Glomerular Filtration
The first step of urine formation takes place in the renal corpuscles. Here, the liquid part of the blood plasma passes through the filtration barrier (glomerular filtration barrier) from the glomerular capillaries into the capsular lumen.
The main driving force for filtration is the high hydrostatic pressure in the glomerular capillaries, which reaches 50–60 mm Hg in the cortical nephrons.
As a result, primary urine (the ultrafiltrate) is formed. Only about 10% of the blood volume flowing through the kidneys is filtered into the capsule.
Composition of primary urine:
- Water and inorganic ions ($Na^+$, $K^+$, $Cl^-$).
- Low-molecular-weight organics: glucose, amino acids.
- Metabolic waste products: urea, creatinine, uric acid.
- Small plasma proteins (albumins, some globulins).
Tubular Reabsorption
Since a huge volume of valuable substances and water is lost in the primary urine (180 L per day!), the body reclaims them back into the blood. This process is called reabsorption and occurs along the renal tubules and collecting ducts.
In the proximal convoluted tubules, obligate (mandatory, hormone-independent) reabsorption takes place. Here, virtually 100% of glucose, amino acids, proteins, and up to 85% of sodium ions are actively reabsorbed with energy expenditure. Following sodium passively (along an osmotic gradient), about 85% of water is also reabsorbed. Proteins are taken up by cells via pinocytosis, while low-molecular-weight substances are transported via secondary active transport coupled with sodium ions.
In the distal tubules and collecting ducts, facultative (optional) reabsorption occurs. It is regulated by hormones according to the current needs of the body.
Tubular Secretion
Concurrently with reabsorption, secretion takes place in the distal parts of the nephron and collecting ducts—this is the active transport of additional substances from the blood into the tubular lumen (forming the urine).
This process is closely linked to sodium reabsorption:
- In the distal tubules, $K^+$ and $H^+$ ions are secreted into the urine in exchange for the reabsorption of $Na^+$ into the blood.
- In the collecting ducts, in addition to hydrogen ions, ammonia is secreted (as $NH_4^+$).
Hormonal Regulation of Urine Formation
Fine-tuning of the volume and composition of the final urine occurs under the influence of hormones:
- Aldosterone stimulates the synthesis of transport proteins in the distal segments. It enhances the reabsorption of $Na^+$ and $Cl^-$ (retaining them in the body) while simultaneously stimulating the secretion of $K^+$ and $H^+$.
- Antidiuretic hormone (ADH, vasopressin) regulates the volume of water excreted. Normally, the walls of the distal tubules and collecting ducts are impermeable to water. ADH makes them permeable by stimulating the insertion of water channels (aquaporins, specifically Aquaporin-2) into the apical membranes of collecting duct cells, leading to enhanced water reabsorption into the blood and a decreased urine volume.