Urine formation involves three main processes namely:
- Glomerular Filtration,
- Reabsorption and
The glomerular capillary blood pressure causes filtration of blood through 3 layers, i.e., the endothelium of glomerular blood vessels, the epithelium of Bowman’s capsule and a basement membrane between these two layers.
The epithelial cells of Bowman’s capsule called podocytes are arranged in an intricate manner so as to leave some minute spaces called filtration slits or slit pores.
The diameter of efferent arteriole (arteriole bringing blood out of the glomerulus) is less than the diameter of afferent arteriole (arteriole taking blood inside the glomerulus). This difference in diameters creates a pressure which facilitates the filtration. Blood is filtered so finely through these membranes, that almost all the constituents of the plasma, except the proteins pass onto the lumen of the Bowman’s capsule. Therefore, it is considered as a process of ultra filtration.
Glomerular Filtration Rate (GFR)
The amount of the filtrate formed by the kidneys per minute is called glomerular filtration rate (GFR). GFR in a healthy individual is approximately 125 ml/minute, i.e., 180 litres per day. On an average, 1100-1200 ml of blood is filtered by the kidneys per minute which constitute roughly 1/5th of the blood pumped out by each ventricle of the heart in a minute.
Regulation of GFR: The kidneys have built-in mechanisms for the regulation of glomerular filtration rate. One such efficient mechanism is carried out by juxta glomerular apparatus (JGA). JGA is a special sensitive region formed by cellular modifications in the distal convoluted tubule and the afferent arteriole at the location of their contact. A fall in GFR can activate the JG cells to release renin which can stimulate the glomerular blood flow and thereby the GFR back to normal.
Reabsorption: A comparison of the volume of the filtrate formed per day (180 litres per day) with that of the urine released (1.5 litres), suggest that nearly 99 per cent of the filtrate has to be reabsorbed by the renal tubules. This process is called reabsorption.
The tubular epithelial cells in different segments of nephron perform this either by active or passive mechanisms. For example, substances like glucose, amino acids, Na+, etc., in the filtrate are reabsorbed actively whereas the nitrogenous wastes are absorbed by passive transport.
Reabsorption of water also occurs passively in the initial segments of the nephron. During urine formation, the tubular cells secrete substances like H+, K+ and ammonia into the filtrate.
Tubular secretion is also an important step in urine formation as it helps in the maintenance of ionic and acid base balance of body fluids.
Function of Tubules
Proximal Convoluted Tubule (PCT)
PCT is lined by simple cuboidal brush border epithelium which increases the surface area for reabsorption. Nearly all of the essential nutrients, and 70-80 per cent of electrolytes and water are reabsorbed by this segment. PCT also helps to maintain the pH and ionic balance of the body fluids by selective secretion of hydrogen ions, ammonia and potassium ions into the filtrate and by absorption of HCO3- from it.
Reabsorption in this segment is minimum. However, this region plays a significant role in the maintenance of high osmolarity of medullary interstitial fluid. The descending limb of loop of Henle is permeable to water but almost impermeable to electrolytes. This concentrates the filtrate as it moves down. The ascending limb is impermeable to water but allows transport of electrolytes actively or passively. Therefore, as the concentrated filtrate pass upward, it gets diluted due to the passage of electrolytes to the medullary fluid.
Distal Convoluted Tubule (DCT)
Conditional reabsorption of Na+ and water takes place in this segment. DCT is also capable of reabsorption of HCO and selective secretion of hydrogen and potassium ions and NH3 to maintain the pH and sodium potassium balance in blood.
This long duct extends from the cortex of the kidney to the inner parts of the medulla. Large amounts of water could be reabsorbed from this region to produce concentrated urine. This segment allows passage of small amounts of urea into the medullary interstitium to keep up the osmolarity. It also plays a role in the maintenance of pH and ionic balance of blood by the selective secretion of H+ and K+ ions
Mechanism of Concentration of Filtrare
The Henle’s loop and vasa recta play a significant role in this. The flow of filtrate in the two limbs of Henle’s loop is in opposite directions and thus forms a counter current. The flow of blood through the two limbs of vasa recta is also in a counter current pattern. The proximity between the Henle’s loop and vasa recta, as well as the counter current in them help in maintaining an increasing osmolarity towards the inner medullary interstitium, i.e., from 300 mOsmolL–1 in the cortex to about 1200 mOsmolL–1 in the inner medulla. This gradient is mainly caused by NaCl and urea. NaCl is transported by the ascending limb of Henle’s loop which is exchanged with the descending limb of vasa recta. NaCl is returned to the interstitium by the ascending portion of vasa recta.
Similarly, small amounts of urea enter the thin segment of the ascending limb of Henle’s loop which is transported back to the interstitium by the collecting tubule.
The above described transport of substances facilitated by the special arrangement of Henle’s loop and vasa recta is called the counter current mechanism. This mechanism helps to maintain a concentration gradient in the medullary interstitium. Presence of such interstitial gradient helps in an easy passage of water from the collecting tubule thereby concentrating the filtrate (urine). Human kidneys can produce urine nearly four times concentrated than the initial filtrate formed.