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The Microcirculation and the Lymphatic System: Capillary Fluid Exchange, Interstitial Fluid and Lymph Flow

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Function of the circulation (microcirculation): transport of nutrients to the tissues and removal of cell excreta.

Structure of the Microcirculation and Capillary System

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The arterioles are highly muscular and their diameters can change manyfold. The metarterials (the terminal arterioles) donot have a continuous muscular coat, but smooth muscle fibers encircle the vessel at intermittent points.

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At the point where each true capillary originates from a metarteriole, a smooth muscle fiber usually encircles the capillary(precapillary sphincter) which can open and close the entrance to the capillary.

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Venules are larger than the arterioles and have a much weaker muscular coat but the pressure in the venules is much lessthan that in the arterioles (still contract considerably)

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Structure of the Capillary Wall

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Pores in the Capillary Membrane

Small passageways connecting the interior of the capillary with the exterior.

Intercellular cleft : thin-slit, curving channel that lies between adjacent endothelial cells; fluid can percolatefreely through the cleft

Plasmalemmal vesicles : form at one surface of the cell by imbibing small packets of plasma/ECF andmove slowly through the endothelial cell

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Special Types of Pores Occur in the Capillaries of Certain Organs

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Brain: junctions between capillary endothelial cells are tight junctions

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Liver: clefts between endothelial cells are wide open

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GI: midway between liver and muscles

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Kidney: fenestrae (small oval windows)

Flow of Blood in the Capillaries—Vasomotion

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Blood does not flow continuously through the capillaries and instead flows intermittently because of intermittentcontraction of the metarterioles and precapillary sphincters.

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Regulation of Vasomotion

[Oxygen] is the most important factor to affect degree of opening and closing of themetarterioles/precapillary sphincters.

Low [oxygen]: intermittent periods of capillary blood flow occur more often and the duration of each period of flow lasts longer (capillary blood then carries increased quantities of oxygen and other nutrientsto the tissue)

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Average Function of the Capillary System

The average functions are in reality the functions of literally billions of individual capillaries, eachoperating intermittently in response to local conditions in the tissues.

Exchange of Water, Nutrients, and Other Substances Between the Blood and Interstitial Fluid

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Diffusion Through the Capillary Membrane

The most important means by which substances are transferred between the plasma and the interstitialfluid is diffusion which results from thermal motion of the water molecules and dissolved substances in thefluid.

Lipid-soluble Substances Can Diffuse Directly Through the Cell Membranes of the Capillary Endothelium

Water-soluble, Non-lipid-soluble Substances Diffuse Only Through Intercellular Pores in the CapillaryMembrane

Effect of Molecular Size on Passage Through the Pores

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The permeability of different substances varies according to their molecular diameters.

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The capillaries in different tissues have extreme differences in their permeabilities.

Effect of Concentration Difference on Net Rate of Diffusion Through the Capillary Membrane

The Interstitium and Interstitial Fluid

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Interstitium : spaces between cells filled with fluid (interstitial fluid)

Two major types of solid structures: collagen fiber bundles and proteoglycan filaments

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“Gel” In the Interstitium

Combination of proteoglycan filaments and fluid entrapped with them with the characteristics of a gel(interstitial fluid is entrapped mainly in the minute spaces among the proteoglycan filaments)

Because of the large number of proteoglycan filaments, it is difficult for fluid to flow through the gel andinstead it mainly diffuses through the gel.

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“Free” Fluid in the Interstitium

Occasionaly small rivulets of free fluid and small free fluid vesicles are also present and can flow freely

Edema: small pockets and rivulets of free fluid expand tremendously

Fluid Filtration Across Capillaries is Determined by Hydrostatic and Colloid Osmotic Pressures,and Capillary Filtration Coefficient

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Hydrostatic pressure forces fluid/dissolved substances through the capillary pores into the interstitial spaces. Osmotic pressure caused by plasma proteins causes fluid movement osmosis from the interstitial spaces into the blood (prevents lossof fluid volume from the blood into the interstitial spaces)

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Lymphatic system: returns to the circulation the small amounts of excess protein and fluid that leak from the blood into theinterstitial spaces

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Four Primary Hydrostatic and Colloid Osmotic Forces Determine Fluid Movement Through the Capillary Membrane

Capillary Pressure : tends to force fluid outward through the capillary membrane

Interstitial Fluid Pressure : tends to force fluid inward through the capillary membrane when it is positive(outward when it’s negative)

Plasma Colloid Osmotic Pressure : causes osmosis of fluid inward through the capillary membrane

Interstitial Fluid colloid osmotic pressure : tends to cause osmosis of fluid outward through the capillarymembrane

If the sum of these forces (net filtration pressure) is positive, there will be a net fluid filtration across thecapillaries. If the sum of the forces is negative, there will be a net fluid absorption from the interstitialspaces into the capillaries.

Net filtration pressure = Pc – Pif -

p +

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Slightly positive under normal conditions (net filtration of fluid across the capillaries into theinterstitial space in most organs)

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Capillary Hydrostatic Pressure

Micropipette Method for Measuring Capillary Pressure

Isogravimetric Method for Indirectly Measuring Functional Capillary Pressure

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Gut held up by one arm of a gravimetric balance and blood perfused through the vessels of the gut wall.

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Arterial pressure decreased

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decrease in capillary pressure

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osmotic pressure of plasma proteins causes absorption of fluid out of the gut wall

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weight of gut decreases

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venous pressure is increased an amount sufficient to overcome the effect of decreasing the arterial pressure

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Capillary pressure is kept constant while simultaneously decreasing arterial pressure andincreasing venous pressure

Why is the Functional Capillary Pressure Lower than Capillary Pressure Measured by the MicropipetteMethod?

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Interstitial Fluid Hydrostatic Pressure

Measurement of Interstitial Fluid Pressure Using the Micropipette

Measurement of Interstitial Free Fluid Pressure in Implanted Perforated Hollow Capsules

Measurement of Interstitial Free Fluid Pressure by Means of a Cotton Wick

Interstitial Fluid Pressures in Tightly Encased Tissues

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The pressure exerted on the skin is atmospheric pressure (zero pressure)

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normal interstitialfluid pressure is usually several mm Hg negative with respect to the pressure that surroundseach tissue.

Is the True Interstitial Fluid Pressure in Loose Subcutaneous Tissue Subatmospheric?

Summary: An Average Value for Negative Interstitial Fluid Pressure in Loose Subcutaneous Tissue

Pumping by the Lymphatic System is the Basic Cause of the Negative Interstitial Fluid Pressure

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Plasma Colloid Osmotic Pressure

Proteins in the Plasma Cause Colloid Osmotic Pressure

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Because the proteins are the only dissolved constituents in the plasma and interstitial fluidsthat do not readily pass through the capillary pores, it is the proteins of the plasma andinterstitial fluids that are responsible for the osmotic pressures on the two sides of thecapillary membrane

Normal Values for Plasma Colloid Osmotic Pressure

Effect of the Different Plasma Proteins on Colloid Osmotic Pressure

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Interstitial Fluid Colloid Osmotic Pressure

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Exchange of Fluid Volume Through the Capillary Membrane

The average capillary pressure at the arterial ends of the capillaries is greater than at the venous ends

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fluid filters out of the capillaries at their arterial ends but at their venous ends fluid is reabsorbed back intothe capillaries

Analysis of the Forces Causing Filtration at the Arterial End of the Capillary

Analysis of Reabsorption at the Venous end of the Capillary

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Venous capillaries are more numerous and more permeable than the arterial capillaries, so thatless reabsorption pressure is required to cause inward movement of fluid.

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The reabsorption pressure causes about 9/10 of the fluid that has filtered out of the arterialends of the capillaries to be reabsorbed at the venous ends. The remaining 1/10 flows into thelymph vessels and returns to the circulating blood.

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Starling Equilibrium Capillary Exchange

The amount of fluid filtering outward from the arterial ends of capillaries equals almost exactly the fluidreturned to the circulation by absorption

Slight excess of filtration is called net filtration and it is the fluid that must be returned to the circulationthrough the lymphatics.

Filtration Coefficient

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Effect of Abnormal Imbalance of Forces at the Capillary Membrane

Lymphatic System

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Accessory route through which fluid can flow from the interstitial spaces into the blood; can carry proteins and large particulate matter away from the tissue spaces

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Lymph Channels of the Body

Terminal Lymphatic Capillaries and Their Permeability

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Endothelial cells of the lymphatic capillary are attached by anchoring filaments to thesurrounding connective tissue. At the junctions of adjacent endothelial cells, the edge of oneendothelial cell overlaps the edge of the adjacent cell in such a way that the overlapping edgeis free to flap inward

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forms a minute valve that opens to the interior of the lymphaticcapillary.

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Formation of Lymph

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Rate of Lymph Flow

Effect of Interstitial Fluid Pressure on Lymph Flow

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Any factor that increases interstitial fluid pressure also increases lymph flow if the lymphvessels are functioning normally

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Elevated capillary pressure

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Decreased plasma colloid osmotic pressure

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Increased interstitial fluid colloid osmotic pressure

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Increased permeability of the capillaries

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Increasing tissue pressure not only increases entry of fluid into the lymphatic capillaries butalso compresses the outside surfaces of the larger lymphatics, thus impeding lymph flow athigher pressures (maximum lymph flow rate)

Lymphatic Pump Increases Lymph Flow

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When a collecting lymphatic vessel becomes stretched with fluid, the smooth muscle in thewall of the vessel automatically contracts and each segment of the lymph vessel betweensuccessive valves functions as a separate automatic pump.

Pumping Caused by External Intermittent Compression of the Lymphatics

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Any external factor that intermittently compresses the lymph vessel also can cause pumping.

Lymphatic Capillary Pump

Summary of Factors That Determine Lymph Flow

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The rate of lymph flow is determined by the product of interstitial fluid pressure times theactivity of the lymphatic pump.

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Role of the Lymphatic System in Controlling Interstitial Fluid Protein Concentration, Interstitial Fluid Volume, andInterstitial Fluid Pressure