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P SITE 1 Toxicant: rotenone e
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e P SITE 2 Toxicant: actimycin A e SITE 3 Toxicant: cyanide 4H + O2 P 2H2O ADP + P ATP P P P e P P P
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ATP Synthetase
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Figure 11.7 Electron (e ) transport along the inner mitochondrial membrane resulting in the pumping of protons (P) out of the mitochondrial matrix. Protons are shuttled back into the matrix through the ATP synthetase complex where ATP is generated. Sites of toxicant action are indicated.
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Inhibitors of Cellular Respiration
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Cellular respiration is the process whereby energy, in the form of ATP, is generated in the cell while molecular oxygen is consumed. The process occurs along respiratory assemblies that are located in the inner mitochondrial membrane. Electrons derived from NADH or FADH2 are transferred along a chain of electron carrier proteins. This step-by-step transfer leads to the pumping of protons out of the mitochondrial matrix, resulting in the generation of a membrane potential across the inner mitochrondrial membrane. Protons are pumped out of the mitochrondrial matrix at three locations along the respiratory chain. Site 1 consists of the NADH-Q reductase complex, site 2 consists of the QH2 -cytochrome c reductase complex, and site 3 is the cytochrome c oxidase complex. ATP is generated from ADP when protons ow back across the membrane through an ATP synthetase complex to the mitochrondrial matrix. The transfer of electrons culminates with the reduction of molecular oxygen to water. Many chemicals can interfere with cellular respiration by binding to the cytochromes that constitute the electron transport chain and inhibiting the ow of electrons along this protein complex. The pesticide rotenone speci cally inhibits electron transfer early in the chain with inhibition of proton transport beginning at site 1. Actimycin A inhibits electron transfer and proton pumping at site 2. Cyanide, hydrogen sul de, and azide inhibit electron ow between the cytochrome oxidase complex and O2 preventing the generation of a proton gradient at site 3. Symptoms of toxicity from the inhibition of
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respiratory chain include excess salivation, giddiness, headache, palpitations, respiratory distress, and loss of consciousness. Potent inhibitors such as cyanide can cause death due to respiratory arrest immediately following poisoning. Some chemicals do not interfere with electron transport leading to the consumption of molecular oxygen but rather interfere with the conversion of ADP to ATP. These uncouplers of oxidative phosphorylation function by leaking protons across the inner membrane back to the mitochondrial matrix. As a result a membrane potential is not generated, and energy required for the phosphorylation of ADP to ATP is lost. The uncoupling of oxidative phosphorylation results in increased electron transport, increased oxygen consumption, and heat production. The controlled uncoupling of oxidative phosphorylation is a physiologically relevant means of maintaining body temperature by hibernating animals, some newborn animals, and in some animals that inhabit cold environments. Chemicals known to cause uncoupling of oxidative phosphorylation include 2,4-dinitrophenol, pentachlorophenol, and dicumarol. Symptoms of intoxication include accelerated respiration and pulse, ushed skin, elevated temperature, sweating, nausea, coma, and death.
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Joy, R. M. Neurotoxicology: Central and peripheral. In Encyclopedia of Toxicology, vol. 2, P. Wexler, ed. New York: Academic Press, 1998, pp. 389 413. Stryer, L. Biochemistry, 4th ed. San Francisco: W. H. Freeman, 1999. Eaton, D. L., and C. D. Klaassen. Principles of toxicology In Casarrett and Doull s Toxicology: The Basic Science of Poisons, 6th ed. C. D. Klaassen, ed. New York: McGraw-Hill, 2001, pp. 11 34. Calabrese, E. J., and L. A. Baldwin. U-shaped dose-responses in biology, toxicology, and public health. An. Rev. Public Health 22: 15 33, 2001.
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Chemical Carcinogenesis
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Carcinogenesis is the process through which cancer develops. Chemical carcinogenesis is the study of the mechanisms through which chemical carcinogens induce cancer and also involves the development/utilization of experimental systems aimed at determining whether a substance is a potential human carcinogen. An important aspect of toxicology is the identi cation of potential human carcinogens. To begin to appreciate the complexity of this subject, it is important to rst have some understanding of cancer and its etiologies. Cancer is not a single disease but a large group of diseases, all of which can be characterized by the uncontrolled growth of an abnormal cell to produce a population of cells that have acquired the ability to multiply and invade surrounding and distant tissues. It is this invasive characteristic that imparts its lethality on the host. Epidemiology studies have revealed that the incidence of most cancers increase exponentially with age (Figure 12.1). Epidemiologist have interpreted this exponential increase in cancer incidence to denote that three to seven critical mutations or hits within a single cell are required for cancer development. Molecular analyses of human tumors have con rmed the accumulation of mutations in critical genes in the development of cancer. These mutations can be the result of imperfect DNA replication/repair, oxidative DNA damage, and/or DNA damage caused by environmental carcinogens. Most cancers are monoclonal in origin (derived from a single cell) and do not arise from a single critical mutation but from the accumulation of sequential critical mutations in relevant target genes within a single cell (Figure 12.2). Initially a somatic mutation occurs in a critical gene, and this provides a growth advantage to the cell and results in the expansion of the mutant clone. Each additional critical mutation provides a further selective growth advantage resulting in clonal expansion of cells with mutations in multiple critical genes. It often requires decades for a cell clone to accumulate multiple critical mutations and for the progeny of this cell to clonally expand to produce a clinically detectable cancer. Thus the time required for accumulation of mutations in critical genes within a cell is likely related to the observation that cancer incidence increases exponentially with age.
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A Textbook of Modern Toxicology, Third Edition, edited by Ernest Hodgson ISBN 0-471-26508-X Copyright 2004 John Wiley & Sons, Inc.
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