Chromosomal Elements:- DNA Molecules Are Much Longer Than the Cellular Packages That Contain Them
Chromosomal DNAs are often many orders of magnitude longer than the cells or viruses in which they are found ( Table 24–1). This is true of every class of organism or parasite.
Viruses are not free-living organisms; rather, they are infectious parasites that use the resources of a host cell to carry out many of the processes they require to propagate. Many viral particles consist of no more than a genome (usually a single RNA or DNA molecule) surrounded by a protein coat. Almost all plant viruses and some bacterial and animal viruses have RNA genomes. These genomes tend to be particularly small. For example, the genomes of mammalian retroviruses such as HIV are about 9,000 nucleotides long, and that of the bacteriophage Q has 4,220 nucleotides. Both types of viruses have single stranded RNA genomes. The genomes of DNA viruses vary greatly in size (Table 24–1). Many viral DNAs are circular for at least part of their life cycle. During viral replication within a host cell, specific types of viral DNA called replicative forms may appear; for example, many linear DNAs become circular and all single-stranded DNAs become double-stranded. A typical medium-sized DNA virus is bacteriophage (lambda), which infects E. coli. In its replicative form inside cells, DNA is a circular double helix. This double-stranded DNA contains 48,502 bp and has a contour length of 17.5 m. Bacteriophage X174 is a much smaller DNA virus; the DNA in the viral particle is a single-stranded circle, and the double-stranded replicative form contains 5,386 bp. Although viral genomes are small, the contour lengths of their DNAs are much greater than the long dimensions of the viral particles that contain them. The DNA of bacteriophage T4, for example, is about 290 times longer than the viral particle itself (Table 24–1).
Bacteria A single E. coli cell contains almost 100 times as much DNA as a bacteriophage particle. The chromosome of an E. coli cell is a single double-stranded circular DNA molecule. Its 4,639,221 bp have a contour length of about 1.7 mm, some 850 times the length of the E. coli cell (Fig. 1). In addition to the very large, circular DNA chromosome in their nucleoid, many bacteria contain one or more small circular DNA molecules that are free in the cytosol. These extrachromosomal elements are called plasmids (Fig. 1). Most plasmids are only a few thousand base pairs long, but some contain more than 10,000 bp. They carry genetic information and undergo replication to yield daughter plasmids, which pass into the daughter cells at cell division. Plasmids have been found in yeast and other fungi as well as in bacteria. In many cases plasmids confer no obvious advantage on their host, and their sole function appears to be self-propagation. However, some plasmids carry genes that are useful to the host bacterium. For example, some plasmid genes make a host bacterium resistant to antibacterial agents. Plasmids carrying the gene for the enzyme -lactamase confer resistance to -lactam antibiotics such as penicillin and amoxicillin (see Box 20–1). These and similar plasmids may pass from an antibiotic-resistant cell to an antibiotic-sensitive cell of the same or another bacterial species, making the recipient cell antibiotic resistant. The extensive use of antibiotics in some human populations has served as a strong selective force, encouraging the spread of antibiotic resistance–coding plasmids (as well as transposable elements, described below, that harbor similar genes) in disease-causing bacteria and creating bacterial strains that are resistant to several antibiotics. Physicians are becoming increasingly reluctant to prescribe antibiotics unless a clear clinical need is confirmed. For similar rea sons, the widespread use of antibiotics in animal feeds is being curbed.
Eukaryotes A yeast cell, one of the simplest eukaryotes, has 2.6 times more DNA in its genome than an E. coli cell (Table 24–2). Cells of Drosophila, the fruit fly used in classical genetic studies, contain more than 35 times as much DNA as E. coli cells, and human cells have almost 700 times as much. The cells of many plants and amphibians contain even more. The genetic material of eukaryotic cells is apportioned into chromosomes, the diploid (2n) number depending on the species (Table 24–2). A human somatic cell, for example, has 46 chromosomes (2). Each chromosome of a eukaryotic cell, such as that shown in Figure 2a, contains a single, very large, duplex DNA molecule. The DNA molecules in the 24 different types of human chromo somes (22 matching pairs plus the X and Y sex chromosomes) vary in length over a 25-fold range. Each type of chromosome in eukaryotes carries a characteristic set of genes. Interestingly, the number of genes does not vary nearly as much as does genome size (see Chapter 9 for a discussion of the types of sequences, besides genes, that contribute to genome size). The DNA of one human genome (22 chromosomes plus X and Y or two X chromosomes), placed end to end, would extend for about a meter. Most human cells are diploid and each cell contains a total of 2 m of DNA. An adult human body contains approximately 1014 cells and thus a total DNA length of 2 X 1011 km. Compare this with the circumference of the earth (4X104 km) or the distance between the earth and the sun (1.5 X 108 km)—a dramatic illustration of the extraordinary degree of DNA compaction in our cells.
Eukaryotic cells also have organelles, mitochondria (Fig. 3) and chloroplasts, that contain DNA. Mitochondrial DNA (mtDNA) molecules are much smaller than the nuclear chromosomes. In animal cells, mtDNA contains fewer than 20,000 bp (16,569 bp in human mtDNA) and is a circular duplex. Each mitochondrion typically has two to ten copies of this mtDNA molecule, and the number can rise to hundreds in certain cells when an embryo is undergoing cell differentiation. In a few organisms (trypanosomes, for example) each mitochondrion contains thousands of copies of mtDNA, or ganizedinto a complex and interlinked matrix known as a kinetoplast. Plant cell mtDNA ranges in size from 200,000 to 2,500,000 bp. Chloroplast DNA (cpDNA) also exists as circular duplexes and ranges in size from 120,000 to 160,000 bp. The evolutionary origin of mitochondrial and chloroplast DNAs has been the subject of much speculation. A widely accepted view is that they are vestiges of the chromosomes of ancient bacteria that gained access to the cytoplasm of host cells and became the precursors of these organelles .
Mitochondrial DNA codes for the mitochondrial tRNAs and rRNAs and for a few mitochondrial proteins. More than 95% of mitochondrial proteins are encoded by nu clear DNA. Mitochondria and chloroplasts divide when the cell divides. Their DNA is replicated before and during division, and the daughter DNA molecules pass into the daughter organelles.
FIGURE 24–4 DNA from a lysed E. coli cell. In this electron micrograph several small, circular plasmid DNAs are indicated by white arrows. The black spots and white specks are artifacts of the preparation.
FIGURE 24–5 Eukaryotic chromosomes. (a) A pair of linked and condensed sister chromatids from a human chromosome. Eukaryotic chromosomes are in this state after replication and at metaphase during mitosis. (b) A complete set of chromosomes from a leukocyte from one of the authors. There are 46 chromosomes in every normal human somatic cell.
FIGURE 24–6 A dividing mitochondrion. Some mitochondrial proteins and RNAs are encoded by one of the copies of the mitochondrial DNA (none of which are visible here). The DNA (mtDNA) is replicated each time the mitochondrion divides, before cell division.
