Academic literature on the topic 'Phloem flow'

Abstract Xylem and phloem sap flows in an intact, young Japanese zelkova tree (Zelkova serrata (Thunb.) Makino) growing outdoors were measured using magnetic resonance imaging (MRI). Two propagator-based sequences were developed for q-space imaging: pulse field gradient (PFG) with spin echo (PFG-SE) and stimulated echo (PFG-STE), which were used for xylem and phloem flow measurements, respectively. The data evaluation methods were improved to image fast xylem flow and slow phloem flow. Measurements were taken every 2–3 h for several consecutive days in August 2016, and diurnal changes in xylem and phloem sap flows in a cross-section of the trunk were quantified at a resolution of 1 mm2. During the day, apparent xylem flow volume exhibited a typical diurnal pattern following a vapor pressure deficit. The velocity mapping of xylem sap flow across the trunk cross section revealed that the greatest flow volume was found in current-year earlywood that had differentiated in April–May. The combined xylem flow in the 1- and 2-year-old annual rings also contributed to one-third of total sap flow. In the phloem, downward sap flow did not exhibit diurnal changes. This novel application of MRI in visualization of xylem and phloem sap flow by MRI is a promising tool for in vivo study of water transport in mature trees.

According to the Münch hypothesis, solution flow through the phloem is driven by a hydrostatic pressure gradient. At the source, a high hydrostatic pressure is generated in the collection phloem by active loading of solutes, which causes a concomitant passive flow of water, generating a high turgor pressure. At the sink, solute unloading from the phloem keeps the turgor pressure low, generating a source-to-sink hydrostatic pressure gradient. Localised changes in loading and unloading of solutes along the length of the transport phloem can compensate for small, short-term changes in phloem loading at the source, and thus, maintain phloem flow to the sink tissue. We tested directly the hydrostatic pressure regulation of the sieve tube by relating changes in sieve tube hydrostatic pressure to changes in solute flow through the sieve tube. A sudden phloem blockage was induced (by localised chilling of a 1-cm length of stem tissue) while sieve-tube-sap osmotic pressure, sucrose concentration, hydrostatic pressure and flow of recent photosynthate were observed in vivo both upstream and downstream of the block. The results are discussed in relation to the Münch hypothesis of solution flow, sieve tube hydrostatic pressure regulation and the mechanism behind the cold-block phenomenon.

The phloem plays a central role in transporting resources and signalling molecules from fully expanded leaves to provide precursors for, and to direct development of, heterotrophic organs located throughout the plant body. We review recent advances in understanding mechanisms regulating loading and unloading of resources into, and from, the phloem network; highlight unresolved questions regarding the physiological significance of the vast array of proteins and RNAs found in phloem saps; and evaluate proposed structure/function relationships considered to account for bulk flow of sap, sustained at high rates and over long distances, through the transport phloem.

Neck shrivel is a quality disorder of European plum (Prunus × domestica L.). It has been suggested that backflow in the xylem (from fruit to tree) could contribute to the incidence of neck shrivel in plum. The objective was to quantify rates of xylem, phloem and of transpiration flow in developing plum fruit. Using linear variable displacement transducers, changes in fruit volume were recorded 1) in un-treated control fruit, 2) in fruit that had their pedicels steam-girdled (phloem interrupted, xylem still functional) and 3) in detached fruit, left in the canopy (xylem and phloem interrupted). Xylem flow rates were occasionally negative in the early hours after sunrise, indicating xylem sap backflow from fruit to tree. Later in the day, xylem flows were positive and generally higher in daytime and lower at night. Significant phloem flow occurred in daytime, but ceased after sunset. During stage II (but not during stage III), the rates of xylem flow and transpiration were variable and closely related to atmospheric vapor pressure deficit. The relative contribution of xylem inflow to total sap inflow averaged 79% during stage II, decreasing to 25% during stage III. In contrast, phloem sap inflow averaged 21% of total sap inflow during stage II, increasing to 75% in stage III. Our results indicate that xylem backflow occurs early in the day. However, xylem backflow rates are considered too low to significantly contribute to the incidence of neck shrivel.

A pressure chamber was used to increase suddenly the hydrostatic pressure in the upper shoot of a Phaseolus vulgaris plant while observing phloem transport of 11C-labelled photoassimilate. Phloem transport in the stem towards the chamber stopped immediately when pressure was applied and then recovered within about 5 min. If the pressure was then released, flow increased again. The results support the hypothesis that flow of photoassimilate in the stem phloem was driven by a hydrostatic pressure gradient.

Abstract In a series of experiments with Ricinus communis, abscisic acid (ABA) concentrations in tissues and transport saps, its de novo biosynthesis, long-distance transport, and metabolism (degradation) were affected by nutritional conditions, nitrogen (N) source, and nutrient limitation, or salt stress. In the present study these data were statistically re-evaluated, and new correlations presented that underpin the importance of this universal phytohormone. The biggest differences in ABA concentration were observed in xylem sap. N source had the strongest effect; however, nutrient limitation (particularly phosphorus limitation) and salt also had significant effects. ABA was found in greater concentration in phloem sap compared with xylem sap; however, the effect of treatment on ABA concentration in phloem was lower. In the leaves, ABA concentration was most variable compared with the other tissues. This variation was only affected by the N source. In roots, ABA was significantly decreased by nutrient limitation. Of the compartments in which ABA was quantified, xylem sap ABA concentration was most significantly correlated with leaf stomatal conductance and leaf growth. Additionally, ABA concentration in xylem was significantly correlated to that in phloem, indicating a 6-fold concentration increase from xylem to phloem. The ABA flow model showed that biosynthesis of ABA in roots affected the xylem flow of ABA. Moreover, ABA concentration in xylem affected the degradation of the phytohormone in shoots and also its export from shoots via phloem. The role of phloem transport is discussed since it stimulates ABA metabolism in roots.

According to the Münch hypothesis, a flow of solution through the sieve tubes is driven by a hydrostatic pressure difference between the source (or collection) phloem and the sink (or release) phloem. A high hydrostatic pressure is maintained in the collection phloem by the active uptake of sugar and other solutes, with a concomitant inflow of water. A lower pressure is maintained in the release phloem through solute unloading. In this work we directly test the role of solute uptake in creating the hydrostatic pressure associated with phloem flow. Solute loading into the phloem of mature leaves of barley and sow thistle was reduced by replacing the air supply with nitrogen gas. Hydrostatic pressure in adjacent sieve elements was measured with a sieve-element pressure probe, a cell pressure probe glued to the exuding stylet of aphids that had been feeding from the phloem. Sieve element sap was sampled by aphid stylectomy; sap osmotic pressure was determined by picolitre osmometry and its sugar concentration by enzyme-linked fluorescence assays. Samples were taken with a time resolution of ~2–3 min. In accordance with Münch’s proposal a drop in osmotic and hydrostatic pressure in the source phloem following treatment of the source leaf with N2 was observed. A decrease in sugar concentration was the major contributor to the change in osmotic pressure. By observing these variables at a time resolution of minutes we have direct observation of the predictions of Münch.

Most growth and storage organs (sinks) of higher plants import assimilates in solution by bulk flow through the phloem, driven by differences in hydrostatic pressure. These differences in pressure, located between the ends of the interconnecting phloem path, are generated by osmotic water movement, driven in turn by membrane transport of solutes. Sucrose, amino-nitrogen compounds and potassium represent the osmotically important solutes found in phloem contents of most species. Phloem loading and unloading events of these assimilate species play central roles in determining phloem translocation rates and partitioning of assimilates and water. Depending on plant species, leaf vein order and sink type, phloem loading and unloading may follow apoplasmic or symplasmic routes. Irrespective of the cellular pathway followed, assimilates are transported across plasma and organellar membranes. Aquaporins, amino-nitrogen transporters, sucrose transporters and potassium channels have been detected in key sites along the source–phloem–sink transport pathway. Reverse genetics has demonstrated that sucrose/proton symporters are important in transport events necessary for phloem loading in Solanaceousplant species. Drawing on circumstantial evidence, we review possible functions the remaining membrane transporters and channels may serve in driving phloem translocation of assimilates and water from source to sink.

Vascular plants transport energy in the form of sugars from the leaves where they are produced to sites of active growth. The mass flow of sugars through the phloem vascular system is determined by the sap flow rate and the sugar concentration. If the concentration is low, little energy is transferred from source to sink. If it is too high, sap viscosity impedes flow. An interesting question is therefore at which concentration is the sugar flow optimal. Optimization of sugar flow and transport efficiency predicts optimal concentrations of 23.5 per cent (if the pressure differential driving the flow is independent of concentration) and 34.5 per cent (if the pressure is proportional to concentration). Data from more than 50 experiments (41 species) collected from the literature show an average concentration in the range from 18.2 per cent (all species) to 21.1 per cent (active loaders), suggesting that the phloem vasculature is optimized for efficient transport at constant pressure and that active phloem loading may have developed to increase transport efficiency.

The ambition of the work presented in this thesis is to address the need for nondestructive, repeatable measurements of long-distance transport of photosynthates through the phloem vascular tissue of woody plants. Quantification of long-distance phloem transport is believed to be able to provide information relevant to the forest carbon cycle as well as to a plant's response to changes in its environment. However, due to the fragility of the tissue, invasive techniques such as used routinely for the monitoring of water flow through the xylem are not applicable. Consequently, very little field data on phloem transport rates and patterns are currently available.

Thesis (PhD (Biochemistry))--University of Stellenbosch, 2009.
ENGLISH ABSTRACT: The sugarcane stalk, besides being the main structural component of the plant, is also the major storage organ for carbohydrates. Sucrose forms the bulk of stored carbohydrates. Previous studies have modelled the sucrose accumulation pathway in the internodal storage parenchyma of sugarcane using kinetic models cast as systems of ordinary differential equations. Typically, results were analysed with methods such as metabolic control analysis. The present study extends those original models within an advection-diffusion-reaction framework, requiring the use of partial differential equations to model sucrose metabolism coupled to phloem translocation. Let N be a stoichiometric matrix, v a vector of reaction rates, s a vector of species concentrations and r the gradient operator. Consider a coupled network of chemical reactions where the species may be advected with velocities, U, or diffuse with coefficients, D, or both. We propose the use of the dynamic system, s + r (Us) + r (Drs) = Nv; for a kinetic model where species can exist in different compartments and can be transported over long distances in a fluid medium, or involved in chemical reactions, or both. Darcy’s law is used to model fluid flow and allows a simplified, phenomenological approach to be applied to translocation in the phloem. Similarly, generic reversible Hill equations are used to model biochemical reaction rates. These are also phenomenological equations, where all the parameters have operationally defined interpretations. Numerical solutions to this formulation are demonstrated with time-courses of two toy models. The first model uses a simple “linear” pathway definition to study the impact of the system geometry on the solutions. Although this is an elementary model, it is able to demonstrate the up-regulation of photosynthesis in response to a change in sink demand. The second model elaborates on the reaction pathway while keeping the same geometry definition as the first. This pathway is designed to be an abstracted model of sucrose metabolism. Finally, a realistic model of sucrose translocation, metabolism and accumulation is presented, spanning eight internodes and four compartments. Most of the parameters and species concentrations used as initial values were obtained from experimental measurements. To analyse the models, a method of sensitivity analysis called the Fourier Amplitude Sensitivity Test (FAST) is employed. FAST calculates the contribution of the possible variation in a parameter to the total variation in the output from the model, i.e. the species concentrations and reaction rates. The model predicted that the most important factors affecting sucrose accumulation are the synthesis and breakdown of sucrose in futile cycles and the rate of cross-membrane transport of sucrose. The models also showed that sucrose moves down a concentration gradient from the leaves to the symplast, where it is transported against a concentration gradient into the vacuole. There was a net gain in carbohydrate accumulation in the realistic model, despite an increase in futile cycling with internode maturity. The model presented provides a very comprehensive description of sucrose accumulation and is a rigorous, quantitative framework for future modelling and experimental design.
AFRIKAANSE OPSOMMING: Benewens sy strukturele belang, is die suikerrietstingel ook die primêre bergingsorgaan vir koolhidrate. Die oorgrote meerderheid van hierdie koolhidrate word as sukrose opgeberg. Studies tot dusver het die metabolisme rondom sukroseberging in die parenchiem van die onderskeie stingellitte as stelsels gewone differensiaalvergelykings gemodelleer. Die resultate is ondermeer met metaboliese kontrole-analise geanaliseer. Hierdie studie brei uit op die oorspronklike modelle, deur gebruik te maak van ’n stromings-diffusie-reaksie-raamwerk. Parsiële differensiaalvergelykings is geformuleer om die metabolisme van sukrose te koppel aan die vloei in die floëem. Gestel N is ’n stoichiometriese matriks, v ’n vektor van reaksiesnelhede, s ’n vektor van spesie-konsentrasies en r die differensiaalvektoroperator. Beskou ’n netwerk van gekoppelde reaksies waar die onderskeie spesies stroom met snelhede U, of diffundeer met koëffisiënte D, of onderhewig is aan beide prosesse. Dit word voorgestel dat die dinamiese stelsel, _s + r (Us) + r (Drs) = Nv; gebruik kan word vir ’n kinetiese model waar spesies in verskeie kompartemente kan voorkom en vervoer kan word oor lang afstande saam met ’n vloeier, of kan deelneem aan chemiese reaksies, of albei. Darcy se wet word gebruik om die vloeier te modeller en maak dit moontlik om ’n eenvoudige, fenomenologiese benadering toe te pas op floëem-vervoer. Eweneens word generiese, omkeerbare Hill-vergelykings gebruik om biochemiese reaksiesnelhede te modelleer. Hierdie vergelykings is ook fenomenologies van aard en beskik oor parameters met ’n duidelike fisiese betekenis. Hierdie omvattende raamwerk is ondermeer gedemonstreer met behulp van numeriese oplossings van twee vereenvoudigde modelle as voorbeelde. Die eerste model het bestaan uit ’n lineêre reaksienetwerk en is gebruik om die geometrie van die stelsel te bestudeer. Alhoewel hierdie ’n eenvoudige model is, kon dit die toename in fotosintese as gevolg van ’n verandering in metaboliese aanvraag verklaar. Die tweede model het uitgebrei op die reaksieskema van die eerste, terwyl dieselfde stelselgeometrie behou is. Hierdie skema is ontwerp as ’n abstrakte weergawe van sukrosemetabolisme. Ten slotte is ’n realistiese model van sukrosevervoer, metabolisme en berging ontwikkel wat agt stingellitte en vier kompartemente omvat. Die meeste parameters en konsentrasies van biochemiese spesies wat as aanvanklike waardes in die model gebruik is, is direk vanaf eksperimentele metings verkry. Die Fourier Amplitude Sensitiwiteits-Toets (FAST) is gebruik om die modelle te analiseer. FAST maak dit moontlik om die bydrae van parameters tot variasie in modeluitsette soos reaksiesnelhede en die konsentrasies van chemiese spesies te bepaal. Die model het voorspel dat sintese en afbraak van sukrose in ’n futiele siklus, asook transmembraan sukrosevervoer, die belangrikste faktore is wat sukrose-berging beïnvloed. Die model het ook getoon dat sukrose saam met ’n konsentrasiegradiënt beweeg vanaf die blare tot by die stingelparenchiem-sitoplasma, van waar dit teen ’n konsentrasiegradiënt na die vogselholte (vakuool) vervoer word. Volgens die realistiese model was daar ’n netto toename in die totale hoeveelheid koolhidrate, ten spyte van ’n toename in die futile siklus van sukrose in die ouer stingellitte. Die model wat in hierdie proefskrif voorgestel word verskaf ’n uitgebreide, omvattende beskrywing van sukroseberging. Voorts stel dit ’n rigiede kwantitatiewe raamwerk daar vir toekomstige modellering en eksperimentele ontwerp.

Potato is one of the most important agricultural crops and there is an attempt to increase and improve yields of tubers, among other things, by elucidation of the mechanisms that regulate the process of tuber induction. Potato tuberization is a morphogenetic process in which the tubers are formed from the underground parts of the stem - stolons. The correct timing of this process is controlled by a complex regulatory network and influenced by many internal and external factors. Under favourable conditions, an inductive signal is generated in the leaves and it is transported to the stolon by a "phloem information superhighway" driven by carbohydrates flow. The signal triggers cell division, expansion, and changes in the cell growth orientation in the stolon. The development of tubers is influenced by number of biochemical and morphological processes driven by a regulatory network of genes that are expressed in different parts of plants. This work was focused on Solanum tuberosum, Lada cultivar and its derived D69 mutant line with lacking isoform of manganese-stabilizing protein (MSP), which is so far the only dissimilarity identified under all tested conditions. I aimed to map the processes related to the production of carbohydrates in leaves (photosynthetic characteristics - rate of photosynthesis...

AbstractWe developed an imaging method utilizing the available RIs. We developed two types of real-time RI imaging systems (RRIS), one for macroscopic imaging and the other for microscopic imaging. The principle of visualization was the same, converting the radiation to light by a Cs(Tl)I scintillator deposited on a fiber optic plate (FOS). Many nuclides were employed, including 14C, 18F, 22Na, 28Mg, 32P 33P, 35S, 42K, 45Ca, 48V, 54Mn, 55Fe, 59Fe, 65Zn, 86Rb, 109Cd, and 137Cs.Since radiation can penetrate the soil as well as water, the difference between soil culture and water culture was visualized. 137Cs was hardly absorbed by rice roots growing in soil, whereas water culture showed high absorption, which could provide some reassurance after the Fukushima Nuclear Accident and could indicate an important role of soil in firmly adsorbing the radioactive cesium.28Mg and 42K, whose production methods were presented, were applied for RRIS to visualize the absorption image from the roots. In addition to 28Mg and 42K, many nuclides were applied to image absorption in the roots. Each element showed a specific absorption speed and accumulation pattern. The image analysis of the absorption of Mg is presented as an example. Through successive images of the element absorption, phloem flow in the aboveground part of the plant was analyzed. The element absorption was visualized not only in the roots but also in the leaves, a basic study of foliar fertilization.In the case of the microscopic imaging system, a fluorescence microscope was modified to acquire three images at the same time: a light image, fluorescent image, and radiation image. Although the resolution of the image was estimated to be approximately 50 μm, superposition showed the expression site of the transporter gene and the actual 32P-phosphate absorption site to be the same in Arabidopsis roots.

Plant transport mechanisms are of interest in developing micropump for engineering devices. We present a two-dimensional phloem loading and transport model incorporating protein level mechanics with cellular level fluid mechanics. Governing Navier-Stokes, continuity, and Nernst-Planck equations are numerically solved to determine fluid flow and sugar transport. Phloem loading mechanics for active loading is incorporated through a six-state proton sucrose pump kinetic model. The influence of binding rates constants, concentrations, and membrane electrical potential differences on resulting sucrose transport is studied. Numerical results show that increasing rates of the sucrose transporter will noticeably increase outflow. Simulation result also show that a lower leaf sieve sucrose concentration improves outflow. In addition, a more negative membrane electrical potential difference will increase outflow. This numerical model offers insight on parameters that may be significant for implementing plant transport mechanisms in microfluidic devices.