Deep chlorophyll maximum and nutricline in the Mediterranean Sea: emerging properties from a multi-platform assimilated biogeochemical model experiment

Data assimilation has led to advancements in biogeochemical modelling and scientific understanding of the ocean. The recent operational availability of data from BGC-Argo (biogeochemical Argo) floats, which provide valuable insights into key vertical biogeochemical processes, stands to further improve biogeochemical modelling through assimilation schemes that include float observations in addition to traditionally assimilated satellite data. In the present work, we demonstrate the feasibility of joint multi-platform assimilation in realistic biogeochemical applications by presenting the results of 1-year simulations of Mediterranean Sea biogeochemistry. Different combinations of satellite chlorophyll data and BGC-Argo nitrate and chlorophyll data have been tested, and validation with respect to available independent non-assimilated and assimilated (before the assimilation) observations showed that assimilation of both satellite and float observations outperformed the assimilation of platforms considered individually. Moreover, the assimilation of BGC-Argo data impacted the vertical structure of nutrients and phytoplankton in terms of deep chlorophyll maximum depth, intensity, and nutricline depth. The outcomes of the model simulation assimilating both satellite data and BGC-Argo data provide a consistent picture of the basin-wide differences in vertical features associated with summer stratified conditions, describing a relatively high variability between the western and eastern Mediterranean, with thinner and shallower but intense deep chlorophyll maxima associated with steeper and narrower nutriclines in the western Mediterranean.

Siderophores as an iron source for Prochlorococcus in deep chlorophyll maximum layers of the oligotrophic ocean

Prochlorococcus is one of the most abundant photosynthesizing organisms in the oligotrophic oceans. Gene content variation among Prochlorococcus populations in separate ocean basins often mirrors the selective pressures imposed by the region's distinct biogeochemistry. By pairing genomic datasets with trace metal concentrations from across the global ocean, we show that the genomic capacity for siderophore-mediated iron uptake is widespread in low-light adapted Prochlorococcus populations from iron-depleted regions of the oligotrophic Pacific and S. Atlantic oceans: Prochlorococcus siderophore consumers were absent in the N. Atlantic ocean (higher iron flux) but constituted up to half of all Prochlorococcus genomes from metagenomes in the N. Pacific (lower iron flux). Prochlorococcus siderophore consumers, like many other bacteria with this trait, also lack siderophore biosynthesis genes indicating that they scavenge exogenous siderophores from seawater. Statistical modeling suggests that the capacity for siderophore uptake is endemic to remote ocean regions where atmospheric iron fluxes are the smallest, particularly at deep chlorophyll maximum and primary nitrite maximum layers. We argue that abundant siderophore consumers at these two common oceanographic features could be a symptom of wider community iron stress, consistent with prior hypotheses. Our results provide a clear example of iron as a selective force driving the evolution of Prochlorococcus.

Variability of Microbial Particulate ATP Concentrations in Subeuphotic Microbes Due to Underlying Metabolic Strategies in the South Pacific Ocean

Adenosine triphosphate (ATP) is the primary energy storage molecule in metabolic pathways. It is common in marine studies to use particulate ATP (PATP) concentrations as representative of microbial biomass. However, there is growing evidence from culture studies, models, and transcriptional data that PATP concentration varies across microbes and conditions, thus compromising interpretations in environmental settings. Importantly, there is a lack of open ocean studies assessing variations in PATP concentrations and thus a deficiency of information on the key biogeochemical drivers for variability in microbial PATP independent of biomass. In sampling the U.S. GO-SHIP P06E zonal transect (32.5°S) across the eastern South Pacific, from the subtropical gyre to the upwelling waters off Chile, we conducted the first comprehensive transect survey quantifying PATP. PATP concentrations increased toward the upwelling region of the transect, but varied vertically when normalized against three measures of biomass: particulate phosphorus, microbial abundance, and microbial biovolume. Generally, greater biomass-normalized PATP concentrations were observed below the deep chlorophyll maximum. Subdividing the P06E transect into four biogeochemical regimes highlighted distinct metabolic strategies used by microbes. Between these regimes, we found PATP concentrations were representative of biomass in upper surface waters. However, below the deep chlorophyll maximum we observed higher biomass normalized PATP concentrations that we hypothesize were due to less availability of energy sources in those subeuphotic zone waters and abundances of chemoautotrophs in the microbial community. This finding suggests that stored energy was more important for these deeper microbes.

Deep chlorophyll maximum and nutricline in the Mediterranean Sea: emerging properties from a multi-platform assimilated biogeochemical model experiment

Data assimilation has had a positive impact on biogeochemical modelling in a number of oceanographic applications. The recent operational availability of data from BGC-Argo floats, which provide valuable insights into key vertical biogeochemical processes, can lead to further improvements in biogeochemical modelling through assimilation schemes that include float observations in addition to traditionally assimilated satellite data. In the present work, we demonstrate the feasibility of joint multi-platform assimilation in realistic biogeochemical applications by presenting the results of one-year simulations of Mediterranean Sea biogeochemistry. Different combinations of satellite chlorophyll data and BGC-Argo nitrate and chlorophyll data have been tested, and validation with respect to available independent and semi-independent (before assimilation) observations showed that assimilation of both satellite and float observations outperformed the assimilation of platforms considered individually. Moreover, the assimilation of BGC-Argo data impacted the vertical structure of nutrients and phytoplankton in terms of deep chlorophyll maximum depth and intensity and nutricline depth. The outcomes of the model simulation assimilating both satellite data and BGC-Argo data have been used to explore the basin-wide differences in vertical features associated with summer stratified conditions, describing a relatively high variability between the western and eastern Mediterranean, with thinner and shallower but intense deep chlorophyll maxima associated with steeper and narrower nutriclines in the western Mediterranean.

Spatial distribution of the summer subsurface chlorophyll maximum in the North South China Sea

Based on the biological, nutrients and hydrological data in August 2018, the vertical chlorophyll a (Chl-a) concentration profiles and the relationship among surface Chl-a (Chl-a(0)) concentration, maximum Chl-a (Chl-a(m)) concentration and depth-integrated Chl-a (Chl-a(int)) concentration were studied in the Northern South China Sea (NSCS). The results indicate that there are 4 different patterns in the vertical Chl-a profiles in the NSCS: (i) Chl-a increases with depth from the surface (e.g. station 1); (ii) there exists subsurface chlorophyll maximum (SCM), with low Chl-a on the surface and at the bottom layers respectively (e.g. station 5); (iii) there is no SCM, only with high Chl-a on the surface and in the bottom (e.g. station 14); (iv) the 4th pattern is similar to (ii), with the higher Chl-a(0) (e.g. station 28). The SCM is observed at 95% stations in the NSCS and is not detected only at a few stations near the Pearl River (PR) estuary. These patterns are mainly regulated by alternative limitation of nutrients and light from the surface to the bottom of euphotic layer. For the pattern 1 (e.g. station 1), light is not a limited factor, and Chl-a and nutrients increase with depth. The pattern 2 (e.g. station 5) exists with the limitation of surface nutrients in offshore region. The nutrients increases with depth and the nutrients limitation turns to light limitation gradually from surface to bottom. And the SCM appears in the layer which need of the light and nutrients is roughly equivalent. Compared with that the offshore SCM, the nutrients for the pattern 3 (e.g. station 14) are rich on the surface with nutrients concentration and light irradiance. Therefore, it is seawater intrusion from the bottom that brings the higher nutrients concentration. The reason for the high Chl-a(0) on the pattern 4 (e.g. station 28) is terrestrial matter from the nearshore. High correlation (R2 = 0.5206, p<0.01) between the depth of SCM (Depth(m)) and Chl-a(0) indicates that the SCM depth is regulated by light masking effect of surface phytoplankton, generally with shallow nutriclines and fast light attenuation for high Chl-a(0) and vice versa low Chl-a(0) brings deeper nutriclines and light attenuate slowly with less shading effect. Further research results shows that Chl-a(int) and Chl-a(m) have a good correlation(R2 = 0.6397, p<0.01). However, the correlation between Chl-a(int) and Chl-a(0) is relative weak (R2 = 0.3202, p<0.01). That could be attributed to the availability of nutrients playing an important role in growth of phytoplankton, with high nutrients at upper euphotic layers for the stations with high Chl-a(0).

18S rRNA gene sequences of leptocephalus gut contents, particulate organic matter, and biological oceanographic conditions in the western North Pacific

Eel larvae apparently feed on marine snow, but many aspects of their feeding ecology remain unknown. The eukaryotic 18S rRNA gene sequence compositions in the gut contents of four taxa of anguilliform eel larvae were compared with the sequence compositions of vertically sampled seawater particulate organic matter (POM) in the oligotrophic western North Pacific Ocean. Both gut contents and POM were mainly composed of dinoflagellates as well as other phytoplankton (cryptophytes and diatoms) and zooplankton (ciliophoran and copepod) sequences. Gut contents also contained cryptophyte and ciliophoran genera and a few other taxa. Dinoflagellates (family Gymnodiniaceae) may be an important food source and these phytoplankton were predominant in gut contents and POM as evidenced by DNA analysis and phytoplankton cell counting. The compositions of the gut contents were not specific to the species of eel larvae or the different sampling areas, and they were most similar to POM at the chlorophyll maximum in the upper part of the thermocline (mean depth: 112 m). Our results are consistent with eel larvae feeding on marine snow at a low trophic level, and feeding may frequently occur in the chlorophyll maximum in the western North Pacific.