Coccolithophore responses to environmental variability in the South China Sea: species composition and calcite content

Coccolithophore contributions to the global marine carbon cycle are regulated by the calcite content of their scales (coccoliths) and the relative cellular levels of photosynthesis and calcification rates. All three of these factors vary between coccolithophore species and with response to the growth environment. Here, water samples were collected in the northern basin of the South China Sea (SCS) during summer 2014 in order to examine how environmental variability influenced species composition and cellular levels of calcite content. Average coccolithophore abundance and their calcite concentration in the water column were 11.82 cells mL−1 and 1508.3 pg C mL−1, respectively, during the cruise. Water samples can be divided into three floral groups according to their distinct coccolithophore communities. The vertical structure of the coccolithophore community in the water column was controlled by the trophic conditions, which were regulated by mesoscale eddies across the SCS basin. The evaluation of coccolithophore-based calcite in the surface ocean also showed that three key species in the SCS (Emiliania huxleyi, Gephyrocapsa oceanica, Florisphaera profunda) and other larger, numerically rare species made almost equal contributions to total coccolith-based calcite in the water column. For Emiliania huxleyi biometry measurements, coccolith size positively correlated with nutrients (nitrate, phosphate), and it is suggested that coccolith length is influenced by light and nutrients through the regulation of growth rates. Larger-sized coccoliths were also linked statistically to low pH and calcite saturation states; however, it is not a simple cause and effect relationship, as carbonate chemistry was strongly co-correlated with the other key environmental factors (nutrients, light)

Phenological characteristics of global coccolithophore blooms

Coccolithophores are recognized as having a significant influence on the global carbon cycle through the production and export of calcium carbonate (often referred to as particulate inorganic carbon or PIC). Using remotely sensed PIC and chlorophyll data, we investigate the seasonal dynamics of coccolithophores relative to a mixed phytoplankton community. Seasonal variability in PIC, here considered to indicate changes in coccolithophore biomass, is identified across much of the global ocean. Blooms, which typically start in February–March in the low-latitude (~30°) Northern Hemisphere and last for ~6–7 months, get progressively later (April–May) and shorter (3–4 months) moving poleward. A similar pattern is observed in the Southern Hemisphere, where blooms that generally begin around August–September in the lower latitudes and which last for ~8 months get later and shorter with increasing latitude. It has previously been considered that phytoplankton blooms consist of a sequential succession of blooms of individual phytoplankton types. Comparison of PIC and chlorophyll peak dates suggests instead that in many open ocean regions, blooms of coccolithophores and other phytoplankton can co-occur, conflicting with the traditional view of species succession that is thought to take place in temperate regions such as the North Atlantic

Spring phytoplankton communities of the Labrador Sea (2005–2014): pigment signatures, photophysiology and elemental ratios

The Labrador Sea is an ideal region to study the biogeographical, physiological, and biogeochemical implications of phytoplankton community composition due to sharp transitions between distinct water masses across its shelves and central basin. We have investigated the multi-year (2005–2014) distributions of late spring and early summer (May to June) phytoplankton communities in the various hydrographic settings of the Labrador Sea. Our analysis is based on pigment markers (using CHEMTAX analysis), and photophysiological and biogeochemical characteristics associated with each phytoplankton community. Diatoms were the most abundant group, blooming first in shallow mixed layers of haline-stratified Arctic shelf waters. Along with diatoms, chlorophytes co-dominated at the western end of the section (particularly in the polar waters of the Labrador Current (LC)), whilst Phaeocystis co-dominated in the east (modified polar waters of the West Greenland Current (WGC)). Pre-bloom conditions occurred in deeper mixed layers of the central Labrador Sea in May, where a mixed assemblage of flagellates (dinoflagellates, prasinophytes, prymnesiophytes, particularly coccolithophores, and chrysophytes/pelagophytes) occurred in low-chlorophyll areas, succeeding to blooms of diatoms and dinoflagellates in thermally stratified Atlantic waters in June. Light-saturated photosynthetic rates and saturation irradiance levels were highest at stations where diatoms were the dominant phytoplankton group ( >  70 % of total chlorophyll a), as opposed to stations where flagellates were more abundant (from 40 up to 70 % of total chlorophyll a). Phytoplankton communities from the WGC (Phaeocystis and diatoms) had lower light-limited photosynthetic rates, with little evidence of photoinhibition, indicating greater tolerance to a high light environment. By contrast, communities from the central Labrador Sea (dinoflagellates and diatoms), which bloomed later in the season (June), appeared to be more sensitive to high light levels. Ratios of accessory pigments (AP) to total chlorophyll a (TChl a) varied according to phytoplankton community composition, with polar phytoplankton (cold-water related) having lower AP  :  TChl a. Polar waters (LC and WGC) also had higher and more variable particulate organic carbon (POC) to particulate organic nitrogen (PON) ratios, suggesting the influence of detritus from freshwater input, derived from riverine, glacial, and/or sea ice meltwater. Long-term observational shifts in phytoplankton communities were not assessed in this study due to the short temporal frame (May to June) of the data. Nevertheless, these results add to our current understanding of phytoplankton group distribution, as well as an evaluation of the biogeochemical role (in terms of C  :  N ratios) of spring phytoplankton communities in the Labrador Sea, which will assist our understanding of potential long-term responses of phytoplankton communities in high-latitude oceans to a changing climate

The origin and rise of complex life:progress requires interdisciplinary integration and hypothesis testing

Understanding of the triggers and timing of the rise of complex life ca 2100 to 720 million years ago has expanded dramatically in recent years. This theme issue brings together diverse and novel geochemical and palaeontological data presented as part of the Royal Society ‘The origin and rise of complex life: integrating models, geochemical and palaeontological data’ discussion meeting held in September 2019. The individual papers offer prescient insights from multiple disciplines. Here we summarize their contribution towards the goal of the meeting; to create testable hypotheses for the differing roles of changing climate, oceanic redox, nutrient availability, and ecosystem feedbacks across this profound, but enigmatic, transitional period

Environmental drivers of coccolithophore abundance and calcification across Drake Passage (Southern Ocean)

Although coccolithophores are not as numerically common or as diverse in the Southern Ocean as they are in subpolar waters of the North Atlantic, a few species, such as Emiliania huxleyi, are found during the summer months. Little is actually known about the calcite production (CP) of these communities or how their distribution and physiology relate to environmental variables in this region. In February 2009, we made observations across Drake Passage (between South America and the Antarctic Peninsula) of coccolithophore distribution, CP, primary production, chlorophyll a and macronutrient concentrations, irradiance and carbonate chemistry. Although CP represented less than 1% of total carbon fixation, coccolithophores were widespread across Drake Passage. The B/C morphotype of E. huxleyi was the dominant coccolithophore, with low estimates of coccolith calcite ( 0.01 pmol C coccolith-/ from biometric measurements. Both cell-normalised calcification (0.01–0.16 pmol C cell-1 d-1/ and total CP (< 20 μmol C m-1 d-1/were much lower than those observed in the subpolar North Atlantic where E. huxleyi morphotype A is dominant. However, estimates of coccolith production rates were similar (0.1–1.2 coccoliths cell-1 h-1/ to previous measurements made in the subpolar North Atlantic. A multivariate statistical approach found that temperature and irradiance together were best able to explain the observed variation in species distribution and abundance (Spearman’s rank correlation D0.4, p < 0.01). Rates of calcification per cell and coccolith production, as well as community CP and E. huxleyi abundance, were all positively correlated (p < 0.05) to the strong latitudinal gradient in temperature, irradiance and calcite saturation states across Drake Passage. Broadly, our results lend support to recent suggestions that coccolithophores, especially E. huxleyi, are advancing polewards. However, our in situ observations indicate that this may owe more to sea-surface warming and increasing irradiance rather than increasing CO2 concentrations

Geographical CO2 sensitivity of phytoplankton correlates with ocean buffer capacity

Accumulation of anthropogenic CO2 is significantly altering ocean chemistry. A range of biological impacts resulting from this oceanic CO2 accumulation are emerging, however the mechanisms responsible for observed differential susceptibility between organisms and across environmental settings remain obscure. A primary consequence of increased oceanic CO2 uptake is a decrease in the carbonate system buffer capacity, which characterises the system's chemical resilience to changes in CO2, generating the potential for enhanced variability in pCO2 and the concentration of carbonate [CO32‐], bicarbonate [HCO3‐] and protons [H+] in the future ocean. We conducted a meta‐analysis of 17 shipboard manipulation experiments performed across three distinct geographical regions that encompassed a wide range of environmental conditions from European temperate seas to Arctic and Southern oceans. These data demonstrated a correlation between the magnitude of natural phytoplankton community biological responses to short‐term CO2 changes and variability in the local buffer capacity across ocean basin scales. Specifically, short‐term suppression of small phytoplankton (<10 μm) net growth rates were consistently observed under enhanced pCO2 within experiments performed in regions with higher ambient buffer capacity. The results further highlight the relevance of phytoplankton cell size for the impacts of enhanced pCO2 in both the modern and future ocean. Specifically, cell‐size related acclimation and adaptation to regional environmental variability, as characterised by buffer capacity, likely influences interactions between primary producers and carbonate chemistry over a range of spatio‐temporal scales

Ocean warming, not acidification, controlled coccolithophore response during past greenhouse climate change

Current carbon dioxide emissions are an assumed threat to oceanic calcifying plankton (coccolithophores) not just due to rising sea-surface temperatures, but also because of ocean acidification (OA). This assessment is based on single species culture experiments that are now revealing complex, synergistic, and adaptive responses to such environmental change. Despite this complexity, there is still a widespread perception that coccolithophore calcification will be inhibited by OA. These plankton have an excellent fossil record, and so we can test for the impact of OA during geological carbon cycle events, providing the added advantages of exploring entire communities across real-world major climate perturbation and recovery. Here we target fossil coccolithophore groups (holococcoliths and braarudosphaerids) expected to exhibit greatest sensitivity to acidification because of their reliance on extracellular calcification. Across the Paleocene-Eocene Thermal Maximum (56 Ma) rapid warming event, the biogeography and abundance of these extracellular calcifiers shifted dramatically, disappearing entirely from low latitudes to become limited to cooler, lower saturation-state areas. By comparing these range shift data with the environmental parameters from an Earth system model, we show that the principal control on these range retractions was temperature, with survival maintained in high-latitude refugia, despite more adverse ocean chemistry conditions. Deleterious effects of OA were only evidenced when twinned with elevated temperatures

Day length as a key factor moderating the response of coccolithophore growth to elevated pCO2

The fate of coccolithophores in the future oceans remains uncertain, in part due to key factors having not been standardized across experiments. A potentially moderating role for differences in day length (photoperiod) remains largely unexplored. We therefore cultured four different geographical isolates of the species Emiliania huxleyi, as well as two additional species, Gephyrocapsa oceanica (tropical) and Coccolithus braarudii (temperate), to test for interactive effects of pCO2 with the light : dark (L : D) cycle. We confirmed a general regulatory effect of photoperiod on the pCO2 response, whereby growth and particulate inorganic carbon and particulate organic carbon (PIC : POC) ratios were reduced with elevated pCO2 under 14 : 10 h L : D, but these reductions were dampened under continuous (24 h) light. The dynamics underpinning this pattern generally differed for the temperate vs. tropical isolates. Reductions in PIC : POC with elevated pCO2 for tropical taxa were largely through reduced calcification and enhanced photosynthesis under 14 : 10 h L : D, with differences dampened under continuous light. In contrast, reduced PIC : POC for temperate strains reflected increases of photosynthesis that outpaced increases in calcification rates under 14 : 10 h L : D, with both responses again dampened under continuous light. A multivariate analysis of 35 past studies of E. huxleyi further demonstrated that differences in photoperiod account for as much as 40% (strain B11/92) to 55% (strain NZEH) of the variance in reported pCO2-induced reductions to growth but not PIC : POC. Our study thus highlights a critical role for day length in moderating the effect of ocean acidification on coccolithophore growth and consequently how this response may play out across latitudes and seasons in future oceans

Species-specific calcite production reveals Coccolithus pelagicus as the key calcifier in the Arctic Ocean

Through the production and export of their calcite coccoliths, coccolithophores form a key component of the global carbon cycle. Despite this key role, very little is known about the biogeochemical role of different coccolithophore species in terms of calcite production, and how these species will respond to future climate change and ocean acidification. Here, we present the first study to estimate species-specific calcite production, from samples collected in the Arctic Ocean and subarctic Iceland Basin in June 2012. We show that although the coccolithophorid Coccolithus pelagicus comprised only a small fraction of the total community in terms of abundance (2%), our estimates indicate that it was the major calcite producer in the Arctic Ocean and Iceland Basin (57% of total calcite production). In contrast, Emiliania huxleyi formed 27% of the total abundance and was responsible for only 20% of the calcite production. That C. pelagicus was able to dominate calcite production was due to its relatively high cellular calcite content compared with the other species present. Our results demonstrate, for the first time, the importance of investigating the complete coccolithophore community when considering pelagic calcite production, as relatively rare but heavily calcified species such as C. pelagicus can be the key calcite producers in mixed communities. Therefore, the response of C. pelagicus to ocean acidification and climate change has the potential to have a major impact on carbon cycling within the North Atlantic and Arctic Ocean

Algal plankton turn to hunting to survive and recover from end-Cretaceous impact darkness

The end-Cretaceous bolide impact triggered the devastation of marine ecosystems. However, the specific kill mechanism(s) are still debated, and how primary production subsequently recovered remains elusive. We used marine plankton microfossils and eco-evolutionary modeling to determine strategies for survival and recovery, finding that widespread phagotrophy (prey ingestion) was fundamental to plankton surviving the impact and also for the subsequent reestablishment of primary production. Ecological selectivity points to extreme postimpact light inhibition as the principal kill mechanism, with the marine food chain temporarily reset to a bacteria-dominated state. Subsequently, in a sunlit ocean inhabited by only rare survivor grazers but abundant small prey, it was mixotrophic nutrition (autotrophy and heterotrophy) and increasing cell sizes that enabled the eventual reestablishment of marine food webs some 2 million years later.</p