A polarographic oxygen determination, with tissue in direct contact with a stationary platinum electrode, has been used to measure the photosynthetic response of marine algae. These were exposed to monochromatic light, of equal energy, at some 35 points through the visible spectrum (derived from a monochromator). Ulva and Monostroma (green algae) show action spectra which correspond very closely to their absorption spectra. Coilodesme (a brown alga) shows almost as good correspondence, including the spectral region absorbed by the carotenoid, fucoxanthin. In green and brown algae, light absorbed by both chlorophyll and carotenoids seems photosynthetically effective, although some inactive absorption by carotenoids is indicated. Action spectra for a wide variety of red algae, however, show marked deviations from their corresponding absorption spectra. The photosynthetic rates are high in the spectral regions absorbed by the water-soluble "phycobilin" pigments (phycoerythrin and phycocyanin), while the light absorbed by chlorophyll and carotenoids is poorly utilized for oxygen production. In red algae containing chiefly phycoerythrin, the action spectrum closely resembles that of the water-extracted pigment, with peaks corresponding to its absorption maxima (495, 540, and 565 mµ). Such algae include Delesseria, Schizymenia, and Porphyrella. In the genus Porphyra, there is a series P. nereocystis, P. naiadum, and P. perforata, with increasingly more phycocyanin and less phycoerythrin: the action spectra reflect this, with increasing activity in the orange-red region (600 to 640 mµ) where phycocyanin absorbs. In all these red algae, photosynthesis is almost minimal at 435 mµ and 675 mµ, where chlorophyll shows maximum absorption. Although the chlorophylls (and carotenoids) are present in quantities comparable to the green algae, their function is apparently not that of a primary light absorber; this role is taken over by the phycobilins. In this respect the red algae (Rhodophyta) appear unique among photosynthetic plants.
The photosynthetic light-harvesting complex, peridinin-chlorophyll a-protein, was isolated from several marine dinoflagellates including Glenodinium sp. by Sephadex and ion-exchange chromatography. The carotenoid (peridinin)-chlorophyll a ratio in the complex is estimated to be 4:1. The fluorescence excitation spectrum of the complex indicates that energy absorbed by the carotenoid is transferred to the chlorophyll a molecule with 100% efficiency. Fluorescence lifetime measurements indicate that the energy transfer is much faster than fluorescence emission from chlorophyll a. The four peridinin molecules within the complex appear to form two allowed exciton bands which split the main absorption band of the carotenoid into two circular dichronic bands (with negative ellipticity band at 538 nm and positive band at 463 nm in the case of peridinin-chlorophyl a-protein complex from Glenodinium sp.). The fluorescence polarization of chlorophyll a in the complex at 200 K is about 0.1 in both circular dichroic excitation bands of the carotenoid chromophore. From these circular dichroic and fluorescence polarization data, a possible molecular arrangement of the four peridinin and chlorophyll molecules has been deduced for the complex. The structure of the complex deduced is also consistent with the magnitude of the exciton spliting (ca. greater than 3000 cm-1) at the intermolecular distance in the dimer pair of peridinin (ca. 12 A). This structural feature accounts for the efficient light-harvesting process of dinoflagellates as the exciton interaction lengthens the lifetime of peridinin (radiative) and the complex topology increases the energy transfer probability. The complex is, therefore, a useful molecular model for elucidating the mechanism and efficiency of solar energy conversion in vivo as well as in vitro.
SUMMARY The chlorophylls and carotenoids of 22 species of dinoflagellates were analysed by thin layer chromatography, using 2‐dimensional sucrose plates, and 1‐dimensional polyethylene plates for chlorophylls c1 and c2. Peridinin was the major carotenoid in 19 of the species, while fucoxanthin was the major carotenoid in 3. In the peridinin‐containing species, 5 carotenoid fractions, constituting more than 95% of the total carotenoids, were always present. These were peridinin (± neo‐peridinin), averaging 64% of the total carotenoid, diadinoxanthin, dinoxanthin, β‐carotene and a polar, unidentified pink xanthophyll. Six other carotenoid fractions occurred in minor or trace quantities among the species, but were not identified. Two of these had, a wide distribution; the other 4 were restricted to one or 2 species. The chlorophyll content of the dinoflagellate cultures ranged from 1–141 μg chlorophyll a + c/106 cells, a pattern which was broadly correlated with cell size. In the peridinin‐containing species the ratio of chlorophyll a to c on a molar basis was approximately 2 (range 1.60–4.39); in the fucoxanthin‐containing species this ratio was approximately 4 (range 2.65–5.73). Both chlorophylls c1 and c2 occurred in the fucoxanthin‐containing dinoflagellates, and only chlorophyll c2 (one exception) occurred in the peridinin‐containing dinoflagellates. These patterns of chlorophyll c and major carotenoid correspond to patterns previously observed in the Pyrrhophyta and the Chrysophyta, suggesting different phylogenetic origins for the “dinoflagellate” chloroplasts.
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