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This study examined the effects of temperature and irradiance on photosynthetic characteristics of the macroscopic sporophyte (SPO) and microscopic gametophyte (GAM) stages of a subarctic brown alga, Saccharina japonica var. japonica (Laminariales) from Hokkaido, Japan. In vitro measurements under short- and long-term exposures were carried out by using optical dissolved oxygen sensors and the pulse amplitude modulation (PAM)-chlorophyll fluorometer, respectively. The heteromorphic life history stages of S. japonica showed photosynthetic optima at 23–23.3 °C, derived from the gross photosynthesis–temperature model. Maximum quantum yields (Fv/Fm) of SPO and GAM after 72 h of temperature exposures in the dark were reduced to near zero above 24 °C, indicating PSII inactivation. Such similarity in their temperature characteristics suggests the co-occurrence of both generations in the habitat despite the seasonal growth and reproduction of the species. Net photosynthesis–irradiance experiments in the two life history stages at 8, 16, and 24 °C revealed similarity in their light-saturated photosynthetic rates (NPmax = 3.02–4.41 μg O2 gww−1 min−1, SPO; 2.87–3.73 μg O2 gww−1 min−1, GAM), but saturation irradiances of SPO (Ek = 81–102 μmol photons m−2 s−1) were higher than those of GAM (48–69 μmol photons m−2 s−1). A slight decrease in net photosynthetic rates of GAM above 500 μmol photons m−2 s−1 was likewise observed. This difference may be related to the light regime of their natural habitat, suggesting the low irradiance adaptation of the microscopic stage that settles on rock crevices beneath algal canopies.

This study investigated the effects of different light spectral qualities and temperature on the photosynthesis and pigment content of a subtidal edible red alga, Meristotheca papulosa. Photosynthesis–irradiance (P–E) experiments were carried out under red (660 nm), blue (450 nm), green (525 nm, light-emitting diodes), and white light (visible light, metal halide lamp), and at 12, 20, and 28 °C, respectively. Maximum net photosynthetic rates (NPmax) were highest under green light. Other P–E parameter estimates were similar among algae under red, blue, and green light, including their lower initial slope (α) and higher saturation irradiances (Ek) as compared to those under white light. Additionally, NPmax and Ek under white light were highest at 28 °C, and lowest at 12 °C with characteristic photoinhibition at irradiances greater than 150 μmol photons m−2 s−1. Photosynthesis–temperature (P–T) experiment revealed that the maximum gross photosynthetic rate (GPmax) occurred at 22.1 °C, which was within the optimal temperature range of Fv/Fm (21.5–23.6 °C). Exposures to the different light qualities at 100 μmol photons m−2 s−1 for 7 days showed increased phycoerythrin (PE) concentration of algae under blue and green light, while chlorophyll-a and phycocyanin (PC) showed little variation in all light qualities. Therefore, considering future management prospects for M. papulosa mariculture, we suggest that green light could be utilized to enhance photosynthesis. Furthermore, if the aim is to achieve high PE content for an improved reddish-color fresh product, exposure to blue or green light could be a good alternative.

The combined effects of photosynthetically active radiation (PAR) and temperature on the photosynthesis of a temperate Japanese brown alga, Sargassum patens (Fucales), were determined by field and laboratory measurements. Underwater measurements of the natural population of this alga in Kagoshima, Japan, revealed that the effective quantum yield (ΦPSII) declined with increasing incident PAR, with minimum ΦPSII occurring during noon to early afternoon. ΦPSII recovered in the evening, indicating dynamic photoinhibition. In laboratory experiments, ΦPSII was also negatively correlated with PAR, which decreased after 12 h of continuous exposure to 200 (low) and 1000 (high) μmol photons m−2 s−1 at 8, 20, and 28 °C. Maximum quantum yield (Fv/Fm) at 8 °C with low PAR failed to recover after 12 h of dark acclimation, suggesting the influence of low temperature in chronic photoinhibition. Photosynthesis–irradiance (P–E) curves likewise revealed lower net photosynthetic rates and photoinhibition at 8 °C. Gross photosynthesis and dark respiration experiments determined over a range of temperatures (8–40 °C) revealed that the maximum gross photosynthetic rate (GPmax) occurred at 26.9 °C. Fv/Fm after 72 h of temperature exposures was stable at 8–32 °C; but it was deactivated at 36 and 40 °C. This species is well-adapted to the current range of temperature in the temperate region of Japan (7–28 °C). However, the combined effects of low temperature and PAR may influence algal photosynthetic efficiency and so may be limiting at the marginal region of northern distribution of this temperate species.

The effects of temperature, irradiance, and desiccation on the photosynthesis of a cultivated Japanese green alga Caulerpa lentillifera (Caulerpaceae) were determined by a pulse amplitude modulation (PAM)-chlorophyll fluorometer and dissolved oxygen sensors. The photochemical efficiency in the photosystem II (Fv/Fm and ΔF/Fm') during the 72-h temperature exposures (8, 12, 16, 20, 24, 28, 32, 36, and 40°C) was generally stable at 16–32°C but quickly dropped at lower and higher temperatures. The photosynthesis–temperature curve at 200 μmol photons m−2 s−1 also revealed that the maximum gross photosynthesis (GPmax) occurred at 30.7°C (30.5–30.9, 95% highest density credible intervals). Photosynthesis–irradiance curves at 16, 24, and 32°C quickly saturated, then expressed photoinhibition, and revealed that the maximum net photosynthetic rates (NPmax) and saturation irradiance (Ek) were highest at 32°C and lowest at 16°C. Continuous 6-h exposure to irradiances of 200 (low) and 400 (high) μmol photons m−2 s−1 at 16, 24, and 32°C expressed greater declines in their ΔF/Fm' at 16°C, revealing chronic chilling-light stress. The response to continuous desiccation (~480 min) under 50% humidity at 24°C showed that ΔF/Fm' dropped to zero at 480-min aerial exposure, and the treatments of more than 60-min desiccation did not return to the initial level even after 24-h subsequent rehydration in seawater. Likewise, ΔF/Fm' fell when the absolute water content (AWC) of the frond dropped below AWC of 90% and mostly did not return to the initial level even after 24-h subsequent rehydration in seawater, signifying a low tolerance to desiccation.

We examined the effects of four stressors, irradiance, temperature, desiccation, and salinity on the photosynthesis of a red alga, Agarophyton vermiculophyllum (= Gracilaria vermiculophylla, Gracilariales, Rhodophyta) from its native distributional range in Hokkaido and Kagoshima, Japan. Photosynthesis–irradiance (P–E) curves at 8, 16, and 28°C showed that the maximum net-photosynthetic rates (NPmax) and saturation irradiance (Ek) were highest at 28°C for both strains. Gross-photosynthesis determined at 8–40°C at 200 μmol photons m-2 s-1 showed that the maximum gross-photosynthetic rate (GPmax) occurred at 25.3°C for Hokkaido and 28.0°C for Kagoshima (ToptGP), which is almost consistent with the summer-time seawater temperature at each habitat. The temperature responses (4–40°C) of effective quantum yields (ΔF/Fm') of photosystem II during 7-day exposures were similar to that of oxygenic photosynthesis and the optimum temperature (ToptΔF/Fm′) was 20.0°C for Hokkaido and 30.1°C for Kagoshima. In the desiccation experiment, the ΔF/Fm' decreased with decreasing absolute water content (AWC); nevertheless, for samples with an AWC above 20%, ΔF/Fm' retuned to initial levels after subsequent 1-day rehydration in seawater, suggesting relatively strong tolerance to desiccation. This alga also showed a broad range of tolerance to salinity ranging from 20 to 60 psu in 7-day exposures, and the ΔF/Fm' tolerated 0 psu at 3-day exposure. The adaptations of A. vermiculophyllum from its native range in Japan to relatively high irradiance, a broad range of temperature, and strong osmotic (desiccation and salinity) tolerance explain its potentially high invasive capacity.

We determined the effects of irradiance, temperature, and desiccation on the photosynthesis of a brown alga, Sargassum muticum (Fucales), from its native distributional range in Japan by using a pulse amplitude modulation (PAM)-chlorophyll fluorometer and optical dissolved oxygen sensors. Photosynthesis–irradiance curves at three temperatures (8, 20, and 28°C) showed that the maximum net photosynthetic rates (NPmax) and saturation irradiance were highest at 28°C. Gross photosynthesis determined at 8–36°C (every four increments) and 300 μmol photons m−2 s−1 showed that the maximum gross photosynthetic rate (GPmax) occurred at 19.5°C (ToptGP), which is consistent with the seawater temperature at its peaked abundance in Japan. The maximum quantum yields (Fv/Fm) of photosystem II (PSII) during the 72-h temperature exposures were above 0.60 at 8–28°C but dropped at higher temperatures. Continuous exposure (12 h) to irradiance of 200 (low) and 1000 (high) μmol photons m−2 s−1 at three temperatures showed remarkable decline in the effective quantum yields (ΔF/Fm') of PSII under high irradiance at 8°C only; the Fv/Fm measured after 12-h dark acclimation also did not recover to initial values, signifying its sensitivity to photoinhibition at 8°C. Furthermore, the alga exhibited tolerance to 2 h of desiccation with 80% of water loss from the thallus, and ΔF/Fm' recovered after 24 h of rehydration in seawater, suggesting potential of photosynthetic recovery of this alga at such low hydration threshold. In conclusion, the adaptation of S. muticum to relatively high irradiance, to broad range of temperature (8–28°C), and to desiccation explains its potentially high invasive capacity.

The chronic effects of three different stressors, irradiance, temperature, and desiccation, on Photosystem II (PSII) photochemical efficiency were investigated in two heteromorphic life-history stages of a cultivated red alga, Pyropia yezoensis f. narawaensis (Bangiales) from Japan. The maximum quantum yields of PSII (Fv/Fm) over the range of temperature between 4 and 36 °C for 96-h exposure showed different responses in the two life-history stages. In the macroscopic gametophyte, the highest value (0.47) occurred at 12.0 °C; however, Fv/Fm quickly decreased at higher temperatures. In contrast, Fv/Fm in the microscopic sporophyte were less sensitive to temperature, suggesting adaptation to the broad range of temperature with a high value (0.40) that occurred at 21.1 °C. Continuous 6-h exposures to the 50, 100, and 1000 μmol photons m−2 s−1 revealed greater decline in their effective quantum yields of PSII (ΔF/Fm′) in the sporophyte. As known in the photoprotective response and its PSII repair system in the photosynthesis, the recovery of quantum yield in two life-history stages after subsequent acclimation (darkness and dim-light treatments) was more accelerated in dim-light than those of darkness. The response to continuous desiccation (240-min aerial exposure) in two life-history stages was also different, and the ΔF/Fm′ in the gametophyte returned to initial values after re-immersion in seawater. In contrast, those in the sporophyte dropped to zero after a 5-min aerial exposure and did not return to initial values, suggesting that farming protocols for the conchospore seeding in Nori-net cultivation need to pay careful attention to desiccation to ensure successful production.

The effects of temperature on the photosynthesis and growth of a subtropical red alga, Phycocalidia tanegashimensis (= Pyropia tanegashimensis, Bangiaceae, Bangiales) from Tanegashima Island, Japan, were determined to better understand the ecology of the macroscopic gametophyte. Net photosynthesis–irradiance (P–E) curves, determined at 12, 20, and 28°C, revealed that the maximum net photosynthetic rates occurred at 28°C. A gross photosynthesis–temperature (P–T) curve, determined at temperatures between 8 and 36°C, supported this result and indicated that optimal temperature (ToptGP) for maximum gross photosynthetic rates was 27.3°C (25.9–28.8 95% highest density credible intervals, HDCI). However, after 4 days of incubation at constant temperature, the effective quantum yields (ΔF/Fm') of photosystem II, determined between 4 and 40°C under 50 μmol photons m−2 s−1 (12L:12D), were greatest at 22.1°C (20.9–23.4 HDCI) and declined quickly below 20°C and 32°C. The relative growth rate (RGR) was highest at 25.8°C (22.5–28.1 HDCI) at the conclusion of a 6-day culture (4 to 36°C) under 50 μmol photons m−2 s−1 (12L:12D). Compared to other temperate species of Bangiaceae, P. tanegashimensis appears to be well adapted to relatively high temperatures. The macroscopic gametophyte stage of Bangiaceae is regarded to be adapted to cold temperatures to overwinter; nevertheless, the significance of our study is that a year-round occurrence of P. tanegashimensis is enabled by its high optimal temperatures for photosynthesis and growth.

Phenology, irradiance, and temperature characteristics of an edible brown alga, Undaria pinnatifida (Laminariales), were examined from the southernmost natural population in Japan, both by culturing gametophytes and examining the photosynthetic activity of sporophytes using dissolved oxygen sensors and pulse amplitude-modulated chlorophyll fluorometer (IMAGING-PAM). Our surveys confirmed that sporophytes were present between winter and early summer, but absent by July. IMAGING-PAM experiments were used to measure maximum effective quantum yield (ΦII at 0 μmol photons m⁻² s⁻¹) for each of 14 temperatures (8–36 °C). Oxygen production was also determined over a coarser temperature gradient. Net photosynthesis and ΦII (at 0 μmol photons m⁻² s⁻¹) were observed to be temperature-dependent; the maximum ΦII was estimated to be 0.67, occurred at 21.2 °C, and was nearly identical to the optimal temperature of the net photosynthetic rate (21.7 °C). A net photosynthesis–irradiance (P–E) model revealed that saturation irradiance (E ₖ) was 119.5 μmol photons m⁻¹ s⁻¹, and the compensation irradiance (E c) was 17.4 μmol photons m⁻¹ s⁻¹. Culture experiments on the gametophytes revealed that most individuals could not survive temperatures over 28 °C and that growth rates were severely inhibited. Based on our observations, temperatures greater than 20 °C are likely to influence photosynthetic activity and gametophyte survival, and therefore, it is possible that this species might become locally extinct if seawater temperatures in this region continue to rise.

Toxic crabs of the family Xanthidae contain saxitoxins (STXs) and/or tetrodotoxin (TTX), but the toxin ratio differs depending on their habitat. In the present study, to clarify within reef variations in the toxin profile of xanthid crabs, we collected specimens of the toxic xanthid crab Zosimus aeneus and their sampling location within a single reef (Yoshihara reef) on Ishigaki Island, Okinawa Prefecture, Japan, in 2018 and 2019. The STXs/TTX content within the appendages and viscera or stomach contents of each specimen was determined by instrumental analyses. Our findings revealed the existence of three zones in Yoshihara reef; one in which many individuals accumulate extremely high concentrations of STXs (northwestern part of the reef; NW zone), another in which individuals generally have small amounts of TTX but little STXs (central part of the reef; CTR zone), and a third in which individuals generally exhibit intermediate characteristics (southeastern part of the reef; SE zone). Furthermore, light microscopic observations of the stomach contents of crab specimens collected from the NW and CTR zones revealed that ascidian spicules of the genus Lissoclinum were dominant in the NW zone, whereas those of the genus Trididemnum were dominant in the CTR zone. Although the toxicity of these ascidians is unknown, Lissoclinum ascidians are considered good candidate source organisms of STXs harbored by toxic xanthid crabs.