Phaeocystis: A Global Enigma.

The genus Phaeocystis is globally distributed, with blooms commonly occurring on continental shelves. This unusual phytoplankter has two major morphologies: solitary cells and cells embedded in a gelatinous matrix. Only colonies form blooms. Their large size (commonly 2 mm but up to 3 cm) and mucilaginous envelope allow the colonies to escape predation, but data are inconsistent as to whether colonies are grazed. Cultured Phaeocystis can also inhibit the growth of co-occurring phytoplankton or the feeding of potential grazers. Colonies and solitary cells use nitrate as a nitrogen source, although solitary cells can also grow on ammonium. Phaeocystis colonies might be a major contributor to carbon flux to depth, but in most cases, colonies are rapidly remineralized in the upper 300 m. The occurrence of large Phaeocystis blooms is often associated with environments with low and highly variable light and high nitrate levels, with Phaeocystis antarctica blooms being linked additionally to high iron availability. Emerging results indicate that different clones of Phaeocystis have substantial genetic plasticity, which may explain its appearance in a variety of environments. Given the evidence of Phaeocystis appearing in new systems, this trend will likely continue in the near future.

Elucidating colony bloom formation mechanism of a harmful alga Phaeocystis globosa (Prymnesiophyceae) using metaproteomics.

Phaeocystis is a globally distributed Prymnesiophyte genus and usually forms massive harmful colony blooms, which impact marine ecosystem, mariculture, human health, and even threaten coastal nuclear power plant safety. However, the mechanisms behind the colony formation from the solitary cells remain poorly understood. Here, we investigated metabolic processes of both solitary and non-flagellated colonial cells of Phaeocystis globosa at different colony bloom stages in the subtropical Beibu Gulf using a metaproteomic approach. Temperature was significantly correlated with Phaeocystis colony bloom formation, and the flagellated motile solitary cells with abundant flagellum-associated proteins, such as tubulin and dynein, were the exclusive cellular morphotype at the solitary cell stage featured with temperatures ≥21 °C. When the temperature decreased to <21 °C, tiny colonies appeared and the flagellum-associated proteins were down-regulated in both solitary and non-flagellated colonial cells, while proteins involved in biosynthesis, chain polymerization and aggregation of glycosaminoglycan (GAG), a key constituent of gelatinous matrix, were up-regulated, indicating the central role of active GAG biosynthesis during the colony formation. Furthermore, light utilization, carbon fixation, nitrogen assimilation, and amino acid and protein synthesis were also enhanced to provide sufficient energy and substrates for GAG biosynthesis. This study highlighted that temperature induced re-allocation of energy and substances toward GAG biosynthesis is essential for colony bloom formation of P. globosa.

The impact assessment of desalination plant discharges on heavy metal pollution in the coastal sediments of the Persian Gulf.

In recent decades the development of desalination plants (DPs) for desalination of seawater has increased dramatically, while little attention has been paid to the effects of this activity on the accumulation of heavy metals (HMs) in the sediments of affected ecosystems. The present study was implemented to evaluate (1) heavy metal accumulation in sediments impacted by DPs discharges, (2) spatial and temporal changes of HMs and the contamination degree by different types of pollution indexes (single and integrated indices), and (3) ecological risk assessment of cadmium (Cd), lead (Pb), zinc (Zn) and copper (Cu) in sediments affected by DPs discharges. A total of 288 sediment samples were collected seasonally at 24 stations from November 2019 to October 2020. Analysis of HMs concentrations in sediments near the desalination plant discharge provided evidence of local contamination. Maximum concentration of Cu and Pb elements were found in sediments near the desalination plant discharge point. Hierarchical cluster analysis revealed clear segregation of stations impacted by desalination plant discharges and away from discharges. The values of PLI index in sediments of all sampling stations were < 1, indicating that there was no metal pollution by this index. The potential ecological risk index (PERI) ranged from 5.33 ± 0.51 to 11.81 ± 4.98 in sampling sediments and were classified as "low potential ecological risk". These results demonstrate that the DPs discharge increased HMs concentrations in the sediments in close proximity to outlets. The necessary and practical regulations and policies regarding the rejection of the DPs discharge and disposal of chemical compounds must be implemented and enforced.

Bacterial Composition Associated With Giant Colonies of the Harmful Algal Species Phaeocystis globosa.

The cosmopolitan algae Phaeocystis globosa forms harmful algal blooms frequently in a number of tropical and subtropical coastal regions in the past two decades. During the bloom, the giant colony, which is formed by P. globosa, is the dominant morphotype. However, the microenvironment and the microbial composition in the intracolonial fluid are poorly understood. Here, we used high-throughput 16S rRNA amplicon sequencing to examine the bacterial composition and predicted functions in intracolonial fluid. Compared with the bacterial consortia in ambient seawater, intracolonial fluids possessed the lower levels of microbial richness and diversity, implying selectivity of bacteria by the unique intracolonial microenvironment enclosed within the P. globosa polysaccharide envelope. The bacterial consortia in intracolonial fluid were dominated by Balneola (48.6% of total abundance) and Labrezia (28.5%). The bacteria and microbial function enriched in intracolonial fluid were involved in aromatic benzenoid compounds degradation, DMSP and DMS production and consumption, and antibacterial compounds synthesis. We suggest that the P. globosa colonial envelope allows for the formation of a specific microenvironment; thus, the unique microbial consortia inhabiting intracolonial fluid has close interaction with P. globosa cells, which may benefit colony development.

Seasonal resource pulses and the foraging depth of a Southern Ocean top predator.

Seasonal resource pulses can have enormous impacts on species interactions. In marine ecosystems, air-breathing predators often drive their prey to deeper waters. However, it is unclear how ephemeral resource pulses such as near-surface phytoplankton blooms alter the vertical trade-off between predation avoidance and resource availability in consumers, and how these changes cascade to the diving behaviour of top predators. We integrated data on Weddell seal diving behaviour, diet stable isotopes, feeding success and mass gain to examine shifts in vertical foraging throughout ice break-out and the resulting phytoplankton bloom each year. We also tested hypotheses about the likely location of phytoplankton bloom origination (advected or produced in situ where seals foraged) based on sea ice break-out phenology and advection rates from several locations within 150 km of the seal colony. In early summer, seals foraged at deeper depths resulting in lower feeding rates and mass gain. As sea ice extent decreased throughout the summer, seals foraged at shallower depths and benefited from more efficient energy intake. Changes in diving depth were not due to seasonal shifts in seal diets or horizontal space use and instead may reflect a change in the vertical distribution of prey. Correspondence between the timing of seal shallowing and the resource pulse was variable from year to year and could not be readily explained by our existing understanding of the ocean and ice dynamics. Phytoplankton advection occurred faster than ice break-out, and seal dive shallowing occurred substantially earlier than local break-out. While there remains much to be learned about the marine ecosystem, it appears that an increase in prey abundance and accessibility via shallower distributions during the resource pulse could synchronize life-history phenology across trophic levels in this high-latitude ecosystem.

Prevention is better than cure: Persian Gulf biodiversity vulnerability to the impacts of desalination plants.

Due to extremely high rates of evaporation and low precipitation in the Persian Gulf, discharges from desalination plants (DPs) can lead to ecological stresses by increasing water temperatures, salinities, and heavy metal concentrations, as well as decreasing dissolved oxygen levels. We discuss the potential ecological impacts of DPs on marine organisms and propose mitigating measures to reduce the problems induced by DPs discharges. The daily capacity of DPs in the Persian Gulf exceeds 11 million m3 per day, which is approximately half of global daily freshwater production; multistage flash distillation (MSF) is the dominant desalination process. Results from field and laboratory studies indicate that there are potentially serious and chronic threats to marine communities following exposure to DP discharges, especially within the zoobenthos, echinodermata, seagrasses, and coral reefs. DP discharges can lead to decreases in sensitive species, plankton abundance, hard substrate epifauna, and growth rates of seagrasses. However, the broad applicability of any one of these impacts is currently hard to scale because of the limited number of studies that have been conducted to assess the ecological impacts of DP discharge on Persian Gulf organisms. Even so, available data suggest that appropriately sited, designed, and operated DPs combined with current developments in impingement and entrainment reduction technology can mitigate many of the negative environmental impacts of DPs.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota.

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

The oceanography and ecology of the Ross Sea.

The continental shelf of the Ross Sea exhibits substantial variations in physical forcing, ice cover, and biological processes on a variety of time and space scales. Its circulation is characterized by advective inputs from the east and exchanges with off-shelf regions via the troughs along the northern portions. Phytoplankton biomass is greater there than anywhere else in the Antarctic, although nitrate is rarely reduced to levels below 10 µmol L(-1). Overall growth is regulated by irradiance (via ice at the surface and by the depths of the mixed layers) and iron concentrations. Apex predators reach exceptional abundances, and the world's largest colonies of Adélie and emperor penguins are found there. Krill are represented by two species (Euphausia superba near the shelf break and Euphausia crystallorophias throughout the continental shelf region). Equally important and poorly known is the Antarctic silverfish (Pleuragramma antarcticum), which is also consumed by most upper-trophic-level predators. Future changes in the Ross Sea environment will have profound and unpredictable effects on the food web.

Survival and recovery of Phaeocystis antarctica (Prymnesiophyceae) from prolonged darkness and freezing.

The colony-forming haptophyte Phaeocystis antarctica is an important primary producer in the Ross Sea, and must survive long periods of darkness and freezing temperature in this extreme environment. We conducted experiments on the responses of P. antarctica-dominated phytoplankton assemblages to prolonged periods of darkness and freezing. Chlorophyll and photosynthetic capacity of the alga declined nonlinearly and independently of each other in the dark, and darkness alone would potentially reduce photosynthetic capacity by only 60 per cent over 150 days (approximately the length of the Antarctic winter in the southern Ross Sea). The estimated reduction of colonial mucous carbon is higher than that of colonial cell carbon, suggesting metabolism of the colonial matrix in the dark. The alga quickly resumed growth upon return to light. Phaeocystis antarctica also survived freezing, although longer freezing durations lengthened the lag before growth resumption. Particulate dimethylsulfoniopropionate relative to chlorophyll increased upon freezing and decreased upon darkness. Taken together, the abilities of P. antarctica to survive freezing and initiate growth quickly after darkness may provide it with the capability to survive in both the ice and the water column, and help explain its repeated dominance in austral spring blooms in the Ross Sea and elsewhere in the Southern Ocean.

Photosynthetic maximum quantum yield increases are an essential component of the Southern Ocean phytoplankton response to iron.

It is well established that an increase in iron supply causes an increase in total oceanic primary production in many regions, but the physiological mechanism driving the observed increases has not been clearly identified. The Southern Ocean iron enrichment experiment, an iron fertilization experiment in the waters closest to Antarctica, resulted in a 9-fold increase in chlorophyll (Chl) concentration and a 5-fold increase in integrated primary production. Upon iron addition, the maximum quantum yield of photosynthesis (phi(m)) rapidly doubled, from 0.011 to 0.025 mol C.mol quanta(-1). Paradoxically, this increase in light-limited productivity was not accompanied by a significant increase in light-saturated productivity (P(max)(b)). P(max)(b), maximum Chl normalized productivity, was 1.34 mg C.mg Chl(-1).h(-1) outside and 1.49 mg C.mg Chl(-1).h(-1) inside the iron-enriched patch. The importance of phi(m) as compared with P(max)(b) in controlling the biological response to iron addition has vast implications for understanding the ecological response to iron. We show that an iron-driven increase in phi(m) is the proximate physiological mechanism affected by iron addition and can account for most of the increases in primary production. The relative importance of phi(m) over P(max)(b) in this iron-fertilized bloom highlights the limitations of often-used primary productivity algorithms that are driven by estimates of P(max)(b) but largely ignore variability in phi(m) and light-limited productivity. To use primary productivity models that include variability in iron supply in prediction or forecasting, the variability of light-limited productivity must be resolved.

COLONY SIZE OF PHAEOCYSTIS ANTARCTICA (PRYMNESIOPHYCEAE) AS INFLUENCED BY ZOOPLANKTON GRAZERS(1).

The haptophyte Phaeocystis antarctica G. Karst. is a dominant phytoplankton species in the Ross Sea, Antarctica, and exists as solitary cells and mucilaginous colonies that differ by several orders of magnitude in size. Recent studies with Phaeocystis globosa suggest that colony formation and enlargement are defense mechanisms against small grazers. To test if a similar grazer-induced morphological response exists in P. antarctica, we conducted incubation experiments during the austral summer using natural P. antarctica and zooplankton assemblages. Dialysis bags that allowed exchange of dissolved chemicals were used to separate P. antarctica and zooplankton during incubations. Geometric mean colony size decreased by 35% in the control, but increased by 30% in the presence of grazers (even without physical contact) over the 15 d incubation. The estimated colonial-to-solitary cell carbon ratio was significantly higher in the grazing treatment. These results suggest that P. antarctica colonies would grow larger in the presence of indigenous zooplankton and skew the carbon partitioning significantly toward the colonial phase. While these observations show that the colony size of P. antarctica was affected by a chemical signal related to grazers, the detailed nature and ecological significance of this signal remain unknown.

Trophic interactions within the Ross Sea continental shelf ecosystem.

The continental shelf of the Ross Sea is one of the Antarctic's most intensively studied regions. We review the available data on the region's physical characteristics (currents and ice concentrations) and their spatial variations, as well as components of the neritic food web, including lower and middle levels (phytoplankton, zooplankton, krill, fishes), the upper trophic levels (seals, penguins, pelagic birds, whales) and benthic fauna. A hypothetical food web is presented. Biotic interactions, such as the role of Euphausia crystallorophias and Pleuragramma antarcticum as grazers of lower levels and food for higher trophic levels, are suggested as being critical. The neritic food web contrasts dramatically with others in the Antarctic that appear to be structured around the keystone species Euphausia superba. Similarly, we suggest that benthic-pelagic coupling is stronger in the Ross Sea than in most other Antarctic regions. We also highlight many of the unknowns within the food web, and discuss the impacts of a changing Ross Sea habitat on the ecosystem.