Module 4 - Lecture 6

Urine Formation

  • Concept of Urine Formation:

    • Involves altering the composition of blood and filtrate that pass through nephrons.

    • Achieved through renal blood flow:

    • Renal Blood Flow: Approximately 1.2 L/min, ~25% of cardiac output, with kidneys weighing only 1-2% of body weight.

    • Ensures high plasma filtration rate.

    • Key Formula:

    • Urinary excretion = Filtration – Reabsorption + Secretion

    • Filtration Rates:

    • Entire plasma volume ~3 L filtered ~60 times a day through the kidneys.

    • Glomerular filtration = 125 mL/min or 180 L/day.


Three Renal Processes that Produce Urine

  1. Glomerular Filtration:

    • Produces cell- and protein-free filtrate.

  2. Tubular Reabsorption:

    • Selectively reabsorbs 99% of substances from filtrate back into blood within renal tubules and collecting ducts.

  3. Tubular Secretion:

    • Selectively moves substances from blood to filtrate.

Glomerular Filtration Rate (GFR)

  • Definition:

    • Volume of plasma filtered per minute by all glomeruli.

    • An important indicator of kidney function.

    • GFR declines in kidney disease.

  • Filtration Fraction:

    • The fraction of renal plasma flow that is filtered in the glomerulus during a single pass through the kidney, calculated as:
      Filtration Fraction=GFRRenal Plasma Flow\text{Filtration Fraction} = \frac{GFR}{\text{Renal Plasma Flow}}

    • Of plasma volume entering the afferent arteriole, 20% is filtered by the glomerulus, and 80% continues back through the efferent arteriole into capillaries.

    • >99% of filtrate is typically reabsorbed.


Glomerular Filtration Process

Filtration Membrane

  • Components:

    • Separates blood in glomerular capillaries from filtrate in glomerular capsule.

    • Porous membrane allows free passage of water and solutes smaller than proteins.

    • Composed of three layers:

    • Fenestrated Endothelium: of glomerular capillaries.

    • Basement Membrane: Prevents protein passage.

    • Podocytes: Specialized cells with foot processes.

  • Filtrate Composition:

    • Blood plasma minus proteins.

  • Process Type:

    • Glomerular filtration is a passive, non-selective process.

  • Driving Force:

    • Hydrostatic pressure forces fluids and solutes out of glomerular capillaries into the glomerular capsule through the filtration membrane.

Pressures Affecting Glomerular Filtration

  • Net Filtration Pressure (NFP):

    • Responsible for filtrate formation, resulting from a balance of outward and inward pressures.

  • Outward Pressures:

    • Promote filtrate formation.

    • Hydrostatic Pressure in Glomerular Capillaries (HPgc):

    • Typical value: 55 mmHg, higher than usual for capillaries (28 mmHg).

  • Inward Pressures:

    • Inhibit filtrate formation.

    • Hydrostatic Pressure in Capsular Space (HPcs) and Colloid Osmotic Pressure in Glomerular Capillaries (OPgc):

    • OPgc exerts pressure by proteins in blood, 'sucking' water into capillaries.

  • NFP Calculation:

    • Main controllable factor determining GFR.


Regulation of Glomerular Filtration

  • Purpose:

    • Maintain filtration and systemic blood pressure.

  • Determination of GFR:

    • Regulated by the volume of blood (BV) flowing into the glomerulus, which affects glomerular hydrostatic pressure.

  • Changes in GFR:

    • Affect filtrate formation, urine output, and systemic blood pressure.

    • Increasing GFR will increase urinary output, thus decreasing blood volume and blood pressure.

Intrinsic Mechanisms of Regulation

  1. Myogenic Stretch of Afferent Arteriole Smooth Muscle:

    • Fluid volume entering arterioles alters arteriole diameter.

    • Vascular smooth muscle contracts when stretched and relaxes when not stretched.

    • High blood volume/stretch triggers vasoconstriction → decreases GFR; low blood volume/stretch triggers vasodilation.

  2. Tubuloglomerular Feedback:

    • Macula densa cells monitor filtrate [NaCl].

    • High GFR reduces reabsorption time; results in high NaCl in filtrate.

    • Vasoactive secretions from macula densa constrict afferent arterioles, which decreases GFR.


Extrinsic Mechanisms of Regulation

  1. Renin-Angiotensin-Aldosterone System (RAAS):

    • Primary mechanism for increasing blood pressure.

    • If blood volume is abnormally low (e.g., due to hemorrhage or dehydration), several pathways activate granular cells to release renin:

      • Activated Macula Densa: Low blood pressure correlates to low GFR, meaning slow filtrate flow in tubules → more NaCl reabsorption; decreased [NaCl] signals granular cells.

      • Reduced Stretch: Granular cells act as mechanoreceptors that secrete renin under low pressure.

  2. Sympathetic Nervous System (SNS) & Baroreceptor Reflex:

    • In resting state, renal arterioles are typically dilated.

    • If blood volume is low, the baroreceptor reflex is inhibited (normally promotes vasodilation); SNS gets activated.

    • Results in vasoconstriction to increase total peripheral resistance, constriction of afferent arterioles decreases blood flow and GFR, leading to increased blood volume and blood pressure.


Tubular Reabsorption of Filtrate

  • Process:

    • Active transport is used to facilitate Na+ reabsorption across apical and basolateral membranes.

    • Substances such as glucose, amino acids, vitamins, and ions (e.g., K+, Cl-) are co-transported alongside Na+ transporters.

    • Osmotic diffusion of water occurs via aquaporins present at the proximal convoluted tubule (PCT).

Varying Reabsorption Across Tubules

  • Proximal Convoluted Tubule:

    • Reabsorbs all glucose, amino acids, and approximately two-thirds of Na+ and water.

  • Nephron Loop:

    • Water is reabsorbed from the descending limb and not the ascending limb (aquaporins are present only in descending limb).

    • Solutes are reabsorbed in the ascending limb but not in descending limb.

  • Distal Convoluted Tubule & Collecting Duct:

    • Regulated reabsorption occurs, based on the body's needs.

    • Antidiuretic Hormone (ADH, or vasopressin) increases water reabsorption by inserting aquaporins in the collecting duct.

    • Aldosterone promotes Na+ reabsorption in DCT and collecting duct, increasing water reabsorption due to osmosis following Na+.


Urine Characteristics and Formation

  • Urine Composition:

    • 95% water, 5% solutes including:

    • Urea: Product of amino acid breakdown

    • Uric Acid: Waste product of amino acid metabolism

    • Creatinine: Metabolite of creatinine phosphate.

  • Concentration Levels in Urine:

    • Urea > Na+ > K+ > creatinine > uric acid

  • Physical and Chemical Characteristics of Urine:

    • Healthy urine is clear and varies from pale to deep yellow due to urochrome (pigment from hemoglobin breakdown).

    • Aromatic when fresh, often develops an ammonia odor due to bacterial metabolism of urea.

    • Slightly acidic (~pH 6.0); high-protein diets lead to more acidity while vegetarian diets lead to more alkalinity.

Mechanism of Osmotic Gradient in Urine Formation

  • Osmotic Gradients:

    • Determined by water movement and solute concentration in the medulla of the kidneys.

    • Countercurrent mechanisms affect urine volume and concentration.

    • Countercurrent Mechanisms:

    • The direction of fluid flow in adjacent segments of a connected tube (descending & ascending loops).

    • Countercurrent multiplier first concentrates and then dilutes the filtrate.

    • Changes in filtrate concentration in the descending limb facilitate water loss to interstitial fluid, affecting solute secretion in the ascending limb.

    • Urea recycling from the collecting duct contributes to high osmolality in the medulla.

Dilute vs Concentrated Urine

  • Well-Hydrated State:

    • The body can dilute urine to $\frac{1}{6}$ the concentration of blood plasma.

  • Dehydrated State:

    • Up to 99% water reabsorbed yielding 0.5 L of concentrated urine at 1200 mOsmol/L.


Countercurrent Exchange via Vasa Recta

  • Function:

    • The vasa recta, a series of capillaries that surround the nephron loops, perform countercurrent exchange, preserving osmotic gradients in the renal medulla.

    • Capillaries exchange solutes and water in opposing directions, maintaining equilibrium between capillaries and interstitial fluid at each section, upholding the concentration gradient throughout the renal medulla.