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Cell Structure and Evolutionary History Structure, p. 22 • All microbial cells share certain basic structures in common, such as cytoplasm, a cytoplasmic membrane, ribosomes, and (usually) a cell wall. • Two structural types of cells are recognized: the prokaryote and the eukaryote. Prokaryotic cells have a simpler internal structure than eukaryotic cells, lacking membrane-enclosed organelles (Figure 2.1). Prokaryotic cell Eukaryotic cell • Viruses are not cells but depend on cells for their replication (Figure 2.3c). Comparison of cells and viruses • Ribosomes—the cell's protein-synthesizing factories—are particulate structures composed of RNA (ribonucleic acid) and various proteins suspended in the cytoplasm. • Ribosomes interact with several cytoplasmic proteins and messenger and transfer RNAs in the key process of protein synthesis (translation) (Figure 1.4). Arrangement of DNA in Microbial Cells • Genes govern the properties of cells, and a cell's complement of genes is called its genome. DNA is arranged in cells to form chromosomes. In prokaryotes, there is usually a single circular chromosome; whereas in eukaryotes, several linear chromosomes exist. • Plasmids are circular extrachromosomal genetic elements (DNA), nonessential for growth, found in prokaryotes. • The nucleus is a membrane-enclosed structure that contains the chromosomes in eukaryotic cells. The nucleoid, in contrast, is the aggregated mass of DNA that constitutes the chromosome of cells of Bacteria and Archaea (Figure 2.4). LM of E. coli EM of Nucleod of E. coli The Tree of Life • Comparative ribosomal RNA sequencing has defined the three domains of life: Bacteria, Archaea, and Eukarya. • Although species of Bacteria and Archaea share a prokaryotic cell structure, they differ dramatically in their evolutionary history. • Evolution is the change in a line of descent over time leading to new species or varieties. The evolutionary relationships between life forms are the subject of the science of phylogeny. • In addition to the genome in the chromosomes of the nucleus, mitochondria and chloroplasts of eukaryotes contain their own genomes (DNA arranged in circular fashion, as in Bacteria) and ribosomes. • Using ribosomal RNA sequencing technology (Figure 2.6), these organelles have been shown to be highly derived ancestors of specific lineages of Bacteria (Figure 2.7). he Phylogenetic Tree of Life as defined by rRNA sequencing • Mitochondria and chloroplasts were thus once free-living cells that established stable residency in cells of Eukarya eons ago. The process by which this stable arrangement developed is known as endosymbiosis. Physiological Diversity of Microorganisms • All cells need carbon and energy sources. Chemoorganotrophs obtain their energy from the oxidation of organic compounds. Chemolithotrophs obtain their energy from the oxidation of inorganic compounds. Phototrophs contain pigments that allow them to use light as an energy source. (Figure 2.8) • Autotrophs use carbon dioxide as their carbon source, whereas heterotrophs use organic carbon. • Extremophiles thrive under environmental conditions in which higher organisms cannot survive. Table 2.1 gives classes and examples of extremophiles. Prokaryotic Diversity • Several lineages are present in the domains Bacteria and Archaea, and an enormous diversity of cell morphologies and physiologies are represented there. • Retrieval and analysis of ribosomal RNA genes from cells in natural samples have shown that many phylogenetically distinct but as yet uncultured prokaryotes exist in nature. • The Proteobacteria is the largest division (called a phylum) of Bacteria (Figure 2.9). Phylogenetic tree of bacteria • The Cyanobacteria (Figure 2.12) are phylogenetic relatives of gram-positive bacteria and are oxygenic phototrophs. Filamentous cyanobacteria Oscillatoria Spirulina • There are two lineages of Archaea, the Euryarchaeota and the Crenarchaeota (Figure 2.18). Phylogenetic tree of archaea Eukaryotic Microorganisms • Microbial eukaryotes are a diverse group that includes algae, protozoa, fungi, and slime molds (Figure 2.22). Phylogenetic tree of eukaryotes • Collectively, microbial eukaryotes are known as the Protista. Some protists, such as the algae, are phototrophic. • Cells of algae and fungi have cell walls, whereas the protozoa do not. • Some algae and fungi have developed mutualistic associations called lichens.