Nitrate transceptor(s) in plants

The availability of mineral nutrients in the soil dramatically fluctuates in both time and space. In order to optimize their nutrition, plants need efficient sensing systems that rapidly signal the local external concentrations of the individual nutrients. Until recently, the most upstream actors of the nutrient signalling pathways, i.e. the sensors/receptors that perceive the extracellular nutrients, were unknown. In Arabidopsis, increasing evidence suggests that, for nitrate, the main nitrogen source for most plant species, a major sensor is the NRT1.1 nitrate transporter, also contributing to nitrate uptake by the roots. Membrane proteins that fulfil a dual nutrient transport/signalling function have been described in yeast and animals, and are called 'transceptors'. This review aims to illustrate the nutrient transceptor concept in plants by presenting the current evidence indicating that NRT1.1 is a representative of this class of protein. The various facets, as well as the mechanisms of nitrate sensing by NRT1.1 are considered, and the possible occurrence of other nitrate transceptors is discussed.     

疏叶骆驼刺根系对土壤异质性和种间竞争的响应

近年来, 植物根系对土壤异质性的响应和植物根系之间的相互作用一直是研究的热点。过去的研究主要是针对一年生短命植物进行的, 而且多是在人工控制的温室条件下进行的。而对于多年生植物根系对养分异质性和竞争的综合作用研究很少。该文对塔里木盆地南缘多年生植物疏叶骆驼刺(Alhagi sparsifolia)根系生长对养分异质性和竞争条件的响应途径与适应策略进行了研究, 结果表明: (1)在无竞争的条件下, 疏叶骆驼刺根系优先向空间大的地方生长, 即使另一侧有养分斑块存在, 其根系也向着空间大的一侧生长; (2)在有竞争的条件下, 疏叶骆驼刺根系生长依然是优先占领空间大的一侧, 但是竞争者的存在抑制了疏叶骆驼刺的生长, 导致其枝叶生物量和根系生物量都明显减少(p < 0.01), 而养分斑块的存在促进了疏叶骆驼刺根系的生长; (3)疏叶骆驼刺根系的生长不仅需要养分, 也需要足够的空间, 空间比养分更重要; (4)有竞争者存在的时候, 两株植物的根系都先长向靠近竞争者一侧的空间, 即先占据“共有空间”。研究结果对理解植物根系觅食行为和植物对环境的适应策略有重要意义。     

Endophyte roles in nutrient acquisition,root system architecture development and oxidative stress tolerance

Plants associate with communities of microbes (bacteria and fungi) that play critical roles in plant development, nutrient acquisition and oxidative stress tolerance. The major share of plant microbiota is endophytes which inhabit plant tissues and help them in various capacities. In this article, we have reviewed what is presently known with regard to how endophytic microbes interact with plants to modulate root development, branching, root hair formation and their implications in overall plant development. Endophytic microbes link the interactions of plants, rhizospheric microbes and soil to promote nutrient solubilization and further vectoring these nutrients to the plant roots making the soil-plant-microbe continuum. Further, plant roots internalize microbes and oxidatively extract nutrients from microbes in the rhizophagy cycle. The oxidative interactions between endophytes and plants result in the acquisition of nutrients by plants and are also instrumental in oxidative stress tolerance of plants. It is evident that plants actively cultivate microbes internally, on surfaces and in soils to acquire nutrients, modulate development and improve health. Understanding this continuum could be of greater significance in connecting endophytes with the hidden half of the plant that can also be harnessed in applied terms to enhance nutrient acquisition through the development of favourable root system architecture for sustainable production under stress conditions.     

Explore less to control more: why and when should plants limit the horizontal exploration of soil by their roots?

In ecosystems limited by soil nutrients, some plants show a restricted horizontal distribution of their roots. We explored the hypothesis that this particular pattern is a foraging strategy emerging from tradeoffs between soil exploration (that increases the pool of nutrients available for plants) and the local control of nutrient cycling within the soil that we call soil occupation. We developed two general analytical models of the cycling of a limiting nutrient in a plant population that is not limited by water. They allowed to explore how plant productivity is affected when roots do not exploit the whole soil available and to determine the conditions for which plant nutrient stock is maximized when plants limit their exploration of soil. We predict that a restricted exploration strategy can be beneficial when 1) there is at least one tradeoff between a nutrient cycling parameter and soil exploration, 2) nutrient availability in the unexplored soil is poor and 3) the area of soil explored by plants is stable over time. The exploration limitation strategy results in spatially heterogeneous and nutrient‐conservative ecosystems. Our results should apply well to perennial tussock grasses within tropical nutrient‐limited ecosystems and raises interesting cues for the construction of more sustainable agro‐ecosystems. Overall, our study underlines the importance of considering the multiplicity of root–soil interactions and of their scales when considering root foraging strategies.     

宁南山区典型植物根系分解特征及其对土壤养分的影响

根系分解是陆地生态系统碳和养分循环的重要地下生态过程,研究宁南山区典型植物根系分解特征及其对土壤养分的影响,能够丰富和完善陆地生态系统的物质和能量循环机制,为我国黄土高原植被恢复过程中植物与土壤之间的养分循环提供依据。连续2年研究了宁南山区3种典型植物(长芒草、铁杆蒿和百里香)根系的分解特征及其对土壤养分的影响。结果表明,长芒草、铁杆蒿和百里香根系年分解指数(K)分别0.00891、0.01128、0.01408,分解速率依次表现为百里香铁杆蒿长芒草。分解16个月后3种典型植物根系释放大量养分,其中碳的释放量在57.05—124.39 g/kg;氮的释放量在0.12—0.47 g/kg。3种典型植物根系对土壤养分的影响主要表现为:试验结束时,0—5 cm表层土壤有机碳含量提高了0.17—0.35 g/kg,5—20 cm土层土壤有机碳含量提高了0.26—0.35 g/kg。相关性分析可知,植物根系养分释放量与土壤养分含量之间存在一定的负相关关系,当土壤养分含量较低时,根系会增加养分释放量进行补充。由此可知,根系分解提高了土壤养分含量,有效的促进了养分在根系-土壤中的循环。     

Water and Nutrient Dynamics in Surface Roots and Soils are not Modified by Short-term Flooding of Phreatophytic Plants in a Hyperarid Desert

Little is known of the mechanisms employed by woody plants to acquire key resources such as water and nutrients in hyperarid environments. For phreatophytic plants, deep roots are necessary to access the water table, but given that most nutrients in many desert ecosystems are stored in the upper soil layers, viable shallow roots may be equally necessary for nutrient uptake. We sought to better understand the interaction between water and nutrient uptake from soil horizons differing in the relative abundance of these resources. To this end, we monitored plant water and nutrient status before and after applying flood irrigation to four phreatophytic perennial plant species in the remote hyperarid Taklamakan desert in western China. Sap flow in the roots of five plants of the perennial desert species Alhagi sparsifolia Shap., Karelina caspica (Pall.) Less., Calligonum caput medusea Schrenk, and Eleagnus angustifolia Hill. was monitored using the heat ratio method (HRM). Additionally we measured predawn and midday water potential, foliar nitrate reductase activity (NRA), xylem sap nutrient concentration and the concentration of total solutes in the leaves before, 12 and 96 h after flooding to investigate possible short-term physiological effects on water and nutrient status. Rates of sap flow measured during the day and at night in the absence of transpiration did not change after flooding. Moderately high rates of sap flow (HRM heat pulse velocity, 5–25 cm h⁻¹) detected during the day in soils that had a near zero water content at the surface indicated that all species had contact to groundwater. There was no evidence from sap flow data that plants had utilised flood water to increase maximum rates of transpiration under similar climatic conditions, and there was no evidence of a process to improve the efficiency of water or nutrient uptake, such as hydraulic redistribution (i.e. the passive movement of water from moist soil to very dry soil via roots). Measurements of plant water status, xylem sap nutrient status, foliar NRA and the concentration of osmotically active substances were also unaffected by flood irrigation. Our results clearly show that groundwater acts as the major source of water and nutrients for these plants. The inability of plants to utilise abundant surface soil–water or newly available nutrients following irrigation was attributed to the absence of fine roots in the topsoil layer.     

植物根系向地性感应的分子机理与养分吸收

植物根系向地性是决定根系空间生长趋势的主要因素之一,对于养分吸收具有重要影响.认识根系向地性感应和根系生长变化的分子机理及其与养分吸收的关系,可为遗传改良根系性状、提高植物养分吸收效率提供理论依据.本文从重力感应、信号转导和生长素非对称分布等方面总结了植物根系向地性感应的分子机理,探讨了根系在养分胁迫下(特别是磷胁迫下)向地性变化的生理基础及其与养分吸收(特别是磷吸收)的关系,最后对根系向地性研究的若干问题进行了展望.     

Root-targeted biotechnology to mediate hormonal signalling and improve crop stress tolerance

Since plant root systems capture both water and nutrients essential for the formation of crop yield, there has been renewed biotechnological focus on root system improvement. Although water and nutrient uptake can be facilitated by membrane proteins known as aquaporins and nutrient transporters, respectively, there is a little evidence that root-localised overexpression of these proteins improves plant growth or stress tolerance. Recent work suggests that the major classes of phytohormones are involved not only in regulating aquaporin and nutrient transporter expression and activity, but also in sculpting root system architecture. Root-specific expression of plant and bacterial phytohormone-related genes, using either root-specific or root-inducible promoters or grafting non-transformed plants onto constitutive hormone producing rootstocks, has examined the role of root hormone production in mediating crop stress tolerance. Root-specific traits such as root system architecture, sensing of edaphic stress and root-to-shoot communication can be exploited to improve resource (water and nutrients) capture and plant development under resource-limited conditions. Thus, root system engineering provides new opportunities to maintain sustainable crop production under changing environmental conditions.     

Effect of heterogeneous distribution of nutrients on root growth, ABA content and drought resistance of wheat plants

Heterogeneous distribution of nutrients influenced morphology of wheat roots; they were shorter and had more laterals at the site of high nutrient content. The roots outside the nutrient-rich patch were longer and penetrated deeper in the soil than those of the plants supplied with the same quantity of nutrients but having them homogeneously distributed. The ABA content of roots sampled from the site of high nutrient content was greater than in those elsewhere. It is suggested that this might be important for inhibition of their extension growth and stimulation of laterals in the nutrient rich patch. Heterogeneously distributed nutrients beneficially influenced accumulation of ¹⁵N and assimilation of CO₂ by the plants, enabling an increased supply of nutrients to the parts of the root system beyond the nutrient rich patch and sustained their rapid extension. Enhanced penetration of roots into the deeper soil layers could contribute to the observed increase in drought resistance of plants grown with localised placement of fertilisers under field conditions.     

Biomass and nutrient allocation strategies in a desert ecosystem in the Hexi Corridor,northwest China

The allocation of biomass and nutrients in plants is a crucial factor in understanding the process of plant structures and dynamics to different environmental conditions. In this study, we present a comprehensive scaling analysis of data from a desert ecosystem to determine biomass and nutrient (carbon (C), nitrogen (N), and phosphorus (P)) allocation strategies of desert plants from 40 sites in the Hexi Corridor. We found that the biomass and levels of C, N, and P storage were higher in shoots than in roots. Roots biomass and nutrient storage were concentrated at a soil depth of 0–30 cm. Scaling relationships of biomass, C storage, and P storage between shoots and roots were isometric, but that of N storage was allometric. Results of a redundancy analysis (RDA) showed that soil nutrient densities were the primary factors influencing biomass and nutrient allocation, accounting for 94.5% of the explained proportion. However, mean annual precipitation was the primary factor influencing the roots biomass/shoots biomass (R/S) ratio. Furthermore, Pearson’s correlations and regression analyses demonstrated that although the biomass and nutrients that associated with functional traits primarily depended on soil conditions, mean annual precipitation and mean annual temperature had greater effects on roots biomass and nutrient storage.     

The Roots of Carnivorous Plants

Carnivorous plants may benefit from animal-derived nutrients to supplement minerals from the soil. Therefore, the role and importance of their roots is a matter of debate. Aquatic carnivorous species lack roots completely, and many hygrophytic and epiphytic carnivorous species only have a weakly devel-oped root system. In xerophytes, however, large, extended and/or deep-reaching roots and sub-soil shoots develop. Roots develop also in carnivorous plants in other habitats that are hostile, due to flood-ing, salinity or heavy metal occurance. Information about the structure and functioning of roots of car- nivorous plants is limited, but this knowledge is essential for a sound understanding of the plants’ physiology and ecology. Here we compile and summarise available information on: (1) The morphology of the roots. (2) The root functions that are taken over by stems and leaves in species without roots or with poorly developed root systems; anchoring and storage occur by specialized chlorophyll-less stems; water and nutrients are taken up by the trap leaves. (3) The contribution of the roots to the nutrient supply of the plants; this varies considerably amongst the few investigated species. We compare nutrient uptake by the roots with the acquisition of nutri-ents via the traps. (4) The ability of the roots of some carnivorous species to tolerate stressful conditions in their habitats; e.g., lack of oxygen, saline conditions, heavy metals in the soil, heat during bushfires, drought, and flooding     

Supply pre-emption, not concentration reduction, is the mechanism of competition for nutrients

Concentration reduction theory is the leading theory regarding the mechanism of competition for nutrients in soils among plants, yet it has not been rigorously tested. Here we used a spatially explicit, fine-scale grid-based model that simulated diffusion and plant uptake of nutrients by plants in soil to test whether concentration reduction theory was appropriate for terrestrial plant competition for nutrients. In the absence of competition, increasing the rate of diffusion allows a plant to maintain positive growth rates below the lowest average concentration to which it can reduce nutrients in soil solution (R*). As such, differences among plants in the reduction of soil moisture, which here primarily affects nutrient diffusion, can cause R* to predict competitive success incorrectly. The stronger competitor for nutrients captures the largest proportion of the nutrient supply by ensuring nutrients contact its roots before those of a competitor. Although the metric derived from concentration reduction theory, R*, might have predictive power for competitive outcomes in terrestrial ecosystems, this evidence suggests that plants outcompete other plants for nutrients by pre-empting the supply, not reducing the average concentration.     

Testing experimentally the effect of soil resource mobility on plant competition

Aims The volume of soil beyond a plant's roots from which that plant is able to acquire a particular nutrient depends upon the mobility of that nutrient in the soil. For this reason it has been hypothesized that the strength of competitive interactions between plants vary with soil nutrient mobility. We aimed to provide an experimental test of this hypothesis.Methods We devised two experimental systems to investigate specifically the effect of nutrient transport rates upon intraspecific competition. In the first, the exchange of rhizosphere water and dissolved nutrients between two connected pots, each containing one plant, was manipulated by alternately raising and lowering the pots. In the second experiment, the roots systems of two competing plants were separated by partitions of differing porosity, thereby varying the plants' access to water and nutrients in the other plant′s rhizosphere. In this second experiment, we also applied varying amounts of nutrients to test whether higher nutrient input would reduce competition when competition for light is avoided, and applied different water levels to affect nutrient concentrations without changing nutrient supply.Important findings In both experiments, lower mobility reduced competitive effects on plant biomass and on relative growth rate (RGR), as hypothesized. In the second experiment, however, competition was more intense under high nutrient input, suggesting that low nutrient supply rates reduced the strength of the superior competitor. Competitive effects on RGR were only evident under the low water level, suggesting that under lower nutrient concentrations, competitive effects might be less pronounced. Taken together, our results provide the first direct experimental evidence that a reduction in nutrient mobility can reduce the intensity of competition between plants.     

The modelled growth of mycorrhizal and non-mycorrhizal plants under constant versus variable soil nutrient concentration

We studied the response of mycorrhizal and non-mycorrhizal plants to variation in soil nutrient concentration. A model for the relative growth rate (RGR) of plant biomass was constructed with soil nutrients as an explanatory variable. A literature survey was carried out to find the relative magnitudes of parameter values for mycorrhizal and non-mycorrhizal plants. Mycorrhizal plants had higher RGR at low nutrient concentrations and non-mycorrhizal plants at high nutrient concentrations. The RGR of mycorrhizal and non-mycorrhizal plants at constant versus log-normally distributed soil nutrient concentration were compared to see the effect of mycorrhizal status on responses to variation. Variation in nutrient concentration generally reduced RGR, especially in mycorrhizal plants. The RGR of a non-mycorrhizal plant may increase with variation where a growth function threshold exists, i.e. a soil nutrient concentration that must be exceeded to allow growth. Mycorrhizal plants appeared more sensitive to variation in nutrient concentration than non-mycorrhizal plants due to the higher affinity of mycorrhizal roots at low nutrient levels. However, this prediction may be reversed if mycorrhizal symbiosis considerably stabilises flow of nutrients to plant physiological processes, such that mycorrhizal plants experience less variation in soil nutrient concentration than non-mycorrhizal plants. Our results also attain broader significance by suggesting a general trade-off between competitive ability in a constant versus variable resource availability.     

Focus Issue on Roots: Plant Nutrition: Root Transporters on the Move

Nutrient and water uptake from the soil is essential for plant growth and development. In the root, absorption and radial transport of nutrients and water toward the vascular tissues is achieved by a battery of specialized transporters and channels. Modulating the amount and the localization of these membrane transport proteins appears as a way to drive their activity and is essential to maintain nutrient homeostasis in plants. This control first involves the delivery of newly synthesized proteins to the plasma membrane by establishing check points along the secretory pathway, especially during the export from the endoplasmic reticulum. Plasma membrane-localized transport proteins are internalized through endocytosis followed by recycling to the cell surface or targeting to the vacuole for degradation, hence constituting another layer of control. These intricate mechanisms are often regulated by nutrient availability, stresses, and endogenous cues, allowing plants to rapidly adjust to their environment and adapt their development.Plants take up nutrients and water from the soil and transport them to the leaves to support photosynthesis and plant growth. However, most soils around the world do not provide optimal conditions for plant colonization. Consequently, plants have evolved sophisticated mechanisms to adjust to deficiency or excess of nutrients and water supply. Membrane transport proteins, including channels and transporters, play crucial roles in the uptake of nutrients and water from the soil and in their radial transport to the root vasculature. Newly synthesized membrane transport proteins have to be properly targeted to a defined compartment, usually the plasma membrane, to efficiently ensure their function. The trafficking of membrane transport proteins along the secretory pathway is tightly controlled and involves the recognition of exit signals by gatekeeper protein complexes. After reaching the plasma membrane, membrane transport proteins can be endocytosed and subsequently recycled to the cell surface or targeted to the vacuole for degradation. Because the subcellular localization of proteins directly influences their activity, modulating the localization of membrane transport proteins constitutes a powerful way to control nutrient and water uptake in plants. This review discusses the fundamental mechanisms at stake in membrane protein secretion and endocytosis, with a specific focus on membrane transport proteins, and how endogenous and exogenous cues affect their dynamics to integrate uptake of nutrients and water to plant growth conditions.     

细菌菌膜的成分、调控及其与植物的关系

菌膜是细菌群落发展的一种高度组织化的群体状态。在菌膜形成过程中,细菌胞外物质EPS(Exopolysaccharides)、eDNA(Extracellular DNA)、胞外蛋白等都参与菌膜的形成,它们为菌膜提供机械稳定性,帮助细菌粘附到物体表面,促进菌膜中不同细菌间物质的循环及基因的水平转移。菌膜形成涉及到群体感应、C-di-GMP(Cyclic diguanylate monophosphate)和sRNA等一系列调控机制。土壤环境中栖息着大量的微生物,许多土壤微生物定殖于植物根际,从而与植物发生着密切的相互作用;菌膜的形成是细菌稳定定殖于植物根际的关键因素,有助于植物促生菌或致病菌在根际更好的生存。本文就菌膜的成分、调控及其与植物的关系等三个方面的内容进行综述。     

Herbivorous insect decreases plant nutrient uptake: the role of soil nutrient availability and association of below‐ground symbionts

1. Plants take nutrients from the rhizosphere via two pathways: (i) by absorbing soil nutrients directly via their roots and (ii) indirectly via symbiotic associations with nutrient‐providing microbes. Herbivorous insects can alter these pathways by herbivory, adding their excrement to the soil, and affecting plant–microbe associations. 2. Little is known, however, about the effects of herbivorous insects on plant nutrient uptake. Greenhouse experiments with soybean, aphids, and rhizobia were carried out to examine the effects of aphids on plant nutrient uptake. 3. First, the inorganic soil nitrogen and the sugar in aphid honeydew between aphid‐infected and ‐free plants were compared. It was found that aphid honeydew added 41 g m^?2 of sugar to the soil, and that aphids decreased the inorganic soil nitrogen by 86%. This decrease may have been caused by microbial immobilisation of soil nitrogen followed by increased microbial abundance as a result of aphid honeydew. 4. Second, nitrogen forms in xylem sap between aphid‐infected and ‐free plants were compared to examine nitrogen uptake. Aphids decreased the nitrogen uptake via both pathways, and strength of the impact on direct uptake via plant roots was greater than indirect uptake via rhizobia. The reduced nitrogen uptake by the direct pathway was as a result of microbial immobilisation, and that by the indirect pathway was probably because of the interaction of microbial immobilisation and carbon stress, which was caused by aphid infection. 5. The present results demonstrate that herbivorous insects can negatively influence the two pathways of plant nutrient uptake and alter their relative importance.     

Heterogeneous distribution of phosphorus and potassium in soil influences wheat growth and nutrient uptake

Heterogeneous distribution of mineral nutrients in soil profiles is a norm in agricultural lands, but its influence on nutrient uptake and crop growth is poorly documented. In this study, we examined the effects of varying phosphorus (P) and potassium (K) distribution on plant growth and nutrient uptake by wheat (Triticum aestivum L.) grown in a layered or split soil culture in glasshouse conditions. In the layered pot system the upper soil was supplied with P and either kept watered or allowed to dry or left P-deficient but watered, whereas the lower soil was watered and fertilised with K. Greater reductions in shoot growth, root length and dry weight in the upper soil layer occurred in −P/wet than in +P/dry upper soil treatment. Shoot P concentration and total P content were reduced by P deficiency but not by upper soil drying. Genotypic responses showed that K-efficient cv. Nyabing grew better and took up more P and K than K-inefficient cv. Gutha in well-watered condition, but the differences decreased when the upper soil layer was dry. In the split-root system, shoot dry weight and shoot P and K contents were similar when P and K were applied together in one compartment or separated into two compartments. In comparison, root growth was stimulated and plants took up more P and K in the treatment with the two nutrients supplied together compared with the treatment in which the two nutrients were separated. Roots proliferated in the compartment applied with either P or K at the expense of root growth in the adjoining compartment with neither P nor K. Heterogeneous nutrient distribution has a direct decreasing effect on root growth in deficient patches, and nutrient redistribution within the plant is unlikely to meet the demand of roots grown in such patches.