Makiko Kosugi

Responses to desiccation stress in lichens are different from those in their photobionts.

In order to clarify the role of symbiotic association in desiccation tolerance of photosynthetic partners in lichens, responses to air-drying and hypertonic treatments in a green-algal lichen (a chlorolichen, Ramalina yasudae Räsänen) and its green algal photobiont (freshly released and cultured Trebouxia sp.) were studied. Responses to dehydration in the isolated Trebouxia sp. were different from those in the lichen, R. yasudae, i.e. (i) the PSII reaction was totally inhibited in R. yasudae when photosynthesis was completely inhibited by desiccation, but it remained partially active in isolated Trebouxia sp; (ii) dehydration-induced quenching of PSII fluorescence was less in the isolated Trebouxia sp. compared with that in R. yasudae, suggesting that a substance(s) or a mechanism(s) to dissipate absorbed light energy to heat was lost by the isolation of the photobiont; and (iii) the air-dried isolated Trebouxia sp. showed a higher sensitivity to photoinhibition than R. yasudae. These results support the idea that association of the photobionts with the mycobionts increases tolerance to photoinhibition under drying conditions.

Arabitol Provided by Lichenous Fungi Enhances Ability to Dissipate Excess Light Energy in a Symbiotic Green Alga under Desiccation

Lichens are drought-resistant symbiotic organisms of mycobiont fungi and photobiont green algae or cyanobacteria, and have an efficient mechanism to dissipate excess captured light energy into heat in a picosecond time range to avoid photoinhibition. This mechanism can be assessed as drought-induced non-photochemical quenching (d-NPQ) using time-resolved fluorescence spectroscopy. A green alga Trebouxia sp., which lives within a lichen Ramalina yasudae, is one of the most common green algal photobionts. This alga showed very efficient d-NPQ under desiccation within the lichen thallus, whereas it lost d-NPQ ability when isolated from R. yasudae, indicating the importance of the interaction with the mycobiont for d-NPQ ability. We analyzed the water extracts from lichen thalli that enhanced d-NPQ in Trebouxia. Of several sugar compounds identified in the water extracts by nuclear magnetic resonance (NMR), mass spectrometry (MS) and gas chromatography (GC) analyses, only d-arabitol recovered d-NPQ in isolated Trebouxia to a level similar to that detected for R. yasudae thallus. Other sugar compounds did not help the expression of d-NPQ at the same concentrations. Thus, arabitol is essential for the expression of d-NPQ to dissipate excess captured light energy into heat, protecting the photobiont from photoinhibition. The relationship between mycobionts and photobionts is, therefore, not commensalism, but mutualism with each other, as shown by d-NPQ expression.

COMPARATIVE STUDY ON THE PHOTOSYNTHETIC PROPERTIES OF PRASIOLA (CHLOROPHYCEAE) AND NOSTOC (CYANOPHYCEAE) FROM ANTARCTIC AND NON‐ANTARCTIC SITES1

The terrestrial cyanobacterium Nostoc commune Vaucher ex Bornet et Flahault occurs worldwide, including in Japan and on the Antarctic continent. The terrestrial green alga Prasiola crispa (Lightf.) Kütz. is also distributed in Antarctica. These two species need to acclimate to the severe Antarctic climate including low ambient temperature and desiccation under strong light conditions. To clarify this acclimation process, the physiological characteristics of the photosynthetic systems of these two Antarctic terrestrial organisms were assessed. The relative rate of photosynthetic electron flow in N. commune collected in Japan and in Antarctica reached maxima at 900 and 1,100 μmol photons · m−2 · s−1, respectively. The difference seemed to reflect the presence of high amounts of UV‐absorbing substances within the Antarctic cyanobacterium. On the other hand, the optimal temperatures for photosynthesis at the two locations were 30°C–35°C and 20°C–25°C, respectively. This finding suggested a decreased photosynthetic thermotolerance in the Antarctic strain. P. crispa exhibited desiccation tolerance and dehydration‐induced quenching of PSII fluorescence. Re‐reduction of the photooxidized PSI reaction center, P700, was also inhibited at fully dry states. Photosynthetic electron flow in P. crispa reached a maximum at 20°C–25°C and at a light intensity of 700 μmol photons ḃ m−2 ḃ s−1. Interestingly, the osmolarity of P. crispa cells suggested that photosynthesis is performed using water absorbed in a liquid form rather than water absorbed from the air. Overall, these data suggest that these two species have acclimated to optimally photosynthesize under conditions of the highest light intensity and the highest temperature for their habitat in Antarctica.

Ideal osmotic spaces for chlorobionts or cyanobionts are differentially realized by lichenized fungi

Lichenized fungi create a suitable habitat for their photobionts to ensure desiccation tolerance by differentially adjusting their cellular osmolarity to match those of chlorobionts or cyanobionts. Lichens result from symbioses between a fungus and either a green alga or a cyanobacterium. They are known to exhibit extreme desiccation tolerance. We investigated the mechanism that makes photobionts biologically active under severe desiccation using green algal lichens (chlorolichens), cyanobacterial lichens (cyanolichens), a cephalodia-possessing lichen composed of green algal and cyanobacterial parts within the same thallus, a green algal photobiont, an aerial green alga, and a terrestrial cyanobacterium. The photosynthetic response to dehydration by the cyanolichen was almost the same as that of the terrestrial cyanobacterium but was more sensitive than that of the chlorolichen or the chlorobiont. Different responses to dehydration were closely related to cellular osmolarity; osmolarity was comparable between the cyanolichen and a cyanobacterium as well as between a chlorolichen and a green alga. In the cephalodium-possessing lichen, osmolarity and the effect of dehydration on cephalodia were similar to those exhibited by cyanolichens. The green algal part response was similar to those exhibited by chlorolichens. Through the analysis of cellular osmolarity, it was clearly shown that photobionts retain their original properties as free-living organisms even after lichenization.

A comparative study of wavelength-dependent photoinactivation in photosystem II of drought-tolerant photosynthetic organisms in Antarctica and the potential risks of photoinhibition in the habitat

Background and Aims All photosynthetic organisms are faced with photoinhibition, which would lead to death in severe environments. Because light quality and light intensity fluctuate dynamically in natural microenvironments, quantitative and qualitative analysis of photoinhibition is important to clarify how this environmental pressure has impacted ecological behaviour in different organisms. Methods We evaluated the wavelength dependency of photoinactivation to photosystem II (PSII) of Prasiola crispa (green alga), Umbilicaria decussata (lichen) and Ceratodon purpureus (bryophyte) harvested from East Antarctica. For evaluation, we calculated reaction coefficients, Epis, of PSII photoinactivation against energy dose using a large spectrograph. Daily fluctuation of the rate coefficient of photoinactivation, kpi, was estimated from Epis and ambient light spectra measured during the summer season. Key Results Wavelength dependency of PSII photoinactivation was different for the three species, although they form colonies in close proximity to each other in Antarctica. The lichen exhibited substantial resistance to photoinactivation at all wavelengths, while the bryophyte showed sensitivity only to UV-B light (<325 nm). On the other hand, the green alga, P. crispa, showed ten times higher Epi to UV-B light than the bryophyte. It was much more sensitive to UV-A (325-400 nm). The risk of photoinhibition fluctuated considerably throughout the day. On the other hand, Epis were reduced dramatically for dehydrated compared with hydrated P. crispa. Conclusions The deduced rate coefficients of photoinactivation under ambient sunlight suggested that P. crispa needs to pay a greater cost to recover from photodamage than the lichen or the bryophyte in order to keep sufficient photosynthetic activity under the Antarctic habitat. A newly identified drought-induced protection mechanism appears to operate in P. crispa, and it plays a critical role in preventing the oxygen-evolving complex from photoinactivation when the repair cycle is inhibited by dehydration.

Stereochemical assignment of the unique electron acceptor 5′-hydroxyphylloquinone, a polar analog of vitamin K1 in photosystem I

A unique electron-accepting analog of vitamin K1 found in photosystem I in several species of oxygenic photosynthetic microorganisms was confirmed to be 5′-hydroxyphylloquinone (1) through stereo-uncontrolled synthesis. Furthermore, the stereochemistry of 1 obtained from Synechococcus sp. PCC 7942 was assigned to be 5′S using proline-catalyzed stereocontrolled reactions. Polar vitamin K1 (1) was synthesized. Stereochemistry of natural 1 was determined to be (5′S) after a derivation, where the same derivative was also synthesized.