown at the molecular level. Here, we show that the expression of five genes that encode root NO(3)(-), NH(4)(+), and SO(4)(2-) transporters in Arabidopsis is regulated diurnally and stimulated by sugar supply. We also provide evidence that one Pi and one K(+) transporter also are sugar inducible. Sucrose, glucose, and fructose are able to induce expression of the ion transporter genes but not of the carboxylic acids malate and 2-oxoglutarate. For most genes investigated, induction by light and induction by sucrose are strongly correlated, indicating that they reflect the same regulatory mechanism (i.e., stimulation by photosynthates). The functional importance of this control is highlighted by the phenotype of the atnrt2 mutant of Arabidopsis. In this mutant, the deletion of the sugar-inducible NO(3)(-) transporter gene AtNrt2.1 is associated with the loss of the regulation of high-affinity root NO(3)(-) influx by light and sugar. None of the sugar analogs used (3-O-methylglucose, 2-deoxyglucose, and mannose) is able to mimic the inducing effect of sugars. In addition, none of the sugar-sensing mutants investigated (rsr1-1, sun6, and gin1-1) is altered in the regulation of AtNrt2.1 expression. These results indicate that the induction of AtNrt2.1 expression by sugars is unrelated to the main signaling mechanisms documented for sugar sensing in plants, such as regulation by sucrose, hexose transport, and hexokinase (HXK) sensing activity. However, the stimulation of AtNrt2.1 transcript accumulation by sucrose and glucose is abolished in an antisense AtHXK1 line, suggesting that HXK catalytic activity and carbon metabolism downstream of the HXK step are crucial for the sugar regulation of AtNrt2.1 expression.

Regulation of root ion transporters by photosynthesis: functional importance and relation with hexokinase.

The role of AtNrt2.1 and AtNrt2.2 genes, encoding putative NO 3 − transporters in Arabidopsis, in the regulation of high-affinity NO 3 − uptake has been investigated in the atnrt2 mutant, where these two genes are deleted. Our initial analysis of the atnrt2 mutant (S. Filleur, M.F. Dorbe, M. Cerezo, M. Orsel, F. Granier, A. Gojon, F. Daniel-Vedele [2001] FEBS Lett 489: 220–224) demonstrated that root NO 3 − uptake is affected in this mutant due to the alteration of the high-affinity transport system (HATS), but not of the low-affinity transport system. In the present work, we show that the residual HATS activity in atnrt2 plants is not inducible by NO 3 − , indicating that the mutant is more specifically impaired in the inducible component of the HATS. Thus, high-affinity NO 3 − uptake in this genotype is likely to be due to the constitutive HATS. Root 15 NO 3 − influx in the atnrt2 mutant is no more derepressed by nitrogen starvation or decrease in the external NO 3 − availability. Moreover, the mutant also lacks the usual compensatory up-regulation of NO 3 − uptake in NO 3 − -fed roots, in response to nitrogen deprivation of another portion of the root system. Finally, exogenous supply of NH 4 + in the nutrient solution fails to inhibit 15 NO 3 − influx in the mutant, whereas it strongly decreases that in the wild type. This is not explained by a reduced activity of NH 4 + uptake systems in the mutant. These results collectively indicate that AtNrt2.1 and/or AtNrt2.2 genes play a key role in the regulation of the high-affinity NO 3 − uptake, and in the adaptative responses of the plant to both spatial and temporal changes in nitrogen availability in the environment.

/pdf/major-alterations-of-the-regulation-of-root-no3-uptake-are-58lg36rbbx.pdf

Miguel Cerezo

Papers

An arabidopsis T-DNA mutant affected in Nrt2 genes is impaired in nitrate uptake.

Regulation of root ion transporters by photosynthesis: functional importance and relation with hexokinase.

Major Alterations of the Regulation of Root NO3− Uptake Are Associated with the Mutation of Nrt2.1 and Nrt2.2 Genes in Arabidopsis