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Water in micro- and nanofluidics systems described using the water potential.

This Tutorial Review shows the behaviour of water in micro- and nanofluidic systems. The chemical potential of water ('water potential') conveniently describes the energy level of the water at different locations in and around the system, both in the liquid and gaseous state. Since water moves from high to low potential, the water potential enables us to predict and describe the direction of water movement inside systems and between systems and their surroundings. Practical examples of microfluidic devices illustrate the different contributions to the water potential (capillary, gravitational, entropic (osmotic) in liquid water; the partial vapour pressure in atmospheric water and van der Waals forces in water films) and the resulting water movement.

Microfluidics↗

Transcellular water transport in lung alveolar epithelium through mercury-sensitive water channels.

The movement of water between the air space and capillary compartments is important for the maintenance of air space hydration during respiration and for reabsorption of excess alveolar fluid. We have obtained immunocytochemical and functional evidence that plasma-membrane water channels are responsible for water transport in the intact lung. Northern and quantitative immunoblot analysis showed high expression of CHIP28 (channel-forming integral membrane protein of 28 kDa) water channels in rat lung; immunocytochemistry showed CHIP28 localization to epithelial cell plasma membranes. Stopped-flow light scattering measurements of osmotic water permeability (Pf) in freshly isolated rat alveolar type II epithelial cells indicated a high Pf of 0.015 +/- 0.002 cm/s (10 degrees C) that was weakly temperature-dependent (activation energy, 4 kcal/mol) and reversibly inhibited by 78 +/- 4% by 0.5 mM HgCl2. An in situ-perfused sheep lung model was used to determine the route for water movement in intact lung. Blood-to-air-space water transport was measured by sampling air space fluid after instillation into distal air spaces of hyperosmolar saline (900 mOsm) containing radioiodinated albumin and [14C]mannitol. In seven sets of experiments, air space osmolality and radioiodinated albumin equilibrated with a t1/2 of 0.85 +/- 0.1 min. In the contralateral lung perfused with 0.5 mM HgCl2, t1/2 increased to 2.7 +/- 0.4 min; the inhibitory effect of HgCl2 was fully reversed by 5 mM 2-mercaptoethanol. These results provide direct evidence for transcellular movement of water across the alveolar epithelium in intact lung through mercury-sensitive water channels.

Animals↗

Sieving of electrolytes at capillary wall of cat skeletal muscle by osmotic water flow.

To test the hypothesis that a significant proportion of transcapillary water flow occurs through solute-restricting channels, we investigated the effects of transcapillary water movement on plasma electrolytes in isolated perfused cat skeletal muscle. The lower hindlimbs of anesthetized cats were perfused with a plasma-albumin solution and were weighed to determine transcapillary water movement. Osmolality was increased 60-70 mosmol/kgH2O with sucrose, creating water fluxes of 8-10 ml.min-1.100 g-1, and the changes in the venous concentrations of sodium, potassium, and chloride were determined. The ion concentrations were all reduced by 6-7% with no significant difference between them. The amount of reduction was quantitatively explained by the flow of ion-free water from the interstitial space into plasma and the diffusion of electrolyte in the same direction. These findings support the hypothesis that important water-only transcapillary channels exist in mammalian skeletal muscle. The observations may also explain some of the electrolyte changes seen in intense exercise.

Animals↗

Paracellular and transcellular routes for water and solute movements across insect epithelia.

Because the frontal area of the intercellular clefts in Malpighian tubules is small, and the osmotic permeability of the cell membranes is large, the route for transepithelial water movement during fluid secretion is transcellular. Water movements appear to be a passive response to osmotic gradients of a few mosmol 1(-1) produced in the cells and in he lumen by active ion transport. The excretory functions of Malpighian tubules are discussed in relation to recent analyses of the routes of passive permeation for non-electrolytes. Uncharged molecules smaller than a disaccharide appear to move at significant rates through the cells whereas molecules as large as inulin traverse the epithelium by a paracellular path. In addition there are specific active transport mechanisms for a variety of organic molecules. The routes and mechanisms proposed for water and solute movements are discussed in relation to comparable studies in other epithelia.

Animals↗

Localization and quantification of plasma membrane aquaporin expression in maize primary root: a clue to understanding their role as cellular plumbers.

Water movement across root tissues occurs by parallel apoplastic, symplastic, and transcellular pathways that the plant can control to a certain extent. Because water channels or aquaporins (AQPs) play an important role in regulating water flow, studies on AQP mRNA and protein expression in different root tissues are essential. Here, we quantified and localized the expression of Zea mays plasma membrane AQPs (ZmPIPs) in primary root tip using in situ and quantitative RT-PCR and immunodetection approaches. All ZmPIP genes except ZmPIP2;7 were expressed in primary roots. Expression was found to be dependent on the developmental stage of the root, with, in general, an increase in expression towards the elongation and mature zones. Two genes, ZmPIP1;5 and ZmPIP2;5, showed the greatest increase in expression (up to 11- and 17-fold, respectively) in the mature zone, where they accounted for 50% of the total expressed ZmPIPs. The immunocytochemical localization of ZmPIP2;1 and ZmPIP2;5 in the exodermis and endodermis indicated that they are involved in root radial water movement. In addition, we detected a polar localization of ZmPIP2;5 to the external periclinal side of epidermal cells in root apices, suggesting an important role in water uptake from the root surface. Finally, protoplast swelling assays showed that root cells display a variable, but globally low, osmotic water permeability coefficient (Pf < 10 microm/s). However, the presence of a population of cells with a higher Pf (up to 26 microm/s) in mature zone of the root might be correlated with the increased expression of several ZmPIP genes.

Amino Acid Sequence↗

Osmotic water permeability of plasma and vacuolar membranes in protoplasts II: theoretical basis.

Water permeability of the plasma membrane (PM) and the vacuolar membrane (VM) is important for intracellular and transcellular water movement in plants, because mature plant cells have large central vacuoles. We have developed a new method for measuring the osmotic water permeability of the PM and VM (P ( f1) and P ( f2), respectively) in individual plant cells. Here, the theoretical basis and procedure of the method are discussed. Protoplasts isolated from higher plant tissues are used to measure P ( f1) and P ( f2). Because of the semi-permeability (selective permeability) of cellular membranes, protoplasts swell or shrink under hypotonic or hypertonic conditions. A theoretical three-compartment model is presented for simulating time-dependent volume changes in the vacuolar and cytoplasmic spaces in a protoplast during osmotic excursions. The model describes the theoretical relationships between P ( f1), P ( f2) and the bulk osmotic water permeability of protoplasts (P ( f(bulk))). The procedure for measuring the osmotic water permeability is: (1) P ( f(bulk)) is calculated from the time when half of the total change in protoplast volume is completed, by assuming that the protoplast has a single barrier to water movement across it (two-compartment model); (2) P ( f2) of vacuoles isolated from protoplasts is obtained in the same manner; and (3) P ( f1) is determined from P ( f(bulk)) and P ( f2) according to the three-compartment model. The theoretical relationship between P ( fl ) (m s(-1)) and L ( Pl ) (hydraulic conductivity, l=1, 2) (m s(-1) Pa(-1)) is also discussed.

Cell Membrane Permeability↗

Maturational changes in rabbit renal brush border membrane vesicle osmotic water permeability.

We have recently shown that the osmotic water permeability (Pf) of proximal tubules from neonatal rabbits is higher than that of adults (AJP 271:F871-F876, 1996). The developmental change in Pf could be due to differences in one or more of the components in the path for transepithelial water transport. The present study examined developmental changes in water transport characteristics of the proximal tubule apical membrane by determining Pf and aquaporin 1 (AQP1) expression in neonatal (10-14 days old) and adult rabbit renal brush border membrane vesicles (BBMV). AQP1 abundance in the adult BBMV was higher than the neonatal BBMV. At 25 degrees C the Pf of neonatal BBMV was found to be significantly lower than the adult BBMV at osmotic gradients from 50 to 250 mOsm/kg water. The activation energy for osmotic water movement was higher in the neonatal BBMV than the adult BBMV (9.19 +/- 0.37 vs. 5.09 +/- 0.57 kcal . deg-1 . mol-1, P < 0.005). Osmotic water movement in neonatal BBMV was inhibited 17.9 +/- 1.3% by 1 mm HgCl2 compared to 34.3 +/- 3.8% in the adult BBMV (P < 0.005). These data are consistent with a significantly greater fraction of water traversing the apical membrane lipid bilayer in proximal tubules of neonates than adults. The lower Pf of the neonatal BBMV indicates that the apical membrane is not responsible for the higher transepithelial Pf in the neonatal proximal tubule.

Animals↗

New method to measure water permeability in emptied-out Xenopus oocytes controlling conditions on both sides of the membrane.

Membrane water permeability is habitually calculated from volume changes in Xenopus laevis oocytes during external osmotic challenges. Nevertheless, this approach is limited by the uncertainty on the oocyte internal composition. To circumvent this limitation a new experimental set up is introduced where the cell membrane of an emptied-out oocyte was mounted as a diaphragm between two chambers. In its final configuration the oocyte membrane was part of a closed compartment and net water movements induced swelling or shrinking of it. Volume changes were followed by video-microscopy and digitally recorded. In this manner, water movements could be continuously monitored while controlling chemical composition and hydrostatic pressure on both sides of the membrane. Using this novel experimental approach an increasing hydrostatic pressure gradient was applied to both mature (stage VI) and immature (stage IV) oocytes. The relative maximal volume change tolerated before disruption was similar in both cases (1.26+/-0.07 and 1.27+/-0.03 respectively) and similar to those previously reported under maximal osmotic stress. Nevertheless the osmotic permeability coefficient (P(OSM)) in mature oocytes ((1.72+/-0.58) x 10(-3) cm s(-1); n=6) was significantly lower than in immature oocytes ((5.18+/-0.59) x 10(-3) cm s(-1), n=5; p<0.005).

Animals↗

Osmotic water permeabilities of human placental microvillous and basal membranes.

Literature data suggest that water accumulation by the human fetus is driven by osmotic gradients of small solutes. However, the existence of such gradients has not been supported by prior measurements. Attempts to estimate the size of the gradient necessary to drive net water movement have been seriously hampered by the lack of permeability data for the syncytiotrophoblast membranes. Stopped-flow light scattering techniques were employed to measure the osmotic water permeability (Pf) of microvillous (MVM) and basal membrane (BM) vesicles isolated from human term placenta. At 37 degrees C, the Pf was determined to be 1.9 +/- 0.06 x 10(-3) cm/sec for MVM and 3.1 +/- 0.20 x 10(-3) cm/sec for BM (mean +/- SD, n = 6). At 23 degrees C, Pf was reduced to 0.7 +/- 0.04 x 10(-3) cm/sec in MVM and 1.6 +/- 0.05 x 10(-3) cm/sec in BM. These Pf values are comparable to those observed in membranes where water has been shown to permeate via a lipid diffusive mechanism. Arrhenius plots of Pf over the range 20-40 degrees C were linear, with activation energies of 13.6 +/- 0.6 kcal/mol for MVM and 12.9 +/- 1.0 kcal/mol for BM. Water permeation was not affected by mercurial sulfhydryl agents and glucose transport inhibitors. These data clearly suggest that water movement across human syncytiotrophoblast membranes occurs by a lipid diffusion pathway. As noted in several other epithelial tissues, the basal membrane has a higher water permeability than the microvillous membrane.(ABSTRACT TRUNCATED AT 250 WORDS)

4,4'-Diisothiocyanostilbene-2,2'-Disulfonic Acid↗

The mechanisms of sodium absorption in the human small intestine.

The present studies were designed to characterize sodium transport in the jejunum and ileum of humans with respect to the effects of water flow, sodium concentration, addition of glucose and galactose, and variations in aniomic composition of luminal fluid. In the ileum, sodium absorption occurred against very steep electrochemical gradients (110 mEq/liter, 5-15 mv), was unaffected by the rate or direction of water flow, and was not stimulated by addition of glucose, galactose, or bicarbonate. These findings led to the conclusion that there is an efficiently active sodium transport across a membrane that is relatively impermeable to sodium. In contrast, jejunal sodium (chloride) absorption can take place against only the modest concentration gradient of 13 mEq/liter, was dramatically influenced by water movement, and was stimulated by addition of glucose, galactose, and bicarbonate. The stimulatory effect of glucose and galactose was evident even when net water movement was inhibited to zero by mannitol. These observations led to the conclusion that a small fraction of jejunal sodium absorption was mediated by active transport coupled either to active absorption of bicarbonate or active secretion of hydrogen ions. The major part of sodium absorption, i.e. sodium chloride absorption, appeared to be mediated by a process of bulk flow of solution along osmotic pressure gradients. The stimulatory effect of glucose and galactose, even at zero water flow, was explained by a model in which the active transport of monosaccharide generates a local osmotic force for the absorption of solution (NaCl and water) from the jejunal lumen, which, in the presence of mannitol, is counterbalanced by a reverse flow of pure solvent (H(2)O) through a parallel set of channels which are impermeable to sodium. Support for the model was obtained by the demonstration that glucose and bicarbonate stimulated the absorption of the nonactively transported solute urea even when net water flow was maintained at zero by addition of mannitol to luminal contents.

Adult↗

Changes in intracellular sodium during the hydroosmotic response to vasopressin.

During vasopressin (VP)-induced water movement, toad urinary bladder epithelial cells undergo unique morphological changes. The osmolality within these responding cells remains relatively stable despite the large transcellular transport of water. We hypothesized that the hydroosmotic response to VP may be associated with a net increase in sodium either as an aid in maintaining the intracellular osmolality or as part of a Na-Ca exchange process. Changes in intracellular sodium (Nai) were monitored over time in individual hemibladders using 23Na NMR. Hemibladders were mounted as bags on glass pipets and filled with deionized water. During NMR studies, the serosal bath consisted of aerated 2.4 mM HCO3 amphibian Ringer's (pH 8.1) made up with 15% D2O containing the shift reagent, dysprosium tripolyphosphate (1 mM). This reagent allowed for visualization of Nai by shifting the extracellular Na signal; it did not affect basal or VP stimulated water flow, short-circuit current, or high energy phosphate metabolism as seen by 31P NMR. Changes in Nai were determined by integrating the area under the unshifted Na peak at each measurement and expressing differences as a ratio relative to baseline. The initial Nai signal from unstimulated hemibladders remained stable in these tissues over at least 180 minutes. Within 30 minutes of VP (20 mU/ml) exposure, however, the Nai peak increased 2.47 times above pretreatment baseline (N = 16, P less than 0.001). The Nai signal returned toward baseline values with removal of VP from the serosal bath but only after approximately 90 minutes. When change in cell shape and water movement were prevented by having isotonic sorbitol in the mucosal bath, VP produced no change in the Nai signal (N = 10).(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗

Role of 5-hydroxytryptamine in intestinal water and electrolyte movement during gut anaphylaxis.

Exposure of sensitised intestine to specific allergen is known to produce appreciable reduction in water and electrolyte absorption. The mediators participating in this process have not been fully characterised. The effects of the 5-hydroxytryptamine2 (5-HT2) and 5-HT3 receptor antagonists, ketanserin and granisetron, respectively, on water movement during intestinal anaphylaxis were studied. Hooded Lister rats (120-150 g) were sensitised to ovalbumen and 14 days later, intestinal water and electrolyte movement was assessed at 10 minute intervals by in situ jejunal perfusion with a plasma electrolyte solution (PES) or PES containing 20 mg/l ovalbumen. Within 20 minutes of exposure to PES+ovalbumen, net water secretion that could be completely prevented by the mast cell stabilising agent doxantrazole occurred compared with absorption with PES alone (median -20 microliters/min/g (interquartile range -43 to -5), n = 11), v (107 (86 to 113), n = 10; p < 0.01). Pre-treatment with subcutaneous ketanserin 200 micrograms/kg (n = 7) or granisetron 300 micrograms/kg (n = 8) partially inhibited the secretory response to PES+ovalbumen (18 (11 to 48) and 13 (6 to 32) respectively; both p < 0.01 compared with PES+ovalbumen control). After 40 minutes perfusion with PES+ovalbumen, the changes in water movement were less pronounced 24 (-3 to 43) and neither ketanserin or granisetron had any effect (ketanserin: 48 (28 to 87), granisetron: 41 (32 to 83); NS). In all experiments, sodium and chloride movement paralleled that of water. Thus, the profound water secretion that occurs in the early stages of intestinal anaphylaxis is partly 5-HT dependent because it can be reversed by 5-HT2 and 5-HT3 receptor antagonists. Other mediators must also be involved, especially in the late phase of anaphylaxis.

Anaphylaxis↗

[Principal stages in the Cenozoic diversification of shallow-water molluscan faunas in the North Pacific].

Cluster analysis of bivalve species recorded in Cenozoic deposits in Sakhalin Island, western Kamchatka, Hokkaido, and California was used to determine geological age of the modem North Pacific biogeographic region and its constituent subregions (Japan-Mandchurian, Beringian, and Oregon-Sitkan). The North Pacific region developed during the Paleogene-Neogene transition due to Drake Passage opening to deep-water movement, formation of the deep-water Antarctic Circumpolar Current, and the change in climate from greenhouse to psychospheric. Differentiation of the three subregions within the North Pacific Region seems to have occurred in late Miocene-early Pliocene, about 5.6 millions years ago and was probably due to the flooding of the Bering Land Bridge and development of the present configuration of circulation in the North Pacific. In the Northwest Pacific, during Paleogene and early Neogene, the faunal diversification occurred more rapidly and was more extensive than in the Northeast Pacific.

Animals↗

Diel Patterns of Water Potential Components for the Crassulacean Acid Metabolism Plant Opuntia ficus-indica when Well-Watered or Droughted.

Under well-watered conditions, chlorenchyma acidity in cladodes of Opuntia ficus-indica increased substantially at night, fully accounting for the 0.26-megapascal nocturnal increase in osmotic pressure in the outer 2 millimeters. Osmotic pressure in the inner part of the chlorenchyma and in the water-storage parenchyma did not change significantly over 24-hour periods. Three months of drought decreased nocturnal acid accumulation by 73% and essentially abolished transpiration; also, 27% of the chlorenchyma water and 61% of the parenchyma water was lost during such drought, but the average tissue osmotic pressure was little affected. Turgor pressure was maintained in the chlorenchyma after 3 months of drought, although it decreased sevenfold in the water-storage parenchyma compared with the well-watered condition. Moreover, the nocturnal increases in turgor pressure of about 0.08 megapascal in the outer part of the chlorenchyma was also unchanged by such drought. The water potential magnitudes favored water movement from the parenchyma to the chlorenchyma at the end of the night and in the reverse direction during the late afternoon. Experiments with tritiated water support this pattern of water movement, which is also in agreement with predictions based on electric-circuit analog models for Crassulacean acid metabolism plants.

Journal Article↗

The effect of nitrogen mustard intoxication on glucose absorption from the small intestine of the rat.

Methyl bis (beta-chlorethyl) amine (HN(2)) was administered as a single intraperitoneal dose to male Sprague-Dawley rats. Animals were given either a median lethal dose and perfused one day after treatment or were given a larger dose and perfused two days after treatment. Intestinal segments were perfused in vivo and samples of effluent were collected and measured for glucose using the glucose oxidase method. Animals treated with the larger dose of HN(2) displayed significant reduction in intestinal dry weight. The average water content of intestinal segments derived from treated animals did not differ significantly from that in controls.Steady-state glucose absorption, obtained from data collected during the last 40 minutes of the perfusion period, was found to be significantly reduced in animals treated with drugs when intestinal absorption for control and treated animals was adjusted for water movement but not for dry intestinal weight and length. However, when glucose absorption was adjusted for dry tissue mass and length as well as for water movement, no significant differences in intestinal glucose absorption between control and treated rats were observed. Alterations can therefore be attributed to loss of intestinal tissue rather than to loss in the ability of the intestinal mass to absorb glucose.

Animals↗

Modeling macropore flow effects on pesticide leaching: inverse parameter estimation using microlysimeters.

Macropore flow is a key factor determining pesticide fate, but models accounting for this process need parameters that cannot be easily measured. This study was conducted to investigate the use of inverse techniques to estimate parameters controlling macropore flow and pesticide fate in the dual-permeability model MACRO. Undisturbed columns were sampled at three landscape positions (hilltop, slope, hollow) with contrasting texture and organic carbon content. Transient leaching experiments were performed for an anionic tracer and the herbicide MCPA (4-chloro-2methylphenoxy acetic acid) during a 4-mo period, first under natural rainfall, and then under controlled irrigation in the laboratory. The tracer breakthrough for the liner-textured soil from the hilltop showed strong evidence of macropore flow, resulting in a rapid leaching of MCPA, while leaching was minimal from the organic-rich hollow soil, since macropore flow was weaker and adsorption stronger. The MACRO model was linked to the inverse modeling program SUFI (Sequential Uncertainty Fitting) to enable calibration of nine key model parameters. Based on calculated model efficiencies, MACRO-SUFI gave generally good predictions of water movement and tracer and pesticide transport, although some errors were attributed to difficulties in simulating the effects of soil moisture on degradation and the timing of water outflows. Even after calibration, significant uncertainties remained for some key parameters controlling macropore flow. Nevertheless, the parameter estimates were significantly different between landscape positions and could also be related to basic soil properties. The posterior uncertainty ranges could probably be reduced with a more exhaustive sampling of the parameter space and improved experimental designs.

2-Methyl-4-chlorophenoxyacetic Acid↗

Physiologic determinants of biliary calcium secretion in the dog.

Studies were carried out to define the origins and physiologic determinants of biliary calcium secretion in the dog. Canalicular bile flow was varied by infusions of several bile acids or a canalicular choleretic agent, SC2644. Three natural bile acids (cholyltaurine, chenodeoxycholyltaurine, and ursodeoxycholyltaurine) that have a low critical micellar concentration and a synthetic hydroxy-oxo bile acid (12-dehydrocholyltaurine) with a much higher critical micellar concentration were used. Calcium output, bile acid output, and bile flow were determined; canalicular bile flow was assessed by measurement of erythritol clearance. Calcium output increased linearly with bile acid output for all three natural bile acids (r = 0.94; 0.026-0.035 mumol delta calcium/mumol delta bile acid), but chenodeoxycholyltaurine and ursodeoxycholyltaurine infusions induced a slightly greater secretion of calcium for a given bile acid output and bile flow than cholyltaurine. Calcium output also increased linearly with bile flow, whether bile flow was induced by the canalicular choleretic SC2644 (r = 0.93; 0.0018 mumol delta calcium/microliter delta water) or the hydroxy-oxo bile acid (r = 0.99; 0.0021 mumol delta calcium/microliter delta water). Because the calcium output induced by the three micelle-forming bile acids was greater than that which could be explained by the induced increase in bile flow per se, it was deduced that canalicular calcium has two origins: a micellar component that is dependent on the presence of micelles or micelle-forming bile acids and an osmotic component that is dependent on osmotic forces, e.g., caused by secretion induced by SC2644 or bile acid molecules and possibly bile acid micelles. A small bile acid-independent output of calcium was also detected and should contribute to the osmotic fraction. To determine whether ductular modification of bile flow influenced biliary calcium secretion, ductular secretion was induced by secretin and ductular absorption was induced by somatostatin. Changes in calcium output followed water movements, increasing with ductular secretion and decreasing with ductular absorption. The relationship between changes in ductular bile flow and calcium output was identical during induced ductular secretion (r = 0.83; 0.0016 mumol delta calcium/microliter delta water) or induced ductular absorption (r = 0.91; 0.0019 mumol delta calcium/microliter delta water). Thus calcium movements in relation to water movements were identical for the osmotic component of canalicular bile and the ductular component of bile.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals↗