Oxygenic photosynthetic organisms perform photosynthesis efficiently by distributing captured light energy to photosystems (PSs) at an appropriate balance. Maintaining photosynthetic efficiency under ...changing light conditions requires modification of light-harvesting and energy-transfer processes. In the current study, we examined how green algae regulate their light-harvesting functions in response to different light qualities. We measured low-temperature time-resolved fluorescence spectra of unicellular green algae
Chlamydomonas reinhardtii
and
Chlorella variabilis
cells grown under different light qualities. By observing the delayed fluorescence spectra, we demonstrated that both types of green algae primarily modified the associations between light-harvesting chlorophyll protein complexes (LHCs) and PSs (PSII and PSI). Under blue light,
Chlamydomonas
transferred more energy from LHC to chlorophyll (Chl) located far from the PSII reaction center, while energy was transferred from LHC to PSI via different energy-transfer pathways in
Chlorella
. Under green light, both green algae exhibited enhanced energy transfer from LHCs to both PSs. Red light induced fluorescence quenching within PSs in
Chlamydomonas
and LHCs in
Chlorella
. In
Chlorella
, energy transfer from PSII to PSI appears to play an important role in balancing excitation between PSII and PSI.
CO
2
concentration and temperature for growth of photosynthetic organisms are two important factors to ensure better photosynthetic performance. In this study, we investigated the effects of CO
2
...concentration and temperature on the photosynthetic performance in a marine centric diatom
Chaetoceros gracilis
. Cells were grown under four different conditions, namely, at 25 °C with air bubbling, at 25 °C with a supplementation of 3% CO
2
, at 30 °C with air bubbling, and at 30 °C with the CO
2
supplementation. It was found that the growth rate of cells at 30 °C with the CO
2
supplementation is faster than those at other three conditions. The pigment compositions of cells grown under the different conditions are altered, and fluorescence spectra measured at 77 K also showed different peak positions. A novel fucoxanthin chlorophyll
a
/
c
-binding protein complex is observed in the cells grown at 30 °C with the CO
2
supplementation but not in the other three types of cells. Since oxygen-evolving activities of the four types of cells are almost unchanged, it is suggested that the CO
2
supplementation and growth temperature are involved in the regulation of photosynthetic light-harvesting apparatus in
C. gracilis
at different degrees. Based on these observations, we discuss the favorable growth conditions for
C. gracilis
.
Photosystem I (PSI) is one of the two photosystems functioning in light-energy harvesting, transfer, and electron transfer in photosynthesis. However, the oligomerization state of PSI is variable ...among photosynthetic organisms. We present a 3.8-Å resolution cryo-electron microscopic structure of tetrameric PSI isolated from the glaucophyte alga Cyanophora paradoxa, which reveals differences with PSI from other organisms in subunit composition and organization. The PSI tetramer is organized in a dimer of dimers with a C2 symmetry. Unlike cyanobacterial PSI tetramers, two of the four monomers are rotated around 90°, resulting in a completely different pattern of monomer-monomer interactions. Excitation-energy transfer among chlorophylls differs significantly between Cyanophora and cyanobacterial PSI tetramers. These structural and spectroscopic features reveal characteristic interactions and excitation-energy transfer in the Cyanophora PSI tetramer, suggesting that the Cyanophora PSI could represent a turning point in the evolution of PSI from prokaryotes to eukaryotes.
To maximize the efficiency of photosynthesis, photosynthetic organisms must properly balance their light-harvesting ability and CO2 utilization. However, the molecular mechanisms of light harvesting ...under various CO2 conditions remain unclear. To reveal these mechanisms, we performed new analysis on cells of the green alga Chlamydomonas reinhardtii under different CO2 conditions. The analysis combines three kinds of fluorometries: pulse-amplitude modulated fluorescence, steady-state fluorescence with absolute intensity, and time-resolved fluorescence. Under low CO2 conditions, the main regulatory mechanism was migration of a light-harvesting chlorophyll–protein complex (LHC) II from photosystem (PS) II to PSI. However, under CO2-deficient conditions with carbon supplementation, some of the LHCII separated from the PSI and aggregated with quenching. These different light-harvesting abilities of LHCII may play an important role in the regulation of light harvesting in C. reinhardtii under various CO2 conditions.
Diatoms are a major group of microalgae in marine and freshwater environments. To utilize the light energy in blue to green region, diatoms possess unique antenna pigment–protein complexes, ...fucoxanthin chlorophyll
a
/
c
-binding proteins (FCPs). Depending on light qualities and quantities, diatoms form FCPs with different energies: normal-type and red-shifted FCPs. In the present study, we examined changes in light-harvesting and energy-transfer processes of a diatom
Chaetoceros gracilis
cells grown using white- and single-colored light-emitting diodes (LEDs), by means of time-resolved fluorescence spectroscopy. The blue LED, which is harvested by FCPs, modified energy transfer involving CP47, and suppressed energy transfer to PSI. Under the red-LED conditions, which is absorbed by both FCPs and PSs, energy transfer to PSI was enhanced, and the red-shifted FCP appeared. The red-shifted FCP was also recognized under the green- and yellow-LEDs, suggesting that lack of the shorter-wavelength light induces the red-shifted FCP. Functions of the red-shifted FCPs are discussed.
Excitation-energy transfer in photosystem I (PSI) is changed by a cation formation of a special pair chlorophyll P700 in the PSI core; however, it remains unclear how light-harvesting pigment–protein ...complexes are involved in the P700-related energy-transfer mechanisms. Here, we report effects of the redox changes of P700 on excitation-energy dynamics in diatom PSI-fucoxanthin chlorophyll a/c-binding protein (PSI-FCPI) and PSI core complexes by means of time-resolved fluorescence (TRF) spectroscopy. For the TRF measurements, the PSI-FCPI and PSI were adapted under P700 neutral and cation conditions using chemical reagents. Upon the P700+ formation, fluorescence decay-associated (FDA) spectra constructed from the TRF spectra exhibit a larger fluorescence decay amplitude relative to a fluorescence rise magnitude within 100 ps in each of the PSI-FCPI and PSI. The decay components are shifted to lower wavelengths in each of the P700-cation PSI-FCPI and PSI than in the P700-neutral PSIs. The rapid fluorescence decays upon the P700+ formation are clearly verified by mean lifetimes reconstructed from the FDA spectra. Because the P700-cation PSI does not cause charge-separation reactions, the relatively strong decay components and rapid fluorescence decays observed are likely attributed to excitation-energy quenching. These observations suggest that chlorophylls in PSI and around/within FCPI are involved in the energy-quenching events by the redox changes of P700.
pH influences excitation-energy-relaxation processes in photosynthetic light-harvesting complexes. Here, we report the excitation-energy dynamics by pH changes in fucoxanthin chlorophyll
/
-binding ...proteins (FCPs) isolated from a diatom
, probed by time-resolved fluorescence spectroscopy at 77 K. The fluorescence curve measured at pH 5.0 showed a shorter lifetime component than that measured at pH 6.5 and 8.0. The rapid decay component at pH 5.0 is supported by fluorescence decay-associated (FDA) spectra, where strong fluorescence decays relative to fluorescence rises appear in the pH-5.0 FDA spectrum with 70 ps. These results indicate that the diatom FCPs switch their function from light-harvesting to energy-quenching via arrangements of the energy-transfer pathways under acidic pHs. Based on the crystal structure of the diatom FCPs, we propose a model for the energy-quenching machinery through structural changes of the pigment environments, thus providing insights into the pH-dependent light-harvesting strategy in the diatom FCPs.
Cyanobacteria and red algae control the energy distributions of two photosystems (PSI and PSII) by changing the energy transfer among phycobilisome (PBS), PSI, and PSII. However, whether PSII → PSI ...energy transfer (spillover) occurs in the intact megacomplexes composed of PBS, PSI, and PSII (PBS–PSII–PSI megacomplexes) in vivo remains controversial. In this study, we measured the delayed fluorescence spectra of PBS-selective excitation in cyanobacterial and red algal cells. In the absence of spillover, 7% of the PBS (at most) would combine with PSII, inconsistent with the PBSs’ function as the antenna pigment–protein complexes of PSII. Therefore, we conclude that spillover occurs in vivo in PBS–PSII–PSI megacomplexes of both cyanobacteria and red algae.
Oxygenic photosynthesis converts light energy into chemical energy via electron transport and assimilates CO
2
in the Calvin–Benson cycle with the chemical energy. Thus, high light and low CO
2
...conditions induce the accumulation of electrons in the photosynthetic electron transport system, resulting in the formation of reactive oxygen species. To prevent the accumulation of electrons, oxygenic photosynthetic organisms have developed photoprotection mechanisms, including non-photochemical quenching (NPQ) and alternative electron flow (AEF). There are diverse molecular mechanisms underlying NPQ and AEF, and the corresponding molecular actors have been identified and characterized using a model green alga
Chlamydomonas reinhardtii
. In contrast, detailed information about the photoprotection mechanisms is lacking for other green algal species. In the current study, we examined the photoprotection mechanisms responsive to CO
2
in the green alga
Chlorella variabilis
by combining the analyses of pulse-amplitude-modulated fluorescence, O
2
evolution, and the steady-state and time-resolved fluorescence spectra. Under the CO
2
-limited condition, ΔpH-dependent NPQ occurred in photosystems I and II. Moreover, O
2
-dependent AEF was also induced. Under the CO
2
-limited condition with carbon supplementation, NPQ was relaxed and light-harvesting chlorophyll-protein complex II was isolated from both photosystems. In
C. variabilis
, the O
2
-dependent AEF and the mechanisms that instantly convert the light-harvesting functions of both photosystems may be important for maintaining efficient photosynthetic activities under various CO
2
conditions.
Photosynthetic organisms handle solar energy precisely to achieve efficient photochemical reactions. Because there are a wide variety of light-harvesting antennas in oxyphototrophs, the excitation ...energy transfer mechanisms are thought to differ significantly. In this study, we compared excitation energy dynamics between photosystem I (PSI) cores and a complex between PSI and fucoxanthin chlorophyll (Chl) a/c-binding protein I (PSI−FCPI) isolated from a diatom, Chaetoceros gracilis, by means of picosecond time-resolved fluorescence analyses. Time-resolved spectra measured at 77 K clearly show that low-energy Chls in the FCPI transfer not only most of the excitation energy to the reaction center Chls in the PSI cores but also the remaining energy to carotenoids for quenching. Under room-temperature conditions, the energy in the low-energy Chls is rapidly equilibrated on Chls in the PSI cores by uphill energy transfer within a few tens of picoseconds. These findings provide solid evidence that the low-energy Chls in the FCPI contribute to the photochemical reactions in PSI.