The First Cell Was Probably a Chemoheterotroph

The earliest cells that arose in the rich mixture of organic compounds, the primordial soup of prebiotic times, were almost certainly chemoheterotrophs (Fig. 1-5). The organic compounds they required were originally synthesized from components of the early atmosphere— CO, CO2, N2, CH4, and such—by the nonbiological actions of volcanic heat and lightning. Early heterotrophs gradually acquired the ability to derive energy from compounds in their environment and to use that energy to synthesize more of their own precursor molecules, thereby becoming less dependent on outside sources. A very significant evolutionary event was the development of pigments capable of capturing the energy of light from the sun, which could be used to reduce, or "fix," CO2 to form more complex, organic compounds. The original electron donor for these photosynthetic processes was probably H2S, yielding elemental sulfur or sulfate (SO|_) as the by-product, but later cells developed the enzymatic capacity to use H2O as the electron donor in photosynthetic reactions, eliminating O2 as waste. Cyanobacteria are the modern descendants of these early photosynthetic oxygen-producers.

Because the atmosphere of Earth in the earliest stages of biological evolution was nearly devoid of oxygen, the earliest cells were anaerobic. Under these conditions, chemoheterotrophs could oxidize organic compounds to CO2 by passing electrons not to O2 but to acceptors such as SO|~, yielding H2S as the product. With the rise of O2-producing photosynthetic bacteria, the atmosphere became progressively richer in oxy-gen—a powerful oxidant and deadly poison to anaerobes. Responding to the evolutionary pressure of the "oxygen holocaust," some lineages of microorganisms gave rise to aerobes that obtained energy by passing electrons from fuel molecules to oxygen. Because the transfer of electrons from organic molecules to O2 releases a great deal of energy, aerobic organisms had an energetic advantage over their anaerobic counterparts when both competed in an environment containing oxygen. This advantage translated into the predominance of aerobic organisms in O2-rich environments.

Modern bacteria inhabit almost every ecological niche in the biosphere, and there are bacteria capable of using virtually every type of organic compound as a source of carbon and energy. Photosynthetic bacteria in both fresh and marine waters trap solar energy and use it to generate carbohydrates and all other cell constituents, which are in turn used as food by other forms of life. The process of evolution continues—and in rapidly reproducing bacterial cells, on a time scale that allows us to witness it in the laboratory.

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