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1.
Contribution of solvent drag through intercellular junctions to absorption of nutrients by the small intestine of the rat 总被引:3,自引:0,他引:3
The lumen of the small intestine in anesthetized rats was recirculated with 50 ml perfusion fluid containing normal salts, 25 mM glucose and low concentrations of hydrophilic solutes ranging in size from creatinine (mol wt 113) to Inulin (mol wt 5500). Ferrocyanide, a nontoxic, quadrupally charged anion was not absorbed; it could therefore be used as an osmotically active solute with reflection coefficient of 1.0 to adjust rates of fluid absorption, Jv, and to measure the coefficient of osmotic flow, Lp. The clearances from the perfusion fluid of all other test solutes were approximately proportional to Jv. From Lp and rates of clearances as a function of Jv and molecular size we estimate (a) the fraction of fluid absorption which passes paracellularly (approx. 50%), (b) coefficients of solvent drag of various solutes within intercellular junctions, (c) the equivalent pore radius of intercellular junctions (50 A) and their cross sectional area per unit path length (4.3 cm per cm length of intestine). Glucose absorption also varied as a function of Jv. From this relationship and the clearances of inert markers we calculate the rate of active transport of glucose, the amount of glucose carried paracellularly by solvent drag or back-diffusion at any given Jv and luminal glucose concentration and the concentration of glucose in the absorbate. The results indicate that solvent drag through paracellular channels is the principal route for intestinal transport of glucose or amino acids at physiological rates of fluid absorption and concentration. In the absence of luminal glucose the rate of fluid absorption and the clearances of all inert hydrophilic solutes were greatly reduced. It is proposed that Na-coupled transport of organic solutes from lumen to intercellular spaces provides the principal osmotic force for fluid absorption and triggers widening of intercellular junctions, thus promoting bulk absorption of nutrients by solvent drag. Further evidence for regulation of channel width is provided in accompanying papers on changes in electrical impedance and ultrastructure of junctions during Na-coupled solute transport. 相似文献
2.
J. R. Pappenheimer 《The Journal of membrane biology》1987,100(1):137-148
Summary Isolated intestinal segments from rats or hamsters were recirculated with balanced salt solutions containing fluorocarbon emulsion to provide 6 vpc oxygen. The lumen contained an axial Ag–AgCl electrode, and the serosal surface was surrounded by a cylindrical shell of Ag–AgCl. Transmural impedances were measured at frequencies from 0.01–30 kHz before and after removal of the mucosal epithelium. The resistance of intercellular junctions,R
J
, the distributed resistance of the lateral spaces,R
L
, and the distributed membrane capacitance,C
M
, were computed from the relations between frequency and impedance. Activation of Na-coupled solute transport by addition of glucose, 3-0-methyl glucose, alanine or leucine caused two- to threefold decreases of transepithelial impedance. Typical changes induced by glucose in hamster small intestine wereR
J
3013 ,R
L
2310 , andC
M
820 F (per cm length of segment). Half maximal response occurred at a glucose concentration of 2–3mm. The area per unit path length of the junctions (Ap/x=specific resistance ÷R
J
) in glucose activated epithelium was 3.7 cm in hamster midgut and 6.8 cm in rat. These values are close to the 4.3 cm estimated independently from coefficients of solvent drag and hydrodynamic conductance in glucose-activated rat intestine in vivo. The transepithelial impedance response to Na-coupled solute transport was reversibly dependent upon oxygen tension.It is proposed that activation of Na-coupled solute transport triggers contraction of circumferential actomyosin fibers in the terminal web of the microvillar cytoskeletal system, thereby pulling apart junctions and allowing paracellular absorption of nutrients by solvent drag as described in the previous accompanying paper. Anatomical evidence in support of this hypothesis is presented in the following second accompanying paper. 相似文献
3.
Physiological regulation of transepithelial impedance in the intestinal mucosa of rats and hamsters 总被引:1,自引:0,他引:1
J R Pappenheimer 《The Journal of membrane biology》1987,100(2):137-148
Isolated intestinal segments from rats or hamsters were recirculated with balanced salt solutions containing fluorocarbon emulsion to provide 6 vpc oxygen. The lumen contained an axial Ag-AgCl electrode, and the serosal surface was surrounded by a cylindrical shell of Ag-AgCl. Transmural impedances were measured at frequencies from 0.01-30 kHz before and after removal of the mucosal epithelium. The resistance of intercellular junctions, RJ, the distributed resistance of the lateral spaces, RL, and the distributed membrane capacitance, CM, were computed from the relations between frequency and impedance. Activation of Na-coupled solute transport by addition of glucose, 3-0-methyl glucose, alanine or leucine caused two- to threefold decreases of transepithelial impedance. Typical changes induced by glucose in hamster small intestine were RJ 30----13 omega, RL 23----10 omega, and CM 8----20 microF (per cm length of segment). Half maximal response occurred at a glucose concentration of 2-3 mM. The area per unit path length of the junctions (Ap/delta chi = specific resistance divided by RJ) in glucose activated epithelium was 3.7 cm in hamster midgut and 6.8 cm in rat. These values are close to the 4.3 cm estimated independently from coefficients of solvent drag and hydrodynamic conductance in glucose-activated rat intestine in vivo. The transepithelial impedance response to Na-coupled solute transport was reversibly dependent upon oxygen tension. It is proposed that activation of Na-coupled solute transport triggers contraction of circumferential actomyosin fibers in the terminal web of the microvillar cytoskeletal system, thereby pulling apart junctions and allowing paracellular absorption of nutrients by solvent drag as described in the previous accompanying paper. Anatomical evidence in support of this hypothesis is presented in the following second accompanying paper. 相似文献
4.
Summary Isolated segments of hamster small intestine were perfused with oxygenated salt-fluorocarbon emulsions with or without 10–25mm glucose, alanine or leucine. Resistances of inter-cellular occluding junctions and of lateral spaces and the distributed capacitance of epithelial plasma membranes were estimated from steady-state transepithelial impedances at frequencies from 0.01–10 kHz. The segments were then fixedin situ with isorheic 2.5% glutaraldehyde while continuing to measure impedance. This method of fixation increased the resistance of lateral spaces but had little effect on the resistance of occluding junctions or on membrane capacitance. The large decreases of impedance induced by glucose or amino acids were preserved in fixed tissue and could therefore be correlated with changes in structure. The observed changes of impedance were interpreted as decreased resistance of occluding junctions and lateral spaces together with increased exposed surface of lateral membranes (capacitance). Glucose, alanine or leucine induced expansion of lateral intercellular spaces as seen by light and electron microscopy. Large dilatations within absorptive cell occluding junctions were revealed by electron microscopy. Freeze-fracture analysis revealed that these dilatations consisted of expansions of compartments bounded by strands/grooves. These solute-induced structural alterations were also associated with condensation of microfilaments in the zone of the perijunctional actomyosin ring, typical of enhanced ring tension. Similar anatomical changes were found in epithelia fixedin situ at 38°C during luminal perfusion with glucose in blood-circulated intestinal segments of anesthetized animals. These structural changes support the hypothesis that Na-coupled solute transport triggers contraction of perijunctional actomyosin, thereby increasing junctional permeability and enhancing absorption of nutrients by solvent drag as described in the two accompanying papers. 相似文献
5.
Structural basis for physiological regulation of paracellular pathways in intestinal epithelia 总被引:15,自引:0,他引:15
Isolated segments of hamster small intestine were perfused with oxygenated salt-fluorocarbon emulsions with or without 10-25 mM glucose, alanine or leucine. Resistances of intercellular occluding junctions and of lateral spaces and the distributed capacitance of epithelial plasma membranes were estimated from steady-state transepithelial impedances at frequencies from 0.01-10 kHz. The segments were then fixed in situ with isorheic 2.5% glutaraldehyde while continuing to measure impedance. This method of fixation increased the resistance of lateral spaces but had little effect on the resistance of occluding junctions or on membrane capacitance. The large decreases of impedance induced by glucose or amino acids were preserved in fixed tissue and could therefore be correlated with changes in structure. The observed changes of impedance were interpreted as decreased resistance of occluding junctions and lateral spaces together with increased exposed surface of lateral membranes (capacitance). Glucose, alanine or leucine induced expansion of lateral intercellular spaces as seen by light and electron microscopy. Large dilatations within absorptive cell occluding junctions were revealed by electron microscopy. Freeze-fracture analysis revealed that these dilatations consisted of expansions of compartments bounded by strands/grooves. These solute-induced structural alterations were also associated with condensation of microfilaments in the zone of the perijunctional actomyosin ring, typical of enhanced ring tension. Similar anatomical changes were found in epithelia fixed in situ at 38 degrees C during luminal perfusion with glucose in blood-circulated intestinal segments of anesthetized animals. These structural changes support the hypothesis that Na-coupled solute transport triggers contraction of perijunctional actomyosin, thereby increasing junctional permeability and enhancing absorption of nutrients by solvent drag as described in the two accompanying papers. 相似文献
6.
Summary Activation of protein kinase C has been shown to cause both stimulation and inhibition of transport processes in the brush-border membrane and renal tubule. This study was designed to examine the dose-response nature and time-dependent effect of 4 -phorbol-12-myristate-13-acetate (PMA) on the rates of bicarbonate absorption (J
HCO3) and fluid absorption (J
v) in the proximal convoluted tubule (PCT) of rat kidney. Bicarbonate flux was determined by total CO2 changes between the collected fluid and the original perfusate as analyzed by microcalorimetry. Luminal perfusion of PMA (10–10 10–5 M) within 10 min caused a significant increase ofJ
HCO3 andJ
v. A peaked curve of the dose response was observed with maximal effect at 10–8 M PMA on both bicarbonate and fluid reabsorption, which could be blocked completely by amiloride (10–3
m) and EIPA (10–5 M). On the other hand, with an increase of perfusion time beyond 15 min, PMA (10–8 and 10–6 M) could inhibitJ
HCO3 andJ
v. Amiloride (10–3 M) or EIPA (10–5 M) significantly inhibitsJ
HCO3 andJ
v, while there is no additive effect of PMA and amiloride or EIPA on PCT transport. An inactive phorbol-ester, 4-phorbol, that does not activate protein kinase C, had no effects onJ
HCO3 andJ
v. Capillary perfusion of PMA (10–8 M) significantly stimulate bothJ
HCO3 andJ
v; however, PMA did not affect glucose transport from either the luminal side or basolateral side of the PCT. These results indicate that activation of endogenous protein kinase C by PMA could either stimulate or inhibit both bicarbonate and fluid reabsorption in the PCT dependent on time and dose, and these effects are through the modulation of Na+/H– exchange mechanism. 相似文献
7.
Matsumoto N Tsuruoka S Iwamoto T Tomich JM Ito K Imai M Suzuki M 《The Journal of membrane biology》2003,193(3):195-200
To better understand the process of fluid movement driven by Cl– conductance, a Cl– channel-forming peptide was delivered to the luminal membrane of microperfused rabbit renal proximal tubules. When the peptide (NK4-M2GlyR) was perfused, a significant new conductance was observed within 3 min and stabilized at 10 min. Alteration of the ion composition revealed it to be a Cl–-specific conductance. Reabsorption of Cl– (J
Cl) was increased by NK4-M2GlyR, but not by a scramble NK4-M2GlyR sequence, suggesting that the active peptide formed de novo Cl– channels in the luminal membrane of the perfused tubules. In the presence of the peptide, reabsorption of fluid (J
v) was dramatically increased and J
Na and J
Ca were concomitantly increased. We propose that introduction of the new Cl– conductance in the luminal membrane leads to a coordinated efflux of water across the membrane and an increase in cation translocation via the paracellular pathway, resulting in an increase in J
v. This novel method could prove useful in characterizing mechanisms of fluid transport driven by Cl– gradients. 相似文献
8.
The Na(+) recirculation theory for solute-coupled fluid absorption is an expansion of the local osmosis concept introduced by Curran and analyzed by Diamond & Bossert. Based on studies on small intestine the theory assumes that the observed recirculation of Na(+) serves regulation of the osmolarity of the absorbate. Mathematical modeling reproducing bioelectric and hydrosmotic properties of small intestine and proximal tubule, respectively, predicts a significant range of observations such as isosmotic transport, hyposmotic transport, solvent drag, anomalous solvent drag, the residual hydraulic permeability in proximal tubule of AQP1 (-/-) mice, and the inverse relationship between hydraulic permeability and the concentration difference needed to reverse transepithelial water flow. The model reproduces the volume responses of cells and lateral intercellular space (lis) following replacement of luminal NaCl by sucrose as well as the linear dependence of volume absorption on luminal NaCl concentration. Analysis of solvent drag on Na(+) in tight junctions provides explanation for the surprisingly high metabolic efficiency of Na(+) reabsorption. The model predicts and explains low metabolic efficiency in diluted external baths. Hyperosmolarity of lis is governed by the hydraulic permeability of the apical plasma membrane and tight junction with 6-7 mOsm in small intestine and < or = 1 mOsm in proximal tubule. Truly isosmotic transport demands a Na(+) recirculation of 50-70% in small intestine but might be barely measurable in proximal tubule. The model fails to reproduce a certain type of observations: The reduced volume absorption at transepithelial osmotic equilibrium in AQP1 knockout mice, and the stimulated water absorption by gallbladder in diluted external solutions. Thus, it indicates cellular regulation of apical Na(+) uptake, which is not included in the mathematical treatment. 相似文献
9.
Passive absorption of D-Galactose (in the presence of 0.5 mM phlorizin), 2-deoxy-D-glucose and D-Mannitol by rat jejunum has been measured in vivo by perfusion of an intestinal segment with recirculation, along successive absorption periods of 5 or 10 min duration. In the range of 1 to 40 mM concentrations, the three solutes were absorbed at a very similar rate that varied as a lineal function of the concentrations in the perfusion solution. Absorption of 1 mM solute was not modified by the presence of 40 mM glucose or galactose. Passive absorption kinetics suggests processes of simple diffusion or solvent drag. The use of paracellular way for the passive absorption is supported by the fact that triaminopyrimidine (TAP) and protamine, which decrease the permeability through the tight junctions, also inhibit the absorption, with similar characteristics for both actions: TAP inhibition (53%) is very rapid and can be easily reversed, while that of protamine (30%) requires some time of previous exposure, lasts longer and can be reversed by heparin. The same analogy is shown by two actions that enhance the paracellular permeability: theophylline increases (30%) the passive absorption with lasting effect, while luminal hypertony enhances absorption transitorily. The passive absorption of the assayed solutes could be estimated to take place by the paracellular way in at least 50% and probably 70% or even more. The measure of net fluid fluxes reveals that solute fluxes must be prevailingly explained by simple diffusion, as the solvent drag can only play a very minor role. 相似文献
10.
The involvement of second messengers in modulating Schistocerca gregaria rectal ion and fluid transport was investigated using two in vitro bioassay systems: everted rectal sacs and rectal flat sheets. Various agents known to block or activate specific signal transduction pathways were employed in these bioassays. Cyclic AMP stimulated rectal fluid reabsorption (Jv) and Cl− transport (Isc) to the same extent as aqueous extracts of corpus cardiacum storage lobes. Cyclic GMP also partially (50%) stimulated both rectal Jv and Isc. Exogenous Ca2+ was not required for the maintenance of rectal transport, indeed Ca2+ free conditions increased the amount of stimulatable Jv. There was some evidence indicating intracellular Ca2+ may play a minor role in controlling rectal transport. The phospholipase C mediated signal transduction pathway was not involved with the stimulation of rectal transport, and appeared to have an inhibitory role. Neuroparsins, antidiuretic neuropeptides from Locusta migratoria, showed no activity upon either S. gregaria rectal ion or fluid transport. The findings of this study show that the two closely related insects possess discrete antidiuretic factors which stimulate rectal transport via different signal transduction pathways. © 1996 Wiley-Liss, Inc. 相似文献
11.
Effect of nitrogen stress and abscisic acid on nitrate absorption and transport in barley and tomato
F. Stuart Chapin III David T. Clarkson John R. Lenton Colin H. S. Walter 《Planta》1988,173(3):340-351
The potential of barley (Hordeum vulgare L.) and tomato (Lycopersicon esculentum Mill.) roots for net NO
3
-
absorption increased two-to five fold within 2 d of being deprived of NO
3
-
supply. Nitrogen-starved barley roots continued to maintain a high potential for NO
3
-
absorption, whereas NO
3
-
absorption by tomato roots declined below control levels after 10 d of N starvation. When placed in a 0.2 mM NO
3
-
solution, roots of both species transported more NO
3
-
and total solutes to the xylem after 2 d of N starvation than did N-sufficient controls. However, replenishment of root NO
3
-
stores took precedence over NO
3
-
transport to the xylem. Consequently, as N stress became more severe, transport of NO
3
-
and total solutes to the xylem declined, relative to controls. Nitrogen stress caused an increase in hydraulic conductance (L
p) and exudate volume (J
v) in barley but decrased these parameters in tomato. Nitrogen stress had no significant effect upon abscisic acid (ABA) levels in roots of barley or flacca (a low-ABA mutant) tomato, but prevented an agerelated decline in ABA in wild-type tomato roots. Applied ABA had the same effect upon barley and upon the wild type and flacca tomatoes: L
p and J
v were increased, but NO
3
-
absorption and NO
3
-
flux to the xylem were either unaffected or sometimes inhibited. We conclude that ABA is not directly involved in the normal changes in NO
3
-
absorption and transport that occur with N stress in barley and tomato, because (1) the root ABA level was either unaffected by N stress (barley and flacca tomato) or changed, after the greatest changes in NO
3
-
absorption and transport and L
p had been observed (wild-type tomato); (2) changes in NO
3
-
absorption/transport characteristics either did not respond to applied ABA, or, if they did, they changed in the direction opposite to that predicted from changes in root ABA with N stress; and (3) the flacca tomato (which produces very little ABA in response to N stress) responded to N stress with very similar changes in NO
3
-
transport to those observed in the wild type.Abbreviation and symbols ABA
abscisic acid
- Jv
exudate volume
- Lp
root hydraulic conductance 相似文献
12.
Lumen to bath J
12/C
1
and bath to lumen J
21/ C
2
fluxes per unit concentration of 19 probes with diameters (d
m) ranging from 3.0–30.0 Å (water, urea, erythritol, mannitol, sucrose, raffinose and 13 dextrans with d
m
9.1–30.0 Å) were measured during volume secretion (J
v
) in the upper segment of the Malpighian Tubule of Rhodnius by perfusing lumen and bath with 14C or 3H-labeled probes. J
net=(J
12/C
1 – J
21/C
2) was studied as a function of J
v
· J
v
was varied by using different concentrations of 5-hydroxy tryptamine. J
net for 3H-water was not different from J
v
We found: (i) A strong correlation between J
net
and J
v
for 8 probes d
m
=3.0–11.8 Å (group a probes), indicating that the convective component of J
net is more important than its diffusive component and than unstirred layers effects which are negligible. Therefore group a probes are solvent dragged as they cross the epithelium, (ii) There is no correlation between J
net and J
v
for 11 probes with d
m=11.8–30 Å (group b). Therefore these probes must cross the epithelium by diffusion and not by solvent drag, (iii) In a plot of J
net/J
v
vs. d
m
group a probes show a steep linear relation with a slope = –0.111, while for group b probes the slope is –0.002. Thus there is a break between groups a and b in this plot. We tried to fit the data with models for restricted diffusion and convention through cylindrical or parallel slit pathways. We conclude that (i) group a probes are dragged by water through an 11.0 Å-wide slit, (ii) Most of J
v
must follow an extracellular noncytosolic pathway, (iii) Group b probes must diffuse through a 42 Å-wide slit, (iv) A cylindrical pathway does not fit the data.E.G. is a Visiting Scientist at IVIC. It is a pleasure to thank Drs. A.E. Hill and Bruria Shachar-Hill for their suggestion of the use of dextrans, their instruction and help with the dextran separation technique, and their extensive discussions. Dr. R. Apitz, Mr H. Rojas and Mrs. Fulvia Bartoli were most helpful with suggestions during the course of the experimental work. Mr. Jose Mora was fundamental help with the equipment. Mrs. Lelis Ochoa and Mr. Luis F. Alvarez helped with some of the drawings. This work was partially supported by CONICIT, Fundación Polar and CDCH of UCV. It is a pleasure to thank Dr. H. Passow and Dr. K.J. Ullrich at the Max Planck Institut für Biophysik (Frankfurt/Main) where this work was initiated. 相似文献
13.
Larsen EH Møbjerg N Nielsen R 《Comparative biochemistry and physiology. Part A, Molecular & integrative physiology》2007,148(1):101-116
The theory of Na+ recirculation for isosmotic fluid absorption follows logically from Hertz's convection-diffusion equation applied to the exit of water and solutes from the lateral intercellular space. Experimental evidence is discussed indicating Na+ recirculation based upon the following approaches: (i) An isotope tracer method in small intestine. Simultaneous measurement of water flow and ion transport in toad skin epithelium demonstrating, (ii) occasional hyposmotic absorbates, and (iii) reduced fluid absorption in the presence of serosal bumetanide. (iv) Studies of the metabolic cost of net Na+ absorption demonstrating an efficiency that is lower than the 18 Na+ per O2 consumed given by the stoichiometry of the Na+/K+-pump. Mathematical modeling predicts a significant range of observations such as isosmotic transport, hyposmotic transport, solvent drag, anomalous solvent drag, the residual hydraulic permeability in proximal tubule of AQP1(-/-) mice, the adverse relationship between hydraulic permeability and the concentration difference needed to reverse transepithelial water flow, and in a non-contradictory way the wide range of metabolic efficiencies from above to below 18 Na+/O2. Certain types of observations are poorly or not at all reproduced by the model. It is discussed that such lack of agreement between model and experiment is due to cellular regulations of ion permeabilities that are not incorporated in the modeling. Clarification of these problems requires further experimental studies. 相似文献
14.
《Bioscience, biotechnology, and biochemistry》2013,77(3):583-591
The absorption characteristics of rosmarinic acid (RA) were examined by measuring permeation across Caco-2 cell monolayers using an HPLC-electrochemical detector (ECD) fitted with a coulometric detection system. RA exhibited nonsaturable transport even at 30 mM, and the permeation at 5 mM in the apical-to-basolateral direction, J ap→bl, was 0.13 nmol/min/mg of protein. This permeation rate is nearly the same as that of 5 mM chlorogenic acid (CLA) and gallic acid, which are paracellularly transported compounds. Almost all of the apically loaded RA was retained on the apical side, and J ap→bl was inversely correlated with paracellular permeability. These results indicate that RA transport was mainly via paracelluar diffusion, and the intestinal absorption efficiency of RA was low. Furthermore, RA appeared to be unsusceptible to hydrolysis by mucosa esterase in Caco-2 cells. These results, together with our previous work (J. Agric. Food Chem., 52, 2518–2526 (2004), J. Agric. Food Chem., 52, 6418–6424 (2004)) suggest that the majority of RA is further metabolized and degraded into m-coumaric and hydroxylated phenylpropionic acids by gut microflora, which are then efficiently absorbed and distributed by the monocarboxylic acid transporter (MCT) within the body. The potential of orally administered RA in vivo will be further investigated. 相似文献
15.
During isotonic fluid flow, Necturus gallbladder epithelium mediates net fluxes of paracellular probes by a convective process. We show here that the paracellular system is modeled by permeation through three populations of channels: (i) convective parallel-sided ones of width 7.7 nm (ii) small diffusive ones of radius 0.6 nm, and (ii) large diffusive ones of radius exceeding 50 nm. The reflexion coefficient of the convective channels is very low and the calculated osmotic flow rate is close to zero when compared with the observed fluid absorptive rate of 2 x 10–6 cm/sec. Analysis reveals that the convective channels behave as though closed to back-diffusion of probes; if this is due to solvent drag then very high fluid velocities are required, acting through minute areas. There are no transjunctional gradients that could drive the flow, and so the fluid must be propelled through the channel by components of the junction.We propose a mechanism based upon an active junctional peristalsis which allows discrimination on the basis of molecular size, in which the channels are always occluded at some point and so back-diffusion cannot occur. There is no local gradient of salt distal to the junctions and therefore the osmotic permeability of the membranes is irrelevant. High fluid velocities are not required, and the flow can occur over a substantial fraction of the junction. The mechanism must involve motile and contractile elements associated with the junction for which there is already considerable evidence.Symbols
A
i
filtration area of channel i;i=b (big), s (small) and c (convectional)
-
B
constant for streamline flow
-
C
i
concentration of probe at i
-
D
diffusion coefficient
-
D
o
diffusion coefficient in free solution
-
d
width of junction
-
F
i
diffusive drag factor in channel i
-
g
ionic conductivity
-
G
i
convective drag factor in channel i
-
J
ij
probe flux from i to j
-
J
net
net probe flux
-
J
v
volume flow per cm2 of epithelium
-
l
linear extent of junction per cm2 epithelial plane
-
L
length of junctional channel
-
L
p
hydraulic conductivity
-
N
Avogadro's number
-
q
available filtration area fraction of channel
-
r
s
probe molecular radius
-
r
c
channel radius or half-width
-
S
i
steric factor in channel i
-
V
w,s
partial molar volume of water or salt
-
v
i
fluid velocity in channel i
-
w
dynamic viscosity of water
-
specific conductivity
-
ratio of solute radius to channel radius or half-width
-
diffusive/pressure-driven flow ratio
-
reflexion coefficient 相似文献
16.
The regulation of component processes of photosynthesis in transgenic tobacco with decreased phosphoribulokinase activity 总被引:2,自引:0,他引:2
Dimah Z. Habash Martin A. J. Parry Saroj Parmar Matthew J. Paul Simon Driscoll Jacqueline Knight John C. Gray David W. Lawlor 《Photosynthesis research》1996,49(2):159-167
Tobacco plants (Nicotiana tabacum L.) transformed with an inverted cDNA encoding ribulose 5-phosphate kinase (phosphoribulokinase,PRK; EC 2.7.1.19) were employed to study the in vivo relationship between photosynthetic electron transport and the partitioning of electron transport products to major carbon metabolism sinks under conditions of elevated ATP concentrations and limited ribulose 1,5-bisphosphate (RuBP) regeneration. Simultaneous measurements of room temperature chlorophyll fluorescence and CO2 gas exchange were conducted on intact leaves. Under ambient CO2 concentrations and light intensities above those at which the plants were grown, transformants with only 5% of PRK activity showed down-regulation of PS II activity and electron transport in response to a decrease in net carbon assimilation when compared to wild-type. This was manifested as a decline in the efficiency of PS II electron transport (PS II), an increase in dissipation of excess absorbed light in the antennae of PS II and a decline in: total linear electron transport (J1), electron transport dedicated to carbon assimilation (JA) and electron transport allocated to photorespiration (JL). The transformants showed no alteration in the Rubisco specificity factor measured in vitro and calculated in vivo but had a relatively smaller ratio of RuBP oxygenation to carboxylation rates (vo/vc), due to a higher CO2 concentration at the carboxylation site (Cc). The relationship between PS II and CO
2was similar in transformants and wild-type under photorespiratory conditions demonstrating no change in the intrinsic relationship between PS II function and carbon assimilation, however, a novel result of this study is that this similar relationship occurred at different values of quantum flux, J1, JA, JL and vo/vc in the transformant. For both wild-type and transformants, an assessment was made of the possible presence of a third major sink for electron transport products, beside RuBP oxygenation and carboxylation, the data provided no evidence for such a sink.Abbreviations Cc
CO2 concentration at the site of carboxylation
- Ci
intercellular CO2 concentration
- gm
mesophyll conductance to CO2
- J1
total linear electron flow
- JA
linear electron flow allocated to CO2 assimilation
- Jc
linear electron flow supporting carbon reduction and oxidation cycles
- JL
linear electron flow allocated to photorespiration (RuBP oxygenation and fixation of released photorespiratory CO2)
- PRK
phosphoribulokinase
- qP, qN
coefficients for photochemical and non-photochemical quenching of fluorescence respectively
- Rubisco
ribulose 1,5-bisphosphate carboxylase-oxygenase
- S
Rubisco specificity to CO2/O2
- vc, vo
rates of RuBP carboxylation and RuBP oxygenation, respectively
- CO
2
relative quantum yield of CO2 assimilation
- C
maximum CO
2 under non-photorespiratory conditions
- exc
the efficiency of excitation capture by open PS II centres
- PS II
relative quantum yield of PS II electron transport 相似文献
17.
Philip L. Smith Stephanie A. Orellana Michael Field M.D. 《The Journal of membrane biology》1981,63(3):199-206
Summary Unidirectional fluxes of35SO4 across and into rabbit ileal epithelium were measured under short-circuit conditions, mostly at a medium SO4 concentration of 2.4mm. Unidirectional mucosa (m)-to-serosa (s) ands-to-m fluxes (J
ms,J
sm) were 0.456 and 0.067 moles hr–1 cm–2, respectively.J
ms was 2.7 times higher in distal ileum than in mid-jejunum. Ouabain abolished net SO4 transport (J
net) by reducingJ
ms. Epinephrine, a stimulus of Cl absorption, had no effect on SO4 fluxes. Theophylline, a stimulus of Cl secretion, reducedJ
ms without affectingJ
sm, causing a 33% reduction inJ
net. Other secretory stimuli (8-Br-cAMP, heat-stable enterotoxin, Ca-ionophore A23187) had similar effects. Replacement of all Cl with gluconate markedly reducedJ
net through both a decrease inJ
ms and an increase inJ
sm. The anion-exchange inhibitor, 4-acetoamido-4-isothiocyano-2,2-sulfonic acid stilbene (SITS), when added to the serosal side, reducedJ
ms by 94%, nearly abolishingJ
net. SITS also decreasedJ
sm by 75%. Mucosal SITS (50 m) was ineffective. 4,4-diisothiocyano-2,2-sulfonic acid stilbene (DIDS) had effects similar to SITS but was less potent. Measurements of initial rates of epithelial uptake from the luminal side (J
me) revealed the following: (1)J
me is a saturable function of medium concentration with aV
max of 0.94 moles hr–1 cm–2 and aK
1/2 of 1.3mm; (2) replacing all Na with choline abolishedJ
me; (3) replacing all Cl with gluconate increasedJ
me by 40%; (4) serosal SITS had no effect onJ
me; and (5) stimuli of Cl secretion had no effect onJ
me or increased it slightly. Determination of cell SO4 with35SO4 indicated that, at steady-state, the average mucosal concentration is 1.1 mmoles per liter cell water, less than half the medium concentration. Cell SO4 was increased to 3.0mm by adding SITS to the serosal side. Despite net transport rates greater than 1.4 Eq hr–1 cm–2, neither addition of SO4 to the SO4-free medium nor addition of SITS to SO4-containing medium altered short-circuit current. The results suggest that (1) ileal SO4 absorption consists of Na-coupled influx (symport) across the brush border and Cl-coupled efflux (antiport) across the basolateral membrane; (2) the overall process is electrically neutral; (3) the medium-to-cell Cl concentration difference may provide part of the driving force for net SO4 absorption; and (4) since agents affecting Cl fluxes (both absorptive and secretory) have little effect on SO4 fluxes, the mechanisms for their transcellular transports are under separate regulation. 相似文献
18.
Ali O Shen Z Tietjen U Martens H 《Journal of comparative physiology. B, Biochemical, systemic, and environmental physiology》2006,176(5):477-487
We have studied the transport of acetate across the isolated epithelium of sheep omasum; no net transport was observed (J
ms ≈ J
sm) under Ussing chamber conditions. Low mucosal pH (pH 6.4) significantly enhanced J
ms acetate and the transport rates of acetate increased linearly and significantly (r
2=0.99) with the luminal acetate concentration. The presence of another short chain fatty acid (propionate) did not affect J
ms acetate significantly. Neither addition of 1 mmol l−1 DIDS to the mucosal side nor HCO3 replacement caused changes of J
ms acetate; this does not support the assumption of acetate transport via anion exchange. Addition of 1 mmol l−1 amiloride to the mucosal side significantly decreased acetate fluxes at high mucosal acetate concentration (100 mmol l−1) and low pH (6.4) indicating interaction between acetate uptake in the undissociated form, intracellular release of protons and activation of Na+/H+ exchange (NHE). However, the mutual interaction between Na transport via NHE and acetate transport is asymmetric. Stimulation or inhibition of Na transport via NHE is much more pronounced than the corresponding changes of acetate fluxes. Thus, the obtained results support the conclusion that acetate is transported via simple diffusion and probably predominantly in the protonated form, thereby explaining the positive and mutual interaction between Na transport and short chain fatty acids. 相似文献
19.
Timothy J. Bradley 《Journal of morphology》1990,206(1):65-69
The ultrastructure of scalariform junctions in the Malpighian tubules of the hemipteran Rhodnius prolixus and the dipteran Aedes taeniorhynchus is described. Both autocellular and intercellular scalariform junctions are illustrated. This is the first report of scalariform junctions in the Malpighian tubules of a dipteran. When combined with previous observations by other authors, the presence of scalariform junctions has now been reported in the Malpighian tubules of insects from five orders, including ametabolous, hemimetabolous, and holometabolous forms. The cell types in which scalariform junctions were found in R. prolixus and A. taeniorhynchus differ in the direction of ion and fluid transport. The cells share the capacity to transport KCl. These same cells also possess morphological features promoting close associations of mitochondria and plasma membranes in the apical region of the cell. The possible role of scalariform junctions is discussed in light of these observations. 相似文献
20.
Photoprotective Function of Photorespiration in Several Grapevine Cultivars Under Drought Stress 总被引:4,自引:2,他引:4
Four grapevine cultivars, i.e. Cabernet Sauvignon (a member of the Western Europe cultivar group), Rizamat (a member of the East cultivar group), Red Double Taste (a hybridized cultivar from Vitis vinifera L. and V. labrusca L.), and 1103Paulsen (a hybridized rootstock), were treated by three severity orders of drought stress for 25 d. Then net photosynthetic rate (P
N), maximal photochemical efficiency (Fv/Fm), actual photochemical efficiency (PS2) of photosystem 2, total electron transport rate (JT), and electron transport flows used in carboxylation (JC) and in oxygenation (JO) reactions catalysed by ribulose-1,5-bisphosphate carboxylase/oxygenase were determined. P
N was determined again after re-watering for 2 d by gas exchange measurement. Along with the increase in severity of drought stress, P
N, Fv/Fm, PS2, JT, and JC in all four cultivars decreased. The range of decrease differed among cultivars. JO expressed various trends from cultivar to cultivar. In Rizamat that received slight and moderate drought stress, P
N evidently decreased, but JO markedly increased, thus maintaining high values of JT and PS2. Prior to the moderate drought stress, the Fv/Fm was high in Rizamat, indicating that the photodamage had not happened ahead of the moderate drought stress given. Under the severe drought stress, the photorespiration rate in Rizamat decreased by 70 %, and JT, PS2, and Fv/Fm also dropped to very low values, i.e. the photodamage of photosynthetic apparatus has taken place. This suggested that the photorespiration has consumed the excessive assimilatory power and the photo-protective function of photorespiration is very important for Rizamat. When Cabernet Sauvignon grew under drought stress, its JO decreased in a small range, thus maintaining higher values of JC, JT, PS2, and Fv/Fm; hence no serious photodamage occurred. Despite of the fact that P
N of cv. Red Double Taste decreased markedly under the slight drought stress, JO still increased under the severe drought stress. This suggests that photorespiration is important in photoprotection under drought stress. JO in cv. 1103Paulsen markedly decreased under slight stress. Accordingly, P
N, Fv/Fm, PS2, JT, and JC decreased to extremely low values. Thus photorespiration effectively protects the photosynthetic apparatus from photo-damage under drought, assists in maintaining a relatively high PS2, and helps P
N to be rapidly recovered after re-watering. 相似文献