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Glomerular Filtration

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Abstract

The sections in this article are:

1 Glomerular Filtration of Water
1.1 Composition of the Glomerular Ultrafiltrate
1.2 Determinants of Glomerular Filtration of Water
1.3 Control of Glomerular Filtration by Hormones and Vasoactive Substances
1.4 Angiotensin II
1.5 Other Vasoconstrictor Substances
1.6 Vasodilator Substances
1.7 Other Hormones and Vasoactive Substances
1.8 Autoregulation of Glomerular Filtration Rate
1.9 Neural Regulation of Glomerular Filtration Rate
1.10 Theoretical Models of Glomerular Ultrafiltration
2 Glomerular Permselectivity
2.1 Experimental Approaches
2.2 Effects of Molecular Properties on Macromolecule Filtration
2.3 Theory of Glomerular Permselectivity
2.4 Glomerular Membrane Parameters
2.5 Determinants of Macromolecule Filtration
2.6 Estimation of Filtration Pressure from Sieving Data
2.7 Altered Glomerular Permselectivity
Figure 1. Figure 1.

Hydraulic and colloid osmotic pressure profiles along idealized glomerular capillaries in hydropenic and euvolemic rats. Values shown are mean values from studies listed in Tables and . ΔP = PGC‐PT and ΔΠ = ΠGC‐ΠT, where PGC and PT are hydraulic pressures in glomerular capillary and Bowman's space, respectively, and ΠGC and ΠT are corresponding colloid osmotic pressures. Because ΠT is negligible, ΔΠ essentially equals ΠGCPUF is the ultrafiltration pressure at any point. Area between ΔP and ΔΠ curves represents net ultrafiltration pressure, (PUF). Curves A and B in left panel represent two of many possible ΔΠ profiles under conditions of filtration pressure equilibrium. Curve D, disequilibrium; line C, hypothetical linear ΔΠ profile; QA, glomerular plasma flow ra])G SNGFR, single nephron glomerular filtration rate.

Figure 2. Figure 2.

Relationship between glomerular capillary ultrafiltration coefficient(Kf) and body weight in rats. Each point represents mean values obtained from one study. Open squares and open circles indicate data obtained from Munich‐Wistar rats and include only those studies in which animals were at or near filtration pressure disequilibrium [euvolemic or plasma‐expanded male or male plus female rats from Tables and , (squares) or hydropenic and euvolemic female rats at disequilibrium from Tables and (circles)]. Thus Kf values are unique rather than miminum estimates. Open triangle, study in hydropenic Holtzman rats at equilibrium and thus is minimum value . Solid line, representing open symbols, is given by equation y = 0.021(x) −0.47 (r = 0.67). Filled squares, values obtained in euvolemic Munich‐Wistar rats fasted for 12–24 h (Table ). All other filled symbols were obtained in species without surface glomeruli and hence relied on stop‐flow pressure measurements to estimate (PGC). Included are data from Sprague‐Dawley rats (filled circles) spontaneous hypertensive rats (half‐filled circles), and Wistar‐Kyoto rats (filled triangles) from Table . Dashed line, denoting best fit to filled symbols, is given by equation y = 0.0074(x) + 0.25 (r = 0.80, P<0.01 vs. solid line).

Figure 3. Figure 3.

Relationships of single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF) to isolated changes in glomerular plasma flow rate. Because SNGFR and SNFF can also be affected by changes in 〈ΔP〉, Kf, or ΠA, only studies in which 〈ΔP〉 = 35–40 mm Hg, minimum Kf ≥ 3 nl/min · mm Hg, and Π;A≅18–20 mm Hg are included. Included are studies from Tables – as well as studies under a variety of physiological conditions that fell within these limits . Solid lines denote model predictions based on mean values of 〈ΔP〉, Kf, and ΠA observed in these studies [〈ΔP〉 = 36.4 mm Hg, ΠA = 18.8 mm Hg, and Kf = 4.9 nl/(min · mm Hg)].

Figure 4. Figure 4.

Relationship between single nephron glomerular filtration rate (SNGFR) and glomerular plasma flow rate. Each point represents mean values obtained in one study. Open circles, Munich‐Wistar rats presented in Tables –; closed triangles, Wistar‐Kyoto rats; open triangles, Holtzman tats; closed circles, spontaneous hypertensive rats presented in Table ; open squares, Sprague‐Dawley rats (presented in Table (and refs. ); closed squares, dogs presented in Table ; half‐closed circles, calculated values of SNGFR and glomerular plasma flow rate in humans based on dividing GFR and renal plasma flow by 1 million (the total number of nephrons) for a variety of studies (see Fig. ).

Figure 5. Figure 5.

Relationship between whole‐kidney glomerular filtration rate (GFR) and renal plasma flow rate in dogs (values expressed per kidney) and humans [values shown for one kidney and expressed as ml/(min · 1.73 m2)]. Human data: closed circles, studies conducted following a low‐protein meal , on alternate‐day protein feeding , or following overnight fast (82, 112, 641. Administration of a protein meal (open circles or half‐closed circles) resulted in increased GFR and renal plasma flow in three studies . Lines connect values in same study. Closed squares, no information given regarding diets ; open square, cardiac failure patients . Dog studies: open triangles, variations in GFR and renal plasma flow resulted primarily from variations in renal perfusion pressure, including either normal or reduced arterial pressures; closed triangles, values obtained following hyperoncotic plasma expansion ; response to a saline load .

Figure 6. Figure 6.

Relationship of single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF) to variations in the transcapillary hydraulic pressure gradient ((ΔP)). Each point denotes mean value from one study. To minimize variations in SNGFR and SNFF caused by differences in ΠA and Kf, all studies selected had ΠA values between 16 and 20 mm Hg and had normal values of Kf ≥ 3 nl/(min · mm Hg). Data are compared at two selected ranges of glomerular plasma flow rate (QA) (130–170 nl/min; mean, 147 ± 2 nl/min or 40–70 nl/min; mean, 60 ± 3 nl/min) in which a relatively wide range of 〈ΔP〉 values were obtained. Low QA data were from several studies . High QA data include all studies listed in Tables and that meet guidelines as well as other studies . Dashed lines, model predictions for low QA studies using mean values of (17 mm Hg) and QA (60 nl/min) obtained in these studies, together with mean (unique) value of Kf in high QA experiments, assuming that variations in QA per se do not alter Kf; solid lines, model predictions for high QA data from rats using mean values of ΠA (18 mm Hg), QA (147 nl/min), and Kf [5.4 nl/(min · mm Hg)] obtained in these studies.

Figure 7. Figure 7.

Effects of selective alterations in Kf on single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF). Only studies in which a unique value of Kf was obtained (i.e., ΠE/〈ΔP〉 ≤ 0.95) are included. Data are shown for two ranges of QA: open circles, 90–125 nl/min (mean, QA = 100nl/min); and closed circles, 150–180 nl/min (mean, QA≅ 170 nl/min). For low QA studies, ΠA was limited to 16–20 mm Hg and 〈ΔP〉 from 36 to 42 mm Hg. High flow studies had similar ranges of ΠA (17–22 mm Hg) and 〈ΔP〉 (36–43 mm Hg). Model predictions shown by dashed lines are for low QA values (data from refs. ); solid lines represent predictions for high QA values (data from refs. ).

Figure 8. Figure 8.

Theoretical relationships between plasma oncotic pressure (ΠA) and both single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF). Dashed line, model predictions for euvolemic Munich‐Wistar rats based on values of QA, 〈ΔP〉, and Kf of 151 nl/min, 35 mm Hg, and 5.6 nl/(min · mm Hg), respectively (Table ); solid line, hydropenia in which QA and 〈ΔP〉 were 79 nl/min and 34 mm Hg (Table ), respectively. Because only a minimum value of Kf can be determined in hydropenic rats (ΠE = 〈ΔP〉, Table ), the unique value of 5.6 nl/min · mm Hg determined in euvolemic rats was used.

Figure 9. Figure 9.

Relationship between Kf and plasma protein concentration (CA). Closed circles, mean values (± S.E.M.) obtained by Baylis et al. under a variety of conditions, with and without volume expansion, designed to alter CA; closed square, a study in which an isovolemic reduction in hematocrit produced filtration pressure disequilibrium without volume expansion ; closed triangles, values obtained in rats with isoncotic plasma volume expansion in which QA was varied by altering renal perfusion pressure by aortic constriction or carotid occlusion .

Modified from Baylis et al.
Figure 10. Figure 10.

Hydraulic pressure profile in rat kidney. Open circles, values obtained in studies of euvolemic Munich‐Wistar rats (Table ); closed circles, studies of hydropenic rats (Table ); closed triangles, values obtained by Casellas and Navar in Sprague‐Dawley rats in unique population of juxtamedullary nephrons at inside cortical surface apposed to pelvic lining and arcuate veins. Hydraulic pressures can be obtained in proximal (Early a.a.) and distal (Late a.a.) portions of afferent arteriole and in proximal (Early e.a.) and distal (Late e.a.) segments of efferent arteriole, and in peritubular capillaries (Pc), interlobular veins (I.V.), and renal vein (R.V.). For micropuncture experiments, systemic arterial pressure ( ) averaged — 110 mm Hg, whereas in study of Casellas and Navar arcuate artery (Arc. Art.) was perfused with whole blood at 96 mm Hg. , hydraulic pressure in glomerular capillaries.

Figure 11. Figure 11.

Pathways of renin synthesis and processing in mouse submaxillary gland. A: proposed sites of cleavage (upward arrows) and Mrs of each compound (far right column). B: synthesis, packaging into granules, and secretion of renin.

From Pratt et al. (497a
Figure 12. Figure 12.

Cellular control of renin release. Occupancy of adenylate cyclase (AC)‐linked receptors by agonists [norepinephrine (NE), histamine (H), and prostaglandins (PG)] results in stimulation of cAMP production and phosphorylation of intermediate protein kinase (PK) that stimulates (+) renin secretion. N, guanine nucleotide regulatory proteins specific for each agonist. Though not shown, angiotensin II (AΠ) and vasopressin (AVP) also act through N (or G) proteins to increase calcium influx as well as calcium release from intracellular storage sites. Membrane depolarization (Dep.) can also increase Ca2+ influx through voltage‐sensitive channels. Increased [Ca2+], leads to formation of the calcium‐calmodulin (Ca‐CM) complex that inhibits (−) renin release. Decreases in [Ca2+]i uncouples the Ca‐CM complex, thus de‐inhibiting renin release. RAΠ, Ravp, RH2, Rβ. and RPG; receptors for angiotensin II, arginine vasopressin, histamine (H2‐type), norepinephrine (β‐type), and prostaglandins, respectively.

From Ballermann et al.
Figure 13. Figure 13.

Effects of intravenous infusion of angiotensin II (200–600 ng/kg/min) determinants of glomerular ultrafiltration in Munich‐Wistar rat. Left, data obtained when renal perfusion pressure ( ) was allowed to increase; right, results obtained when increase in was prevented. Data are means ± S.E. SNGFR, single nephron glomerular filtration rate.

From Myers et al.
Figure 14. Figure 14.

Role of angiotensin II in mediating altered renal function of congestive heart failure induced by myocardial infarction (M.I.). Control euvolemic sham‐operated rats were studied during two periods; before (open circles) and during (filled circles) infusion of angiotensin I‐converting enzyme inhibitor teprotide [SQ20881; ]. SNGFR, single nephron glomerular filtration ra])G SNFF, single nephron filtration fracti])R , renal perfusion pressure.

Figure 15. Figure 15.

Mechanism of action of angiotensin II (AT II) in contraction of renal vasculature and glomerular mesangial cells. R, receptor; Gαβγ, guanine nucleotide‐binding regulatory protein with its three subunits, α, β, and γ; GTP, guanine trisphospha])G PLC, phospholipase C; PlP2, phosphatidylinositol‐4,5‐bisphospha])G IP3, inositol‐1,4,5‐trisphospha])G DAG, 1,2‐diacylglycerol; Ca2+, intracellular free calcium; PKC, protein kinase C; LC, lipocortin; PLA2, phospholipase A2; PL, membrane phospholipids; AA, arachidonic acid; PGE2, prostaglandin E2; cAMP, cyclic adenosine monophospha])G cAMP‐PK, cAMP‐dependent protein kinase; MLCK, myosin light chain kinase; P‐MLCK, phosphorylated MLCK; MLCK‐CAM, MLCK‐calcium‐calmodulin active complex; Pi, phospha])G *, activated form of enzyme. Parts A, B, and C in upper panel show sequence of events leading from occupancy of the receptor to activation of PLC. Lower panel shows sequence of events occurring in cells after PLC activation.

Figure 16. Figure 16.

Lipoxygenase pathway for biosynthesis of vasoactive leukotrienes and cyclooxygenase pathway for synthesis and degradation of prostaglandins, prostacyclin, and thromboxane A2 (TxA2).

Figure 17. Figure 17.

Effects of PGE2, PGI2, and dibutyryl (DB)‐cAMP on the determinants of glomerular filtration in rat. Data include effects on single nephron glomerular filtration rate (SNGFR), panel A; glomerular plasma flow rate (QA), panel B; mean glomerular capillary hydraulic pressure ( ), panel C; glomerular ultrafiltration coefficient (Kf), panel D; and sum of the pre‐ and postglomerular resistances (RTA), panel E. Euvolemic control animals were infused with cyclooxygenase inhibitors meclofenamate (Meclo.) or indomethacin (Indo.) or a cyclooxygenase inhibitor plus angiotensin II competitive antagonist saralasin. Effects of PGE2, PGI2, and DB‐cAMP were first examined during inhibition of endogenous prostaglandin production by Indo. or Meclo. and then with the added inhibition of endogenous angiotensin II by saralasin.

Data are from Schor et al.
Figure 18. Figure 18.

Effects of reduced mean arterial pressure on the measured determinants of glomerular ultrafiltration in hydropenic rats (A) and in mildly plasma‐expanded Munich‐Wistar rats before (B) and during (C) the administration of smooth muscle relaxant papaverine. Data (from ref. and ) are shown as means ± 1 S.E.

From Baylis and Brenner
Figure 19. Figure 19.

Filtrate‐to‐plasma concentration ratio (θ) as function of molecular size for dextran sulfate (DS), neutral dextran (D), and diethylaminoethyl dextran (DEAE). Symbols, mean values ± S.E. measured by Bohrer et al. in normal hydropenic Munich‐Wistar rats; curves are theoretical calculations based on values of QA, 〈ΔP〉, and CA reported in those studies with Kf = 4.8 nl/(min · mm Hg), ro = 47 Å, and Cm= 165 mEq/liter.

From Deen et al.
Figure 20. Figure 20.

Schematic of macromolecular solute (represented as sphere of radius, rs) moving through membrane pore (of radius ro and length l). λ = rs/ro.

Figure 21. Figure 21.

Hindrance factors for convection (W) and diffusion (H) based on hydrodynamic theory for movement of uncharged spherical molecules in cylindrical pores.

From Deen et al.
Figure 22. Figure 22.

Filtrate‐to‐plasma concentration ratio (θ) as function of molecular charge (z) molecular radius (rs). Results are shown for two values of membrane fixed charge concentration (Cm). Other inputs were QA = 75 nl/min, 〈ΔP> = 35 mm Hg, CA = 5.7 g/dl, Kf = 4.8 nl/(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen et al.
Figure 23. Figure 23.

Filtrate‐to‐plasma concentration ratio (θ) for serum albumin as function of membrane fixed charge concentration (Cm) and effective molecular charge (z). Other inputs were QA = 75 nl/min, 〈ΔP〉 = 35 mm Hg, CA = 5.7 g/dl, Kf = 4.8 nl/(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen and Satvat
Figure 24. Figure 24.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and glomerular plasma flow rate (QA). Calculations assumed Cm = 165 mEq/liter, rs = 30 Å, 〈ΔP〉 = 35 mm Hg, CA, = 5.7g/dl, Kf = 4.8 nl/‐(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen and Satvat
Figure 25. Figure 25.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and transmembrane hydraulic pressure difference 〈ΔP〉. Input quantities were Cm=165 mEq/liter, rs = 30Å, QA = 75 nl/min, CA = 5.7g/dl, Kf = 4.8 nl/min · mm Hg), and ro = 50Å.

From Deen and Satvat
Figure 26. Figure 26.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and ultrafiltration coefficient(Kf). Effective pore radius is assumed to remain constant. Input quantities were Cm=165 mEq/liter, rs = 30 Å, QA = 75 nl/min, 〈ΔP〉 = 35 mm Hg, CA = 5.7g/dl, and ro = 50 A.

From Deen and Satvat


Figure 1.

Hydraulic and colloid osmotic pressure profiles along idealized glomerular capillaries in hydropenic and euvolemic rats. Values shown are mean values from studies listed in Tables and . ΔP = PGC‐PT and ΔΠ = ΠGC‐ΠT, where PGC and PT are hydraulic pressures in glomerular capillary and Bowman's space, respectively, and ΠGC and ΠT are corresponding colloid osmotic pressures. Because ΠT is negligible, ΔΠ essentially equals ΠGCPUF is the ultrafiltration pressure at any point. Area between ΔP and ΔΠ curves represents net ultrafiltration pressure, (PUF). Curves A and B in left panel represent two of many possible ΔΠ profiles under conditions of filtration pressure equilibrium. Curve D, disequilibrium; line C, hypothetical linear ΔΠ profile; QA, glomerular plasma flow ra])G SNGFR, single nephron glomerular filtration rate.



Figure 2.

Relationship between glomerular capillary ultrafiltration coefficient(Kf) and body weight in rats. Each point represents mean values obtained from one study. Open squares and open circles indicate data obtained from Munich‐Wistar rats and include only those studies in which animals were at or near filtration pressure disequilibrium [euvolemic or plasma‐expanded male or male plus female rats from Tables and , (squares) or hydropenic and euvolemic female rats at disequilibrium from Tables and (circles)]. Thus Kf values are unique rather than miminum estimates. Open triangle, study in hydropenic Holtzman rats at equilibrium and thus is minimum value . Solid line, representing open symbols, is given by equation y = 0.021(x) −0.47 (r = 0.67). Filled squares, values obtained in euvolemic Munich‐Wistar rats fasted for 12–24 h (Table ). All other filled symbols were obtained in species without surface glomeruli and hence relied on stop‐flow pressure measurements to estimate (PGC). Included are data from Sprague‐Dawley rats (filled circles) spontaneous hypertensive rats (half‐filled circles), and Wistar‐Kyoto rats (filled triangles) from Table . Dashed line, denoting best fit to filled symbols, is given by equation y = 0.0074(x) + 0.25 (r = 0.80, P<0.01 vs. solid line).



Figure 3.

Relationships of single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF) to isolated changes in glomerular plasma flow rate. Because SNGFR and SNFF can also be affected by changes in 〈ΔP〉, Kf, or ΠA, only studies in which 〈ΔP〉 = 35–40 mm Hg, minimum Kf ≥ 3 nl/min · mm Hg, and Π;A≅18–20 mm Hg are included. Included are studies from Tables – as well as studies under a variety of physiological conditions that fell within these limits . Solid lines denote model predictions based on mean values of 〈ΔP〉, Kf, and ΠA observed in these studies [〈ΔP〉 = 36.4 mm Hg, ΠA = 18.8 mm Hg, and Kf = 4.9 nl/(min · mm Hg)].



Figure 4.

Relationship between single nephron glomerular filtration rate (SNGFR) and glomerular plasma flow rate. Each point represents mean values obtained in one study. Open circles, Munich‐Wistar rats presented in Tables –; closed triangles, Wistar‐Kyoto rats; open triangles, Holtzman tats; closed circles, spontaneous hypertensive rats presented in Table ; open squares, Sprague‐Dawley rats (presented in Table (and refs. ); closed squares, dogs presented in Table ; half‐closed circles, calculated values of SNGFR and glomerular plasma flow rate in humans based on dividing GFR and renal plasma flow by 1 million (the total number of nephrons) for a variety of studies (see Fig. ).



Figure 5.

Relationship between whole‐kidney glomerular filtration rate (GFR) and renal plasma flow rate in dogs (values expressed per kidney) and humans [values shown for one kidney and expressed as ml/(min · 1.73 m2)]. Human data: closed circles, studies conducted following a low‐protein meal , on alternate‐day protein feeding , or following overnight fast (82, 112, 641. Administration of a protein meal (open circles or half‐closed circles) resulted in increased GFR and renal plasma flow in three studies . Lines connect values in same study. Closed squares, no information given regarding diets ; open square, cardiac failure patients . Dog studies: open triangles, variations in GFR and renal plasma flow resulted primarily from variations in renal perfusion pressure, including either normal or reduced arterial pressures; closed triangles, values obtained following hyperoncotic plasma expansion ; response to a saline load .



Figure 6.

Relationship of single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF) to variations in the transcapillary hydraulic pressure gradient ((ΔP)). Each point denotes mean value from one study. To minimize variations in SNGFR and SNFF caused by differences in ΠA and Kf, all studies selected had ΠA values between 16 and 20 mm Hg and had normal values of Kf ≥ 3 nl/(min · mm Hg). Data are compared at two selected ranges of glomerular plasma flow rate (QA) (130–170 nl/min; mean, 147 ± 2 nl/min or 40–70 nl/min; mean, 60 ± 3 nl/min) in which a relatively wide range of 〈ΔP〉 values were obtained. Low QA data were from several studies . High QA data include all studies listed in Tables and that meet guidelines as well as other studies . Dashed lines, model predictions for low QA studies using mean values of (17 mm Hg) and QA (60 nl/min) obtained in these studies, together with mean (unique) value of Kf in high QA experiments, assuming that variations in QA per se do not alter Kf; solid lines, model predictions for high QA data from rats using mean values of ΠA (18 mm Hg), QA (147 nl/min), and Kf [5.4 nl/(min · mm Hg)] obtained in these studies.



Figure 7.

Effects of selective alterations in Kf on single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF). Only studies in which a unique value of Kf was obtained (i.e., ΠE/〈ΔP〉 ≤ 0.95) are included. Data are shown for two ranges of QA: open circles, 90–125 nl/min (mean, QA = 100nl/min); and closed circles, 150–180 nl/min (mean, QA≅ 170 nl/min). For low QA studies, ΠA was limited to 16–20 mm Hg and 〈ΔP〉 from 36 to 42 mm Hg. High flow studies had similar ranges of ΠA (17–22 mm Hg) and 〈ΔP〉 (36–43 mm Hg). Model predictions shown by dashed lines are for low QA values (data from refs. ); solid lines represent predictions for high QA values (data from refs. ).



Figure 8.

Theoretical relationships between plasma oncotic pressure (ΠA) and both single nephron glomerular filtration rate (SNGFR) and single nephron filtration fraction (SNFF). Dashed line, model predictions for euvolemic Munich‐Wistar rats based on values of QA, 〈ΔP〉, and Kf of 151 nl/min, 35 mm Hg, and 5.6 nl/(min · mm Hg), respectively (Table ); solid line, hydropenia in which QA and 〈ΔP〉 were 79 nl/min and 34 mm Hg (Table ), respectively. Because only a minimum value of Kf can be determined in hydropenic rats (ΠE = 〈ΔP〉, Table ), the unique value of 5.6 nl/min · mm Hg determined in euvolemic rats was used.



Figure 9.

Relationship between Kf and plasma protein concentration (CA). Closed circles, mean values (± S.E.M.) obtained by Baylis et al. under a variety of conditions, with and without volume expansion, designed to alter CA; closed square, a study in which an isovolemic reduction in hematocrit produced filtration pressure disequilibrium without volume expansion ; closed triangles, values obtained in rats with isoncotic plasma volume expansion in which QA was varied by altering renal perfusion pressure by aortic constriction or carotid occlusion .

Modified from Baylis et al.


Figure 10.

Hydraulic pressure profile in rat kidney. Open circles, values obtained in studies of euvolemic Munich‐Wistar rats (Table ); closed circles, studies of hydropenic rats (Table ); closed triangles, values obtained by Casellas and Navar in Sprague‐Dawley rats in unique population of juxtamedullary nephrons at inside cortical surface apposed to pelvic lining and arcuate veins. Hydraulic pressures can be obtained in proximal (Early a.a.) and distal (Late a.a.) portions of afferent arteriole and in proximal (Early e.a.) and distal (Late e.a.) segments of efferent arteriole, and in peritubular capillaries (Pc), interlobular veins (I.V.), and renal vein (R.V.). For micropuncture experiments, systemic arterial pressure ( ) averaged — 110 mm Hg, whereas in study of Casellas and Navar arcuate artery (Arc. Art.) was perfused with whole blood at 96 mm Hg. , hydraulic pressure in glomerular capillaries.



Figure 11.

Pathways of renin synthesis and processing in mouse submaxillary gland. A: proposed sites of cleavage (upward arrows) and Mrs of each compound (far right column). B: synthesis, packaging into granules, and secretion of renin.

From Pratt et al. (497a


Figure 12.

Cellular control of renin release. Occupancy of adenylate cyclase (AC)‐linked receptors by agonists [norepinephrine (NE), histamine (H), and prostaglandins (PG)] results in stimulation of cAMP production and phosphorylation of intermediate protein kinase (PK) that stimulates (+) renin secretion. N, guanine nucleotide regulatory proteins specific for each agonist. Though not shown, angiotensin II (AΠ) and vasopressin (AVP) also act through N (or G) proteins to increase calcium influx as well as calcium release from intracellular storage sites. Membrane depolarization (Dep.) can also increase Ca2+ influx through voltage‐sensitive channels. Increased [Ca2+], leads to formation of the calcium‐calmodulin (Ca‐CM) complex that inhibits (−) renin release. Decreases in [Ca2+]i uncouples the Ca‐CM complex, thus de‐inhibiting renin release. RAΠ, Ravp, RH2, Rβ. and RPG; receptors for angiotensin II, arginine vasopressin, histamine (H2‐type), norepinephrine (β‐type), and prostaglandins, respectively.

From Ballermann et al.


Figure 13.

Effects of intravenous infusion of angiotensin II (200–600 ng/kg/min) determinants of glomerular ultrafiltration in Munich‐Wistar rat. Left, data obtained when renal perfusion pressure ( ) was allowed to increase; right, results obtained when increase in was prevented. Data are means ± S.E. SNGFR, single nephron glomerular filtration rate.

From Myers et al.


Figure 14.

Role of angiotensin II in mediating altered renal function of congestive heart failure induced by myocardial infarction (M.I.). Control euvolemic sham‐operated rats were studied during two periods; before (open circles) and during (filled circles) infusion of angiotensin I‐converting enzyme inhibitor teprotide [SQ20881; ]. SNGFR, single nephron glomerular filtration ra])G SNFF, single nephron filtration fracti])R , renal perfusion pressure.



Figure 15.

Mechanism of action of angiotensin II (AT II) in contraction of renal vasculature and glomerular mesangial cells. R, receptor; Gαβγ, guanine nucleotide‐binding regulatory protein with its three subunits, α, β, and γ; GTP, guanine trisphospha])G PLC, phospholipase C; PlP2, phosphatidylinositol‐4,5‐bisphospha])G IP3, inositol‐1,4,5‐trisphospha])G DAG, 1,2‐diacylglycerol; Ca2+, intracellular free calcium; PKC, protein kinase C; LC, lipocortin; PLA2, phospholipase A2; PL, membrane phospholipids; AA, arachidonic acid; PGE2, prostaglandin E2; cAMP, cyclic adenosine monophospha])G cAMP‐PK, cAMP‐dependent protein kinase; MLCK, myosin light chain kinase; P‐MLCK, phosphorylated MLCK; MLCK‐CAM, MLCK‐calcium‐calmodulin active complex; Pi, phospha])G *, activated form of enzyme. Parts A, B, and C in upper panel show sequence of events leading from occupancy of the receptor to activation of PLC. Lower panel shows sequence of events occurring in cells after PLC activation.



Figure 16.

Lipoxygenase pathway for biosynthesis of vasoactive leukotrienes and cyclooxygenase pathway for synthesis and degradation of prostaglandins, prostacyclin, and thromboxane A2 (TxA2).



Figure 17.

Effects of PGE2, PGI2, and dibutyryl (DB)‐cAMP on the determinants of glomerular filtration in rat. Data include effects on single nephron glomerular filtration rate (SNGFR), panel A; glomerular plasma flow rate (QA), panel B; mean glomerular capillary hydraulic pressure ( ), panel C; glomerular ultrafiltration coefficient (Kf), panel D; and sum of the pre‐ and postglomerular resistances (RTA), panel E. Euvolemic control animals were infused with cyclooxygenase inhibitors meclofenamate (Meclo.) or indomethacin (Indo.) or a cyclooxygenase inhibitor plus angiotensin II competitive antagonist saralasin. Effects of PGE2, PGI2, and DB‐cAMP were first examined during inhibition of endogenous prostaglandin production by Indo. or Meclo. and then with the added inhibition of endogenous angiotensin II by saralasin.

Data are from Schor et al.


Figure 18.

Effects of reduced mean arterial pressure on the measured determinants of glomerular ultrafiltration in hydropenic rats (A) and in mildly plasma‐expanded Munich‐Wistar rats before (B) and during (C) the administration of smooth muscle relaxant papaverine. Data (from ref. and ) are shown as means ± 1 S.E.

From Baylis and Brenner


Figure 19.

Filtrate‐to‐plasma concentration ratio (θ) as function of molecular size for dextran sulfate (DS), neutral dextran (D), and diethylaminoethyl dextran (DEAE). Symbols, mean values ± S.E. measured by Bohrer et al. in normal hydropenic Munich‐Wistar rats; curves are theoretical calculations based on values of QA, 〈ΔP〉, and CA reported in those studies with Kf = 4.8 nl/(min · mm Hg), ro = 47 Å, and Cm= 165 mEq/liter.

From Deen et al.


Figure 20.

Schematic of macromolecular solute (represented as sphere of radius, rs) moving through membrane pore (of radius ro and length l). λ = rs/ro.



Figure 21.

Hindrance factors for convection (W) and diffusion (H) based on hydrodynamic theory for movement of uncharged spherical molecules in cylindrical pores.

From Deen et al.


Figure 22.

Filtrate‐to‐plasma concentration ratio (θ) as function of molecular charge (z) molecular radius (rs). Results are shown for two values of membrane fixed charge concentration (Cm). Other inputs were QA = 75 nl/min, 〈ΔP> = 35 mm Hg, CA = 5.7 g/dl, Kf = 4.8 nl/(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen et al.


Figure 23.

Filtrate‐to‐plasma concentration ratio (θ) for serum albumin as function of membrane fixed charge concentration (Cm) and effective molecular charge (z). Other inputs were QA = 75 nl/min, 〈ΔP〉 = 35 mm Hg, CA = 5.7 g/dl, Kf = 4.8 nl/(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen and Satvat


Figure 24.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and glomerular plasma flow rate (QA). Calculations assumed Cm = 165 mEq/liter, rs = 30 Å, 〈ΔP〉 = 35 mm Hg, CA, = 5.7g/dl, Kf = 4.8 nl/‐(min · mm Hg), and ro = 50 Å, all representative of the normal hydropenic Munich‐Wistar rat.

From Deen and Satvat


Figure 25.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and transmembrane hydraulic pressure difference 〈ΔP〉. Input quantities were Cm=165 mEq/liter, rs = 30Å, QA = 75 nl/min, CA = 5.7g/dl, Kf = 4.8 nl/min · mm Hg), and ro = 50Å.

From Deen and Satvat


Figure 26.

Relationship between filtrate‐to‐plasma concentration ratio (θ), effective molecular charge (z), and ultrafiltration coefficient(Kf). Effective pore radius is assumed to remain constant. Input quantities were Cm=165 mEq/liter, rs = 30 Å, QA = 75 nl/min, 〈ΔP〉 = 35 mm Hg, CA = 5.7g/dl, and ro = 50 A.

From Deen and Satvat
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David A. Maddox, William M. Deen, Barry M. Brenner. Glomerular Filtration. Compr Physiol 2011, Supplement 25: Handbook of Physiology, Renal Physiology: 545-638. First published in print 1992. doi: 10.1002/cphy.cp080113