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Tamarin: Principles of Genetics, Seventh Edition
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III. Molecular Genetics
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16. Gene Expression: Control in Eukaryotes
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The McGraw Hill Companies, 2001
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Immunogenetics
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Figure 2 AIDS virion structure. Numbers associated with proteins are kilodalton masses (e.g., gp120 is a 120-kilodalton
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(From Nester et al., Microbiology: A Human Perspective, 3rd edition. Copyright 2001 The McGraw-Hill Companies, Inc. Reprinted
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over 33 million people are affected. Most of those who got the disease before 1990 have died. However, the infection rate seems to have peaked in the United States in 1985; the only area in which infections are increasing is through heterosexual sex. HIV GENES As mentioned, a retrovirus minimally contains only the gag (group antigen gene), pol (polymerase), and env (envelope) genes. The viral messenger RNA is translated starting with gag ( g. 3). There is a translation termination signal at the end of the gag gene that is occasionally read through, resulting in a gag-pol protein. The env gene is translated only after the viral RNA is spliced to remove the gag-pol region. The protein products of all three genes are further modi ed by cleavage and other changes (phosphorylation and glycosylation), resulting in core virion proteins from gag, reverse transcriptase, protease, and in-
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tegrase from pol, and envelope glycoproteins from env. The HIV retrovirus is especially complicated. Not only does it have the gag, pol, and env genes, but it
also has six other genes ( g. 4), including two main regulatory genes, tat and rev. One, tat (for trans-activating transcription factor), has a protein product that binds at a sequence
Figure 3 Expression of a retroviral mRNA. Translation begins with the gag gene and occasionally, due to read-through, proceeds through the gag-pol genes. The results are core virion proteins and the enzymes reverse transcriptase, protease, and integrase. Splicing must take place before env can be translated. Cleavage of the primary transcript and some modi cation produces envelope glycoproteins. continued
Tamarin: Principles of Genetics, Seventh Edition
III. Molecular Genetics
16. Gene Expression: Control in Eukaryotes
The McGraw Hill Companies, 2001
Sixteen
Gene Expression: Control in Eukaryotes
BOX 16.3 CONTINUED
in the long terminal repeat named TAR, for trans-activating response element. Tat enhances the processivity of transcription of the proviral DNA and also recruits chromatinremodeling proteins to the promoter. The product of the other regulatory gene, rev (regulation of expression of virion proteins), binds at a region in the env gene called RRE (for rev response element) and enhances the transport of viral messenger RNAs into the cytoplasm. Together, tat and rev are responsible for the major expression of viral structural genes (gag, pol, and env). The four remaining genes vif, vpr, nef, and vpu are called the accessory genes because it rst seemed that their action was not necessary for viral functioning. We now know that each gene produces a protein that has a role in viral replication and infectivity. The Vpr protein (viral protein R) is involved in transporting the viral RNA to the nucleus.Vpr can also induce cell cycle arrest at G2, which may have a role in protecting infected cells from cytotoxic T-cell activities. Vpu (viral protein U ) degrades CD4; this action frees viral surface protein precursors from the endoplasmic reticulum. In addition, degradation of CD4 helps prevent superinfection of cells, keeping them alive longer. The main function of Vif (viral infectivity factor) is to stabilize the virion. Nef (negative factor) was originally thought to be a negative regulator of viral activity, hence its name. However, it is now known that Nef can reduce production of cellular CD4 protein and enhance infection by viruses free in the blood. TESTING AND TREATMENT AIDS testing is done by various techniques, such as western blots, looking for antibodies to the AIDS proteins, usually gp120, gp41, and reverse transcriptase. Initially, dideoxy nucleotides, such as the drug 3 -azido-2 , 3 -dideoxythymidine (AZT, g. 5) and dideoxyinosine were used to treat AIDS. AZT is a thymidine analogue without a 3 -OH group, meaning that it causes chain termination during DNA replication. It seems that during the reverse transcription process, reverse transcriptase preferentially chooses AZT over normal thymidinecontaining nucleotides, whereas mammalian DNA polymerases prefer the opposite. Thus, AZT preferentially prevents the reverse transcription of the HIV RNA, keeping it at levels that are not toxic to the cell. Dideoxyinosine has the same effect and has also been licensed as an AIDS treatment. Unfortunately, the AIDS virus mutates at a high rate, rendering these single-substance treatments ultimately ineffective. In 1996, treatment success improved remarkably when new therapies involving combinations of drugs, including protease inhibitors, were developed. (Dr. David Ho of the Aaron Diamond AIDS Research Center in New York City was named Time Magazine s Man of the Year for his role in this therapy.) Thus, at the moment, optimism is rising that AIDS may be controllable and eventually curable.
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