Chloroplasts and Photosynthesis
All animals and most microorganisms rely on the continual uptake of large amounts of organic compounds from their environment. These compounds are used to provide both the carbon skeletons for biosynthesis and the metabolic energy that drives cellular processes. It is believed that the first organisms on the primitive Earth had access to an abundance of the organic compounds produced by geochemical processes, but that most of these original compounds were used up billions of years ago. Since that time, the vast majority of the organic materials required by living cells have been produced by photosynthetic organisms, including many types of photosynthetic bacteria. The most advanced photosynthetic bacteria are the cyanobacteria, which have minimal nutrient requirements. They use electrons from water and the energy of sunlight when they convert atmospheric CO2 into organic compounds—a process called carbon fixation. In the course of splitting water [in the overall reaction nH2O + nCO2 (CH2O)n + nO2], they also liberate into the atmosphere the oxygen required for oxidative phosphorylation. As we see in this section, it is thought that the evolution of cyanobacteria from more primitive photosynthetic bacteria eventually made possible the development of abundant aerobic life forms. In plants and algae, which developed much later, photosynthesis occurs in a specialized intracellular organelle—the chloroplast. Chloroplasts perform photosynthesis during the daylight hours. The immediate products of photosynthesis, NADPH and ATP, are used by the photosynthetic cells to produce many organic molecules. In plants, the products include a low-molecular-weight sugar (usually sucrose) that is exported to meet the metabolic needs of the many nonphotosynthetic cells of the organism. Biochemical and genetic evidence strongly suggest that chloroplasts are descendants of oxygen-producing photosynthetic bacteria that were endocytosed and lived in symbiosis with primitive eucaryotic cells. Mitochondria are also generally believed to be descended from an endocytosed bacterium. The many differences between chloroplasts and mitochondria are thought to reflect their different bacterial ancestors, as well as their subsequent evolutionary divergence. Nevertheless, the fundamental mechanisms involved in light-driven ATP synthesis in chloroplasts are very similar to those that we have already discussed for respiration-driven ATP synthesis in mitochondria. The Chloroplast Is One Member of the Plastid Family of Organelles Chloroplasts are the most prominent members of the plastid family of organelles. Plastids are present in all living plant cells, each cell type having its own characteristic complement. All plastids share certain features. Most notably, all plastids in a particular plant species contain multiple copies of the same relatively small genome. In addition, each is enclosed by an envelope composed of two concentric membranes. As discussed in Chapter 12 (see Figure 12-3), all plastids develop from proplastids, small organelles in the immature cells of plant meristems (Figure 14-33A). Proplastids develop according to the requirements of each differentiated cell, and the type that is present is determined in large part by the nuclear genome. If a leaf is grown in darkness, its proplastids enlarge and develop into etioplasts, which have a semicrystalline array of internal membranes containing a yellow chlorophyll precursor instead of chlorophyll. When exposed to light, the etioplasts rapidly develop into chloroplasts by converting this precursor to chlorophyll and by synthesizing new membrane pigments, photosynthetic enzymes, and components of the electron-transport chain.
Figure 14-33 Plastid diversity
(A) A proplastid from a root tip cell of a bean plant. Note the double membrane; the inner membrane has also generated the relatively sparse internal membranes present. (B) Three amyloplasts (a form of...
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