Effects of climate change factors on marine macroalgae: A review.

Marine macroalgae, the main primary producers in coastal waters, play important roles in the fishery industry and global carbon cycles. With progressive ocean global changes, however, they are increasingly exposed to enhanced levels of multiple environmental drivers, such as ocean acidification, warming, heatwaves, UV radiation and deoxygenation. While most macroalgae have developed physiological strategies against variations of these drivers, their eco-physiological responses to each or combinations of the drivers differ spatiotemporally and species-specifically. Many freshwater macroalgae are tolerant of pH drop and its diel fluctuations and capable of acclimating to changes in carbonate chemistry. However, calcifying species, such as coralline algae, are very sensitive to acidification of seawater, which reduces their calcification, and additionally, temperature rise and UV further decrease their physiological performance. Except for these calcifying species, both economically important and harmful macroalgae can benefit from elevated CO₂ concentrations and moderate temperature rise, which might be responsible for increasing events of harmful macroalgal blooms including green macroalgal blooms caused by Ulva spp. and golden tides caused by Sargassum spp. Upper intertidal macroalgae, especially those tolerant of dehydration during low tide, increase their photosynthesis under elevated CO₂ concentrations during the initial dehydration period, however, these species might be endangered by heatwaves, which can expose them to high temperature levels above their thermal windows' upper limit. On the other hand, since macroalgae are distributed in shallow waters, they are inevitably exposed to solar UV radiation. The effects of UV radiation, depending on weather conditions and species, can be harmful as well as beneficial to many species. Moderate levels of UV-A (315-400nm) can enhance photosynthesis of green, brown and red algae, while UV-B (280-315nm) mainly show inhibitory impacts. Although little has been documented on the combined effects of elevated CO₂, temperature or heatwaves with UV radiation, exposures to heatwaves during midday under high levels of UV radiation can be detrimental to most species, especially to their microscopic stages which are less tolerant of climate change induced stress. In parallel, reduced availability of dissolved O₂ in coastal water along with eutrophication might favour the macroalgae's carboxylation process by suppressing their oxygenation or photorespiration. In this review, we analyse effects of climate change-relevant drivers individually and/or jointly on different macroalgal groups and different life cycle stages based on the literatures surveyed, and provide perspectives for future studies.