Adaptive strategies of high and low nucleic acid prokaryotes in response to declining resource availability and selective grazing by protozoa

Prokaryotes play a fundamental role in global ocean biogeochemical cycles.However,how the abundance and metabolic activity of ecologically distinct subgroups(i.e.,high nucleic acid(HNA)and low nucleic acid(LNA)cells),and their regulating factors,change in response to changing marine environmental conditions remains poorly understood.Here,we delved into the time-evolving dynamic responses of the HNA and LNA prokaryotic subgroups to declining resource availability and selective grazing by protozoa by conducting a 73-day incubation experiment in a large-volume(117,000 L)macrocosm that facilitates community-level exploration.We found that the metabolic activity of the HNA subgroup was higher than that of the LNA subgroup when the macrocosm was resource replete but that the HNA subgroup declined more rapidly than the LNA subgroup as the resources became increasingly scarce,leading to a steadily increasing contribution of LNA cells to prokaryotic activity.Meanwhile,as resources in the macrocosm became limited,protozoan grazing preference shifted from the HNA to the LNA subgroup and the contributions of the LNA subgroup to the carbon flow within the macrocosm increased.The findings highlight the resilience of LNA cells in resource-limited environments,illuminate the critical role of selective grazing by protozoa in balancing distinct prokaryotic subgroups under changing resource conditions,and demonstrate the complex and adaptive interactions between protozoa and prokaryotes across diverse environmental contexts.

Carbon pools and fluxes in the China Seas and adjacent oceans

The China Seas include the South China Sea, East China Sea, Yellow Sea, and Bohai Sea. Located off the Northwestern Pacific margin, covering 4700000 km^2 from tropical to northern temperate zones, and including a variety of continental margins/basins and depths, the China Seas provide typical cases for carbon budget studies. The South China Sea being a deep basin and part of the Western Pacific Warm Pool is characterized by oceanic features; the East China Sea with a wide continental shelf, enormous terrestrial discharges and open margins to the West Pacific, is featured by strong cross-shelf materials transport; the Yellow Sea is featured by the confluence of cold and warm waters; and the Bohai Sea is a shallow semiclosed gulf with strong impacts of human activities. Three large rivers, the Yangtze River, Yellow River, and Pearl River, flow into the East China Sea, the Bohai Sea, and the South China Sea, respectively. The Kuroshio Current at the outer margin of the Chinese continental shelf is one of the two major western boundary currents of the world oceans and its strength and position directly affect the regional climate of China. These characteristics make the China Seas a typical case of marginal seas to study carbon storage and fluxes. This paper systematically analyzes the literature data on the carbon pools and fluxes of the Bohai Sea,Yellow Sea, East China Sea, and South China Sea, including different interfaces(land-sea, sea-air, sediment-water, and marginal sea-open ocean) and different ecosystems(mangroves, wetland, seagrass beds, macroalgae mariculture, coral reefs, euphotic zones, and water column). Among the four seas, the Bohai Sea and South China Sea are acting as CO_2 sources, releasing about0.22 and 13.86–33.60 Tg C yr^(-1) into the atmosphere, respectively, whereas the Yellow Sea and East China Sea are acting as carbon sinks, absorbing about 1.15 and 6.92–23.30 Tg C yr^(-1) of atmospheric CO_2, respectively. Overall, if only the CO_2 exchange at the sea-air interface is considered, the Chinese marginal seas appear to be a source of atmospheric CO_2, with a net release of 6.01–9.33 Tg C yr^(-1), mainly from the inputs of rivers and adjacent oceans. The riverine dissolved inorganic carbon (DIC) input into the Bohai Sea and Yellow Sea, East China Sea, and South China Sea are 5.04, 14.60, and 40.14 Tg C yr^(-1),respectively. The DIC input from adjacent oceans is as high as 144.81 Tg C yr^(-1), significantly exceeding the carbon released from the seas to the atmosphere. In terms of output, the depositional fluxes of organic carbon in the Bohai Sea, Yellow Sea, East China Sea, and South China Sea are 2.00, 3.60, 7.40, and 5.92 Tg C yr^(-1), respectively. The fluxes of organic carbon from the East China Sea and South China Sea to the adjacent oceans are 15.25–36.70 and 43.93 Tg C yr^(-1), respectively. The annual carbon storage of mangroves, wetlands, and seagrass in Chinese coastal waters is 0.36–1.75 Tg C yr^(-1), with a dissolved organic carbon(DOC) output from seagrass beds of up to 0.59 Tg C yr^(-1). Removable organic carbon flux by Chinese macroalgae mariculture account for 0.68 Tg C yr^(-1) and the associated POC depositional and DOC releasing fluxes are 0.14 and 0.82 Tg C yr^(-1), respectively. Thus, in total, the annual output of organic carbon, which is mainly DOC, in the China Seas is 81.72–104.56 Tg C yr^(-1). The DOC efflux from the East China Sea to the adjacent oceans is 15.00–35.00 Tg C yr^(-1). The DOC efflux from the South China Sea is 31.39 Tg C yr^(-1). Although the marginal China Seas seem to be a source of atmospheric CO_2 based on the CO_2 flux at the sea-air interface, the combined effects of the riverine input in the area, oceanic input, depositional export,and microbial carbon pump(DOC conversion and output) indicate that the China Seas represent an important carbon storage area.