Chlorophyll Metabolism

The two pathways for chlorophyll biosynthesis and catabolism of plants are located in distinct cellular compartments. While the entire biosynthesis of chlorophyll from the first committed metabolic precursor, 5-aminolaevulinic acid, occurs in plastids, chlorophyll breakdown starts in plastids and the final non-fluorescent breakdown products are deposited in vacuoles. The current understanding about both pathways in a diverse range of photosynthetic organisms including photosynthetic bacteria, algae and higher plants is outlined in the following report. It is quite obvious that the photosynthetic organisms evolved a complex control of these two pathways to enable specific adaptation to various environmental conditions.

Keywords: pigment biosynthesis; photosynthesis; light acclimation; photosensitization; senescence; primary metabolism

Figure 1. Structure of chlorophyll a and chlorophyll b (left side) and bacteriochlorophyll a and bacteriochlorophyll b (right side). The major esterifying alcohol (5 R) is phytol in chlorophyll a and b, geranylgeraniol in bacteriochlorophyll a, and phytadienol in bacteriochlorophyll b.
Figure 2. Flow chart of the metabolic pathway of tetrapyrrole biosynthesis towards the end products chlorophyll and protohaem. Potential feedback control mechanisms in the metabolic pathway of tetrapyrroles are indicated. The synthesis of 5-aminolaevulinate from glutamate is the rate-limiting step of tetrapyrrole biosynthesis. Feedback regulation on the enzymes of the 5-aminolaevulinic acid pathway can start out from various sites: at the level of haem formation, at the level of protochlorophyllide reduction and at the beginning of the Mg porphyrin branch.
Figure 3. Compartmentation of the biosynthetic pathway of tetrapyrroles and the catabolic pathway of chlorophyll.
Figure 4. The pathway of chlorophyll catabolism. Chlorophyllide a is the central molecule that connects the biosynthetic and the catabolic pathway as well as the ‘chlorophyll cycle’. ALA, 5-aminolaevulinic acid; CAO, chlorophyll(ide) a oxygenase; CAR, 7-hydroxychlorophyll(ide) a reductase; CBR, chlorophyll(ide) b reductase; Chl, chlorophyll; Chlase, chlorophyllase; Chlide, chlorophyllide; CS, chlorophyll synthase; PAO, phaeophorbide a oxygenase; pFCC, primary fluorescent chlorophyll catabolite; Phaeide a, phaeophorbide a; Pchlide, protochlorophyllide; RCC, red chlorophyll catabolite; RCCR, RCC reductase.
Figure 5. Structure and absorption spectra of three selected chlorophyll catabolites: (a) phaeophorbide a, (b) primary fluorescent chlorophyll catabolite and (c) non-fluorescent chlorophyll catabolite.
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 References
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    book Blankenship RE (2002) Molecular Mechanisms of Photosynthesis. Oxford: Blackwell Science.
    book Smith AG and Witty M (eds) (2001) Heme, Chlorophyll and Bilins – Methods and Protocols. New Jersey, USA: Humana Press.
    Tanaka A and Tanaka R (2006) Chlorophyll metabolism. Current Opinions in Plant Biology 9: 1–8.
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Eckhardt, Ulrich, and Grimm, Bernhard(Sep 2007) Chlorophyll Metabolism. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0020084]