Rhizosphere Oxidation in Rice and other Species: A Mathematical Model Based on the Oxygen Flux Component

A mathematical model is presented from which one can predict the likely dimensions of oxidised rhizospheres due to oxygen diffusing from roots into anaerobic media such as wet soil. The analysis applies Fick's law of diffusion to the diffusion of oxygen from a cylindrical object (the root) into a sink (the soil) which is absorbing oxygen at a constant rate M. Solution of the final equation gives the dimensions of the oxygenated rhizosphere, i.e. the distance from the root at which the oxygen concentration becomes zero. The results obtained supoprt the view that oxygen diffusing from roots will produce a rhizosphere ‘sheath’ charged with oxygen. Furthermore, some of the predicted limits for this oxygen sheath correspond with those of ferric iron sheaths around roots in waterlogged soils. Calculations also show that oxygenated rhizospheres around the roots of plants very tolerant of reducing conditions. e.g. Menyanthes trifoliata, Oryza sativa and Eriophorum angustifolium, should be from two to three times as broad as for Molinia coerulea which is a species intolerant of strong reducing conditions. Consequently if oxidation reactions are not instantaneous but occur only gradually as quantities of reduced products enter the oxygenated zone, then the plants with the larger rhizospheres will be significantly better protected from absorbing large amounts of reduced products than will those plants with smaller oxygenated zones. This may in part explain the reason for intervarietal defferences in physiological disease resistance in rice. Finally the observation is made that small lateral roots can be expected to oxygenate a zone nearly as large as, or probably larger, than the primary root from which they have arisen.     

Metabolic acclimation of source and sink tissues to salinity stress in bermudagrass (Cynodon dactylon)

Salinity is one of the major environmental factors affecting plant growth and survival by modifying source and sink relationships at physiological and metabolic levels. Individual metabolite levels and/or ratios in sink and source tissues may reflect the complex interplay of metabolic activities in sink and source tissues at the whole‐plant level. We used a non‐targeted gas chromatography–mass spectrometry (GC‐MS) approach to study sink and source tissue‐specific metabolite levels and ratios from bermudagrass under salinity stress. Shoot growth rate decreased while root growth rate increased which lead to an increased root/shoot growth rate ratio under salt stress. A clear shift in soluble sugars (sucrose, glucose and fructose) and metabolites linked to nitrogen metabolism (glutamate, aspartate and asparagine) in favor of sink roots was observed, when compared with sink and source leaves. The higher shifts in soluble sugars and metabolites linked to nitrogen metabolism in favor of sink roots may contribute to the root sink strength maintenance that facilitated the recovery of the functional equilibrium between shoot and root, allowing the roots to increase competitive ability for below‐ground resource capture. This trait could be considered in breeding programs for increasing salt tolerance, which would help maintain root functioning (i.e. water and nutrient absorption, Na⁺ exclusion) and adaptation to stress.     

Microsensor Analysis of Oxygen in the Rhizosphere of the Aquatic Macrophyte Littorella uniflora (L.) Ascherson

Oxygen released by the roots of submerged plants may oxidize organic compounds from the roots and reduced substances continuously supplied by diffusion from the surrounding anoxic hydrosoil. We provide here the first visualization of this gradient environment obtained by microsensor analysis of oxygen in the rhizosphere of the freshwater plant Littorella uniflora (L.) Ascherson. The plants were rooted in an agar medium, in which amorphous FeS provided the main oxygen sink. The oxygen concentration at the root surface ranged from 20 to 450 [mu]M (atmospheric saturation = 280 [mu]M) between darkness and saturating light, and the oxic shell surrounding the roots varied from about 0.5 to 5 mm in thickness. The oxygen flux from the roots was a saturating function of the incident light intensity on the leaves, and the oxygen released was consumed mainly at the fluctuating oxic/anoxic interface. The oxic zones around individual roots are under dynamic control by light, root morphology, root density, and sediment reducing capacity, and, therefore, oxygen concentrations should be subject to substantial diurnal fluctuations in dense Littorella populations in nutrient-poor sediments.     

The developmental and organ specific expression of sucrose cleaving enzymes in sugar beet suggests a transition between apoplasmic and symplasmic phloem unloading in the tap roots

Sucrose utilisation in sink tissues depend on its cleavage and is mediated by two different classes of enzymes, invertase and sucrose synthase, which determine the mechanism of phloem unloading. Cloning of two extracellular (BIN35 and BIN46) and one vacuolar invertase (BIN44) provided the basis for a detailed molecular analysis of the relative contribution of the sucrose cleaving enzymes to the sink metabolism of sugar beets (Beta vulgaris) during development. The determination of the steady state levels of mRNAs has been complemented by the analysis of the corresponding enzyme activities. The present study demonstrates an inverse regulation of extracellular invertase and sucrose synthase during tap root development indicating a transition between functional unloading pathways. Extracellular cleavage by invertase is the dominating mechanism to supply hexoses via an apoplasmic pathway at early stages of storage root development. Only at later stages sucrose synthase takes over the function of the key sink enzyme to contribute to the sink strength of the tap root via symplasmic phloem unloading. Whereas mRNAs for both extracellular invertase BIN35 and sucrose synthase were shown to be induced by mechanical wounding of mature leaves of adult plants, only sucrose synthase mRNA was metabolically induced by glucose in this source organ supporting the metabolic flexibility of this species.     

Intervention de l'oxygène atmosphériqoe sur l'absorption d'eau chez Helianthus annuus

The role of atmospheric oxygen on root water absorption in Helianthus annuus . The effect of atmospheric anoxia on root water absorption was studied. The experiments were carried out on intact young sunflowers in controlled temperature, light and gas environment; roots were kept in aerated nutrient solution at constant temperature. The evolution of root water absorption and transpiration rate was measured continuously. Before the experiment, the plant was preconditioned at a high transpiration rate by illumination or by CO₂ free air in darkness. Then the atmospheric oxygen was suppressed for 1 h, after which the normal conditions were restored.
In anoxia and darkness, the root water absorption cannot balance transpiration, so that an important water stress develops in the plant; the light compensates this effect through the photosynthetic oxygen. The supply of oxygen, in darkness or in light, immediately removes inhibition of stomatal closure and of root water absorption. Two mechanisms control water absorption by roots: the fast one occurs in the leaves and the slower one cannot develop without the root system.     

STUDIES ON PHOTOSYNTHESIS : SOME EFFECTS OF LIGHT OF HIGH INTENSITY ON CHLORELLA

1. The effect on oxygen evolution of Chlorella vulgaris produced by light intensities up to about 40,000 f.-c. has been studied by the use of the Warburg technique. 2. Above a certain critical intensity, which is determined by the previous history of the cells, the rate of oxygen evolution decreases from the maximum to another constant rate. This depression is at first a completely reversible effect. 3. With a sufficiently high intensity this constant rate represents an oxygen uptake greater than the rate of dark respiration. During such a constant rate of oxygen uptake a progressive injury to the photosynthetic mechanism takes place. After a given oxygen consumption the rate falls off, approaching zero, and the cells are irreversibly injured. 4. The constant rate of oxygen evolution (2 and 3) decreases in a continuous manner with increasing light intensities, approaching a value which is approximately constant for all lots of cells regardless of previous history. 5. Two alternative hypotheses have been presented to explain the observed phenomena. The more acceptable of these proposes quick inactivation of the photosynthetic mechanism, the extent of inhibition depending on the light intensity. 6. In Chlorella vulgaris solarization is influenced by the previous history of the cells.     

Plant sulphate transporters: co-ordination of uptake, intracellular and long-distance transport

Proton/sulphate co-transport in the plasma membrane of root cells is the first step for the uptake of sulphate from the environment by plants. Further intracellular, cell-to-cell and long-distance transport must fulfil the requirements for sulphate assimilation and source/sink demands within the plant. A gene family of sulphate transporters, which may be subdivided into five groups, has been identified with examples from many different plant species. For at least two groups, proton/sulphate co-transport activity has been confirmed. It appears that each group represents sulphate transporters with distinct kinetic properties, patterns of expression, and cell/tissue specificity related to specific roles in the uptake and allocation of sulphate. High-affinity sulphate uptake and low-affinity vascular transport, as well as vacuolar efflux, are controlled by the nutritional status of the plant. Most notably there is an apparent increase in capacity for cellular sulphate uptake and vacuolar efflux when sulphur supply is limiting. Within the groups, the individual sulphate transporters may be further subdivided by differences in temporal, cellular and tissue expression. Many of the transporters are regulated by the nutritional status of the individual tissues, to optimize sulphate movement within and between sink and source organs.     

Adventitious rooting in hypocotyls of sunflower (Helianthus annuus) seedlings

Removal of the main root system of sunflower ( Helianthus annuus ) initiates adventitious root development on the lower portion of the hypocotyl. The first cytological changes (enlarged nuclei in the interfascicular parenchymatous cells adjacent to the phloem and some cell divisions) are observed 24 h after root excision. On the basis of experiments in which (a) roots, apical buds and various amounts of cotyledonary tissue were removed, (b) cuttings were subjected to various light regimes, (c) benzyladenine oas applied to cotyledons to create an artificial sink, it was concluded that the roots normally produce factors inhibiting to adventitious rooting and might be a sink for stimulatory substances produced in the shoots. The cotyledons seem to be the major source of these stimulators. Application of aqueous and ethanolic extracts of cotyledons and hypocotyls to cuttings promoted adventitious rooting.     

Comparing different approaches to calculate the effects of heterogeneous root distribution on nutrient uptake: a case study on subsoil nitrate uptake by a barley root system

Simulation models of nutrient uptake of root systems starting with one-dimensional single root approaches up to complex three-dimensional models are increasingly used for examining the interacting of root distribution and nutrient uptake. However, their accuracy was seldom systematically tested. The objective of the study is to compare one-dimensional and two-dimensional modelling approaches and to test their applicability for simulation of nutrient uptake of heterogeneously distributed root systems giving particular attention to the impact of spatial resolution. Therefore, a field experiment was carried out with spring barley (Hordeum vulgare L. cv. Barke) in order to obtain data of in situ root distribution patterns as model input. Results indicate that a comparable coarse spatial resolution can be used with sufficient modelling results when a steady state approximation is applied to the sink cells of the two-dimensional model. Furthermore, the accuracy of the model was clearly improved compared to a simple zero sink approach assuming both near zero concentrations within the sink cell and a linear gradient between the sink cell and its adjacent neighbours. However, for modelling nitrate uptake of a heterogeneous root system a minimum number of grid cells is still necessary. The tested single root approach provided a computational efficient opportunity to simulate nitrate uptake of an irregular distributed root system. Nevertheless, two-dimensional models are better suited for a number of applications (e.g. surveys made on the impact of soil heterogeneity on plant nutrient uptake). Different settings for the suggested modelling techniques are discussed.     

Autoregulation of root nodule formation: signals of both symbiotic partners studied in a split-root system of Vicia sativa subsp. nigra

Inhibition of root nodule formation on leguminous plants by already induced or existing root nodules is called autoregulation of root nodule formation (AUT). Optimal conditions for AUT were determined using a split-root technique newly developed for Vicia sativa subsp. nigra. Infection of a root A with nodulating Rhizobium leguminosarum bv. viciae bacteria systemically inhibited nodulation of a spatially separated root B inoculated 2 days later with the same bacteria. This treatment gives complete AUT (total absence of nodules on root B). Only partial AUT of root B was obtained by incubation of root A with mitogenic nodulation (Nod) factors or with a noninfective strain producing normal mitogenic Nod factors. Nonmitogenic Nod factors did not evoke AUT. We identified two systemic plant signals induced by Rhizobium bacteria. Signal 1 (at weak buffering) was correlated with sink formation in root A and induced acidification of B-root medium. This signal is induced by treatment of root A with (i) nodulating rhizobia, (ii) mitogenic Nod factors, (iii) nonmitogenic Nod factors, or (iv) the cytokinin zeatin. Signal 2 (at strong buffering) could only be evoked by treatment with nodulating rhizobia or with mitogenic Nod factors. Most probably, this signal represents the specific AUT signal. Induction of complete AUT appears to require actively dividing nodule cells in nodule primordia, nodule meristems, or both of root A.     

Hypocotyl of seedlings of the large-seeded species Araucaria angustifolia: an important underground sink of the seed reserves

Araucaria angustifolia exhibits cryptogeal germination, where the root–hypocotyl axis emerges first and penetrates into the soil. In Araucaria bidwillii, the whole process of transferring reserves from the seed to the seedling takes place before shoot emergence, and there is a major storage of these reserves in the underground hypocotyl, which assumes a tuberous form. In A. angustifolia, the shoot emerges before seed reserves are depleted. Though it does not grow like a tuber, the hypocotyl of A. angustifolia grows thicker than the adjacent taproot during initial growth, and we hypothesize that it may act as a major sink for seed reserves during this stage. The study tests this hypothesis by evaluating changes in the mass of different plant parts during initial growth. Four harvests were conducted during a ~6-month period to compare the dry mass of different fractions (attached seed, seedling, its shoot and root and the hypocotyl) of seedlings growing under darkness and high light. While seed reserves were still being depleted, the hypocotyl mass showed an initial increase and then a reduction. This was more abrupt when light was available. After seed mass had stabilized, the mass of the hypocotyl continued to decrease in the dark-grown seedlings, but showed a second increase in the light-grown ones. Results confirm the hypothesis that the hypocotyl represents a major sink for the seed reserves of A. angustifolia, acting as an underground storage structure for the growing seedling. Its reserves seem to be important for sustaining initial shoot growth and might also act as a storage sink for photosynthates.     

On the mechanism of mitochondrial permeability transition induction by glycyrrhetinic acid

Glycyrrhetinic acid (GE), the aglycone of glycyrrhizic acid, a triterpene glycoside which represents one of the main constituents of licorice root, induces an oxidative stress in liver mitochondria responsible for the induction of membrane permeability transition. In fact, GE, by interacting with the mitochondrial respiratory chain, generates hydrogen peroxide which in turn oxidizes critical thiol groups and endogenous pyridine nucleotides leading to the opening of the transition pore. Most likely the reactive group of GE is the carbonyl oxygen in C-11 which, by interacting mainly with a Fe/S centre of mitochondrial complex I, generates an oxygen-centered radical responsible for the pro-oxidant action.     

On the mechanism of mitochondrial permeability transition induction by glycyrrhetinic acid

Glycyrrhetinic acid (GE), the aglycone of glycyrrhizic acid, a triterpene glycoside which represents one of the main constituents of licorice root, induces an oxidative stress in liver mitochondria responsible for the induction of membrane permeability transition. In fact, GE, by interacting with the mitochondrial respiratory chain, generates hydrogen peroxide which in turn oxidizes critical thiol groups and endogenous pyridine nucleotides leading to the opening of the transition pore. Most likely the reactive group of GE is the carbonyl oxygen in C-11 which, by interacting mainly with a Fe/S centre of mitochondrial complex I, generates an oxygen-centered radical responsible for the pro-oxidant action.     

Distorted phytochrome action spectra in green plants

An evaluation was made of the extent which a Münch-type pressure flow mechanism (i.e., osmotically-generated pressure flow) might contribute to phloem transport in soybean. Estimates of sucrose concentrations in source (leaf) and sink (root) sieve tubes were obtained by a negativestaining procedure. Water potential measurements of the leaf and of the nutrient solution allowed calculation of the turgor pressures in source and sink sieve tubes. The turgor difference between source and sink sieve tubes was compared to that required to drive translocation at the observed velocity between the source and sink, as measured by [¹⁴C] photosynthate movement. Sieve-tube conductivity was calculated from the sieve-tube dimensions, assuming an essentially unobstructed pathway. In three experiments, the sucrose concentration was consistently higher in source sieve tubes (an average of 11.5%) than in sink sieve tubes (an average of 5.3%). The ratio of these values (2.3:1) agreed reasonably well with an earlier ratio for source/sink sieve tube concentrations of 1.8:1, obtained by quantitative microautoradiography. The resulting calculated turgor difference (an average of 4.1 bars) was adequate to drive a pressure flow mechanism at the observed translocation velocities (calculated to require a turgor difference of 1.2 to 4.6 bars). No other force need be presumed to be involved.This work was presented in part at a joint U.S.-Australian Conference on Transport and Transfer Processes in Plants, Canberra, Australia, December 15–20, 1975; see Fisher (1976)     

Effect of percolation rate on nutrient kinetics and rice yield in tropical rice soils

A model is presented with which the contribution of longitudinal oxygen diffusion to total oxygen requirement of a root can be estimated. Oxygen transport in and respiration of the soil are taken into account. Given the air-filled root porosity, root diameter, coefficient of oxygen transfer between root and soil, root and soil respiration rate, and the coefficient for oxygen diffusion in the soil, the maximum length a root can attain with an adequate oxygen supply to the root tip can be calculated. Results show the importance of root porosity for root aeration, also in unsaturated soils. For thick roots (radius >0.03 cm), diffusion along the internal pathway can provide 50–75% of the total oxygen requirement even, at modest values of the root porosity.     

Modelling the influence of assimilate availability on root growth and architecture

A model has been designed to simulate rubber seedling root development as related to assimilate availability. Each root of the system is defined both as an element of a network of axes, characterized by its order, position and connections and as an individual sink competing for assimilates. At each time step, the growth of each root is calculated as a function of its own growth potential and of assimilate availability calculated within the whole plant. The potential elongation rate of a root is estimated by its apical diameter, which reflects the size of the meristem. When a root is initiated, the apical diameter depends on root type, but it varies thereafter according to assimilate availability. Thus, the latter controls both current and potential elongation. The model was able to simulate periodicity in root development as related to shoot growth and to reproduce differences in sensitivity to assimilate availability related to root type. It thereby validated the hypothesis that root growth but also root system architecture depend on assimilate allocation and that apical diameter is a good indicator of root growth potential. Provided that specific calibration is done, this model may be used for other species.     

TheCaesalpinia peltophoroides cell cycle under different aeration conditions

Summary Cell cycle parameters were studied inCaesalpinia peltophoroides meristems proliferating under different oxygen tensions. This species has been selected for mixed planting in degraded areas in Brazil, some of which are occasionally flooded. As the species’ adaptation to oxygen deprivation during flooding is not fully understood, the objective of this study was to characterize the meristematic activity of root cells under various oxygen regimes. Synchronous binucleate cells, induced by a pulse of caffeine, showed a cell-cycle time constant under both control (5.6 mg of O₂ per l) and oxygenated conditions (7.9 and 3.2 mg of O₂ per l). The whole cell cycle lasted 10 h, although the relative duration of metaphase and anaphase/early telophase increased in more hypoxic conditions. The species appeared to utilise oxygen diffusing from the shoot to the root system to maintain cell division and root growth.     

Effects of source-sink relations on photosynthetic acclimation to elevated CO2

Abstract. While photosynthesis of C₃ plants is stimulated by an increase in the atmospheric CO₂ concentration, photosynthetic capacity is often reduced after long-term exposure to elevated CO₂. This reduction appears to be brought about by end product inhibition, resulting from an imbalance in the supply and demand of carbohydrates. A review of the literature revealed that the reduction of photosynthetic capacity in elevated CO₂ was most pronounced when the increased supply of carbohydrates was combined with small sink size. The volume of pots in which plants were grown affected the sink size by restricting root growth. While plants grown in small pots had a reduced photosynthetic capacity, plants grown in the field showed no reduction or an increase in this capacity. Pot volume also determined the effect of elevated CO₂ on the root/shoot ratio: the root/shoot ratio increased when root growth was not restricted and decreased in plants grown in small pots. The data presented in this paper suggest that plants growing in the field will maintain a high photosynthetic capacity as the atmospheric CO₂ level continues to rise.