Bacteria are prokaryotes, lacking well-defined nuclei and membrane-bound organelles, and with chromosomes composed of a single closed DNA circle. They come in many shapes and sizes, from minute spheres, cylinders and spiral threads, to flagellated rods, and filamentous chains.
In the late 1600s, Antoni van Leeuwenhoek became the first to study bacteria under the microscope. During the nineteenth century, the French scientist Louis Pasteur and the German physician Robert Koch demonstrated the role of bacteria as pathogens (causing disease). The twentieth century saw numerous advances in bacteriology, indicating their diversity, ancient lineage, and general importance. Most notably, a number of scientists around the world made contributions to the field of microbial ecology, showing that bacteria were essential to food webs and for the overall health of the Earth's ecosystems. The discovery that some bacteria produced compounds lethal to other bacteria led to the development of antibiotics, which revolutionized the field of medicine.
There are two different ways of grouping bacteria. They can be divided into three types based on their response to gaseous oxygen. Aerobic bacteria require oxygen for their health and existence and will die without it. Anerobic bacteria can't tolerate gaseous oxygen at all and die when exposed to it. Facultative aneraobes prefer oxygen, but can live without it.
The second way of grouping them is by how they obtain their energy. Bacteria that have to consume and break down complex organic compounds are heterotrophs. This includes species that are found in decaying material as well as those that utilize fermentation or respiration. Bacteria that create their own energy, fueled by light or through chemical reactions, are autotrophs.
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Capsule - Some species of bacteria have a third protective covering, a capsule made up of polysaccharides (complex carbohydrates). Capsules play a number of roles, but the most important are to keep the bacterium from drying out and to protect it from phagocytosis (engulfing) by larger microorganisms. The capsule is a major virulence factor in the major disease-causing bacteria, such as Escherichia coli and Streptococcus pneumoniae. Nonencapsulated mutants of these organisms are avirulent, i.e. they don't cause disease.
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Cell Envelope - The cell envelope is made up of two to three layers: the interior cytoplasmic membrane, the cell wall, and -- in some species of bacteria -- an outer capsule.
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Cell Wall - Each bacterium is enclosed by a rigid cell wall composed of peptidoglycan, a protein-sugar (polysaccharide) molecule. The wall gives the cell its shape and surrounds the cytoplasmic membrane, protecting it from the environment. It also helps to anchor appendages like the pili and flagella, which originate in the cytoplasm membrane and protrude through the wall to the outside. The strength of the wall is responsible for keeping the cell from bursting when there are large differences in osmotic pressure between the cytoplasm and the environment.
Cell wall composition varies widely amongst bacteria and is one of the most important factors in bacterial species analysis and differentiation. For example, a relatively thick, meshlike structure that makes it possible to distinguish two basic types of bacteria. A technique devised by Danish physician Hans Christian Gram in 1884, uses a staining and washing technique to differentiate between the two forms. When exposed to a gram stain, gram-positive bacteria retain the purple color of the stain because the structure of their cell walls traps the dye. In gram-negative bacteria, the cell wall is thin and releases the dye readily when washed with an alcohol or acetone solution.
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Cytoplasm - The cytoplasm, or protoplasm, of bacterial cells is where the functions for cell growth, metabolism, and replication are carried out. It is a gel-like matrix composed of water, enzymes, nutrients, wastes, and gases and contains cell structures such as ribosomes, a chromosome, and plasmids. The cell envelope encases the cytoplasm and all its components. Unlike the eukaryotic (true) cells, bacteria do not have a membrane enclosed nucleus. The chromosome, a single, continuous strand of DNA, is localized, but not contained, in a region of the cell called the nucleoid. All the other cellular components are scattered throughout the cytoplasm.
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Cytoplasmic Membrane - A layer of phospholipids and proteins, called the cytoplasmic membrane, encloses the interior of the bacterium, regulating the flow of materials in and out of the cell. This is a structural trait bacteria share with all other living cells; a barrier that allows them to selectively interact with their environment. Membranes are highly organized and asymmetric having two sides, each side with a different surface and different functions. Membranes are also dynamic, constantly adapting to different conditions.
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Flagella - Flagella (singular, flagellum) are hairlike structures that provide a means of locomotion for those bacteria that have them. They can be found at either or both ends of a bacterium or all over its surface. The flagella beat in a propeller-like motion to help the bacterium move toward nutrients; away from toxic chemicals; or, in the case of the photosynthetic cyanobacteria; toward the light.
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Nucleoid - The nucleoid is a region of cytoplasm where the chromosomal DNA is located. It is not a membrane bound nucleus, but simply an area of the cytoplasm where the strands of DNA are found. Most bacteria have a single, circular chromosome that is responsible for replication, although a few species do have two or more. Smaller circular auxiliary DNA strands, called plasmids, are also found in the cytoplasm.
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Pili - Many species of bacteria have pili (singular, pilus), small hairlike projections emerging from the outside cell surface. These outgrowths assist the bacteria in attaching to other cells and surfaces, such as teeth, intestines, and rocks. Without pili, many disease-causing bacteria lose their ability to infect because they're unable to attach to host tissue. Specialized pili are used for conjugation, during which two bacteria exchange fragments of plasmid DNA.
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Ribosomes - Ribosomes are microscopic "factories" found in all cells, including bacteria. They translate the genetic code from the molecular language of nucleic acid to that of amino acids—the building blocks of proteins. Proteins are the molecules that perform all the functions of cells and living organisms. Bacterial ribosomes are similar to those of eukaryotes, but are smaller and have a slightly different composition and molecular structure. Bacterial ribosomes are never bound to other organelles as they sometimes are (bound to the endoplasmic reticulum) in eukaryotes, but are free-standing structures distributed throughout the cytoplasm. There are sufficient differences between bacterial ribosomes and eukaryotic ribosomes that some antibiotics will inhibit the functioning of bacterial ribosomes, but not a eukaryote's, thus killing bacteria but not the eukaryotic organisms they are infecting.
One of those components, plasmids, are small, extrachromosomal genetic structures carried by many strains of bacteria. Like the chromosome, plasmids are made of a circular piece of DNA. Unlike the chromosome, they are not involved in reproduction. Only the chromosome has the genetic instructions for initiating and carrying out cell division, or binary fission, the primary means of reproduction in bacteria. Plasmids replicate independently of the chromosome and, while not essential for survival, appear to give bacteria a selective advantage.
Plasmids are passed on to other bacteria through two means. For most plasmid types, copies in the cytoplasm are passed on to daughter cells during binary fission. Other types of plasmids, however, form a tubelike structure at the surface called a pilus that passes copies of the plasmid to other bacteria during conjugation, a process by which bacteria exchange genetic information. Plasmids have been shown to be instrumental in the transmission of special properties, such as antibiotic drug resistance, resistance to heavy metals, and virulence factors necessary for infection of animal or plant hosts. The ability to insert specific genes into plasmids have made them extremely useful tools in the fields of molecular biology and genetics, specifically in the area of genetic engineering.
Bacteria are the smallest and most versatile independently living cells known. Bacterial structure is responsible for some of their unique pathogenicity.
Because of the simplicity of bacteria relative to larger organisms and the ease with which they can be manipulated experimentally, the cell structure of bacteria has been well studied, revealing many biochemical principles that have been subsequently applied to other organisms.
Gross Morphology
Bacteria have characteristic shapes (cocci, rods, spirals, etc.) and often occur in characteristic aggregates (pairs, chains, tetrads, clusters, etc.).
These traits are usually typical for a genus and are diagnostically useful.
Overall Cell Structure
Bacteria (prokaryotes) have a nucleoid (nuclear body) rather than an enveloped nucleus and lack membrane-bound cytoplasmic organelles. The plasma membrane in prokaryotes performs many of the functions carried out by membranous organelles in multicelled organisms(eukaryotes).
Multiplication is by binary fission.
Bacterial Wall
Bacteria are protected by a rigid cell wall composed of peptidoglycans. A cell wall is a layer located outside the cell membrane. A peptidoglycan cell wall composed of disaccharides and amino acids. The bacterial cell wall is often a target for antibiotic treatment.
Flagella: The flagella of motile bacteria differ in structure from eukaryotic flagella. A basal body anchored in the plasma membrane and cell wall gives rise to a cylindrical protein filament. The flagellum moves by whirling about its long axis. The number and arrangement of flagella on the cell are diagnostically useful.
Pili (Fimbriae): Pili are slender, hairlike, proteinaceous appendages on the surface of many (particularly Gram-negative) bacteria. They are important in adhesion to host surfaces.
Capsules: Some bacteria form a thick outer capsule of high-molecular-weight, viscous polysaccharide gel; others have more amorphous slime layers. Capsules confer resistance to phagocytosis.
Important Chemical Components of Surface Structures
Cell Wall Peptidoglycans: Both Gram-positive and Gram-negative bacteria possess cell wall peptidoglycans, which confer the characteristic cell shape and provide the cell with mechanical protection. Peptidoglycans are unique to prokaryotic organisms and consist of a glycan backbone of muramic acid and glucosamine (both N-acetylated), and peptide chains highly cross-linked with bridges in Gram-positive bacteria (e.g., Staphylococcus aureus) or partially cross-linked in Gram-negative bacteria (e.g., Escherichia coli). The cross-linking transpeptidase enzymes are some of the targets for b-lactam antibiotics.
Teichoic Acids: Teichoic acids are polyol phosphate polymers bearing a strong negative charge. They are covalently linked to the peptidoglycan in some Gram-positive bacteria. They are strongly antigenic, but are generally absent in Gram-negative bacteria.
Lipoteichoic Acids: Lipoteichoic acids as membrane teichoic acids are polymers of amphiphitic glycophosphates with the lipophilic glycolipid and anchored in the cytoplasmic membrane. They are antigenic, cytotoxic and adhesins (e.g., Streptococcus pyogenes).
Lipopolysaccharides: One of the major components of the outer membrane of Gram-negative bacteria is lipopolysaccharide (endotoxin), a complex molecule consisting of a lipid A anchor, a polysaccharide core, and chains of carbohydrates. Sugars in the polysaccharide chains confer serologic specificity.
Wall-Less Forms: Two groups of bacteria devoid of cell wall peptidoglycans are the Mycoplasmaspecies, which possess a surface membrane structure, and the L-forms that arise from either Gram-positive or Gram-negative bacterial cells that have lost their ability to produce the peptidoglycan structures.
Cytoplasmic Structures
Plasma Membrane: The bacterial plasma membrane is composed primarily of protein and phospholipid (about 3:1). It performs many functions, including transport, biosynthesis, and energy transduction.
Organelles: The bacterial cytoplasm is densely packed with 70S ribosomes. Other granules represent metabolic reserves (e.g., poly-β-hydroxybutyrate, polysaccharide, polymetaphosphate, and metachromatic granules).
Endospores: Bacillus and Clostridium species can produce endospores: heat-resistant, dehydrated resting cells that are formed intracellularly and contain a genome and all essential metabolic machinery. The endospore is encased in a complex protective spore coat.
Multiplication
The process whereby a cell divides asexually to produce two daughter cells.
All bacteria have a peptidoglycan cell wall.
Bacteria are classified as gram positive or gram negative. The Gram stain is used to tell whether a bacterium is gram negative (red) or gram positive (blue). The structure of the bacterial cell wall explains why bacteria are either gram positive or gram negative. . In gram-positive bacteria it is thick, whereas in gram-negative bacteria it is thin. Importantly, gram-negative bacteria have an additional membrane outside the cell wall, consisting mainly of lipopolysaccharide (LPS) . It is this LPS layer that prevents the blue dye applied during a Gram stain from being trapped in the cell wall, unlike gram-positive bacteria, which take up the blue dye. The process of Gram staining consists of applying the blue (crystal violet) dye, washing away the extra with alcohol, then applying a red (safranin) counterstain, which will adhere to anything not already stained by the blue dye (i.e., gram-negative bacteria). Hence, gram-positive bacteria appear blue, and gram-negative bacteria appear red.
Besides being important for Gram staining, the LPS layer of gram-negative bacteria can activate a cytokine-mediated immune response in a host, one of the major causes of septic shock. Specifically, the component of LPS called lipid A is an endotoxin (“endo” because it is found on the innermost portion of the LPS membrane). When the immune system lyses these bacteria in the blood, lipid A activates the cytokine cascade. Gram-positive bacteria and fungi can also cause septic shock.
“The lipid A component of LPS is a major cause of septic shock.”
Importantly, the peptidoglycan cell wall consists of glycoproteins covalently bonded together by the enzyme transpeptidase , also called penicillin-binding protein because it is the enzyme inhibited by penicillin.
Bacteria shape is also important for distinguishing among the various species. They can be cocci (spherical), bacilli (rods), coccobacilli (short rods), spiral (comma shaped, S shaped, spiral shaped), or pleomorphic (lacking a characteristic shape).
Metabolic characteristics are also key to distinguishing among the various species. In fact, oxygen can be toxic through the formation of superoxide radicals, and only bacteria containing enzymes to break these down ( catalase, peroxidase, or superoxide dismutase ) can survive in oxygen. Obligate aerobes require oxygen to survive; facultative anaerobes prefer oxygen but can live without it through fermentation; microaerophilic bacteria can tolerate low amounts of oxygen but prefer to ferment their energy; and obligate anaerobes can’t live in the presence of oxygen.
Finally, there is a category of bacteria that are obligate intracellular organisms. These bacteria cannot make their own adenosine triphosphate (ATP) and must rely on the host cell machinery for survival. These bacteria only live within other cells, providing us with challenges for detection and treatment. Facultative intracellular organisms don’t need to live in cells but have the ability to survive and replicate in macrophages after being phagocytosed because of enzymes protecting them against superoxide radical digestion. These organisms are Listeria monocytogenes, Salmonella typhi, Yersinia, Francisella tularensis, Brucella, Legionella, and Mycobacterium.
Some bacterial structures are important to know about:
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Flagella are used for movement and come in a variety of shapes.
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Capsules are an additional sugar protective layer around some bacteria ( Bacillus anthracis is the only bacterium with an amino acid capsule). The capsules aid in immune evasion, and encapsulated bugs are particularly dangerous in individuals without spleens (e.g., sickle cell disease) because that is where most phagocytosis of opsonized encapsulated bugs occurs. Opsonization is where the bacteria is coated with antibodies and therefore marked for phagocytosis and destruction. To detect an encapsulated bug, one must use either the India ink stain or the Quellung reaction (in which antibodies that bind to capsules are used to appreciate a swelling of the capsule under the microscope; Fig. 5-3 ).
Figure 5-3Micrograph showing swelling of the capsule of Klebsiella pneumonia after antibody administration, demonstrating the Quellung reaction.(From Greenwood D, Slack R, Peitherer J, Barer M. Medical Microbiology . 17th ed. Philadelphia: Elsevier; 2007.) - ❍
Spores are found in a few species of bacteria and are a protective form of the bacteria that allow them to survive harsh conditions until a host is found.
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Biofilms are similar to capsules in that they are made of a lattice of polysaccharides, in this case allowing the bacteria to bind to prosthetic devices (found, e.g., in Staphylococcus epidermidis ).
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Toxins are an important part of the disease-causing mechanisms of many bacteria. We can divide toxins into the organ system they infect (e.g., neurotoxins and enterotoxins), where they are found in the bacteria ( exotoxins are released by all gram-positive bacteria except Listeria, which releases an endotoxin, and endotoxins such as lipid A, which was previously discussed), or by the effects they cause ( pyrogenic exotoxins cause fever, tissue invasive exotoxins allow bacteria to tunnel through tissue, and miscellaneous exotoxins do various things tailored for certain bacteria).
What sort of individuals are susceptible to infection by encapsulated bacteria?
Because the spleen normally sequesters encapsulated bacteria, patients who have undergone splenectomy are at a greater risk for incurring infection by encapsulated bacteria.