ABSTRACT). We hypothesize that upper and lower bounds for phloem flow velocity may exist: when phloem flow velocity is too high, parietal organelles may be stripped away from sieve tube walls; when sap flow is too slow or is highly variable, phloem-borne signalling could become unpredictable.
Sieve elements are one of the least understood cell types in plants. Translocation velocities and volume flow to supply sinks with photoassimilates greatly depend on the geometry of the microfluidic sieve tube system and especially on the anatomy of sieve plates and sieve plate pores. Several models for phloem translocation have been developed, but appropriate data on the geometry of pores, plates, sieve elements, and flow parameters are lacking. We developed a method to clear cells from cytoplasmic constituents to image cell walls by scanning electron microscopy. This method allows high-resolution measurements of sieve element and sieve plate geometries. Sieve tube-specific conductivity and its reduction by callose deposition after injury was calculated for green bean (Phaseolus vulgaris), bamboo (Phyllostachys nuda), squash (Cucurbita maxima), castor bean (Ricinus communis), and tomato (Solanum lycopersicum). Phloem sap velocity measurements by magnetic resonance imaging velocimetry indicate that higher conductivity is not accompanied by a higher velocity. Studies on the temporal development of callose show that small sieve plate pores might be occluded by callose within minutes, but plants containing sieve tubes with large pores need additional mechanisms.
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.
(1)H NMR relaxation times (T(1) and T(2)) in parenchyma tissue of apple can identify three populations of water with different relaxation characteristics. By following the uptake of Mn(2+) ions in the tissue it is shown that the observed relaxation times originate from particular water compartments: the vacuole, the cytoplasm, and the cell wall/extracellular space.Proton exchange between these compartments is controlled by the plasmalemma and tonoplast membranes. During the Mn(2+) penetration experiment, conditions occur that cause the relaxation times of protons of cytoplasmic water to be much shorter than their residence time in the cytoplasm. Then the tonoplast permeability coefficient P(d) for water can be calculated from the vacuolar T(1) and T(2) values to be 2.44 10(-5) m.s(-1).
The flow field dynamics in open and packed segments of capillary columns has been studied by a direct motion encoding of the fluid molecules using pulsed magnetic field gradient nuclear magnetic resonance. This noninvasive method operates within a time window that allows a quantitative discrimination of electroosmotic against pressure-driven flow behavior. The inherent axial fluid flow field dispersion and characteristic length scales of either transport mode are addressed, and the results demonstrate a significant performance advantage of an electrokinetically driven mobile phase in both open-tubular and packed-bed geometries. In contrast to the parabolic velocity profile and its impact on axial dispersion characterizing laminar flow through an open cylindrical capillary, a pluglike velocity distribution of the electroosmotic flow field is revealed in capillary electrophoresis. Here, the variance of the radially averaged, axial displacement probability distributions is quantitatively explained by longitudinal molecular diffusion at the actual buffer temperature, while for Poiseuille flow, the preasymptotic regime to Taylor-Aris dispersion can be shown. Compared to creeping laminar flow through a packed bed, the increased efficiency observed in capillary electrochromatography is related to the superior characteristics of the electroosmotic flow profile over any length scale in the interstitial pore space and to the origin, spatial dimension, and hydrodynamics of the stagnant fluid on the support particles' external surface. Using the Knox equation to analyze the axial plate height data, an eddy dispersion term smaller by a factor of almost 2.5 than in capillary high-performance liquid chromatography is revealed for the electroosmotic flow field in the same column.
The role of stagnant zones in hydrodynamic dispersion is studied for creeping flow through a fixed bed of spherical permeable particles, covering several orders of characteristic time and length scales associated with fluid transport. Numerical simulations employ a hierarchical model to cope with the different temporal and spatial scales, showing good agreement with our experimental results on diffusionlimited mass transfer, transient, and asymptotic longitudinal dispersion. These data demonstrate that intraparticle liquid holdup in macroscopically homogeneous porous media clearly dominates over contributions caused by the intrinsic flow field heterogeneity and boundary-layer mass transfer. DOI: 10.1103/PhysRevLett.88.234501 PACS numbers: 47.15.Gf, 05.60. -k, 47.55.Mh A detailed understanding of transport in porous media over the intrinsic temporal and spatial scales is important in many technological and environmental processes [1]. For example, natural and industrial materials such as soil, rock, filter cakes, or catalyst pellets often contain lowpermeability zones with respect to hydraulic flow of liquid through the medium or even stagnant regions which then remain purely diffusive. The relevance of stagnant zones stems from their influence on dispersion: Fluid molecules entrained in the deep diffusive pools cause a substantial holdup contribution and thereby affect the time scale of transient dispersion, as well as the value of the asymptotic dispersion coefficient (if the asymptotic long-time limit can be reached at all) [2][3][4]. Consequently, the associated kinetics of mass transfer between fluid percolating through the medium and stagnant fluid becomes rate limiting in a number of dynamic processes, including the separation and reaction efficiency of chromatographic columns and reactors, or economic oil recovery from a reservoir.In this respect, transport phenomena observed in model systems such as random packings of spheres may help to characterize materials with a higher disorder [5][6][7]. For random packings of nonporous (impermeable) particles, for example, the long-time longitudinal dispersion coefficient is dominated by the boundary-layer contribution (due to the no-slip condition at the solid-liquid interface) or by medium and large-scale velocity fluctuations in the flow field depending on the actual disorder of the medium and the Peclet number, Pe u ay d p D m (with u ay , the average velocity; d p , particle diameter; and D m , the molecular diffusivity) [6,8]. This behavior contrasts with random packings of porous (permeable) particles. In that case, liquid holdup associated with stagnant zones inside the particles may dominate dispersion when convective times t c uayt dp significantly exceed the dimensionless time for diffusion, t d. In many situations, however, both a macroscopic flow heterogeneity and solute trapping in stagnant zones contribute to transient and asymptotic dispersion [3,7,9].Despite numerous theoretical, experimental, and numerical studies (e.g., [1,7,8,[10][11][12]),...
1 H NMR relaxometry is used in earth science as a non-destructive and time-saving method to determine pore size distributions (PSD) in porous media with pore sizes ranging from nm to mm. This is a broader range than generally reported for results from X-ray computed tomography (X-ray CT) scanning, which is a slower method. For successful application of 1 H NMR relaxometry in soil science, it is necessary to compare PSD results with those determined from conventional methods. The PSD of six disturbed soil samples with various textures and soil organic matter (SOM) content were determined by conventional soil water retention at matric potentials between −3 and −390 kPa (pF 1.5-3.6). These PSD were compared with those estimated from transverse relaxation time (T 2 ) distributions of water in soil samples at pF 1.5 using two different approaches. In the first, pore sizes were estimated using a mean surface relaxivity of each soil sample determined from the specific surface area. In the second and new approach, two surface relaxivities for each soil sample, determined from the T 2 distributions of the soil samples at different matric potentials, were used. The T 2 distributions of water in the samples changed with increasing soil matric potential and consisted of two peaks at pF 1.5 and one at pF 3.6. The shape of the T 2 distributions at pF 1.5 was strongly affected by soil texture and SOM content (R 2 = 0.51 − 0.95). The second approach (R 2 = 0.98) resulted in good consistency between PSD, determined by soil water retention, and 1 H NMR relaxometry, whereas the first approach resulted in poor consistency. Pore sizes calculated from the NMR data ranged from 100 μm to 10 nm. Therefore, the new approach allows 1 H NMR relaxometry to be applied for the determination of PSD in soil samples and for studying swelling of SOM and clay and its effects on pore size in a fast and non-destructive way. This is not, or only partly, possible by conventional soil water retention or X-ray CT.
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