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PubMed1978

Examination of transport equations pertaining to permeable elastic tubules such as Henle's loop.

The transport equations applicable to loops of Henle and similar elastic permeable tubules were re-examined to assess the effect of radial transport resistance in the lumen and tubule geometry on solute transport. Active transport at the wall as well as external gradients equivalent to a 2--1,000-fold concentration increase per centimeter of distance were considered. Wall permeabilities and active transport constants were varied up to 2 . 10(-2) cm/s. It is shown that for conditions applicable to the loop of Henle, resistance to radial solute transfer in the lumen is negligible, both for passive and active transmural transport with concomitant water flux, and that axial dispersion further reduces that resistance. These conclusions apply equally to conical and elliptical geometries likely to arise in loop operation. The validity of Poiseuille's equation for these geometries is discussed. Ii is concluded that the one-dimensional transport equations are a valid representation of loop operation.

Biological Transport, Active↗
PubMed1984

ADH-like effects of calcitonin on electrolyte transport by Henle's loop of rat kidney.

The effects of physiological doses of human calcitonin (HCT) on renal excretion and tubular transport of water and electrolytes were investigated in hormone-deprived rats, i.e., homozygous DI Brattleboro rats with reduced levels of circulating glucagon, parathyroid hormone, and thyrocalcitonin, as these hormones are believed, together with ADH, to stimulate the same cells of the thick ascending limb. The experimental design was similar to the one used in a preceding study aimed at determining the effects of ADH in hormone-deprived rats [C. de Rouffignac et al. Am. J. Physiol. 244 (Renal Fluid Electrolyte Physiol. 13): F156-F164, 1983]. In the present experiments, HCT consistently increased the reabsorption of Mg, Ca, and K and, to a lesser extent, Na and Cl in the loop of Henle, but phosphate transport did not rise. The urinary excretion rate of Mg and Ca fell significantly. These data are very similar to the findings obtained with ADH on hormone-deprived rats. It is concluded that, in vivo, administration of HCT 1) stimulates reabsorption of Na, Cl, Mg, Ca, and K by the thick ascending limb, and 2) consistently enhances Mg and Ca reabsorption by the whole kidney by enhancing reabsorption in the loop of Henle. The similarity of the physiological responses elicited by ADH and calcitonin on the thick ascending limb supports the hypothesis of multiple hormonal control of electrolyte transport by the thick ascending limb.

Animals↗
PubMed1983

Cellular mechanism of the action of loop diuretics on the thick ascending limb of Henle's loop.

During the passed few years the cellular mechanisms responsible for the NaCl reabsorption in the thick ascending limb of the Henle loop of mammalian nephron and of the early distal tubule of amphibian kidney have been extensively studied. From these studies a new type of secondarily active transport mechanism, i.e. the Na+--2Cl- --K+ symport has emerged. Meanwhile it has been recognized that this system is also present in many other epithelia. All these epithelia share in common that they are sensitive to the so called loop diuretics. The present article summarizes our current knowledge of how the loop diuretics, by reversible interaction with the above cotransport system, inhibit the NaCl reabsorption in the thick ascending limb of Henle's loop. It is shown that these drugs transfer the thick ascending limb cell to a state in which not only transepithelial NaCl reabsorption ceases but in which also very little energy is consumed since then K+ and Cl- "relax" to passive distribution across both cell membranes and Na+ entry into the cell is blocked.

Absorption↗
PubMed1979

Effects of lumen volume transit time and pressure on loop of Henle function.

Experiments on Henle's loops were designed to demonstrate the relationships of absorption to distal pressure, transit time, and luminal diameter. Loops of superficial nephrons in hydropenic rats, isolated from the rest of the nephron by oil or solid paraffin blocks, were microperfused at 13.6--20 nl/min. Two samples of fluid were collected from the early distal tubule--one with suction in order to lower distal pressure and reduce luminal volume, the other without suction so that the lumen was distended. Transit times were 30 +/- 2 s without and 19 +/- 2 s with suction. Proximal tubule pressure and perfusion rate were not altered by collection with suction. Absolute absorption, however, descreased from 10.6 +/- 0.4 to 8.4 +/- 0.4 nl/min (P less than 0.001). When salt transport was inhibited by 10(-4) M furosemide in the perfusate, water absorption was 7.8 +/- 0.7 nl/min without suction and 6.1 +/- 0.8 nl/min with suction (P less than 0.01). Computer simulation of Henle's loop shows that these observations cannot be explained by changes in transit time, hydrostatic pressure, or unstirred layers. The observations are simulated when radial fluxes depend on wall thickness and surface area in the descending thin limb.

Animals↗
PubMed1977

Evidence for a concentration gradient favoring outward movement of sodium from the thin loop of Henle.

Recent models of the urinary concentrating mechanism have postulated that urea in the medullary interstitium creates a transtubular concentration gradient for sodium between fluid at the end of the descending limb of Henle's loop and the medullary interstitium, favoring the passive outward movement of sodium from Henle's thin ascending limb. These experiments were designed to determine whether such a gradient normally exists. Young nondiuretic Munich-Wistar rats were prepared for micropuncture of the exposed left renal papilla. Samples of loop of Henle fluid and vasa recta plasma (assumed to reflect the composition of interstitial fluid) were obtained from adjacent sites. Loop fluid values in 21 comparisons from 18 rats (mean +/- SE) were: sodium 344 +/- 12 meq/liter; potassium, 26 +/- 2 meq/liter; osmolality, 938 +/- 37 mosmol/kg H23. Vasa recta plasma values (in corresponding units of measurement) were: sodium, 284 +/- 11; potassium, 34 +/- 2; osmolality, 935 +/- 34. Mean values of paired differences (loop fluid minus vasa recta plasma) were: delta sodium, 60 +/- 11.1 (P less than 0.001); delta potassium, -8.0 +/- 2.1 (P less than 0.001); delta osmolality, 4 +/- 16 (NS). Corrected for plasma water, the loop fluid minus vasa recta differences (in milliequivalents per kilogram H2O) were: delta sodium, 40 +/- 11.4 (P less than 0.005); delta potassium, -9.7 +/- 1.9 (P less than 0.001). We interpret these findings to indicate that in the papilla of nondiuretic rats, a significant difference in sodium concentration exists across the thin loop of Henle favoring outward movement of sodium, which confirms a key requirement of the passive models. A concentration difference for potassium in the reverse direction was also observed.

Animals↗
PubMed1993

Simulation of lithium transport along the thin segments of Henle's loop.

Although the renal clearance of lithium is widely used as an index of sodium and water delivery to the end of the proximal tubule, micropuncture studies of superficial nephrons suggest that lithium may be reabsorbed additionally in the loop of Henle. In order to examine the possibility of lithium transport in the thin loop segments of deep nephrons, we conducted a computer simulation study using a modification of the mathematical model reported previously. The data for the model were obtained from measurements of osmolality and electrolyte concentrations in systemic plasma and renal papillary interstitial fluid of control and furosemide-treated rats. Papillary interstitium/plasma Li+ concentration ratios were 3.23 and 1.48 in the control and the furosemide group, respectively. Assuming that solute concentration in the renal medulla increases as an exponential function, and applying phenomenological coefficients obtained from hamsters, the transport profiles of Li+ along the thin loop segments were calculated to be very similar to those of Na+. In the control group, about 82% of delivered Li+ was reabsorbed along the entire thin loop segments, with all segments contributing equally. This value may represent the highest possible capacity of the thin loop segments. In the furosemide group, Li+ reabsorption in the thin loop was reduced to 31% of delivered Li+. From these analyses, we conclude that an appreciable amount of lithium may be reabsorbed in the thin loop segments of long-looped nephrons by passive mechanisms. Furosemide inhibits Li+ reabsorption by dissipating the osmotic concentration gradient in the renal medulla.(ABSTRACT TRUNCATED AT 250 WORDS)

Animals↗
PubMed1986

Distribution of Henle's loops may enhance urine concentrating capability.

A simple mathematical model that was developed by Charles S. Peskin (unpublished manuscript) for a single nephron is introduced and then extended to reflect the decreasing loop of Henle population as a function of increasing medullary depth. In the model, if all the loops turn at the same depth, the concentrating capability is limited by a factor of e over plasma osmolality. However, a decreasing loop population causes a multiplier effect that greatly enhances the concentrating capability. Using the loop distribution of the rat, the model produces a sigmoidal osmolality profile similar to the profiles found in tissue-slice studies of rat kidneys. These model calculations suggest that the decreasing nephron population found in vivo may be an important factor in the concentrating mechanism of the mammalian kidney.

Animals↗
PubMed1981

Freeze-fracture studies on the thin limbs of Henle's loop in Psammomys obesus.

The thin limbs of short and long loops of Henle of the desert rodent Psammomys obesus were studied by freeze-fracture techniques. Intercellular junctions and internal membrane characteristics of thin-limb epithelia are of interest with regard to the high urine-concentrating capacity of this animal. The epithelium of the descending thin limbs of short loops is remarkably undifferentiated and equipped with multistrand tight junctions. In the descending thin limb of long loops, two segments are to be distinguished. The upper parts are characterized by an extensive cellular interdigitation and single-strand tight junctions. Thus, the paracellular pathways are prominent from two aspects: the junctional belt is elongated by interdigitation, and its apico-basal depth is shallow. The transition from the upper to the lower part appears to be abrupt, as indicated by the change in intramembrane particle density. The lower parts are characterized by a noninterdigitating epithelium with junctions consisting of few, but always more than two, strands. In addition, this thin-limb segment is characterized by regularly distributed infoldings of the basal cell membrane. The ascending thin limbs are established by an interdigitation epithelium, with junctions generally consisting of one strand. Once again, the elongated junctional belt is shallow. This study presents further evidence that remarkable species difference occur among thin-limb epithelia, especially concerning the descending thin limbs of long loops. Those differences may well explain discrepant functional findings concerning the transport properties of this segment in various species.

Animals↗
PubMed2002

Bicarbonate reabsorption and NHE-3 expression: abundance and activity are increased in Henle's loop of remnant rats.

BACKGROUND: The bulk of bicarbonate reabsorption along the loop of Henle (LOH) is localized at the level of the thick ascending limb (TAL) and is mainly dependent on the presence of luminal Na+-H+ exchanger (NHE-3). We investigated whether the reduction of renal mass is associated with alterations in LOH bicarbonate transport coupled to changes in NHE-3 gene expression and in vivo activity. METHODS: Sham-operated and remnant rats (4/6 nephrectomy) were studied 15 days after the surgery. To measure net bicarbonate reabsorption (JHCO3-) superficial loops were perfused by in vivo micropuncture. Perfusate was an end-like proximal solution containing 3H-methoxy-inulin. NHE-3 gene expression was quantified by competitive PCR using an internal standard of cDNA that differed from the wild-type NHE-3 by a deletion of 76 bp. Western blot experiments were performed on TAL suspension using anti-NHE-3 antibodies. RESULTS: At various LOH bicarbonate loads, JHCO3- was constantly larger in remnant rats as compared to sham-operated animals. NHE-3 mRNA abundance was estimated to be 0.339 +/- 0.031 attomoles (amol)/ng-1 total RNA in sham-operated (N = 5) and it increased to 0.465 +/- 0.023 in remnant rats (N = 5, P < 0.01). Western blot experiments showed a significant increase of NHE-3 protein abundance in TAL of remnant rats as compared to sham-operated animals. Finally, by means of a specific NHE-3 inhibitor, S-3226, in vivo microperfusion experiments demonstrated that NHE-3 in vivo activity along the LOH was substantially increased in remnant rats in addition to the non-NHE-3 bicarbonate transport. CONCLUSIONS: These data indicate that the reduction of renal mass increases mRNA, protein abundance and in vivo activity of NHE-3 along the TAL. This may explain, at least in part, the augmented transepithelial bicarbonate transport along the LOH. Such an effect will counterbalance the increased glomerular bicarbonate load, thus preventing urinary bicarbonate loss and mitigating the ensuing metabolic acidosis.

Acid-Base Equilibrium↗
PubMed2001

Chloride channels in the loop of Henle.

Cl- transport in the loop of Henle is responsible for reclamation of 25-40% of the filtered NaCl load and for the formation of dilute urine. Our understanding of the physiologic and molecular mechanisms responsible for Cl- reabsorption in both the thin ascending limb and thick ascending limb of Henle's loop has increased greatly over the last decade. Plasma membrane Cl- channels are known to play an integral role in transcellular Cl- transport in both the thin and thick ascending limbs. This review focuses on the functional characteristics and molecular identities of these Cl- channels, as well as the role of these channels in the pathophysiology of disease.

Amino Acid Sequence↗
PubMed1983

Effect of "high ceiling" diuretics on active salt transport in the cortical thick ascending limb of Henle's loop of rabbit kidney. Correlation of chemical structure and inhibitory potency.

The group of "high ceiling" diuretics consists of a variety of chemically different potent diuretic and saluretic substances. Appart from a few exemptions direct evidence for an action of these substances in the thick ascending limb of the loop of Henle (TAL) is still lacking. For furosemide, we have reported recently that it inhibits most likely the Na+-2 Cl--K+ cotransport system present in the lumen membrane of the TAL. The present study tests: 1. whether other "high ceiling" diuretics have a similar site and mode of action, and 2. how modifications of the furosemide molecule alter the inhibitory potency. Isolated cortical TAL (cTAL) segments (n = 185) of rabbit kidneys were perfused in vitro. The equivalent short circuit current (Isc = transepithelial PD/transepithelial resistance), as a measure of active salt transport was correlated to the dose of 64 substances. Several diuretics, such as 2-aminomethyl-4-(1,1-dimethyl-ethyl)-6-iodophenol hydrochloride (MK 447), hydrochlorothiazide, muzolimine, etozoline, tizolimide, amiloride, and triamterene were ineffective both from the lumen and basolateral side at concentrations as high as 10(-4) - 10(-3) mol X 1(-1). The phenoxyacetic acids ethacrynic acid, indacrinone (MK 196), and to less an extend tienilic acid were inhibitory active. They differed from furosemide in one or more of the following criteria: delayed onset, incomplete reversibility, stronger action from the bath, different slope of the dose response curve. Similarly, 1-ozolinone acted stronger from the bath. In contrast, the diuretics of the furosemide type and related compounds (bumetanide and piretanide) showed rapid onset and complete reversibility of inhibition. These substances acted stronger from the lumen. The individual positions in the benzyl ring of the diuretics were differently affected by substitutions, leading to parallel shifts in the dose response curves with halfmaximal inhibition at concentrations ranging between 8 X 10(-8) to greater than 10(-4) mol X 1(-1). For these substances the calculated Hill coefficients were close to unity: 0.96 +/- 0.05. We conclude that the so called "high ceiling" or "loop" diuretics consist of at least 3 groups: 1. drugs that do not interfere with the active salt transport in the cTAL segment, 2. drugs that interfere by so far not characterised mechanisms, and 3. drugs of the furosemide type which inhibit the Na+-2 Cl--K+ cotransport system in the lumen membrane of the cTAL segment.

Animals↗
PubMed1982

A cytotoxin of Pseudomonas aeruginosa acts directly on the cortical thick ascending limb of Henle's loop of rabbit kidney.

The present study examines directly the effect of a cytotoxin of Pseudomonas aeruginosa on the in vitro perfused rabbit cortical thick ascending limb of the loop of Henle (cTAL). 25 cTAL segments were perfused at high rate. The open circuit transepithelial electrical PD (PDte) and the specific electrical transepithelial resistance (Rt) were recorded continuously. From PDte/Rt the equivalent short circuit current (Isc) was calculated. The Isc was 214 +/- 30 mu A.cm-2 under control conditions, and decreased significantly to 74 +/- 34 mu A.cm-2 60 s after the addition of toxin (2 mg.1(-1)) to the lumen perfusate. Microscopic observation and photographs taken at that time clearly indicated swelling of the cTAL cells. Thereafter inhibition of active transport proceeded further, Rt fell progressively, and cells started to desquamate from the basement membrane. This effect of the toxin was dose dependent, and was half maximal at approximately 1.2 mg.1(-1). From the bath side the effect was less marked and higher doses of toxin had to be used (half maximal effect at 5 mg.1(-1)). We conclude that this toxin of Pseudomonas aeruginosa exerts its toxic effect on the cTAL segment by increasing primarily the permeability of the lumen membrane.

Animals↗
PubMedDate not supplied

Acid-base transport in Henle's loop: the effects of reduced renal mass and diabetes.

The loop of Henle (LOH) is an important site of renal acidification. Using the in vivo microperfusion technique of LOH combined with quantitative polymerase chain reaction (PCR) performed on isolated thick ascending limbs (TAL), we demonstrated that the Na + -H + exchanger is the main transport mechanism involved, although a small, but significant contribution from the H+-ATPase also occurs. Among the various Na+-H+ exchanger isoforms we have evidenced that NHE3 is expressed and functionally active along the TAL. Since the LOH is exposed to osmotic stress, bicarbonate transport was also measured under medullary hypotonicity conditions, which led to the stimulation of bicarbonate reabsorption. We demonstrated that the LOH can participate in the tubular adaptation to an increased filtered bicarbonate load by increasing net LOH bicarbonate transport. In this setting, at the molecular level, mRNA and protein abundance of NHE3 were also stimulated, and coincided with an increase in NHE3 activity. Finally, NHE3 expression and abundance was highly stimulated in the early phase of diabetes, which is characterized by increased glomerular filtration rate (GFR).

Acid-Base Equilibrium↗
PubMed1998

A decrease in renal medullary tonicity stimulates anion transport in Henle's loop of rat kidneys.

To investigate the effect of reduction in renal medulla osmolality on loop of Henle (LOH) net bicarbonate reabsorption, clearance and microperfusion experiments were performed on Sprague-Dawley rats. The decrease of renal medulla osmolality was induced by intravenous infusion of either a large dose of mannitol (mannitol protocol) or a hypotonic solution (hypotonic protocol) delivered at a rate to match the sodium and bicarbonate load of the control period. During the mannitol protocol, clearance data demonstrated a rise in glomerular filtration rate (GFR), renal plasma flow, urine pH, and fractional bicarbonate excretion. On the contrary, microperfusion experiments, performed in the absence of mannitol in the tubular perfusate, revealed a significant increase both in the absolute and fractional LOH bicarbonate transport. During the hypotonic protocol, there was a decrease in GFR, associated with an increase in fractional excretion of bicarbonate. In the microperfusion experiments, hypotonic saline, similar to mannitol, stimulated absolute and fractional LOH bicarbonate transport. Net reabsorption of chloride, measured under the same experimental conditions, was also found to be activated. Therefore, the intravenous infusion of hypotonic solution affected the LOH transepithelial net reabsorption of both bicarbonate and chloride. We hypothesize that the increase in the transport rate of these two anions, along the same segment and in similar experimental conditions, may be mediated, at least in part, by decreased medullary tonicity, which is one factor common both to hypertonic mannitol and hypotonic saline infusion.

Animals↗
PubMed1986

Development of differentiated characteristics in cultured kidney (thick ascending loop of Henle) cells.

This study describes the characterization of epithelial cells in culture following their isolation from the thick ascending limb of Henle's loop of rabbit kidney, by enzymatic digestion and subsequent purification using density gradient centrifugation. In culture, these cells expressed a variety of morphological, enzymatic and functional parameters expected of such cells in vivo. These cells were polarised, formed tight junctions and exhibited considerable lateral interdigitation between adjacent cells. They also developed characteristically high levels of activity of Na,K-ATPase, comparable to those seen in freshly isolated cells, and also expressed the functionally important Na,K,Cl-co transport system. The development of these systems in culture, however, was not coincident and their activities were reduced upon extended culture. The ability of these cells to develop and express differentiated characteristics in culture indicates that cells derived from defined kidney cell populations should provide valuable models for the study of the factors involved in the development and regulation of kidney cell type-specific characteristics.

Alkaline Phosphatase↗
PubMed2006

Glucocorticoid impairs growth of kidney outer medulla and accelerates loop of Henle differentiation and urinary concentrating capacity in rat kidney development.

In the rat, urinary concentrating ability develops progressively during the third postnatal (P) week and nearly reaches adult level at weaning (P21) governed by a rise in circulating glucocorticoid. Elevated extracellular osmolality can lead to growth arrest of epithelial cells. We tested the hypothesis that supranormal exposure of rat pups to glucocorticoid before the endogenous surge enhances urinary concentrating ability but inhibits renomedullary cell proliferation. Proliferating-cell nuclear antigen (PCNA)-positive cells shifted from the nephrogenic zone in the first postnatal week to Tamm-Horsfall-positive thick ascending limb (TAL) cells at the corticomedullary junction at P10-14. Renal PCNA protein abundance was stable in the suckling period and decreased 10-fold after weaning. Renal PCNA protein abundance decreased in response to dexamethasone (DEXA; 100 microg x kg(-1) x day(-1), P8-12). Prolonged administration of DEXA (P1-P11) reduced selectively the area and thickness of the outer medulla and the number of PCNA-positive cells. DEXA (P8-12) increased urinary and papillary osmolality in normohydrated and water-deprived pups and led to osmotic equilibrium between interstitium and urine, whereas apoptotic and GADD153-positive cells increased in the inner medulla. TAL-associated NaCl transporters Na-K-2Cl cotransporter, Na-K-ATPase-alpha(1), Na/H exchanger type 3, and ROMK increased significantly at weaning and in response to DEXA. We conclude that a low level of circulating glucocorticoid is permissive for proliferation of Henle's loop and the outer medulla before weaning. A reduced papillary tonicity is a crucial factor for the reduced capacity to concentrate urine during postnatal kidney development. We speculate that supranormal exposure to glucocorticoid in the suckling period can alter kidney medullary structure and function permanently.

Aging↗
PubMed1980

Loop of Henle during the water-to-antidiuresis transition in Brattleboro rats.

A mathematical model of the renal medulla predicted patterns of NaCl and urea concentration and flow rate when a rat undergoing water diuresis receives antidiuretic hormone (ADH), We tested these predictions with micropuncture and clearance experiments in Brattleboro rats lacking endogenous ADH. The model successfully predicted urine flow rate, urea excretion rate, and ascending vasa recta (AVR) solute concentration patterns. A novel result the model correctly foresaw was that AVR urea concentration lagged behind NaCl, because ADH temporarily interrupted urea recycling. The model suggested that this lag would be carried over to descending limb of Henle's loop (DLH) fluid only if DLH permeabilities allowed solutes as well as water to move across the DLH wall, but not if solute movement was negligible. Experimental results showed the lag, therby suggesting significant solute entry into DLH fluid during the transition, and, by extension, in other conditions as well, As shown in a companion paper (Am. J. Physiol. 239 (Renal Fluid Electrolyte Physiol. 8): F57-F71, 1980), solute permeation across DLH limits hypertonic urine formation in the inner medulla unless thin ascending limbs perform active transport of NaCl.

Animals↗
PubMed1995

Unique localization of mRNA encoding plasma membrane Ca2+ pump isoform 3 in rat thin descending loop of Henle.

In our previous study of plasma membrane Ca2+ pump (PMCA) isozymes in the rat kidney, we found that the mRNA coding isoform PMCA3 was detected primarily in the outer medullary zone of rat kidney tissue. We now investigated the location of the mRNAs coding for isoforms PMCA3 and PMCA4 of Ca2+ pump in the nephron segments that are present in outer medullary parenchyma using the method of reverse transcription and polymerase chain reaction. In mRNA extracted from whole dissected outer medulla we detected mRNA encoding three splice variants (a, b, and c) of isoform PMCA3; isoform PMCA4 was found in the outer medulla almost exclusively as variant b. Analysis of mRNA from microdissected tubule segments show that proximal straight tubules (PST), medullary thick ascending limbs, outer medullary collecting ducts, and descending thin limb of Henle's loop (DTL) all contained mRNA for isoform 4b. In contrast, the mRNA encoding isoform 3 was detected exclusively in DTL and not in other nephron segments. The unique presence of isoform 3 in DTL is rather surprising, since the specific role of this nephron segment in vectorial Ca2+ transport or in intracellular Ca2+ signaling is not known. The data suggest that isoform 3 in cells of the DTL may have a hitherto unrecognized specific role in Ca2+ signaling or transport of Ca2+, which is distinct from the role of the isoforms of the PMCA in all other nephron segments.

Animals↗