Explain how different excretion systems function

Microorganisms and invertebrate animals use more primitive and simple mechanisms to get rid of their metabolic wastes than the mammalian system of kidney and urinary function. Three excretory systems evolved in organisms before complex kidneys: vacuoles, flame cells, and Malpighian tubules.

Contractile Vacuoles in Microorganisms

Figure 1. Some unicellular organisms, such as the amoeba, ingest food by endocytosis. The food vesicle fuses with a lysosome, which digests the food. Waste is excreted by exocytosis.

The most fundamental feature of life is the presence of a cell. In other words, a cell is the simplest functional unit of a life. Bacteria are unicellular, prokaryotic organisms that have some of the least complex life processes in place; however, prokaryotes such as bacteria do not contain membrane-bound vacuoles. The cells of microorganisms like bacteria, protozoa, and fungi are bound by cell membranes and use them to interact with the environment. Some cells, including some leucocytes in humans, are able to engulf food by endocytosis—the formation of vesicles by involution of the cell membrane within the cells. The same vesicles are able to interact and exchange metabolites with the intracellular environment. In some unicellular eukaryotic organisms such as the amoeba, shown in Figure 1, cellular wastes and excess water are excreted by exocytosis, when the contractile vacuoles merge with the cell membrane and expel wastes into the environment. Contractile vacuoles (CV) should not be confused with vacuoles, which store food or water.

Flame Cells of Planaria and Nephridia of Worms

As multi-cellular systems evolved to have organ systems that divided the metabolic needs of the body, individual organs evolved to perform the excretory function. Planaria are flatworms that live in fresh water. Their excretory system consists of two tubules connected to a highly branched duct system. The cells in the tubules are called flame cells (or protonephridia) because they have a cluster of cilia that looks like a flickering flame when viewed under the microscope, as illustrated in Figure 2a. The cilia propel waste matter down the tubules and out of the body through excretory pores that open on the body surface; cilia also draw water from the interstitial fluid, allowing for filtration. Any valuable metabolites are recovered by reabsorption. Flame cells are found in flatworms, including parasitic tapeworms and free-living planaria. They also maintain the organism’s osmotic balance.

Earthworms (annelids) have slightly more evolved excretory structures called nephridia, illustrated in Figure 2b. A pair of nephridia is present on each segment of the earthworm. They are similar to flame cells in that they have a tubule with cilia. Excretion occurs through a pore called the nephridiopore. They are more evolved than the flame cells in that they have a system for tubular reabsorption by a capillary network before excretion.

Figure 2. In the excretory system of the (a) planaria, cilia of flame cells propel waste through a tubule formed by a tube cell. Tubules are connected into branched structures that lead to pores located all along the sides of the body. The filtrate is secreted through these pores. In (b) annelids such as earthworms, nephridia filter fluid from the coelom, or body cavity. Beating cilia at the opening of the nephridium draw water from the coelom into a tubule. As the filtrate passes down the tubules, nutrients and other solutes are reabsorbed by capillaries. Filtered fluid containing nitrogenous and other wastes is stored in a bladder and then secreted through a pore in the side of the body.

Malpighian Tubules of Insects

Malpighian tubules are found lining the gut of some species of arthropods, such as the bee illustrated in Figure 3.

Figure 3. Malpighian tubules of insects and other terrestrial arthropods remove nitrogenous wastes and other solutes from the hemolymph. Na⁺ and/or K⁺ ions are actively transported into the lumen of the tubules. Water then enters the tubules via osmosis, forming urine. The urine passes through the intestine, and into the rectum. There, nutrients diffuse back into the hemolymph. Na⁺ and/or K⁺ ions are pumped into the hemolymph, and water follows. The concentrated waste is then excreted.

They are usually found in pairs and the number of tubules varies with the species of insect. Malpighian tubules are convoluted, which increases their surface area, and they are lined with microvilli for reabsorption and maintenance of osmotic balance. Malpighian tubules work cooperatively with specialized glands in the wall of the rectum. Body fluids are not filtered as in the case of nephridia; urine is produced by tubular secretion mechanisms by the cells lining the Malpighian tubules that are bathed in hemolymph (a mixture of blood and interstitial fluid that is found in insects and other arthropods as well as most mollusks). Metabolic wastes like uric acid freely diffuse into the tubules. There are exchange pumps lining the tubules, which actively transport H⁺ ions into the cell and K⁺ or Na⁺ ions out; water passively follows to form urine. The secretion of ions alters the osmotic pressure which draws water, electrolytes, and nitrogenous waste (uric acid) into the tubules. Water and electrolytes are reabsorbed when these organisms are faced with low-water environments, and uric acid is excreted as a thick paste or powder. Not dissolving wastes in water helps these organisms to conserve water; this is especially important for life in dry environments.

Nitrogenous Wastes

Of the four major macromolecules in biological systems, both proteins and nucleic acids contain nitrogen. During the catabolism, or breakdown, of nitrogen-containing macromolecules, carbon, hydrogen, and oxygen are extracted and stored in the form of carbohydrates and fats. Excess nitrogen is excreted from the body. Nitrogenous wastes tend to form toxic ammonia, which raises the pH of body fluids. The formation of ammonia itself requires energy in the form of ATP and large quantities of water to dilute it out of a biological system. Animals that live in aquatic environments tend to release ammonia into the water. Animals that excrete ammonia are said to be ammonotelic. Terrestrial organisms have evolved other mechanisms to excrete nitrogenous wastes. The animals must detoxify ammonia by converting it into a relatively nontoxic form such as urea or uric acid. Mammals, including humans, produce urea, whereas reptiles and many terrestrial invertebrates produce uric acid. Animals that secrete urea as the primary nitrogenous waste material are called ureotelic animals.

Terrestrial Animals: The Urea Cycle

The urea cycle is the primary mechanism by which mammals convert ammonia to urea. Urea is made in the liver and excreted in urine. The overall chemical reaction by which ammonia is converted to urea is 2 NH₃ (ammonia) + CO₂ + 3 ATP + H₂O → H₂N-CO-NH₂ (urea) + 2 ADP + 4 P_i + AMP.

The urea cycle utilizes five intermediate steps, catalyzed by five different enzymes, to convert ammonia to urea, as shown in Figure 4. The amino acid L-ornithine gets converted into different intermediates before being regenerated at the end of the urea cycle. Hence, the urea cycle is also referred to as the ornithine cycle. The enzyme ornithine transcarbamylase catalyzes a key step in the urea cycle and its deficiency can lead to accumulation of toxic levels of ammonia in the body. The first two reactions occur in the mitochondria and the last three reactions occur in the cytosol. Urea concentration in the blood, called blood urea nitrogen or BUN, is used as an indicator of kidney function.

Figure 4. The urea cycle converts ammonia to urea.

Birds and Reptiles: Uric Acid

Birds, reptiles, and most terrestrial arthropods convert toxic ammonia to uric acid or the closely related compound guanine (guano) instead of urea. Mammals also form some uric acid during breakdown of nucleic acids. Uric acid is a compound similar to purines found in nucleic acids. It is water insoluble and tends to form a white paste or powder; it is excreted by birds, insects, and reptiles. Conversion of ammonia to uric acid requires more energy and is much more complex than conversion of ammonia to urea Figure 5.

Figure 5. Nitrogenous waste is excreted in different forms by different species. These include (a) ammonia, (b) urea, and (c) uric acid. (credit a: modification of work by Eric Engbretson, USFWS; credit b: modification of work by B. “Moose” Peterson, USFWS; credit c: modification of work by Dave Menke, USFWS)

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