Environmental,developmental, and genetic factors controlling root system architecture

A better understanding of the development and architecture of roots is essential to develop strategies to increase crop yield and optimize agricultural land use. Roots control nutrient and water uptake, provide anchoring and mechanical support and can serve as important storage organs. Root growth and development is under tight genetic control and modulated by developmental cues including plant hormones and the environment. This review focuses on root architecture and its diversity and the role of environment, nutrient, and water as well as plant hormones and their interactions in shaping root architecture.     

Dynamics of water and ion transport driven by corn canopy in the Yellow River basin

Water and ion balance in a corn field in the semi-arid region of the upper Yellow River basin (Inner Mongolia, China) was analyzed with special reference to transpiration stream and selective nutrient uptake driven by the crop canopy. During the crop development stage (June 7 to July 17, 2005), crop transpiration and soil evaporation were evaluated separately on a daily basis, and concentrations of NO ₃ ^? , PO ₄ ^3? , K⁺, Na⁺, Ca²⁺, Mg²⁺ and Cl^? ions in the Yellow River water, irrigation water, ground water, soil of the root zone and xylem sap of the crop were analyzed.The crop transpiration accounted for 83.4% of the evapotranspiration during the crop development stage. All ions except for Na⁺ were highly concentrated in the xylem sap due to the active and selective uptake of nutrients by roots. In particular, extremely high concentrations of the major essential nutrients were found in the nighttime stem exudate, while these concentrations in the river water, the irrigation water, the ground water and the root-zone soil were lower. On the other hand, Na⁺, which is not the essential element for crop growth, was scarcely absorbed by roots and was not highly concentrated in the xylem sap. Consequently, Na⁺ remained in the ground water and the root-zone soil at higher concentrations. These results indicate that during the growing season, crop transpiration but not soil evaporation induces the most significant driving force for mass flow (capillary rise) transporting the ground water toward the rhizosphere, where the dynamics of ion balance largely depends on the active and selective nutrient uptake by roots.     

三角叶滨藜根吸水特点与其抗盐性的关系

采用压力室和冰点渗透压计测定了三角叶滨藜在不同浓度NaCl的根系环境溶液中根木质部的压力势和伤流液的渗透势,并利用原子吸收分光光度计测定了植株和伤流液以及环境溶液中Na 含量。结果表明:随着根环境溶液NaCl浓度的增加,三角叶滨藜植株和木质部伤流液中Na 含量虽呈上升趋势,但根系的过滤系数和体内Na 相对累积量逐渐降低,说明三角叶滨藜根细胞对盐分有很强的过滤作用;木质部伤流液的渗透势随着环境溶液渗透势的降低而降低,但根木质部溶液的水势则逐渐高出根外环境溶液的渗透势;表明三角叶滨藜能够利用较低的木质部负压来抵抗根外溶液的低渗透势而反渗透吸水,并利用根细胞对盐分的过滤作用来避免从环境摄取过量的盐分。     

Root-To-Shoot Signalling: Assessing The Roles of ‘Up’ In the Up and Down World of Long-Distance Signalling In Planta

An important mediator of shoot physiological processes can be the supply of signal molecules (other than water and nutrients) from the root system. Root-to-shoot signalling is often considered to be important in regulating shoot growth and water use when soil conditions change without any demonstrable change in shoot water or nutrient status. Changes in xylem sap composition are often thought to be synonymous with changes in root-to-shoot signalling, even though there is considerable re-cycling of compounds between xylem and phloem. Techniques used to collect xylem sap are reviewed. Elucidating the roles of putative root signal molecules in planta has usually taken priority over identifying the sources of signal molecules in xylem sap. The roles of several signal molecules are considered. This choice is selective, and the failure of known signals to account for observed physiological changes in some systems has lead to the conclusions that other novel signals can be important. The efficacy of a given signal molecule can depend on the shoot water and nutrient status, as demonstrated by variation in stomatal responses to abscisic acid. If such variation is widespread in crop species, this may have implications for the increasing intentional use of root-to-shoot signals to modify crop water use and shoot architecture. Research into root-to-shoot signalling may become increasingly reductionist, in trying to evaluate the contribution of root signals versus local processes to observed physiological changes. However, future challenges are to successfully integrate this basic research into improved crop production systems.     

Dynamics of concentrations and nutrient fluxes in the xylem of Ricinus communis– diurnal course, impact of nutrient availability and nutrient uptake

Diurnal courses of nutrient transport in the xylem and their response to external availability of nutrients were studied. In soil culture, maximal concentrations in all analysed substances were observed during night‐time. Over experimental periods of up to 20 d, concentrations of some ions increased, most by accumulation in the soil. Stringent nutrient conditions were established in a novel pressure chamber. An aeroponic nutrient delivery system inside allows the sampling of xylem sap from intact plants under full control of the nutrient conditions at the root. Analysis of xylem transport under these highly defined conditions established that (1) diurnal variations in concentrations and fluxes in the xylem are dominated by plant‐internal processes; (2) concentrations of nutrients in the xylem sap are highly but specifically correlated with each other; (3) nitrate uptake and nitrate flux to the shoot are largely uncoupled; and (4) in continuous light, diurnal variations of xylem sap concentrations vanish. Step changes in nitrate concentrations of the nutrient solution established that (5) the concomitant increase in nitrate concentration and flux in the xylem is delayed by 2–3 h and is only transient. Diurnal variations of xylem sap composition and use of the new technique to elucidate xylem‐transport mechanisms are discussed.     

Hormonal Control of the Shoot-to-Root Ratio Is Not Related to Water Deficiency in Wheat Plants under Mineral Deficiency

We measured the content of hormones, the rate of growth, and some parameters of water regime (water content, transpiration, and stomatal and hydraulic conductivities) one and two days after wheat plant transfer from 10 to 1% Hoagland-Arnon nutrient medium. It was shown that, a day after dilution of nutrient solution, the content of various cytokinin forms decreased in the xylem sap, shoots, and roots. This decrease was most pronounced in the case of zeatin in the xylem sap and zeatin riboside in the mature zone of the first leaf. ABA was found to accumulate in shoots. A day after dilution of nutrient solution, we observed root elongation evidently induced by mineral nutrient deficiency, and this accelerated root growth was maintained later. Two days after dilution of nutrient solution, we observed the slowing of shoot weight accumulation, whereas root weight remained unchanged. Plant growth response could be related to ABA accumulation in shoots and cytokinin depletion in the whole plant. A reduced hydraulic conductivity and water content in the growing leaf zone was detected only two days after dilution of nutrient solution. Thus, changes in the growth rates and hormone contents could not result from disturbances in water regime induced by mineral nutrient deficiency.     

Two-component elements mediate interactions between cytokinin and salicylic acid in plant immunity

Recent studies have revealed an important role for hormones in plant immunity. We are now beginning to understand the contribution of crosstalk among different hormone signaling networks to the outcome of plant-pathogen interactions. Cytokinins are plant hormones that regulate development and responses to the environment. Cytokinin signaling involves a phosphorelay circuitry similar to two-component systems used by bacteria and fungi to perceive and react to various environmental stimuli. In this study, we asked whether cytokinin and components of cytokinin signaling contribute to plant immunity. We demonstrate that cytokinin levels in Arabidopsis are important in determining the amplitude of immune responses, ultimately influencing the outcome of plant-pathogen interactions. We show that high concentrations of cytokinin lead to increased defense responses to a virulent oomycete pathogen, through a process that is dependent on salicylic acid (SA) accumulation and activation of defense gene expression. Surprisingly, treatment with lower concentrations of cytokinin results in increased susceptibility. These functions for cytokinin in plant immunity require a host phosphorelay system and are mediated in part by type-A response regulators, which act as negative regulators of basal and pathogen-induced SA-dependent gene expression. Our results support a model in which cytokinin up-regulates plant immunity via an elevation of SA-dependent defense responses and in which SA in turn feedback-inhibits cytokinin signaling. The crosstalk between cytokinin and SA signaling networks may help plants fine-tune defense responses against pathogens.     

Halophytes as a source of genes for abiotic stress tolerance

Agricultural productivity is majorly impacted due to various abiotic stresses, particularly salinity and drought. Halophytes serve as an excellent resource for identifying and developing new crop systems, as these grow very luxuriously in very high saline soils. Understanding salinity stress tolerance mechanisms in such plants is an important step towards generating crop varieties that can cope with environmental stresses. Use of modern tools of ‘omics’ analyses and small RNA sequencing has helped to gain insights into the complex plant stress responses. Salinity tolerance being a multigenic trait requires a combination of strategies and techniques to successfully develop improved crops varieties. Many transgenic crops are being developed through genetic transformation. Besides marker-assisted breeding/QTL approaches are also being used to improve abiotic stress tolerance. In this review, we focus on the recent developments in the utilization of halophytes as a source of genes for genetic improvement in abiotic stress tolerance of crops.     

植物对逆境交叉适应的分子机制

交叉适应(cross-adaptation)是植物应答复合逆境的主要表现形式,它涉及环境刺激、信号转导、基因表达及细胞代谢调节等.为明确交叉适应的分子机制,本文从活性氧、激素、促细胞分裂原激活性蛋白激酶等方面进行了综述,以深入系统地阐明植物对逆境的响应,为作物抗逆栽培提供经济、高效的途径.     

Seasonal dynamics of nutrients in leaves and xylem sap of coffee plants as related to different soil compartments

The northwestern province of Costa Rica is a marginal coffee growing area. At the onset of the rainy season low redox potentials probably induce the mobilization of soil Mn resulting in enhanced plant uptake of Mn. To test this hypothesis we monitored from April to the end of June 1995 the mobile Mn in the soil and nutrient and Mn concentrations in leaves and xylem sap of coffee plants. Every 2 weeks we took aggregate and bulk soil samples. The aggregates were mechanically separated into interior and exterior, air-dried and all soil samples were extracted with 1 M NH₄NO₃. We also extracted the field moist soil with distilled water. In addition, the 3rd and the youngest pair of coffee leaves and xylem sap were sampled and analyzed. According to the results of leaf analyses the nutrient supply of the coffee plants in general seemed to be balanced. However, Mn concentrations of 223 mg kg^-1 in the 3rd leaf pair at 18 April were above the optimum and the youngest leaves indicated Fe deficiency, but senescent leaves accumulated Fe and overcame the deficiency. Manganese concentrations in the xylem sap showed a pronounced maximum 2 weeks prior to a similar maximum of mobile Mn in the aggregate exterior. But in general the temporal variation of nutrient concentrations (especially Ca and Mg) in the plants are well correlated with the easily extractable nutrient concentrations in bulk soil. Probably due to its specific absorption and high rates of redistribution within the plant, K in the soil extracts did not correlate with plant concentrations. Element concentrations of youngest leaves could not be correlated with soil concentrations and are not considered to be an adequate tool for monitoring current nutrient uptake. Since plant element concentrations did not correlate with the aggregate interior, plants probably cannot use that nutrient source efficiently.     

Intact plant MRI for the study of cell water relations, membrane permeability, cell-to-cell and long distance water transport

Water content and hydraulic conductivity, including transport within cells, over membranes, cell-to-cell, and long-distance xylem and phloem transport, are strongly affected by plant water stress. By being able to measure these transport processes non-invasely in the intact plant situation in relation to the plant (cell) water balance, it will be possible explicitly or implicitly to examine many aspects of plant function, plant performance, and stress responses. Nuclear magnetic resonance imaging (MRI) techniques are now available that allow studying plant hydraulics on different length scales within intact plants. The information within MRI images can be manipulated in such a way that cell compartment size, water membrane permeability, water cell-to-cell transport, and xylem and phloem flow hydraulics are obtained in addition to anatomical information. These techniques are non-destructive and non-invasive and can be used to study the dynamics of plant water relations and water transport, for example, as a function of environmental (stress) conditions. An overview of NMR and MRI methods to measure such information is presented and hardware solutions for minimal invasive intact plant MRI are discussed.     

Over-accumulation of abscisic acid in transgenic tomato plants increases the risk of hydraulic failure

Climate change threatens food security, and plant science researchers have investigated methods of sustaining crop yield under drought. One approach has been to overproduce abscisic acid (ABA) to enhance water use efficiency. However, the concomitant effects of ABA overproduction on plant vascular system functioning are critical as it influences vulnerability to xylem hydraulic failure. We investigated these effects by comparing physiological and hydraulic responses to water deficit between a tomato (Solanum lycopersicum) wild type control (WT) and a transgenic line overproducing ABA (sp12). Under well-watered conditions, the sp12 line displayed similar growth rate and greater water use efficiency by operating at lower maximum stomatal conductance. X-ray microtomography revealed that sp12 was significantly more vulnerable to xylem embolism, resulting in a reduced hydraulic safety margin. We also observed a significant ontogenic effect on vulnerability to xylem embolism for both WT and sp12. This study demonstrates that the greater water use efficiency in the tomato ABA overproducing line is associated with higher vulnerability of the vascular system to embolism and a higher risk of hydraulic failure. Integrating hydraulic traits into breeding programmes represents a critical step for effectively managing a crop's ability to maintain hydraulic conductivity and productivity under water deficit.     

Ionic control of the lateral exchange of water between vascular bundles in tomato

Ions can enhance water flow through the xylem via changes in the hydraulic resistance at border pit membranes. Because flow between adjacent xylem vessels occurs primarily via bordered pit fields, it is hypothesized that xylem sap ion concentrations would affect lateral movement of water more than longitudinal flow. Using tomato as a model system, evidence is presented for ion-mediated changes in xylem hydraulic resistance and the lateral transport of water. Water flow between adjacent xylem bundles increased by approximately 50% in the presence of ions while longitudinal flow only increased by approximately 20%. However, the enhancement of lateral exchange due to ions was magnified by the presence of a pressure difference between vascular bundles. These results indicate that the degree of nutrient-sharing among sectors of a plant may depend on both nutrient concentration and the availability of water in the root zone.     

Hormonal Interactions and Stomatal Responses

Both environmental and hormonal factors and their interactions affect stomatal behavior. Methodologies for identifying hormonal interactions affecting stomatal function are reviewed. Although there is abundant evidence that abscisic acid (ABA) closes stomata, evidence that the other classical plant hormones (auxins, cytokinins, ethylene, gibberellins) in isolation alter stomatal response often comes from exogenous applications to detached epidermes and leaves, rather than correlation of endogenous concentrations with stomatal conductance (g_s). Evidence for hormonal interactions comes from isolated tissues with exogenous hormones supplied at nonphysiological concentrations, or from variation in stomatal response to xylem ABA concentration in planta. The roles of hormonal changes in causing stomatal closure following changes in soil environment are considered. Although soil drying induces multiple changes in xylem sap composition, analysis of stomatal responses suggests a dominant role for increased endogenous ABA concentrations and relatively little evidence of roles for other hormones. A similar picture emerges from studies of soil compaction. Although soil flooding decreases ABA export from the root system, there is some evidence that apoplastic ABA accumulation elicits stomatal closure. Stomatal closure following nitrogen deprivation does not appear to involve ABA and may provide a suitable experimental system to investigate roles for other hormones. The availability of mutant or transgenic lines with altered hormone homeostasis or sensitivity provides opportunities to screen for altered stomatal behavior in response to different environments, and may provide new evidence that hormonal interactions are important in the control of stomatal behavior.     

植物抗旱性中的补偿效应及其在农业节水中的应用

在论述植物补偿效应存在类型和研究范畴的基础上,详细评述了植物抗旱性中根系形态结构功能及地上部干物质积累、产量和水分利用效率方面的补偿效应及其影响因素,并对植物抗旱作用中补偿生长的可能生理学机制作了探讨。同时,对补偿效应在提高农业水分利用效率中的应用进行了讨论     

Dynamic root growth and architecture responses to limiting nutrient availability: linking physiological models and experimentation

In recent years the study of root phenotypic plasticity in response to sub-optimal environmental factors and the genetic control of these responses have received renewed attention. As a path to increased productivity, in particular for low fertility soils, several applied research projects worldwide target the improvement of crop root traits both in plant breeding and biotechnology contexts. To assist these tasks and address the challenge of optimizing root growth and architecture for enhanced mineral resource use, the development of realistic simulation models is of great importance. We review this research field from a modeling perspective focusing particularly on nutrient acquisition strategies for crop production on low nitrogen and low phosphorous soils. Soil heterogeneity and the dynamics of nutrient availability in the soil pose a challenging environment in which plants have to forage efficiently for nutrients in order to maintain their internal nutrient homeostasis throughout their life cycle. Mathematical models assist in understanding plant growth strategies and associated root phenes that have potential to be tested and introduced in physiological breeding programs. At the same time, we stress that it is necessary to carefully consider model assumptions and development from a whole plant-resource allocation perspective and to introduce or refine modules simulating explicitly root growth and architecture dynamics through ontogeny with reference to key factors that constrain root growth. In this view it is important to understand negative feedbacks such as plant–plant competition. We conclude by briefly touching on available and developing technologies for quantitative root phenotyping from lab to field, from quantification of partial root profiles in the field to 3D reconstruction of whole root systems. Finally, we discuss how these approaches can and should be tightly linked to modeling to explore the root phenome.     

Different cation stresses affect specifically osmotic root hydraulic conductance, involving aquaporins, ATPase and xylem loading of ions in Capsicum annuum, L. plants

In order to study the effect of nutrient stress on water uptake in pepper plants (Capsicum annuum L.), the excess or deficiency of the main cations involved in plant nutrition (K(+), Mg(2+), Ca(2+)) and two different degrees of salinity were related to the activity of plasma membrane H(+)-ATPase, the pH of the xylem sap, nutrient flux into the xylem (J(s)) and to a number of parameters related to water relations, such as root hydraulic conductance (L(0)), stomatal conductance (g(s)) and aquaporin activity. Excess of K(+), Ca(+) and NaCl produced a toxic effect on L(0) while Mg(2+) starvation produced a positive effect, which was in agreement with aquaporin functionality, but not with ATPase activity. The xylem pH was altered only by Ca treatments. The results obtained with each treatment could suggest that detection of the quality of the nutrient supply being received by roots can be related to aquaporins functionality, but also that each cation stress triggers specific responses that have to be assessed individually.     

Contrasting Effects of GA3 Treatments on Tomato Plants Exposed to Increasing Salinity

The role of plant hormones under saline stress is critical in modulating physiological responses that will eventually lead to adaptation to an unfavorable environment. Nevertheless, the functional level of plant hormones, and their relative tissue concentration, may have a different impact on plant growth and stress tolerance at increasing salinity of the root environment. Vigorous plant growth may counteract the negative effects of salinization. In contrast, low gibberellin (GA) levels have been associated with reduced growth in response to salinity. Based on these facts and considering that the physiological basis of the cause-effect relationship between functional growth control and stress adaptation/survival is still a matter of debate, we hypothesized that exogenous applications of the plant hormone GA₃ may compensate for the salt-induced growth deficiency and consequently facilitate tomato plant adaptation to a saline environment. GA₃ application (0 or 100 mg GA₃ l⁻¹) was compared under four salinity levels, obtained by adding equal increments of NaCl:CaCl₂ (2:1 molar basis) (EC = 2.5, 6.8, 11.7, 16.7 dS m⁻¹) to the nutrient solution. GA₃ treatment reduced stomatal resistance and enhanced plant water use at low salinity. These responses were associated with an increased number of fruit per plant at harvest. However, moderate and high salinity nullified these differences. The fruit carotenoid level was generally lower in GA₃-treated plants, indicating either an inhibitory effect of GA₃ treatment on carotenoid biosynthesis or a reduced perception of the stress environment by GA₃-treated tomato plants.     

Logistics of water and salt transport through the plant: structure and functioning of the xylem

The xylem is a long‐distance transport system that is unique to higher plants. It evolved into a very sophisticated plumbing system ensuring controlled loading/unloading of ions and water and their effective translocation to the required sinks. The focus of this overview will be the intrinsic inter‐relations between structural and functional features of the xylem. Taken together the xylem is designed to prevent cavitation (entry of air bubbles), induced by negative pressures under transpiration and to repair the cavitated vessels. Half‐bordered pits between xylem parenchyma cells and xylem vessels are on the one hand the gates to the vessels but on the other hand a serious ‘bottle‐neck’ for transport. Hence it becomes evident that special transport systems exist at the interface between the cells and vessels, which allow intensive fluxes of ions and water to and out of the xylem. The molecular identification and biophysical/biochemical characterization of these transporters has just started. Paradigms for the sophisticated mechanism of controlled xylem transport under changing environmental conditions are SKOR, a Shaker‐like channel involved in K⁺‐loading and SOS1, a Na⁺/H⁺ antiporter with a proposed dual function in Na⁺ transport. In view of the importance of plant water relations it is not surprising to find that water channels dominate the gate of access to xylem. Future studies will focus on the mechanism(s) that regulate water channels and ion transporters and on their physiological role in, for example, the repair of embolism. Clearly, progress in this specific field of research will greatly benefit from an integration of molecular and biophysical techniques aimed to understand ‘whole‐plant’ behaviour under the ever‐changing environmental conditions in the daily life of all plants.