13.1 Prokaryotic Diversity
Prokaryotes existed for billions of years before plants and animals appeared. Microbial mats are thought to represent the earliest forms of life on Earth, and there is fossil evidence, called stromatolites, of their presence about 3.5 billion years ago. During the first 2 billion years, the atmosphere was anoxic and only anaerobic organisms were able to live. Cyanobacteria began the oxygenation of the atmosphere. The increase in oxygen concentration allowed the evolution of other life forms.
Prokaryotes (domains Archaea and Bacteria) are single-celled organisms lacking a nucleus. They have a single piece of circular DNA in the nucleoid area of the cell. Most prokaryotes have cell wall outside the plasma membrane. Bacteria and Archaea differ in the compositions of their cell membranes and the characteristics of their cell walls.
Bacterial cell walls contain peptidoglycan. Archaean cell walls do not have peptidoglycan. Bacteria can be divided into two major groups: Gram-positive and Gram-negative. Gram-positive organisms have a thick cell wall. Gram-negative organisms have a thin cell wall and an outer membrane. Prokaryotes use diverse sources of energy to assemble macromolecules from smaller molecules. Phototrophs obtain their energy from sunlight, whereas chemotrophs obtain it from chemical compounds.
Infectious diseases caused by bacteria remain among the leading causes of death worldwide. The excessive use of antibiotics to control bacterial infections has resulted in resistant forms of bacteria being selected. Foodborne diseases result from the consumption of contaminated food, pathogenic bacteria, viruses, or parasites that contaminate food. Prokaryotes are used in human food products. Microbial bioremediation is the use of microbial metabolism to remove pollutants. The human body contains a huge community of prokaryotes, many of which provide beneficial services such as the development and maintenance of the immune system, nutrition, and protection from pathogens.
13.2 Eukaryotic Origins
The first eukaryotes evolved from ancestral prokaryotes by a process that involved membrane proliferation, the loss of a cell wall, the evolution of a cytoskeleton, and the acquisition and evolution of organelles. Nuclear eukaryotic genes appear to have had an origin in the Archaea, whereas the energy machinery of eukaryotic cells appears to be bacterial in origin. The mitochondria and plastids originated from endosymbiotic events when ancestral cells engulfed an aerobic bacterium (in the case of mitochondria) and a photosynthetic bacterium (in the case of chloroplasts). The evolution of mitochondria likely preceded the evolution of chloroplasts. There is evidence of secondary endosymbiotic events in which plastids appear to be the result of endosymbiosis after a previous endosymbiotic event.
Protists are extremely diverse in terms of biological and ecological characteristics due in large part to the fact that they are an artificial assemblage of phylogenetically unrelated groups. Protists display highly varied cell structures, several types of reproductive strategies, virtually every possible type of nutrition, and varied habitats. Most single-celled protists are motile, but these organisms use diverse structures for transportation.
The process of classifying protists into meaningful groups is ongoing, but genetic data in the past 20 years have clarified many relationships that were previously unclear or mistaken. The majority view at present is to order all eukaryotes into six supergroups. The goal of this classification scheme is to create clusters of species that all are derived from a common ancestor.
Fungi are eukaryotic organisms that appeared on land over 450 million years ago. They are heterotrophs and contain neither photosynthetic pigments such as chlorophylls nor organelles such as chloroplasts. Because they feed on decaying and dead matter, they are saprobes. Fungi are important decomposers and release essential elements into the environment. External enzymes digest nutrients that are absorbed by the body of the fungus called a thallus. A thick cell wall made of chitin surrounds the cell. Fungi can be unicellular as yeasts or develop a network of filaments called a mycelium, often described as mold. Most species multiply by asexual and sexual reproductive cycles, and display an alternation of generations.
The divisions of fungi are the Chytridiomycota, Zygomycota, Ascomycota, Basidiomycota, and Glomeromycota.
Fungi establish parasitic relationships with plants and animals. Fungal diseases can decimate crops and spoil food during storage. Compounds produced by fungi can be toxic to humans and other animals. Mycoses are infections caused by fungi. Superficial mycoses affect the skin, whereas systemic mycoses spread through the body. Fungal infections are difficult to cure.
Fungi have colonized all environments on Earth but are most often found in cool, dark, moist places with a supply of decaying material. Fungi are important decomposers because they are saprobes. Many successful mutualistic relationships involve a fungus and another organism. They establish complex mycorrhizal associations with the roots of plants. Lichens are a symbiotic relationship between a fungus and a photosynthetic organism, usually an alga or cyanobacterium.
Fungi are important to everyday human life. Fungi are important decomposers in most ecosystems. Mycorrhizal fungi are essential for the growth of most plants. Fungi, as food, play a role in human nutrition in the form of mushrooms and as agents of fermentation in the production of bread, cheeses, alcoholic beverages, and numerous other food preparations. Secondary metabolites of fungi are used in medicine as antibiotics and anticoagulants. Fungi are used in research as model organisms for the study of eukaryotic genetics and metabolism.