Figure 16.5 gives an overview of urine formation, which is divided into these steps: glomerular filtration, tubular reabsorption, and tubular secretion.
Glomerular filtration occurs when whole blood enters the afferent arteriole and the glomerulus. Due to glomerular blood pressure, water and small molecules move from the glomerulus to the inside of the glomerular capsule. This is a filtration process because large molecules and formed elements are unable to pass through the capillary wall. In effect, then, blood in the glomerulus has two portions: the filterable components and the nonfilterable components:
Figure 16.5 Steps in urine formation. Top: The three main steps in urine formation are described in boxes that are color-coded to arrows showing the movement of molecules into or out of the nephron at specific locations. In the end, urine is composed of the substances within the collecting duct (see gray arrow, lower right).
The substances that are not reabsorbed become the tubular fluid, which enters the loop of the nephron.
Tubular secretion is a second way by which substances are removed from blood and added to the tubular fluid. Hydrogen ions, potassium ions, creatinine, and drugs such as penicillin are some of the substances that are moved by active transport from the blood into the distal convoluted tubule. In the end, urine contains (1) substances that have undergone glomerular filtration but have not been reabsorbed, and (2) substances that have undergone tubular secretion.
The glomerular filtrate contains small dissolved molecules in approximately the same concentration as plasma. Small molecules that escape being filtered and the nonfilterable components leave the glomerulus by way of the efferent arteriole.
As indicated in Table 16.1, nephrons in the kidneys filter 180 liters of water per day, along with a considerable amount of small molecules (such as glucose) and ions (such as sodium). If the composition of urine were the same as that of the glomerular filtrate, the body would continually lose water, salts, and nutrients. Therefore, we can conclude that the composition of the filtrate must be altered as this fluid passes through the remainder of the tubule.
Tubular reabsorption occurs as molecules and ions are both passively and actively reabsorbed from the nephron into the blood of the peritubular capillary network. The osmolarity of the blood is maintained by the presence of both plasma proteins and salt. When sodium ions (Na+) are actively reabsorbed, chloride ions (Cl-) follow passively. The reabsorption of salt (NaCl) increases the osmolarity of the blood compared to the filtrate, and therefore water moves passively from the tubule into the blood. About 67% of Na+ is reabsorbed at the proximal convoluted tubule.
Nutrients such as glucose and amino acids also return to the blood at the proximal convoluted tubule. This is a selective process because only molecules recognized by carrier molecules are actively reabsorbed. Glucose is an example of a molecule that ordinarily is completely reabsorbed because there is a plentiful supply of carrier molecules for it. However, every substance has a maximum rate of transport, and after all its carriers are in use, any excess in the filtrate will appear in the urine. For example, as reabsorbed levels of glucose approach 1.8-2 mg/ml plasma, the rest appears in the urine. In diabetes mellitus, excess glucose occurs in the blood, and then in the filtrate, and then in the urine, because the liver and muscles have failed to store glucose as glycogen, and the kidneys cannot reabsorb all of it. The presence of glucose in the filtrate increases its osmolarity compared to that of the blood, and therefore less water is reabsorbed into the peritubular capillary network. The frequent urination and increased thirst experienced by untreated diabetics are due to the fact that water is not being reabsorbed.
We have seen that the filtrate that enters the proximal convoluted tubule is divided into two portions: components that are reabsorbed from the tubule into the blood, and components that are not reabsorbed and continue to pass through the nephron to be further processed into urine: