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Figure 1.7 A mature and functional chloroplast in an immature leaf of bean (Phaseolus vulgaris) with an extensive network of photosynthetic membranes (thylakoids), parts of which are appressed into moderate granal stacks, and suspended in a gel-like matrix (stroma).The chloroplast containing a pair of starch grains (S) is encapsulated in a double membrane (envelope) and suspended within a granular cytoplasmic matrix adjacent to a mitochondrion (M) and in close proximity to the cell wall (CW). Scale bar = 1 µm. (Micrograph courtesy S. Craig and C. Miller)
Chloroplasts are easily recognised under a light microscope in leaf sections as distinctive green organelles suspended in the cytoplasm and usually appressed against cell walls. Chloroplasts are abundant in mesophyll tissue (commonly 200–300 in each palisade cell) and functional organelles can be isolated from homogenates of leaf tissue.
Chloroplasts are surrounded by a double membrane, or envelope, just visible in transmission electron micrographs (Figure 1.7). This envelope encapsulates a soluble (gel-like) stroma which contains all the enzymes necessary for carbon fixation, many enzymes of nitrogen and sulphur metabolism and the chloroplast’s own genetic machinery.
The inner membrane of a chloroplast envelope is an effective barrier between stroma and cytoplasm, and houses transporters for phosphate and metabolites (Section 2.1.8) as well as some of the enzymes for lipid synthesis. By comparison, the outer membrane of the chloroplast envelope is less complex and more permeable to both ions and metabolites.
Suspended within the stroma, and entirely separate from envelope membranes, is an elaborately folded system of photosynthetic membranes or ‘thylakoids’ (literally ‘little sacs’). Embedded within these membranes are the complexes that enable light harvesting and electron flow from H2O molecules to NADP+, thereby converting light energy into chemically usable forms. There are four basic complexes comprising two types of photosystem (with interlinked protein and pigment molecules), cytochrome b/f complexes (pivotal for photosynthetic electron transport) and ATP synthase complexes (responsible for proton egress from thylakoid lumen to stroma, and consequent ATP generation). These complexes are densely packed within the thylakoids. This remarkable transduction of energy, with such profound implications for life as we know it, starts with selective absorption of incoming light by chlorophylls and accessory pigments (certain carotenoids) that operate within both photosystems.