Carotenoids are usually considered to perform two major functions in photosynthesis. They serve as accessory light harvesting pigments, extending the range of wavelengths over which light can drive photosynthesis, and they act to protect the chlorophyllous pigments from the harmful photodestructive reaction which occurs in the presence of oxygen. Drawing upon recent work with photosynthetic bacteria, evidence is presented as to how the carotenoids are organized within both portions of the photosynthetic unit (the light harvesting antenna and the reaction centre) and how they discharge both their functions. The accessory pigment role is a singlet-singlet energy transfer from the carotenoid to the bacteriochlorophyll, while the protective role is a triplet-triplet energy transfer from the bacteriochlorophyll to the carotenoid.

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Carotenoids in Photosynthesis

Energy transfer and trapping in photosynthesis

The aerobic anoxygenic phototrophic bacteria are a relatively recently discovered bacterial group. Although taxonomically and phylogenetically heterogeneous, these bacteria share the following distinguishing features: the presence of bacteriochlorophyll a incorporated into reaction center and light-harvesting complexes, low levels of the photosynthetic unit in cells, an abundance of carotenoids, a strong inhibition by light of bacteriochlorophyll synthesis, and the inability to grow photosynthetically under anaerobic conditions. Aerobic anoxygenic phototrophic bacteria are classified in two marine (Erythrobacter and Roseobacter) and six freshwater (Acidiphilium, Erythromicrobium, Erythromonas, Porphyrobacter, Roseococcus, and Sandaracinobacter) genera, which phylogenetically belong to the α-1, α-3, and α-4 subclasses of the class Proteobacteria. Despite this phylogenetic information, the evolution and ancestry of their photosynthetic properties are unclear. We discuss several current proposals for the evolutionary origin of aerobic phototrophic bacteria. The closest phylogenetic relatives of aerobic phototrophic bacteria include facultatively anaerobic purple nonsulfur phototrophic bacteria. Since these two bacterial groups share many properties, yet have significant differences, we compare and contrast their physiology, with an emphasis on morphology and photosynthetic and other metabolic processes.

Aerobic Anoxygenic Phototrophic Bacteria

In photosynthesis, the harvesting of solar energy and its subsequent conversion into a stable charge separation are dependent upon an interconnected macromolecular network of membraneassociated chlorophyll–protein complexes. Although the detailed structure of each complex has been determined1–4, the size and organization of this network are unknown. Here we show the use of atomic force microscopy to directly reveal a native bacterial photosynthetic membrane. This first view of any multi-component membrane shows the relative positions and associations of the photosynthetic complexes and reveals crucial new features of the organization of the network: we found that the membrane is divided into specialized domains each with a different network organization and in which one type of complex predominates. Two types of organization were found
for the peripheral light-harvesting LH2 complex. In the first, groups of 10–20 molecules of LH2 form light-capture domains that interconnect linear arrays of dimers of core reaction centre (RC)–light-harvesting 1 (RC–LH1–PufX) complexes; in the second they were found outside these arrays in larger clusters. The LH1 complex is ideally positioned to function as an energy collection hub, temporarily storing it before transfer to the RC where photochemistry occurs: the elegant economy of the photosynthetic membrane is demonstrated by the close packing of these linear arrays, which are often only separated by narrow ‘energy conduits’ of LH2 just two or three complexes wide.

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The native architecture of a photosynthetic membrane.

3. Excitation transfer in the PSU 18 3.1 Electronic excitations of BChls 22 3.1.1 Individual BChls 22 3.1.2 Rings of BChls 22 3.1.2.1 Exciton states 22 3.1.3 Effective Hamiltonian 24 3.1.4 Optical properties 25 3.1.5 The effect of disorder 26 3.2 Theory of excitation transfer 29 3.2.1 General theory 29 3.2.2 Mechanisms of excitation transfer 32 3.2.3 Approximation for long-range transfer 34 3.2.4 Transfer to exciton states 35 3.3 Rates for transfer processes in the PSU 37 3.3.1 Car U BChl transfer 37 3.3.1.1 Mechanism of Car U BChl transfer 39 3.3.1.2 Pathways of Car U BChl transfer 40 3.3.2 Efficiency of Car U BChl transfer 40 3.3.3 B800–B850 transfer 44 3.3.4 LH-II U LH-II transfer 44 3.3.5 LH-II U LH-I transfer 45 3.3.6 LH-I U RC transfer 45 3.3.7 Excitation migration in the PSU 46 3.3.8 Genetic basis of PSU assembly 49

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Photosynthetic apparatus of purple bacteria

— The ratio of total bacteriochlorophyll (BChl) to reaction center BChl has been determined in the wild-type and carotenoid-less mutants of Rhodopseudomonas spheroides and Rhodospirillum rubrum with various specific BChl contents. In wild-type R. spheroides the ratio increases as the specific BChl content increases; in the other strains the ratio is almost invariant.



The ability of the wild-type of R. spheroides to alter the size of the photosynthetic unit is apparently correlated with its multi-component BChl spectrum. The amount of reaction center BChl relative to the amount of the longest wavelength component of the spectrum is constant. The variation in the size of the photosynthetic unit is due to variation in the amount of this component relative to total BChl.



The size of the photosynthetic unit does not change as the specific BChl content decreases during growth under aerobic conditions of a culture grown previously under photosynthetic conditions.

Jeanette Aagaard

Papers

Control of synthesis of reaction center bacteriochlorophyll in photosynthetic bacteria