General features of excretory structures and functions
- Related Topics:
- urination
- urine
- secretion
- egestion
- bicarbonate threshold
The physiological process by which an organism disposes of its nitrogenous by-products is called excretion. The mechanisms for that process constitute the excretory systems, particularly such organs of vertebrate animals as elaborate and complicated as the kidney and its associated urinary ducts.
The meaning of excretion is most easily understood in the context of vertebrate physiology. The animal swallows food (ingestion). In the stomach and intestine some of the food is broken down into soluble products (digestion) that are absorbed into the body (assimilation). In the body these soluble products undergo further chemical change (metabolism); some are used by the body for growth, but most provide energy for the various activities of the body. Metabolism involves the uptake of oxygen and the elimination of carbon dioxide in the lungs (respiration). Besides carbon dioxide, compounds of nitrogen arise from metabolism and are eliminated, chiefly by the kidney, in the urine (excretion). Food not digested is eliminated through the anus (defecation).
These processes are characteristic of animals in general, but not of plants. A green plant takes in carbon dioxide from the atmosphere and nitrogen (as nitrate) from the soil. It uses the energy of sunlight to build these nutrients into the materials required for growth and in the process gives out oxygen (see photosynthesis).
In a broad sense animals live on plants, and the by-products of animals are the raw materials on which plants grow. These mutually supporting activities of plants and animals are kept precisely in balance by the activities of bacteria. Bacteria convert the urine and feces of animals (and also the dead bodies of both plants and animals) to carbon dioxide and nitrate. In the living world as a whole, carbon and nitrogen are in continuous circulation, driven by the energy of sunlight (see biosphere). Over most of the earth, for most of time, no by-products accumulate. Occasionally the cycles get out of balance, as they must have done during the prehistoric period when coal was being formed in the earth as a consequence of the failure of bacteria to decompose all the remains of plants.
Products of excretion
Although every type of organism takes in some materials and eliminates others, excretion in the strict sense is a process found only in animals. For the purposes of this article excretion will be taken to mean the elimination of nitrogenous by-products and the regulation of the composition of the body fluids.
The primary excretory product arising naturally in the animal body is ammonia, derived almost entirely from the proteins of the ingested food. In the process of digestion proteins are broken down into their constituent amino acids. Some of the amino-acid pool is then used by the animal to build up its own proteins, but a great deal is used as a source of energy to drive other vital processes. The first step in the mobilization of amino acids for energy production is deamination, the splitting off of ammonia from the amino-acid molecule. The remainder is oxidized to carbon dioxide and water, with the concomitant production of the energy-rich molecules of adenosine triphosphate (ATP; see metabolism).
Since excessive levels of ammonia are highly toxic to most animals, they must be effectively eliminated. This is no problem in small aquatic animals because ammonia rapidly diffuses, is highly soluble in water, and escapes easily into the external medium before its concentration in the body fluids can reach a dangerous level. But in terrestrial animals, and in some of the larger aquatic animals, ammonia is converted into some less harmful compounds (detoxication). In mammals, including humans, it is detoxified to urea, which may be considered as being formed by the condensation of one molecule of carbon dioxide with two molecules of ammonia (though the biochemistry of the process is more complex than that). Urea is highly soluble in water but cannot be excreted in a highly concentrated solution because of the osmotic pressure (see below) it would exert. Because the conservation of water is important for most terrestrial animals, it is not surprising that many of them have evolved more economical methods for disposing of nitrogenous by-products. Birds, reptiles, and terrestrial insects excrete nitrogen in the form of uric acid, which is highly insoluble in water and can be removed from the body as a thick suspension or even as a dry powder.
Excretory mechanisms
Osmotic pressure
In order to understand the advantages of the excretion of uric acid over urea it is necessary to know something about the behaviour of molecules in solution. Molecules of a solute (e.g., salt, sugar) in water tend to move by diffusion from a region where they are in high concentration to one where they are in low concentration, and molecules of water tend to move in the opposite direction. If a porous membrane is placed between these regions, the movements of molecules may be variously restricted depending upon their size in relation to the size of the submicroscopic pores in the membrane. The passage of water molecules from pure water through such a membrane into a solution containing molecules that are too large to pass is called osmosis, a process that takes place spontaneously and does not require energy. This process can be reversed by applying hydrostatic pressure to the solution, a process that does require energy. The level of hydrostatic pressure at which there is no net movement of water in either direction across the membrane is called the osmotic pressure of that particular solution; the greater the concentration of dissolved molecules in the solution the greater is its osmotic pressure and the greater the force needed to remove water from it.
These principles explain why more energy is required to remove water from urine containing urea than from urine containing the same weight of uric acid. The molecule of urea is smaller than that of uric acid, so that with the same weight, there are more molecules of urea to exert osmotic pressure. But an even more important difference is that whereas urea is highly soluble in water, uric acid is not. As water is progressively removed from a solution of urea, the osmotic pressure opposing further removal progressively increases. For the uric acid solution, however, as water is removed, the uric acid comes out of solution, or precipitates, when the solution is at a lower concentration, and, therefore, at a lower osmotic pressure, which does not increase further.
Regulation of water and salt balance
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.
