- Related Topics:
- urination
- urine
- secretion
- bicarbonate threshold
- egestion
The mechanisms of detoxication that animals use are related to their modes of life. This is true, with greater force, of the mechanisms of homeostasis, the ability of organisms to maintain internal stability. A desert-living mammal constantly faces the problem of water conservation; but a freshwater fish faces the problem of getting rid of the water that enters its body by osmosis through the skin. At the level of the individual cell, whether it is the cell that constitutes a unicellular organism or a cell in the body of a multicellular organism, the problems of homeostasis present themselves in similar ways.
To continue its intracellular processes a cell must maintain an intracellular chemical environment in which the concentrations of various ions (see below) are kept constant in the face of changing concentrations in the medium surrounding the cell. This is the task of the cell membrane. In the higher animals the task is easier since cells in the interior of their bodies are bathed in an internal medium—the blood—whose composition is regulated so as to minimize the effects of changes in the external medium. This regulatory function is undertaken by specialized cells or organs such as the kidney, thereby lessening the regulatory burden of the other cells of the body.
The biological necessity for homeostatic mechanisms is particularly urgent for controlling the inorganic components of cells and body fluids. Inorganic salts can exert even greater osmotic pressure against membranes impermeable to them than urea. This is so because, under the conditions in the body, they are almost completely dissociated into their component ions. For example, a molecule of common salt (sodium chloride) is dissociated into two inorganic ions—a positively charged sodium ion and a negatively charged chloride ion—both of which can exert osmotic pressure.
Aside from their osmotic effects, inorganic ions have profound effects upon metabolic processes, which in general will take place only in the presence of appropriate concentrations of these ions. The most important inorganic ions in organisms are the positively charged hydrogen, sodium, potassium, calcium, and magnesium ions, and the negatively charged chloride, phosphate, and bicarbonate ions. The membranes of cells are not completely impermeable to these ions and are in fact endowed with the ability to transport ions between the inside and outside of the cell, whereby they control the concentrations of ions within the cells; when such transport is in the direction that requires a supply of energy, it is called active transport (see cell: The plasma membrane).
Osmotic regulation is the maintenance of the normal concentration of the body fluids; i.e., the total concentration of all dissolved substances (solutes) that would exert osmotic pressure against a membrane impermeable to them. Osmotic regulation controls the amount of water in the body fluids relative to the amount of osmotically active solutes. Ionic regulation is the maintenance of the concentrations of the various ions in the body fluids relative to one another. There is no consistent distinction between the two processes; organs that participate in one process at the same time participate in the other.
Principal excretory structures
Whereas the kidney is the principal organ subserving both nitrogenous excretion and osmotic and ionic regulation in the mammalian body, these functions are not always performed by a single organ in other animals. As indicated earlier, primitive aquatic animals do not require any special provision for nitrogenous excretion. But by reason of their permeable skins they may have serious problems of osmotic and ionic regulation, especially in fresh water, where cells covering the surface of the body have the ability to actively transport salts into or out of the animal. In some cases these nonkidney regulatory activities are performed by certain specialized cells; e.g., in the gills of fishes (see below Vertebrate excretory systems: Fishes). In other cases, specialized cells are assembled into organs of salt uptake or salt elimination; e.g., the salt glands of birds (see below Vertebrate excretory systems: Birds and reptiles).
This dispersal of the regulatory function may be the primitive condition, for it is only in the more highly evolved terrestrial animals that the regulatory function is restricted to an excretory system proper. This is readily understandable in view of the need of terrestrial animals to conserve water. This evolutionary development toward one system reaches its climax in the birds, reptiles, and terrestrial insects, in which all the processes of elimination that might involve loss of water—defecation, nitrogenous excretion, and ionic regulation—converge upon the same final channel.
For the excretory organs of a wide variety of vertebrate and invertebrate animals, there is evidence that the primary process of urine production is nonselective, in that in those animals all substances dissolved in their body fluids, with the possible exception of proteins, are found in the primary urine. In many animals the primary urine is produced by filtration from the blood. At a later stage, substances in the primary urine that are useful to the body are selectively reabsorbed. In addition, a few substances are known to be actively transported (secreted) into the urine.
The nonselective formation of primary urine serves another aspect of excretion: the elimination of foreign substances. Mechanisms of active transport are highly specific to the substances transported. All dissolved constituents of the body fluids pass freely into the primary urine, and then specific reabsorptive mechanisms gather up the “wanted” substances. In this way a natural economy automatically eliminates “unwanted” substances simply by not providing mechanisms for their reabsorption.
Invertebrate excretory systems
In their detoxication mechanisms, so far as they have been investigated, the invertebrates in general conform to the principles applying to all animals, namely, that aquatic forms get rid of ammonia by diffusion through the surface of the body; terrestrial forms convert ammonia to uric acid. This implies that in aquatic forms the excretory organ is principally of importance for the composition of their body fluids. Normally, the body fluids of marine invertebrates have the same concentration as seawater; they usually differ, however, in the proportions of ions, with relatively more potassium and less magnesium than seawater. Furthermore, their urine normally has the same concentration as seawater, but correspondingly it contains less potassium and more magnesium. In freshwater invertebrates the urine is commonly, though not invariably, more dilute than the body fluids. By producing dilute urine a freshwater invertebrate conserves the salt content of its body while eliminating the water that enters its body by osmosis through its water-permeable surface.
Some invertebrates, notably echinoderms, cnidarians, and sponges, have no organs to which an excretory function can be confidently ascribed. Since all of these animals are aquatic, it is reasonable to suppose that they excrete nitrogen (as ammonia) by simple diffusion. Their body fluids (where present) are closely similar to seawater in composition, and it may be presumed that regulation operates only at the cellular level.
The excretory organs of other invertebrates are of diverse evolutionary origin. This is not to say, however, that each invertebrate phylum has evolved its own particular type of excretory organ; rather, there appear to be five main types of invertebrate excretory organ: contractile vacuole, nephridium, renal gland, coxal gland, and malpighian tubule.
The contractile vacuoles of protozoans
Some protozoan animals possess an organelle having the form of an internal sac, or vacuole, which enlarges by the accumulation of a clear fluid and then discharges its contents to the exterior. The cycle of filling and emptying may be repeated as frequently as every half minute. The chief role of the contractile vacuole appears to be in osmotic regulation, not in nitrogen excretion.
Contractile vacuoles occur more frequently and are more active in freshwater species than in closely related marine species. In fresh water, the concentration of dissolved substances in the cell is greater than in the external medium, and the cell takes in water by osmosis. If the contractile vacuole is put out of action, the cell increases in volume. If the concentration of salts in the medium increases—which would have the effect of decreasing the rate of osmosis—the rate of output by the contractile vacuole diminishes. The fluid eliminated by the vacuole is more dilute than the cytoplasm.
The nephridia of annelids, nemertines, flatworms, and rotifers
The word nephridium applies in its strict sense only to the excretory organs of annelids, but it may usefully be extended to include the excretory organs of other phyla having similar characteristics. Annelids are segmented animals that typically contain a pair of nephridia on each segment. Each nephridium has the form of a very fine tubule, often of considerable length; one end usually opens into the body cavity and the other to the exterior. In some annelids, however, the tubule does not open into the body cavity but ends internally in a cluster of cells of a special type known as solenocytes, or flame cells. The possession of solenocytes by some annelids is one of the characteristics that allies them with other nonsegmented phyla that have no true body cavity. They also have a system of tubules opening at the surface and ending internally in flame cells embedded among the other cells of the body. In most cases, there is no regular arrangement of the various parts of the system. Animals belonging to all of these phyla are primarily aquatic, and, in the few cases known, the main excretory product is ammonia. How much of it leaves the body by the nephridia and how much through the body surface is not known.
Few physiological studies have been made on nephridia other than those of the earthworm. Although the earthworm is considered a terrestrial animal, its relationships with its environment are characteristically those of a freshwater animal. The nephridium of the earthworm is longer and more complex than that of marine annelids, four regions being distinguishable. Body fluid enters the nephridium via an internal opening called the nephridiostome. As the fluid passes along the tubule, probably driven by cilia, its composition is modified. In the two lower regions of the tubule the fluid becomes progressively more dilute, presumably as a result of the reabsorption of salts. Finally, a very dilute urine passes into the bladder (an enlarged portion of the tubule) and then to the exterior through the external opening, or nephridiopore. The rate of urine flow for an earthworm may be as much as 60 percent of its body weight in a period of 24 hours.
