Changes in the energy distribution between chlorophyll-protein complexes of thylakoid membranes from pea mutants with modified pigment content. I. Changes due to the modified pigment content

The low-temperature (77 K) emission and excitation chlorophyll fluorescence spectra in thylakoid membranes isolated from pea mutants were investigated. The mutants have modified pigment content, structural organization, different surface electric properties and functions [Dobrikova et al., Photosynth. Res. 65 (2000) 165]. The emission spectra of thylakoid membranes were decomposed into bands belonging to the main pigment protein complexes. By an integration of the areas under them, the changes in the energy distribution between the two photosystems as well as within each one of them were estimated. It was shown that the excitation energy flow to the light harvesting, core antenna and RC complexes of photosystem II increases with the total amount of pigments in the mutants, relative to the that to photosystem I complexes. A reduction of the fluorescence ratio between aggregated trimers of LHC II and its trimeric and monomeric forms with the increase of the pigment content (chlorophyll a, chlorophyll b, and lutein) was observed. This implies that the closer packing in the complexes with a higher extent of aggregation regulates the energy distribution to the PS II core antenna and reaction centers complexes. Based on the reduced energy flow to PS II, i.e., the relative increased energy flow to PS I, we hypothesize that aggregation of LHC II switches the energy flow toward LHC I. These results suggest an additive regulatory mechanism, which redistributes the excitation energy between the two photosystems and operates at non-excess light intensities but at reduced pigment content.     

ENERGY TRANSFER FROM PHYCOBILINS TO CHLOROPHYLL a IN HEAT-STRESSED CELLS OF Anacystis nidulans: CHARACTERIZATION OF THE LOW TEMPERATURE 683 nm FLUORESCENCE EMISSION BAND*

Abstract— Heat-induced changes of the characteristics of fluorescence spectra of Anacystis nidulans cells were studied after 39°C-grown cells were heated at 55°C. Heat-treatment of the cells induced no changes in the absorption properties or photosystem I-catalyzed cytochrome oxidation, but induced a dramatic change in the fluorescence characteristics of the cells. The low temperature fluorescence emission spectra of heated cells showed a large increase of fluorescence emission at683–685 nm (F683) and at 695 nm, while the bands at 660 nm (allophycocyanin) and at 718 nm (chlorophyll a of photosystem I) were not affected when the cells were excited with light absorbed by phycobilins. When the cells were heated for various periods, a progressive increase of the intensity of F683 occurred with the loss in oxygen evolution capacity. The increase of the F683 band was observed prior to the increase of the F695 band. Quenching of emission spectra by the addition of quinones indicates that the F683 band emanated mainly from a long wavelength form of allophycocyanin. Excitation spectra of heated cells measured at 77 K showed that light absorbed by phycobilins was effective in exciting F685, F695, and F715 emission. A possible energy distribution pathway in Anacystis nidulans is discussed.     

EXCITATION ENERGY TRANSFER IN THE LIGHT HARVESTING ANTENNA SYSTEM OF THE RED ALGA Porphyridium cruentum AND THE BLUE-GREEN ALGA Anacystis nidulans: ANALYSIS OF TIME-RESOLVED FLUORESCENCE SPECTRA

Abstract— Time-resolved fluorescence spectra of intact cells of red and blue-green algae Porphyridium cruentum and Anacystis nidulans were measured by means of a ps laser and a time-correlated photon counting system. Fluorescence spectra were observed successively from various pigments in the light harvesting system in the order of phycoerythrin (PE), phycocyanin (PC), allophycocyanin (APC) and chlorophyll a (Chl a ). The spectrum changes with time in the range of0–400 ps in P. cruentum and of0–1000 ps in A. nidulans . The time-resolved spectra were analyzed into components to obtain the rise and decay curve of each fluorescence component. Overall time behaviors of the sequential fluorescence emissions from various pigments can be interpreted with a decay kinetics ofexp(–2 kt ^½). The rate constants of the energy transfer show that the energy transfer takes place much faster in the red alga P. cruentum than in the blue-green alga A. nidulans , particularly in the step PCAPC. Results also indicated that a special form of APC, far-emitting APC, exists in the pigment system of A. nidulans , but it does not mediate a main energy transfer from phycobilisome to Chl a.     

SIMULTANEOUS MEASUREMENT OF ACTION SPECTRA FOR PHOTOREACTIONS I AND II OF PHOTOSYNTHESIS

Abstract— We have devised a method of obtaining simultaneous action spectra for photoreactions I and II by analysis of direct and indirect effects involved in enhancement. The method requires previous determination of the neutral wavelength which gives maximum quantum yield by virtue of equal fractions of open reaction centers ( p and q ) for each photoreaction. A sufficient intensity of the neutral wavelength is used as a constant background. Upon addition of a weak modulated measuring light of intensity I_m and wavelength λ _m two amperometric signals are obtained for rate of oxygen evolution. A modulated signal (AC¯) isolates the direct effect of I_m and gives action of photoreaction II as AC/ I_m . An increment in total rate (ΔDC) also includes an indirect effect of I_m in perturbing reaction center conditions ( p and q ). From analysis of interaction of the two photoreactions, action for photoreaction I can be estimated as (2 ΔDC-AC)/ I_m . The method is applicable to whole cells, properly scales the two action spectra to each other, and removes contribution of the State 1-State 2 phenomena. Action spectra were obtained for Chlorella.     

ENERGY TRANSFER IN SELF-ASSEMBLED CHLOROPHYLL a SYSTEMS *

Abstract Computer deconvolution of the optical absorption spectra has been used to follow the formation and transformation of various chlorophyll species formed by lowering the temperature in solutions of toluene or methylcyclohexane containing a hydrogen-bonding nucleophile such as ethanol. At cryogenic temperatures the self-assembled chlorophyll a-ethanol species has optical properties similar to those of special pair photoreaction center chlorophyll. Analysis of the absorption and fluorescence behavior of the self-assembled species suggests that chlorophyll-chlorophyll species are also formed in the self-assembly process. Fluorescence lifetime measurements indicate that the pathways for dissipation of excitation energy in these multicomponent systems are complex. Selective optical excitation at wavelengths corresponding to absorption of monomer, oligomer, etc. chlorophyll a species has been used to demonstrate the heterogeneous nature of these self-assembled systems.     

PHOTOSYNTHETIC ENERGY TRANSFER REVERSIBLY INHIBITED BY HYDROSTATIC PRESSURE*

Abstract— Hydrostatic pressure is found to affect reversibly the emission spectra of Porphyra perforata. At 1200atm phycoerythrin and phycocyanin fluorescence show a remarkable increase, whereas at the same time chlorophyll a (Chl a ) fluorescence decreases. Upon release of pressure the fluorescence intensities of the individual pigments return to their original levels. This effect indicates that hydrostatic pressure acts as a unique reversible inhibitor of energy transfer between phycobilins and Chi a in the chloroplast.
The effects of pressure and its release are relatively slow (minutes). It is suggested that pressure changes thylakoid membrane structure sufficiently to alter the critical distance between the phycobilisomes and Chl a , thus blocking the inductive resonance transfer of excitation energy.     

ACTION SPECTRA FOR ACCUMULATION OF INORGANIC CARBON IN THE CYANOBACTERIUM, Anabaena variabilis

Abstract— Action spectra for accumulation of inorganic carbon were obtained for Anabaena variabilis , strainM–2, in the presence and absence of photosynthetic CO₂ fixation. The action spectrum for inorganic carbon accumulation in the presence of CO₂ fixation showed a peak around 684 nm, corresponding to chlorophyll a absorption in PS 1, while that for CO₂ fixation showed a peak around 630 nm, corresponding to phycocyanin absorption in PS 2. The action spectra obtained in the presence of iodoacetamide or diuron, which inhibit CO₂ fixation, showed two peaks, one at about 684 nm and the other at 630 nm, with the 630 nm peak height 80 to 90% of the 684 nm peak. These results indicate that inorganic carbon transport in A. variabilis can be driven with near equal efficiency by energy derived from absorption in photosystem 1 alone and with energy transferred to PS 1 after absorption by PS 2.     

Peroxidative reactions attenuate oxygen effect on spectroscopic properties of isolated chloroplasts.

Steady-state absorption and fluorescence excitation spectra have been measured at 25 degrees C in order to elucidate the differences between isolated chloroplasts from pea (chilling-sensitive plant) and bean (chilling-tolerant plant) and their response to oxygen treatment. A weaker light harvesting in bean in comparison with pea chloroplasts is related to higher free fatty acids level and extended peroxidation activities of bean chloroplasts. Peroxidation of free fatty acids in bean chloroplasts results in an accumulation of oxygenated forms of fatty acids demonstrated by a large negative band around 400 nm in absorption difference spectra, while the excitation spectra are not significantly altered. Similar changes have been observed in the lipase-treated pea chloroplasts. In contrast, in both pea and bean chloroplasts exhibiting no peroxidation due to antimycin A treatment, oxygen induces a pronounced absorbance increase in the regions around 435, 470 and 674 nm indicating the chloroplast swelling. A decline of chlorophyll fluorescence excitation caused by oxygen, may result from a decrease in energy transfer from antennae complexes to chlorophyll species emitting at both 680 and 740 nm. The oxygen-induced changes are partially reversed upon restoration of anaerobic conditions. The presented data show for the first time, that in contrast to pea chloroplasts the peroxidation abolishes an oxygen-induced decrease in light harvesting in bean chloroplasts, i.e., a chilling-sensitive plant.     

EXCITATION ENERGY TRANSFER IN THE CRYPTOPHYTES. FLUORESCENCE EXCITATION SPECTRA AND PICOSECOND TIME-RESOLVED EMISSION SPECTRA OF INTACT ALGAE AT 77 K

Excitation spectra of chlorophyll- a (Chl- a ) fluorescence in intact cells of Cryptomonas ovata, Chroomonas pauciplastida and Chroomonas salina were determined at 77 K. For all species the excitation spectra for emission from Chl- a associated with photosystem II (PSII) showed increased contributions by a carotenoid (493 nm) and phycobiliproteins, and decreased contributions by carotenoid (417 nm, 505 nm) and Chl- a (445 nm) as compared to excitation spectra for emission from Chl- a associated with photosystem I (PSI). Excitation spectra of C. salina and C. ovata showed an increased contribution by Chl- c ² to PSII Chl- a fluorescence emission. In all three species the absorbance band positions of Chl- a , as determined from the excitation spectra, were similar to those previously described in green plants. green algae and phycobilisome-containing organisms. Time-resolved 77 K fluorescence emission spectra of C. ovata and C. salina showed successive emission from both phycoerythrin and Chl- c ², PSII Chl- a , and PSI Chl- a. C. pauciplastida showed successive emission from phycocyanin, PSII Chl- a , and PSI Chl- a. Spectral red-shifts with time were observed for the phycobiliprotein peaks in all three species. The fluorescence decay of phycoerythrin in C. ovata and C. salina was faster than that of phycocyanin in C. pauciplastida. The results are discussed in relation to the organization of the antenna pigments of PSII and PSI in the cryptophyte algae.     

SELECTIVE ALTERATION OF THE PIGMENT COMPOSITION OF THE BLUE-GREEN ALGA, Anacystis nidulans

Abstract— Conditions were established which allow selective alteration of the pigment composition of Anacystis cells. Nitrate-starvation at 28°C and 39°C resulted in the preponderance of chlorophyll a (Chi a ) and that of phycocyanin (PC). In the case of starvation at 39°C strong correlation was observed between the disappearance of Chi a absorption and the appearance of an absorption band at 750 nm (P750) as the starvation proceeded. During regeneration the starved cells lost their P750 content. We show that P750 is quantitatively accounted for in the form of Chi a aggregates.     

IS ZEAXANTHIN CAPABLE OF ENERGY TRANSFER TO CHLOROPHYLL a IN PARTIALLY GREENED LEAVES? A STUDY OF FLUORESCENCE EXCITATION SPECTRA DURING VIOLAXANTHIN DEEPOXIDATION

Absorbance spectra and excitation spectra of chlorophyll a fluoresence were recorded during the light-induced deepoxidation of violaxauthin to zeaxanthin in bean leaves (Phaseolus coccineus) greened under intermittent light. Light minus dark fluorescence excitation difference spectra showed distinct minima at 440, 465, and 500 nm corresponding to maxima in the absorbance difference spectra. Both difference spectra were prevented by the deepoxidase inhibitor dithiothreitol and were inverted when zeaxanthin was epoxidized. The fluorescence excitation difference spectra were successfully modeled by considering the absorbance differences between violaxanthin and zeaxanthin, assuming no energy transfer from the two pigments to chlorophyll a, and accounting for light-induced scattering changes. The pigment stoichiometry and the scattering changes of the simulation were in accordance with experimental data. The results indicate that, in the early stage of leaf development, light absorbed by the cycle pigments violaxanthin and zeaxanthin is not transferred to chlorophyll.     

Changes in the energy distribution in mutant thylakoid membranes of pea with modified pigment content. II. Changes due to magnesium ions concentration

Low-temperature (77K) steady-state chlorophyll fluorescence emission spectra, room temperature fluorescence and light scattering of thylakoid membranes isolated from pea mutants were studied as a function of Mg2+ concentration. The mutants have modified pigment content and altered structural organization of the pigment-protein complexes, distinct surface electric properties and functions. The analysis of the 77K emission spectra revealed that Mg2+-depletion of the medium caused not only an increased energy flow toward photosystem I in all investigated membranes but also changes in the quenching of the fluorescence, most probably by internal conversion. The results indicated that the macroorganization of the photosynthetic apparatus of mutants at supramolecular level (distribution and segregation of two photosystems in thylakoid membranes) and at supermolecular level (stacking of photosystem II supercomplexes) required different Mg ion concentrations. The data confirmed that the segregation of photosystems and the stacking of thylakoid membranes are two distinct phenomena and elucidated some features of their mechanisms. The segregation is initiated by changes in the lateral microorganization of light harvesting complexes II, their migration (repulsion from photosystem I) and subsequent separation of the two photosystems. Most likely 3D aggregation and formation of macrodomains, containing only photosystem II antenna complexes, play a certain precursory role for the increasing degree of the membrane stacking and the energy coupling between the light harvesting complexes II and the core complexes of photosystem II in the frame of photosystem II supercomplexes.     

CHLOROPHYLL a FLUORESCENCE OF GONYAULAX POLYEDRA GROWN ON A LIGHT-DARK CYCLE AND AFTER TRANSFER TO CONSTANT LIGHT

Abstract— The chlorophyll a fluorescence properties of Gonyaulax polyedra cells before and after transfer from a lightdark cycle (LD) to constant dim light (LL) were investigated. The latter display a faster fluorescence transient from the level ‘I’ (intermediary peak) to ‘D’ (dip) to ‘P’ (peak) than the former (3 s as compared to 10 s), and a different pattern of decline in fluorescence from ‘I’ to ‘D’ and from ‘P’ to the steady state level with no clearly separable second wave of slow fluorescence change, referred to as ‘s' (quasi steady state)→‘M’ (maximum) →‘T’ (terminal steady state). The above differences are constant features of cells in LD and LL, and are not dependent on the time of day. They are interpreted as evidence for a greater ratio of photosystem II/photosystem I activity in cells in LL. After an initial photoadaptive response following transfer from LD to LL, the cell absorbance at room temperature and fluorescence emission spectra at 77 K for cells in LL and LD are comparable. The major emission peak is at 685–688 nm (from an antenna Chl a 680, perhaps Chl a-c complex), but, unlike higher plants and other algae, the emission bands at 696–698 nm (from Chl a_II complex, Chl a 685, close to reaction center II) and 710–720 nm (from Chl a₁, complexes, Chl a 695, close to reaction center I) are very minor and could be observed only in the fluorescence emission difference spectra of LL minus LD cells and in the ratio spectra of DCMU-treated to non-treated cells. Comparison of emission spectra of cells in LL and LD suggested that, in LL, there is a slightly greater net excitation energy transfer from the light-harvesting peridinin-Chl a (Chl a 670) complex, fluorescing at 675 nm, to the other antenna chlorophyll a complex fluorescing at 685–688 nm, and from the Chl a., complex to the reaction center II. Comparison of excitation spectra of fluorescence of LL and LD cells, in the presence of DCMU, confirmed that cells in LL transfer energy more extensively from the peridinin-Chl a complex to other Chl a complexes than do cells in LD.     

ACTION SPECTRA AGAIN?*

Action spectroscopy has a long history and is of central importance to photobiological studies. Action spectra were among the first assays to point to chlorophyll as the molecule most responsible for plant growth and to DNA as the genetic material. It is useful to construct action spectra early in the investigation of new areas of photobiological research in an attempt to determine the wavelength limits of the radiation region causing the studied response. But due to the severe absorption of ultraviolet (UV) radiation by biological samples, UV action spectra were first limited to small cells (bacteria and fungi). Advances in techniques (e.g. single cell culture) and analysis allowed accurate action spectra to be reported even for mammalian cells. But precise analytical action spectra are often difficult to obtain when large, pigmented, or groups of cells are investigated. Here some action spectra are limited in interpretation and merely supply a wavelength vs effect curve. When polychromatic sources are employed, the interpretation of action spectra is even more complex and formidable. But such polychromatic action spectra can be more directly related to ambient responses. Since precise action spectra usually require the completion of a relatively large number of careful experiments using somewhat sophisticated equipment over a range of at least six wavelengths, they are often not pursued. But they remain central to the elucidation of the effect being studied. The worldwide community has agreed that stratospheric ozone is depleting, with the possibility of a consequent rise in the amount of UV-B (290-320 nm) reaching the earth's surface. It is therefore essential that new action spectra be completed for UV-B effects on a large variety of responses of human, animal, and aquatic plant systems. Combining these action spectra with the known amounts of UV-B reaching the biosphere can give rise to solar UV effectiveness spectra that, in turn, can give rise to estimates of effect. Preliminary estimates suggest that ozone layer depletion may seriously impact such important biological end-points as skin cancer, cataracts, the immune system, crop yields, and oceanic phytoplankton. So action spectra continue to play a central role in important photobiological research.     

Regulation of the excitation energy utilization in the photosynthetic apparatus of chlorina f2 barley mutant grown under different irradiances

Acclimation of the photosynthetic apparatus of chlorophyll b-less barley mutant chlorina f2 to low light (100 micromolm(-2)s(-1); LL) and extremely high light level (1000 micromolm(-2)s(-1); HL) was examined using techniques of pigment analysis and chlorophyll a fluorescence measurements at room temperature and at 77 K. The absence of chlorophyll b in LL-grown chlorina f2 resulted in the reduction of functional antenna size of both photosystem II (by 67%) and photosystem I (by 21%). Chlorophyll fluorescence characteristics of the LL-grown mutant indicated no impairment of the utilization of absorbed light energy in photosystem II photochemistry. Thermal dissipation of excitation energy estimated as non-photochemical quenching of minimal fluorescence (SV(0)) was significantly higher as compared to the wild-type barley grown under LL. Despite impaired assembly of pigment-protein complexes, chlorina f2 was able to efficiently acclimate to HL. In comparison with chlorina f2 grown under LL, HL-grown chlorina f2 was characterized by unaffected maximal photochemical efficiency of photosystem II (F(V)/F(M), doubled content of both beta-carotene and the xanthophyll cycle pigments and considerably reduced efficiency of excitation energy transfer from carotenoids to chlorophyll a. The enormous xanthophyll cycle pool size was however associated with reduced SV(0) capacity. We suggest that the substantial part of the xanthophyll cycle pigments is not bound to the remaining pigment-protein complexes and acts as filter for excitation energy, thereby contributing to the efficient photoprotection of chlorina f2 grown under HL.     

CHLOROPHYLL a FLUORESCENCE YIELD IN LIVE CELLS AND SOLUTIONS

Abstract— Fluorescence yields of chlorophyll a in ether and methanol solutions ( F_s ), on the one hand, and in Chlorella pyrenoidosa and Anacystis nidulans cells (F_c), on the other hand, were determined by excitation at 600 mμ (instead of at 436 mμ, as was done in earlier research in our, and other, laboratories). The ratio of the yields Fs/F_c , was found to be 5.9 for Chlorella (compared to an ether solution of chlorophyll a ), and of 4.5 (as compared to a methanol solution—not too different from the corresponding ratios of fluorescence lifetimes, τ_s /τ_c which were determined earlier as 3.1 and 4, respectively. The much higher values of the yield ratio, previously reported for Chlorella (about 13, compared to chlorophyll a in ether solution), may have been due to disregard of light absorption in carotenoids in live cells; and perhaps also to quenching of chlorophyll excitation by carotenoids. The latter can occur (as suggested in an earlier publication) when chlorophyll is excited to its second singlet excited state.
For Anacystis , the yield ratios were now found to be 5.1 and 3.8, when compared to ether and methanol solutions respectively; while the previously determined lifetime ratios were 3.7 and 4.9, respectively. It remains to be seen whether the remaining differences between the lifetime ratios and the yield ratios are real and significant.     

INCREASED ENERGY TRANSFER FROM PHYCOBILISOMES TO PHOTOSYSTEM II IN HIGH LIGHT ADAPTED Anabaena cylindrica

Excitation energy transfer from phycobilisomes to photosystem II in high-light adapted cells of Anabaena cylindrica was studied by fluorescence spectroscopy and compared to that of low-light adapted cells. Measurements were made on membrane fragments containing phycobilisomes, photosystem I and II, isolated in 0.75 M K-phosphate. Relative efficiency of 430 to 590 nm light in the excitation of F₆₈₀ chlorophyll fluorescence was compared in low and high light adapted cells, respectively. The values indicate that light energy absorbed by phycobilisomes is transferred to photosystem II antenna chlorophylls with higher efficiency in high-light adapted cells than in low-light adapted cells. Partial dissociation and uncoupling of energy transfer caused by low ion concentration were different in the membrane fragments isolated from the two kinds of cells and indicated a higher aggregation state of pigment-protein complexes of phycobilisomes in high-light adapted A. cylindrica cells.     

DCMU-INDUCED STIMULATION OF PHOTOSYSTEM I ELECTRON TRANSPORT. INVOLVEMENT OF MEMBRANE STRUCTURAL CHANGES

DCMU-induced stimulation of the rate of photosystem I (PS I) electron transport in DCIPH₂→ MV photoreaction occurs through the action of DCMU on the rate-limiting step which contains the site of electron donation of DCIPH₂ (Ramanujam et al. , 1981). The magnitude of stimulation of the rate by 50 μ M DCMU decreased with increasing concentration of chlorophyll (Chl), implying that DCMU is stoichiometrically related to Chl with respect to the stimulation of the PS I rate.
DCMU-induced stimulation was sensitive to the ionic condition of the thylakoids, the effect being reduced at low cation concentration. Cation-induced scattering changes in thylakoid suspension were partially reversed by DCMU, and the percent Chl in the 10 K fraction of the thylakoid decreased upon addition of DCMU, indicating that grana structure is disrupted by DCMU. Hydroquinone-mediated reduction of cytochrome f in thylakoids in the dark was accelerated in the presence of DCMU. The DCMU effect was not observed in isolated PS I particles.
It is concluded that DCMU binds to the thylakoid membranes and brings about structural changes leading to unstacking of the thylakoids accompanied by an altered interaction of the electron transfer chain components with the added electron donor. This binding of DCMU must have an affinity lower than the well-known binding of DCMU to photosystem II (PS II), because the concentration required is markedly higher.     

Photoionization cross-section weighted DFT simulations as promising tool for the investigation of the electronic structure of open shell metal-phthalocyanines

The valence band structure of different metal-phthalocyanines was investigated by comparing ultraviolet photoelectron spectra at different excitation energies with simulated spectra that take the different photoionization cross-sections at these energies into account. The Kohn-Sham eigenvalue spectra, derived from density functional theory calculations, using hybrid exchange-correlation functionals, were weighted with the photoionization coefficients in accordance with the used excitation energy. By applying these techniques, the differences in the photoelectron spectra using He I and He II radiation can be reproduced and investigated. It will be shown that the 3d-orbitals of the used metal central atom of these molecules have a major influence. The changes at different excitation energies were studied for Fe, Co, and Cu central atoms to describe the chemical tailoring effects.     

PHOTOSYNTHETIC ACTION SPECTRA AND LIGHT-HARVESTING IN GRIFFITHSIA MONILIS (RHODOPHYTA)

Abstract— In cells of the red alga Griffithsia monilis the action spectrum of photosynthetic oxygen production at low light intensity shows that the phycobilins (including allophycocyanin) are the major light-harvesting pigments. As the light intensity is increased carotenoids and chlorophyll a contribute proportionately more to the spectrum, since the phycobilin activity becomes light-saturated. When action spectra are performed against a background light of various monochromatic wavelengths it can be shown that chlorophyll a increases in its light-harvesting activity. Nevertheless light absorbed at a single wavelength (487 nm) by phycoerythrin (and possibly a carotenoid) still shows the highest photosynthetic activity. Fluorescence measurements at 77K indicate that a chlorophyll a fluorescence is small and that the amount of chlorophyll a _ll (f 693) is very low. A model is proposed in which the phycobilins, in phycobilisomes, pass on absorbed light energy to either photosystem, whereas light absorbed by chlorophyll is passed on mainly to photosystem I.