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Microcirculation of the intestinal mucosa

D. Neil Granger, Peter R. Kvietys, Ronald J. Korthuis, Andre J. Premen

10.1002/cphy.cp060139

Source: Supplement 16: Handbook of Physiology, The Gastrointestinal System, Motility and Circulation

Originally published: 1989

Published online: January 2011

Full Article on Wiley Online Library



Abstract

The sections in this article are:

1 Intrinsic Regulation of Blood Flow and Oxygenation
1.1 Basic Concepts
1.2 Vascular Response to Arterial Pressure Alterations
1.3 Vascular Response to Venous Pressure Elevation
1.4 Reactive Hyperemia
1.5 Vascular Response to Alterations in Arterial Blood Gases and Hematocrit
1.6 Postprandial Intestinal Hyperemia
2 Influence of Blood Flow on Intestinal Transport
2.1 Washout of Flow‐Limited Solutes
2.2 Effects of Oxygen Delivery on Absorption
3 Countercurrent Exchange
3.1 Anatomical Basis
3.2 Theoretical Considerations
3.3 Experimental Evidence
4 Vascular Capacitance
5 Vasoactive Agents and Intestinal Oxygen Uptake
5.1 Relation Between Blood Flow and Oxygen Uptake
5.2 Oxidative Metabolism
5.3 Effective Capillary Density
5.4 Intramural Blood Flow Distribution
6 Angiotensin II and Vasopressin
7 Effects of Experimental Conditions on the Intestinal Circulation
7.1 Anesthesia and Adjuvants
7.2 Respiration and Blood Gases
7.3 Laparotomy and Visceral Manipulation
7.4 Temerature
8 Transcapillary Fluid and Solute Exchange
8.1 Starling Hypothesis
8.2 Rate of Transcapillary Fluid Movement (Lymph Flow)
8.3 Capillary Filtration Coefficient
8.4 Capillary Hydrostatic Pressure
8.5 Interstitial Hydrostatic Pressure
8.6 Osmotic Reflection Coefficient
8.7 Transcapillary Oncotic Pressure Gradient
8.8 Interaction of Capillary and Interstitial Forces
9 Small‐Solute and Macromolecule Exchange
9.1 Small Solutes
9.2 Macromolecules
9.3 Factors Influencing Permeability
9.4 Pathways for Macromolecule Exchange
10 Conclusions
Figure 1. Figure 1.

Metabolic and myogenic theories of intestinal blood flow regulation.
Figure 2. Figure 2.

Dependence of superior mesenteric blood flow (F/Fo, blood flow normalized to control) and autoregulatory closed‐loop gain (Gc) on perfusion pressure (P/Po, arterial pressure normalized to control) in fed and fasted animals.

From Norris et al. 351
Figure 3. Figure 3.

Effects of varying perfusion pressure on total intestinal blood flow (venous outflow) and villous plasma flow (plasma particles).

From Lundgren 311
Figure 4. Figure 4.

Relationships between capillary filtration coefficient, arteriovenous O2 difference, and O2 delivery‐to‐demand ratio in the canine ileum.

From Kvietys et al. 285
Figure 5. Figure 5.

Dependence of intestinal O2 uptake (Vo2), arteriovenous O2 difference (A‐VΔAO2), normalized blood flow (F/Fo), and autoregulatory closed‐loop gain (Gc) on arterial pressure (P/Po) in fed (dashed lines) and fasted (dotted lines) dogs.

From Granger and Norris 192
Figure 6. Figure 6.

Steady‐state relation between intestinal O2 consumption and arterial pressure predicted by mathematical model for a passive system (no regulation of exchange or resistance vessels), resistance vessel regulation only, exchange vessel regulation only, and regulation of both resistance and exchange vessels. Difference to inflection points of O2 consumption‐arterial pressure curves provides a measure of margin of safety against hypoxia afforded by resistance and/or exchange vessel regulation.

From Granger and Granger 152
Figure 7. Figure 7.

Relationship between capillary filtration coefficient and capillary pressure in the cat small intestine. Capillary pressure was altered by venous pressure elevation or arterial pressure reduction. The inverse correlation is believed to result from myogenic control of perfused capillary density.

From Granger and Barrowman 147
Figure 8. Figure 8.

Percentage of total intestinal blood flow in mucosa‐submucosa and muscularis at various venous pressures (Pv); Kf,c, capillary filtration coefficient.

From Granger et al. 178
Figure 9. Figure 9.

Effect of metabolic rate on total and mucosal blood flow payback‐to‐debt ratio after a 60‐s arterial occlusion in the canine intestine.

From Shepherd and Riedel 435
Figure 10. Figure 10.

Effects of arterial hypoxia [partial pressure of O2 () ≦50 mmHg] on hemodynamics and oxygenation in canine intestinal segments perfused either under constant pressure (upper panel) or constant flow (lower panel) conditions. O2 consumption (); (a‐v)O2, arteriovenous O2 difference (A‐VO2); PS, permeability‐surface area product.

From Shepherd 422
Figure 11. Figure 11.

Effects of alterations in arterial blood O2 content (CaO2) on intestinal blood flow (Qi), O2 extraction and O2 consumption () in fetal and neonatal lambs.

From Edelstone and Holzman 97
Figure 12. Figure 12.

Effects of hematocrit on canine intestinal blood flow, O2 consumption (), arterial O2 content, and arteriovenous O2 difference ().

From Shepherd and Riedel 434
Figure 13. Figure 13.

Effects of intraluminal placement of various constituents of chyme on intestinal blood flow.

Modified from Granger et al. 160
Figure 14. Figure 14.

Relationship between intestinal blood flow (upper panel), O2 uptake (lower panel), and various functional activities (absorption, secretion, motility).

From Granger et al. 160
Figure 15. Figure 15.

Changes in intestinal blood flow and O2 extraction produced by intraluminal placement of food in the small bowel. Solid circles, data obtained at a control arteriovenous O2 difference (a–v)O2 ≧ 6 vol%; open circles, data obtained at a control (a–v)O2 ≦ 5 vol%; X, control values for blood flow and O2 extraction.

From Granger and Norris 192
Figure 16. Figure 16.

Role of arteriolar feedback, precapillary sphincter feedback, and passive factors in augmenting transcapillary O2 flux after graded elevations in intestinal O2 demand.

From Granger and Granger 152
Figure 17. Figure 17.

Dependence of passive solute absorption rate on jejunal blood flow in the rat.

Modified from Winne 491
Figure 18. Figure 18.

Patterns of countercurrent exchange of blood‐borne (A) and luminally derived (B) solutes.

Modified from Lundgren 311
Figure 19. Figure 19.

A, time estimated for attainment of diffusion equilibrium between outer surface and center of cylinders of various radii (r); B, mean transit time of plasma in villous vascular loops of the cat versus total intestinal blood flow.

From Lundgren 311
Figure 20. Figure 20.

Relationship between venous pressure and blood volume in an isolated intestinal vein [data from Simon et al. 444] and in an entire intestinal loop

data from Rothe et al. 404
Figure 21. Figure 21.

Effects of venoconstriction on the intestinal pressure‐volume relationship. Vuns, unstressed volume obtained by linear extrapolation of the pressure‐volume curve to the volume at zero pressure; curve A, relationship produced by venoconstriction that causes a reduced compliance (slope of curve) without a change in unstressed volume; curve B, relationship produced by venoconstriction that causes a reduced unstressed volume (Vuns) without a change in compliance.

From Rothe 403
Figure 22. Figure 22.

Examples of blood flow–independent (A) and blood flow–dependent (B) O2 uptake curves for the small intestine. Control value indicated by closed circle.

Adapted from Kvietys et al. (285; curve A) and from Shepherd (426; curve B)
Figure 23. Figure 23.

Effects of intraluminal placement of glucose (upper panel) and reductions in luminal temperature (lower panel) on relation between ileal O2 demand and blood flow. Relations were obtained by altering blood flow with a pump. Asterisks indicate significant (P < 0.05) differences in plateau portion of curves from control values (empty lumen at 39°C).

From Kvietys et al. 285
Figure 24. Figure 24.

Relation between blood flow and O2 uptake and factors that alter this relationship. Note that alterations in tissue oxidative metabolism shift curves vertically, whereas alterations in capillary density shift curves horizontally. Dot represents control blood flow under normal conditions. Pathway A, vasodilator that increases oxidative metabolism; pathway B, vasodilator that either does not affect metabolism or increases capillary density; pathway C, vasodilator that decreases capillary density; pathway D, vasodilator that decreases metabolism; pathway E, is taken by vasoconstrictor that decreases metabolism; pathway F, vasoconstrictor that decreases capillary density; pathway G, vasoconstrictor that does not affect metabolism or capillary density; pathway H, vasoconstrictor that increases capillary density; pathway I, vasoconstrictor that increases metabolism.

From Kvietys and Granger 278
Figure 25. Figure 25.

Relationship among intestinal O2 uptake (A), arteriovenous O2 difference (B), and lumen temperature.

From Kvietys et al. 281
Figure 26. Figure 26.

Distribution of pressures in the microcirculation of the rat small intestine.

From Bohlen and Gore 31
Figure 27. Figure 27.

Steady‐state relationship among intestinal interstitial fluid volume, interstitial fluid pressure, and interstitial hydraulic conductance.

From Granger et al. 162
Figure 28. Figure 28.

Starling forces and capillary membrane parameters in the small intestine under control (nontransporting) conditions. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162
Figure 29. Figure 29.

Safety factors against interstitial edema in the small intestine and the colon for an increment in capillary pressure of 12–13.2 mmHg.

From Granger and Barrowman 147
Figure 30. Figure 30.

Relations among intestinal capillary pressure, precapillary‐to‐postcapillary resistance ratio (Ra/Rv), and arterial pressure. Dotted lines in upper panel represent predicted mean values for capillary pressure, assuming Ra/Rv remained at value obtained at 125 mmHg arterial pressure. Dotted line in lower panel, Ra/Rv required for perfect autoregulation of capillary pressure. Asterisks, statistical significance at P < 0.05 (*) and P < 0.01 (**) levels as compared with values at 125 mmHg arterial pressure.

From Granger et al. 167
Figure 31. Figure 31.

Effects of changes in lymph flow, interstitial fluid pressure, and interstitial oncotic pressure on preventing interstitial dehydration in intestine during reductions in arterial and capillary pressures.

From Granger et al. 167
Figure 32. Figure 32.

Effects of sympathetic stimulation on intestinal trans‐capillary fluid exchange. Pt, interstitial fluid pressure; Δπ, transcapillary oncotic pressure difference; *, P < 0.05.

From Granger et al. 148
Figure 33. Figure 33.

Effects of net fluid absorption rate on intestinal interstitial volume, interstitial hydraulic conductivity, and the excluded volume fraction of albumin.

From Granger et al. 162
Figure 34. Figure 34.

Steady‐state relations between interstitial hydrostatic (Pt) and oncotic (πt) pressures and net fluid absorption rate.

From Granger 146
Figure 35. Figure 35.

Steady‐state relations between rate of removal of absorbed fluid by intestinal capillaries and lymphatics and net fluid absorption rate.

From Granger 146
Figure 36. Figure 36.

Effects of net fluid absorption on Starling forces and capillary membrane parameters in the small intestine. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NAP, net capillary absorptive pressure.

From Granger et al. 162
Figure 37. Figure 37.

Changes in Starling forces and capillary membrane parameters that lead to filtration secretion. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162
Figure 38. Figure 38.

Effects of active (solute‐coupled) fluid secretion on Starling forces and capillary membrane parameters in the small intestine. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162
Figure 39. Figure 39.

Theoretical relationship between lymph‐to‐plasma protein concentration ratio (CL/CP) and lymph flow. The osmotic reflection coefficient (σd) can be estimated from relation σd = 1 – CL/CP when CL/CP is filtration rate independent.

From Granger and Taylor 184
Figure 40. Figure 40.

Experimental data from cat and rat small intestine showing relationship between lymph flow and lymph‐to‐plasma protein concentration ratio (L/P).

From Taylor and Granger 460
Figure 41. Figure 41.

Application of pore‐stripping analysis to osmotic reflection coefficient (σd); dashed lines, data acquired during neurotensin infusion; solid lines, data acquired in control animals; dot/dashed lines, data acquired during fat absorption.

From Harper et al. 218
Figure 42. Figure 42.

Steady‐state relation between intestinal lymph‐to‐plasma total protein concentration ratio (L/P) and lymph flow after cream feeding and intraluminal placement of bile‐oleic acid. Solid line represents control relationship established in fasted animals. Osmotic reflection coefficient (σd) was estimated from relation σd = 1 – L/P when L/P is filtration rate independent.

From Granger et al. 173
Figure 43. Figure 43.

Relation between intestinal transcapillary protein fluxes (net, diffusive, and convective) and transcapillary volume flow when capillaries are in an absorbing state. Positive values denote blood‐to‐interstitium movement, whereas negative values indicate interstitium‐to‐blood movement.

From Granger et al. 172


Figure 1.

Metabolic and myogenic theories of intestinal blood flow regulation.



Figure 2.

Dependence of superior mesenteric blood flow (F/Fo, blood flow normalized to control) and autoregulatory closed‐loop gain (Gc) on perfusion pressure (P/Po, arterial pressure normalized to control) in fed and fasted animals.

From Norris et al. 351


Figure 3.

Effects of varying perfusion pressure on total intestinal blood flow (venous outflow) and villous plasma flow (plasma particles).

From Lundgren 311


Figure 4.

Relationships between capillary filtration coefficient, arteriovenous O2 difference, and O2 delivery‐to‐demand ratio in the canine ileum.

From Kvietys et al. 285


Figure 5.

Dependence of intestinal O2 uptake (Vo2), arteriovenous O2 difference (A‐VΔAO2), normalized blood flow (F/Fo), and autoregulatory closed‐loop gain (Gc) on arterial pressure (P/Po) in fed (dashed lines) and fasted (dotted lines) dogs.

From Granger and Norris 192


Figure 6.

Steady‐state relation between intestinal O2 consumption and arterial pressure predicted by mathematical model for a passive system (no regulation of exchange or resistance vessels), resistance vessel regulation only, exchange vessel regulation only, and regulation of both resistance and exchange vessels. Difference to inflection points of O2 consumption‐arterial pressure curves provides a measure of margin of safety against hypoxia afforded by resistance and/or exchange vessel regulation.

From Granger and Granger 152


Figure 7.

Relationship between capillary filtration coefficient and capillary pressure in the cat small intestine. Capillary pressure was altered by venous pressure elevation or arterial pressure reduction. The inverse correlation is believed to result from myogenic control of perfused capillary density.

From Granger and Barrowman 147


Figure 8.

Percentage of total intestinal blood flow in mucosa‐submucosa and muscularis at various venous pressures (Pv); Kf,c, capillary filtration coefficient.

From Granger et al. 178


Figure 9.

Effect of metabolic rate on total and mucosal blood flow payback‐to‐debt ratio after a 60‐s arterial occlusion in the canine intestine.

From Shepherd and Riedel 435


Figure 10.

Effects of arterial hypoxia [partial pressure of O2 () ≦50 mmHg] on hemodynamics and oxygenation in canine intestinal segments perfused either under constant pressure (upper panel) or constant flow (lower panel) conditions. O2 consumption (); (a‐v)O2, arteriovenous O2 difference (A‐VO2); PS, permeability‐surface area product.

From Shepherd 422


Figure 11.

Effects of alterations in arterial blood O2 content (CaO2) on intestinal blood flow (Qi), O2 extraction and O2 consumption () in fetal and neonatal lambs.

From Edelstone and Holzman 97


Figure 12.

Effects of hematocrit on canine intestinal blood flow, O2 consumption (), arterial O2 content, and arteriovenous O2 difference ().

From Shepherd and Riedel 434


Figure 13.

Effects of intraluminal placement of various constituents of chyme on intestinal blood flow.

Modified from Granger et al. 160


Figure 14.

Relationship between intestinal blood flow (upper panel), O2 uptake (lower panel), and various functional activities (absorption, secretion, motility).

From Granger et al. 160


Figure 15.

Changes in intestinal blood flow and O2 extraction produced by intraluminal placement of food in the small bowel. Solid circles, data obtained at a control arteriovenous O2 difference (a–v)O2 ≧ 6 vol%; open circles, data obtained at a control (a–v)O2 ≦ 5 vol%; X, control values for blood flow and O2 extraction.

From Granger and Norris 192


Figure 16.

Role of arteriolar feedback, precapillary sphincter feedback, and passive factors in augmenting transcapillary O2 flux after graded elevations in intestinal O2 demand.

From Granger and Granger 152


Figure 17.

Dependence of passive solute absorption rate on jejunal blood flow in the rat.

Modified from Winne 491


Figure 18.

Patterns of countercurrent exchange of blood‐borne (A) and luminally derived (B) solutes.

Modified from Lundgren 311


Figure 19.

A, time estimated for attainment of diffusion equilibrium between outer surface and center of cylinders of various radii (r); B, mean transit time of plasma in villous vascular loops of the cat versus total intestinal blood flow.

From Lundgren 311


Figure 20.

Relationship between venous pressure and blood volume in an isolated intestinal vein [data from Simon et al. 444] and in an entire intestinal loop

data from Rothe et al. 404


Figure 21.

Effects of venoconstriction on the intestinal pressure‐volume relationship. Vuns, unstressed volume obtained by linear extrapolation of the pressure‐volume curve to the volume at zero pressure; curve A, relationship produced by venoconstriction that causes a reduced compliance (slope of curve) without a change in unstressed volume; curve B, relationship produced by venoconstriction that causes a reduced unstressed volume (Vuns) without a change in compliance.

From Rothe 403


Figure 22.

Examples of blood flow–independent (A) and blood flow–dependent (B) O2 uptake curves for the small intestine. Control value indicated by closed circle.

Adapted from Kvietys et al. (285; curve A) and from Shepherd (426; curve B)


Figure 23.

Effects of intraluminal placement of glucose (upper panel) and reductions in luminal temperature (lower panel) on relation between ileal O2 demand and blood flow. Relations were obtained by altering blood flow with a pump. Asterisks indicate significant (P < 0.05) differences in plateau portion of curves from control values (empty lumen at 39°C).

From Kvietys et al. 285


Figure 24.

Relation between blood flow and O2 uptake and factors that alter this relationship. Note that alterations in tissue oxidative metabolism shift curves vertically, whereas alterations in capillary density shift curves horizontally. Dot represents control blood flow under normal conditions. Pathway A, vasodilator that increases oxidative metabolism; pathway B, vasodilator that either does not affect metabolism or increases capillary density; pathway C, vasodilator that decreases capillary density; pathway D, vasodilator that decreases metabolism; pathway E, is taken by vasoconstrictor that decreases metabolism; pathway F, vasoconstrictor that decreases capillary density; pathway G, vasoconstrictor that does not affect metabolism or capillary density; pathway H, vasoconstrictor that increases capillary density; pathway I, vasoconstrictor that increases metabolism.

From Kvietys and Granger 278


Figure 25.

Relationship among intestinal O2 uptake (A), arteriovenous O2 difference (B), and lumen temperature.

From Kvietys et al. 281


Figure 26.

Distribution of pressures in the microcirculation of the rat small intestine.

From Bohlen and Gore 31


Figure 27.

Steady‐state relationship among intestinal interstitial fluid volume, interstitial fluid pressure, and interstitial hydraulic conductance.

From Granger et al. 162


Figure 28.

Starling forces and capillary membrane parameters in the small intestine under control (nontransporting) conditions. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162


Figure 29.

Safety factors against interstitial edema in the small intestine and the colon for an increment in capillary pressure of 12–13.2 mmHg.

From Granger and Barrowman 147


Figure 30.

Relations among intestinal capillary pressure, precapillary‐to‐postcapillary resistance ratio (Ra/Rv), and arterial pressure. Dotted lines in upper panel represent predicted mean values for capillary pressure, assuming Ra/Rv remained at value obtained at 125 mmHg arterial pressure. Dotted line in lower panel, Ra/Rv required for perfect autoregulation of capillary pressure. Asterisks, statistical significance at P < 0.05 (*) and P < 0.01 (**) levels as compared with values at 125 mmHg arterial pressure.

From Granger et al. 167


Figure 31.

Effects of changes in lymph flow, interstitial fluid pressure, and interstitial oncotic pressure on preventing interstitial dehydration in intestine during reductions in arterial and capillary pressures.

From Granger et al. 167


Figure 32.

Effects of sympathetic stimulation on intestinal trans‐capillary fluid exchange. Pt, interstitial fluid pressure; Δπ, transcapillary oncotic pressure difference; *, P < 0.05.

From Granger et al. 148


Figure 33.

Effects of net fluid absorption rate on intestinal interstitial volume, interstitial hydraulic conductivity, and the excluded volume fraction of albumin.

From Granger et al. 162


Figure 34.

Steady‐state relations between interstitial hydrostatic (Pt) and oncotic (πt) pressures and net fluid absorption rate.

From Granger 146


Figure 35.

Steady‐state relations between rate of removal of absorbed fluid by intestinal capillaries and lymphatics and net fluid absorption rate.

From Granger 146


Figure 36.

Effects of net fluid absorption on Starling forces and capillary membrane parameters in the small intestine. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NAP, net capillary absorptive pressure.

From Granger et al. 162


Figure 37.

Changes in Starling forces and capillary membrane parameters that lead to filtration secretion. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162


Figure 38.

Effects of active (solute‐coupled) fluid secretion on Starling forces and capillary membrane parameters in the small intestine. Jv,c, rate of transcapillary fluid movement; Kf,c, capillary filtration coefficient; Pc, capillary hydrostatic pressure; Pt, interstitial hydrostatic pressure; σd, osmotic reflection coefficient; πc, plasma oncotic pressure; πt, interstitial oncotic pressure; NFP, net capillary filtration pressure.

From Granger et al. 162


Figure 39.

Theoretical relationship between lymph‐to‐plasma protein concentration ratio (CL/CP) and lymph flow. The osmotic reflection coefficient (σd) can be estimated from relation σd = 1 – CL/CP when CL/CP is filtration rate independent.

From Granger and Taylor 184


Figure 40.

Experimental data from cat and rat small intestine showing relationship between lymph flow and lymph‐to‐plasma protein concentration ratio (L/P).

From Taylor and Granger 460


Figure 41.

Application of pore‐stripping analysis to osmotic reflection coefficient (σd); dashed lines, data acquired during neurotensin infusion; solid lines, data acquired in control animals; dot/dashed lines, data acquired during fat absorption.

From Harper et al. 218


Figure 42.

Steady‐state relation between intestinal lymph‐to‐plasma total protein concentration ratio (L/P) and lymph flow after cream feeding and intraluminal placement of bile‐oleic acid. Solid line represents control relationship established in fasted animals. Osmotic reflection coefficient (σd) was estimated from relation σd = 1 – L/P when L/P is filtration rate independent.

From Granger et al. 173


Figure 43.

Relation between intestinal transcapillary protein fluxes (net, diffusive, and convective) and transcapillary volume flow when capillaries are in an absorbing state. Positive values denote blood‐to‐interstitium movement, whereas negative values indicate interstitium‐to‐blood movement.

From Granger et al. 172
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Article Title: Microcirculation of the intestinal mucosa
 

How to Cite

D. Neil Granger, Peter R. Kvietys, Ronald J. Korthuis, Andre J. Premen. Microcirculation of the intestinal mucosa. Compr Physiol 2011, Supplement 16: Handbook of Physiology, The Gastrointestinal System, Motility and Circulation: 1405-1474. First published in print 1989. doi: 10.1002/cphy.cp060139