the beer brand made by the distributor La Domadora y el León
which is based in Frigiliana on the eastern side of Malaga province
travelled to the city of Brotas in Sao Paolo state in Brazil
where they brewed the first ‘Brazilian-style’ version of their Tropical Pale Ale
a master brewer who has family connections with Frigiliana as his grandmother was born there
Marcio and Javier first met in 2014 through their connections with the brewing industry and now the link has come full circle with Javier and Charo's visit to Brotas
The couple explain that the 'Made in Brazil' La Axarca beer has been brewed using four hops
but three new types of barley ave been added to maintain "the colour
aroma and alcohol and thus get as close as possible to the original beer"
It has been christened #MeGusta and the couple describe the new beer as made with "the same recipe as La Axarca" but "more tropical than ever"
It will be available as a limited edition beer at La Domadora y El León craft beer shop in Frigiliana towards the end of April
Comentar es una ventaja exclusiva para registrados
The leading authority for the Architecture & Design community
Recharge in a Restored Spanish Farmhouse in Menorca
Words: Michael Snyder
Photography: Filippo Bamberghi/Living Inside
When the brothers Humberto and Fernando Campana opened their eponymous design studio Estúdio Campana in São Paulo in 1984, their witty, daring sensibility came as a shock to an architecture scene defined by one of the world’s most robust modernist traditions. Over the last four decades, they’ve invented a language all their own, making furniture with scrap wood and stuffed animals, cast bronze and bubble wrap.
In 2020, the brothers started work on a sprawling, 130-acre park in their rural hometown of Brotas, a 150-mile drive northwest of the city. Since his sibling’s untimely death in 2022 at age 61, Humberto, eight years Fernando’s senior, has thrown himself into that final shared project, slated to soft open in June as a place for conservation and study, but also, like all their work, of provocation and play. As Estúdio Campana looks toward its 40th anniversary, Humberto tells us more.
Humberto Campana: I was very blessed to have been born and raised there because it has such a beautiful landscape. But at the same time, it was extremely boring, so Fernando and I created our own universe. There was a movie theater that screened American westerns and films from Federico Fellini and Pier Paolo Pasolini and we would recreate the scenery in our yard. We made an unconscious vocabulary that we would only discover years later.
We also avoided being contaminated by modernism. Brazilian architecture has such a strong connection to its modern tradition, but Brazil is much more than that. It’s crazy and colorful, full of texture and even kitsch, and we wanted to bring that into our work. We’re maximalists! We should be proud of all the elements of our cultural heritage.
HC: My grandfather had used this property as a coffee plantation. Later, my father rented it out to cattle ranchers. From the time Fernando and I inherited it, we wanted to use it for conservation, but we were nomads, traveling all over the world, so we kept renting because we didn’t want it to sit there abandoned. When the pandemic came, it put us right back in the countryside and we started thinking: We’ve done workshops all over the world, why not in our hometown?
Growing up, it took almost eight hours to get to Sao Paulo on the unpaved roads, and we would see wolves and jaguars and other animals. Nowadays it’s a desert of sugarcane and soy beans. We wanted to seduce people—the families who work in the agri-businesses that are devastating the environment—with poetry, music, and film. We’ve planted over 16,000 trees, and the idea is to plant more, working with agronomists and environmental engineers.
Then we had the idea to create 12 architectural pavilions (there are six, so far) as spaces where people can have classes, meditate, and watch movies and concerts. There will be an educational program, too, both artistic and environmental, and it’s important for us that all the park’s furniture is produced in the countryside with local materials.
I want to create a school to preserve craft traditions with workshops for welding, weaving, painting, embroidery—all the things we used to have in Brotas when I was a child. Across the world, people are finally giving these crafts the respect they deserve. It’s the right moment to invest in the countryside. Life has been so generous to me and, living in a country with such deep social divisions, I feel it’s time to give back.
HC: Right now in the park, all the poetry is there, but none of the logistics, so I’m working with a firm in São Paulo to complete all of that. In the studio, we’re working on a documentary that will launch at the Milan Triennale during the Salone and an exhibition at Friedman Benda in New York. I’m also working on a book about our way of thinking and making. But really, I’m focused on opening the park.
HC: Fernando and I had a wonderful relationship. There was so much trust and respect and intimacy. When I lost him, I felt completely naked, and thought it would be so difficult to keep creating. But I’m actually in a very creative moment right now. Creativity gave me a voice—I came from the countryside, I was supposed to be no one—and now it’s helping me to survive. The park is a memorial, an homage. All the energy I’m investing in it—it’s for him.
ESTÚDIO PLANTAR IDEIAS; LICURÍ PAISAGISMO: LANDSCAPE ARCHITECTURE.
Our Standards: The Thomson Reuters Trust Principles., opens new tab
, opens new tab Browse an unrivalled portfolio of real-time and historical market data and insights from worldwide sources and experts.
, opens new tabScreen for heightened risk individual and entities globally to help uncover hidden risks in business relationships and human networks.
© 2025 Reuters. All rights reserved
Volume 10 - 2023 | https://doi.org/10.3389/fmars.2023.1229692
This article is part of the Research TopicThe Atlantic Meridional Transect Programme (1995-2023)View all 12 articles
The Atlantic Meridional Transect (AMT) program (www.amt-uk.org) provides the perfect opportunity to observe the phytoplankton community size structure over a long latitudinal transect 50oN to 50oS
thereby covering the most important latitude-related basin-scale environmental gradients of the Atlantic Ocean
This work presents cell abundance data of phytoplankton taxa recently collected during cruises AMT28 and 29 (in 2018 and 2019
respectively) using flow cytometer and microscope observations
as well as the pigment composition of the community
to assess the abundance and spatial distribution of taxonomic groups across the Atlantic
The community size structure showed a clear consistency between cruises at large spatial scale
with a dominance of picoplanktonic Cyanobacteria in oceanic gyres
an increase in all groups in the equatorial upwelling region
and high biomass of microplankton size class at higher latitudes
Phytoplankton carbon biomass for oceanographic provinces
ranged from median values of 10 to 47 mg Carbon m-3
Satellite images of total chlorophyll a (as a proxy for phytoplankton biomass) as well as the relative contribution of the three phytoplankton size classes were produced for both cruises
statistically agreed well with in situ size classes estimated as carbon biomass
constituting the first attempt in the literature to match satellite size classes with in situ data derived from cell abundance
The comparison of community structure between recent cruises (2019
2015) and earlier ones (1995-1998) indicates a decrease in the number of diatom-bloom forming species
whereas nitrogen-fixing Trichodesmium abundance in tropical Atlantic remains constant
a relative increase in the median values of picoplankton fraction was seen in SATL region
counterbalanced by a decrease in both nano- and microplankton fractions
this study includes a database of species identified by microscopy
providing a basis for long-term series of phytoplankton species
to understand whether such community shifts and diversity changes are due to natural response to environmental drivers or due to changes arising from climate change
Databases of long temporal series are vital to address these questions
as multiple sets of observations of species variability over a range of different time and space scales could contribute to our better understanding of species dynamics and characterisation of deviations
diazotrophs (filamentous Cyanobacteria and symbiontic diatoms)
heterotrophic dinoflagellates and mixed nanoeukaryotes
coupled with in situ observations are essential
who found that the area of the north and south Atlantic gyres are expanding
This work aims to contribute to the following main goals of the AMT programme (stated in Rees et al., 2017): i) “building multi-decadal ocean time series” and ii) “quantifying the nature of ecological and biological variability in planktonic systems”
The specific objectives of this paper are: 1) to characterise spatial patterns of biomass and community composition variability on a ~50°N to ~50°S transect on the Atlantic in two recent cruises
applying complementary in situ and satellite observations
focusing on biomass estimation (converted into carbon) of the phytoplankton taxonomic groups; and 2) to investigate and compare the most abundant phytoplankton species (within diatoms
filamentous Cyanobacteria) in earlier AMT cruises from the late 1990s with those from recent years (2015
This work comprises data collected on two AMT cruises: AMT28, on board RRS James Clark Ross, which took place from 23rd September to 30th October 2018, and AMT29 onboard RRS Discovery, from 13th October to 25th November 2019. The full reports of the cruises are available at https://www.amt-uk.org/Cruises
Seawater samples were taken from 24 x 20 litre OTE (Ocean Test Equipment) CTD bottles mounted on a stainless steel rosette frame and a Seabird CTD system
The samples were taken from the predawn and noon CTD casts
Chla-Fluorescence was obtained from the CTD sensor Wet Labs ECO-AFL/FL Fluorometer
Samples for Flow Cytometer analyses of phytoplankton groups and HPLC analyses of the concentration of photosynthetic pigments were taken at the surface for 63 stations in AMT28 and 52 in AMT29
Nutrient samples were collected at every depth from each CTD cast according to GO-SHIP protocols
For microscopic enumeration of phytoplankton
surface samples were taken from a water depth between 2-7m for both cruises at all sampling stations
Deep Chlorophyll Maximum (DCM) samples for microscopy were also taken at 19 stations for AMT28
located in northern latitudes in the North and South
and in the area of the Equatorial upwelling
The DCM was assessed during sampling by observing the profile of Chla Fluorescence obtained through the CTD
water samples taken at each CTD cast were sub-sampled into clean (acid-washed) 60 mL HDPE (Nalgene) sample bottles
which were rinsed three times with sample seawater prior to filling and capping
samples were analysed on the ship as soon as possible after sampling and were not stored or preserved
samples were analysed at Plymouth Marine Laboratory (PML)
Samples were kept at -20o C during the cruise
For both cruises Micro-molar nutrient analysis was carried out using a four-channel SEAL analytical AAIII segmented flow nutrient auto-analyser
The colourimetric analysis methods used were: Nitrate (Brewer and Riley, 1965, modified), Nitrite (Grasshoff, 1976), and Phosphate and Silicate (Kirkwood, 1989). The detection limit for Silicate was 0.02 μm and for the three others was 0.01μm. Sample handling and protocols were carried out where possible according to GO-SHIP protocols (Becker et al., 2020)
For each site, a sample of 200 mL was put in amber glass bottles and fixed with acid Lugol’s iodine solution (2% final concentration). In the laboratory, observations were carried out with a Zeiss Axiovert 200 inverted microscope with 10x40 magnification. Cells > 10 µm were counted in a 50 mL chamber, following the Utermöhl method (Utermöhl, 1958)
aiming to count at least a total number of 400 cells; when this value was not reached
Species from the following divisions/classes were identified and counted: Bacillariophyceae (diatoms)
separated into Autotrophic (ADinos) and Heterotrophic dinoflagellates (HDinos)
For Trichodesmium spp (filamentous Cyanobacteria)
the number of cells was estimated for each filament
giving an average number of 100 cells per filament
These groups were defined according to work carried out on AMTs 1 to 10, from which a database of taxa identified by microscopy was published (Sal et al., 2013), and restarted in AMT25 by Brotas et al. (2022)
Coccolithophore species > 10 µm were not assessed by microscopy, as this community was the object of another paper in this special issue (Guerreiro et al., 2023)
coccolithophore species were only assessed by Flow Cytometry
Taxonomic names were checked against the World Register of Marine Species (WoRMS)
Flow cytometery targets the smaller component of phytoplankton cells
from ESD of < 1 µm up to ca 10 µm
whereas microscopy may identify cells with ESD ≥ 10 µm
which has the values of pg Carbon per cell for all microplanktonic species identified)
For taxa measured by flow cytometry, cell numbers of Prochlorococcus and Synechococcus were converted to Cell Carbon by applying conversion factors of 32 and 110 fg Carbon Cell-1, respectively, following Tarran et al. (2001). Cell Carbon content factors for PEUK, NEUK, CRYPTO, and COCCOS were 0.44, 3.53, 23.66 and 33.37 pg Carbon Cell-1, respectively (Tarran et al., 2006)
In order to analyse the concentration in photosynthetic pigments
a volume of 2 to 6 L of seawater was vacuum-filtered through 25 mm diameter GF/F filters onboard
Filters were folded into 2 mL cryovials and stored immediately in the -80oC freezer
Phytoplankton pigments were determined later in the laboratory using High Performance Liquid Chromatography (HPLC). The chromatographic separation of pigments is based on the C8 method developed by Zapata et al. (2000), and adapted by Mendes et al. (2007)
This method is based on a reverse-phase and a pyridine-containing mobile phase
Filters are extracted with 3 ml of 95% cold-buffered methanol (2% ammonium acetate) in a polypropylene test tube
trans-β-apo-8’-carotenal at a concentration of 0.005 mg L-1
for posterior correction of pigment concentrations
The equipment used is a Shimadzu Prominence – I Plus® Lc 2030C 3D Plus
with a monomeric octysilica (OS) C8 column Symmetry® (150mm x 4.6mm dimensions
335 m2 g-1 surface area and 12% carbon load)
The solvent gradient has a flow rate of 1 mL min-1 and a run of 40 min; the sample injection volume is 100 µL
The composition of the solvents is the following: A – Methanol: Aqueous pyridine: Acetonitrile (50:25:25 v/v/v) and B - Methanol: Acetonitrile: Acetone (20:60:20v/v/v)
The HPLC was calibrated using a suite of standards purchased from DHI (Denmark)
Pigments were identified based on retention time and spectral match using a photo-diode array
The three size classes’ abundance was estimated according to Uitz et al. (2006). The weight of each diagnostic pigment followed Brewin et al. (2017), who investigated the pigment ratios and the influence of ambient light on a vast in situ dataset in the Atlantic, updating the initial coefficients proposed by Uitz et al. (2006)
the weighted sum of all diagnostic pigments
equivalent to total Chlorophyll a was computed from the expression C= ∑ WiPi
chlorophyll b + divinyl chlorophyll b and zeaxanthin}
Microplankton class was determined by the sum of fucoxanthin and peridinin (diagnostic pigments diatoms and dinoflagellates
and a non-differentiated assemblage of nano-eukaryotes (represented by 19’hexanoyloxyfucoxanthin
and chlorophyll b + divinyl chlorophyll b and zeaxanthin were used to estimate picoplankton fraction (dominated by the small Cyanobacteria Prochlorococcus and Synechoccocus)
The filamentous Cyanobacteria (which contain the cyanobacteria diagnostic pigments) did not fit into the size class approach
they should be considered microplankton and not picoplankton
Aerosol optical thickness (AOT) at 550nm was obtained from the NOAA Suomi VIIRS sensor at daily
Composite images were produced as simple averages of all daily data for the duration of each cruise
Size class specific Chla was computed using the daily Chla and SST data
Daily matchups were obtained between the satellite results and contemporaneous in situ sampling of biomass for AMT28 and AMT29
Only results from surface samples were considered (2 to 7 m)
Table 1 Dates for the AMT cruises considered for comparison of species
number of stations with microscopy observations
Regression analyses were performed of the matchups of satellite-derived size class Chla and the in situ carbon biomass of the three size classes
estimated from the cellular carbon content described above
A matchup was between the in situ data and the satellite retrievals for the same location and the same day; obviously
these were limited by cloud cover masking the sea-surface from the satellite sensor
Microplankton was computed as the sum of diatoms and dinoflagellates
nanoplankton as the sum of coccolithophores
and picoplankton the sum of Prochlorococcus
There were a total of 15 matchups for AMT28 and 17 for AMT29
All Chla data were logarithm 10 transformed prior to analysis
All these analyses and routines were performed using tools found in PRIMER® 6 software package (PRIMER-E
A principal component analysis (PCA) was performed to understand how environmental patterns (Chla, SST, salinity, macronutrients, depth of the nutricline) shaped the surface distribution of taxonomic groups, using data from both cruises (n=111). Data were standardized (value-average divided by standard deviation) prior to running the analysis. The PCA was run using the sklearn module in python 3.8.8 (Pedregosa et al., 2011)
The tracks of AMT28 and AMT29 across the Atlantic Ocean, plotted over cruise-averaged images of Chla, SST and AOT) are shown in Figure 2
SST showed the expected latitudinal distribution and was very similar between cruises
the one-month lag difference is only noticeable around 40oN
The Chla showed similar patterns for AMT 28 and 29 with low Chla in the oligotrophic gyres in the north and south Atlantic
though with a larger extent on AMT29 reflecting the later timing of this cruise (and
Higher Chla values appeared at higher northerly or southerly latitudes while enhanced values north of the equator are probably associated with equatorial upwelling and the extension offshore to the west of Africa due to eastern boundary upwelling
The AOT 550nm cruise composite data were markedly different between the cruises
with Saharan dust plumes extending offshore between 10°N and 22oN during AMT28 and lower values on AMT29
Figure 2 Composite images of Sea Surface Temperature (SST)
(left): Chla mg m-3 (middle): Aerosol Optical Thickness (AOT)
(right) for AMT28 (top) and AMT29 (bottom)
Figure 3 shows the nutrient concentration between the surface and 200m depth along the transects. The Nutricline was plotted as the isoline of NOx = 1μmol L-1, following Moore et al. (2009)
The North-South nutrient distribution followed the same general pattern in both cruises
with higher concentrations reaching the surface at the higher latitude regions of both hemispheres (from 40°-50°N and 40°-50°S) but more notably in the South Atlantic
The highest concentrations of all macronutrients were observed at greater water depths between 18°N-10°S
where the nutricline was shallower (reaching its shallowest position to the north of the equator ~40 m water depth at 5-15°N) and at the equator (70-80 m water depth)
At the higher-latitude regions during both AMTs
enhanced mixing-induced nutrient supply to the surface was more notable in the southern hemisphere
Figure 3 Concentration of nitrate+nitrite
phosphate and silicate for AMT28 (Left) and AMT29 (Right)
The isoline of 1 µmol L-1 NOx is plotted in all graphs
Phytoplankton relative composition for AMT28 and AMT29 for ten groups, along with the concentration of Chla is illustrated in Figure 4
Biomass (derived from cell concentration) was expressed as mg Carbon m-3
to allow for a comparison of the various groups
it should be highlighted here that values of carbon content per cell are strongly dependent on the cell biovolume
Cell biovolume ranged from 0.11 (Prochlorococcus) to 1.23 x 106 μm3 (the large diatom Rhizosolenia styliformis) hence
cell abundance was determined by microscopy
whereas for coccolithophores < 10 μm
Prochlorococcus and Synechococcus it was determined by Flow Cytometry
had fewer sampling stations in the South Atlantic Gyre (SATL) province and none in South Atlantic Subtropical Convergence (SSTC)
Figure 4 Relative abundance of major phytoplankton groups in surface samples
dinoflagellates and filamentous Cyanobacteria) and estimated in mg Carbon m-3
The correspondence between groups and size classes is given by the colour code: Picoplankton – blue tones
Note that Chla in the 1st station of AMT28 is not plotted as the value was 2.89 mg m-3
Note that AMT29 was sampled only until 35oS
and species from the classes Prasinophyceae and Euglenophyceae
but with very low biomass values across the transect
Coccolithophores abundance only accounted for the cells enumerated by flow cytometry; hence
the biomass contribution of this group in the figure is underestimated
The spatial distribution of the different phytoplankton groups in AMT28 and AMT29 presented the same general pattern with high abundance of diatoms and dinoflagellates (higher than 103 cells L-1) in higher latitudes
and picoplankton carbon biomass (comprising PEUK
Synechococcus and Prochlorococcus) over 40% of total biomass from 20oN to 40oS on both cruises
a clear increase in Chla was noticeable in the region around the Equator (where nutrient concentrations increased within the photic zone
The dominance of picoplankton is clear in the ocean gyres
where Prochlorococcus is the most abundant taxa in the majority of the stations
and it was replaced by Synechococcus when the NOx values were above ~0.1 μmol L-1; PEUK had similar distribution and abundance to Synecoccocus
HDinos and NEUK were the least important groups in terms of carbon contributions
and did not show a clear meridional distribution
occurring both in the oligotrophic gyres where Prochlorococcus dominated
but also in the more productive regions where diatoms and ADinos were more abundant
There are differences in the relative abundance between the two AMTs
as whilst diatoms are more abundant in AMT28
both ADinos and HDinos presented clearly higher biomass in AMT29
with ADinos being the dominant taxa north of 20oN
Another major difference was the occurrence of the bloom of the autotrophic dinoflagellate Prorocentrum cordatum
contributing to a Chla value of 2.83 mg m-3
located in the northern station of AMT28 (49.63oN)
Non-heterocystous filamentous Cyanobacteria included two species: Trichodesmium erythraeum and Oscillatoria limosa, the former was largely more abundant, appearing in both cruises in the Tropical region north of the Equator, on AMT28, from 22°N to 7oN, and on AMT29, from 5°N to 25oN, in the surface samples, but also at the DCM (Figure 5)
Figure 5 Surface and DCM carbon biomass (mg Carbon m-3) of microplankton taxa plotted over Chla values (measured with CTD fluorometer)
The biomass results (in mg Carbon m-3) obtained for diatoms, ADinos, HDinos and filamentous Cyanobacteria, for the stations sampled in surface and DCM, are plotted in Figure 5
superimposed on the integrated Chla profile (measured by CTD fluorescence) down to 200m depth
and showed that biomass of each group was higher at the surface than at the DCM
which allowed for a more detailed analysis of the more abundant species in surface samples only
The common patterns between the two cruises were: i) the overall distribution of Chla
with highest values in higher Northern and Southern latitudes
and an increase in the equatorial region (10oN to 10oS): ii) in higher latitudes
both micro-and nanoplankton contributed to the high Chla values
within the region between 15°N to 10°S
the several small peaks observed are mostly due to an increase in the nanoplankton fraction
iii) the median values in both cruises were almost the same: 0.092 and 0.091 mg Chla m-3 for AMT28 and AMT29
Figure 6 Distribution of TChla (mg m-3) and Chla for each size class: pico-
nano- and microplankton determined by pigment’s concentration for surface stations
The main difference was the high Chla value in AMT28 at 49oN
due to the Prorocentrum cordatum bloom in the English Channel
The taxonomical structure of the community determined by cell enumeration and expressed as carbon biomass was fitted into the three size-class classification as described above. Table 2 discriminates the median values for TCarbon
and the relative distribution of the total community by size classes through the oceanic provinces determined as above
the overall median for both cruises gave the same result: 17 mg Carbon m-3
The results evidence the spatial distribution of phytoplankton community along the North-South track in the Atlantic
and above 18 in North Atlantic Drift (NADR)
The highest median value of 47 mg Carbon m-3 was observed for the SATL region in AMT28
where microplankton carbon biomass fraction reached almost 70%; we split the vast SATL region into two areas
separating the gyre from the more productive southern SATL
The minimum values observed in both cruises were similar: 8 mg Carbon m-3 for AMT28 at 12°S and 7 mg Carbon m-3 for AMT29 at 15°S
and median values of the percentage of micro- nano- and pico-plankton for each biogeographic oceanic province
Using satellite imagery, the following products were derived: the relative contribution of micro-, nano- and picoplankton, as well the contribution of diatoms and dinoflagellates, according to Brewin et al. (2017). Cruise composite images of Chla (mg m-3) for each size class, for the total period of the cruises are displayed in Figure 7
The large-scale nutrient distribution affects all three size classes
as all classes present higher abundance in high-latitude regions and lower in oligotrophic gyres
the response of cells to nutrient enrichment scales up according to cell size
Picoplankton was present in all latitudinal regions
where micro- and nanoplankton attain minimum values
Nanoplankton taxa biomass increased in tropical Atlantic and higher latitudes
whereas Microplankton fraction presented high values only in high northern and southern latitudes
Figure 7 Concentration of phytoplankton size classes
Composite image of Chla (mg m-3) for each size class
The number of daily matchups between satellite and in situ data found was 15 for AMT28 and 17 for AMT29, so the comparison analysis was made on samples of the two cruises together. For the in situ, the total carbon of the three size classes was computed as described above. The results in Table 3 showed significant correlation values (r2) for the three size classes
as well as when considering diatoms and dinoflagellates separated
Table 3 Results of linear regression Type II in log-transformed data of satellite products (Brewin et al., 2017) and in situ carbon
Nano- and Picoplankton and for the diatoms and dinoflagellates
A total of 30 diatom taxa, 156 dinoflagellate taxa and three filamentous Cyanobacteria taxa were identified during AMT28 and AMT29 (see Supplementary Material, Table 1). Our taxonomic criteria followed as much as possible those presented by Sal et al. (2013)
in order to compare phytoplankton species occurrence and abundance with previous AMT studies
Data from diatoms, dinoflagellates and filamentous Cyanobacteria obtained during AMT28, AMT29 presented here were compared to data from AMT25 (Brotas et al., 2022) and from the earlier AMT1, 3, 5, 7, all of which also took place during boreal Autumn. While all cruises followed the same general meridional latitudinal track, AMT25 and AMT28 extended further south compared to the other cruises (Figure 1)
significant track differences occurred in the northern part of NATR
with AMT25 and AMT29 being ca 20o displaced to the east compared to the other track cruises
and thereby following a more open-ocean route
The present analysis is focused only on the most abundant species in these seven cruises
only species which “bloomed” (where a bloom was considered when the cell concentration was higher than 104 cells L-1); it should be noted that for Trichodesmium spp
the bloom was defined not by the number of filaments
and were also frequently observed in our samples
Table 4 Species/Taxa which presented abundances > 104 cell L-1
Cluster analysis, with the SIMPROF test (Figure 8)
which were statistically distinct (at the 5% level)
SIMPER analysis determined the taxon or group of taxa that mostly contributed to each group’s average similarity
The largest group (D) clustered the tropical Atlantic (NATR and WTRA) from AMT1
being defined by the presence of Trichodesmium blooms
Group C’s average similarity was mainly due to Athecate dinoflagellates
and cluster latitudinal regions where up to 6 different species of diatoms were observed
Group A was defined by the Prorocentrum cordatum blooms that occurred twice in the NADR region in AMT28
and at an ad-hoc station of AMT25; both blooms occurred in the English Channel
with this species attaining 99 % of the total cell abundance of the microplankton fraction
group F contained two blooms that share no similarities with other assemblages
namely Gymnodinium galeaeforme (AMT1) and Pennate diatoms (> 40µm)
Figure 8 Dendrogram for hierarchical clustering of phytoplankton abundance samples using group-average linking of Bray–Curtis similarities
Only samples which presented abundances > 104 cell L-1
in the different Longhurst regions were included and data
Red dotted lines indicate nonsignificant clustering (SIMPROF test; significance level: P < 5%)
in the North Atlantic Subtropical Gyre (NAST)
which comprise a significant part of the North and South Atlantic oligotrophic gyres
except for Athecate dinoflagellates and “Pennate group (10-30 μm)” in AMT3
The large-scale relative abundance of phytoplankton groups in Longhurst biogeographic provinces, was comparable, in terms of the dominant groups for each region. The comparison of the median values of TCarbon over the provinces gave very similar results for AMT28, AMT29, and AMT25 (Brotas et al., 2022, their Table 3)
Ocean gyres presented median values of 10-15 mg Carbon m-3
whereas higher latitudes registered higher variability
and median values between 25-47 mg Carbon m-3
We used three complementary methodological approaches to study phytoplankton composition
photosynthetic pigments and cell enumeration
and the third using satellite observations
All three approaches showed a similar pattern for the spatial distribution of phytoplankton community composition in the Atlantic regions for the two cruises
suggesting that environmental conditions of each region play a major role in shaping the relative composition of phytoplankton communities
The group ADinos appeared scattered throughout the space defined by both axes
with values on the right end of axis 1 linked with high Chla and high nutrient values
the Bray-Curtis analysis focused only on microplankton species
clustering the same regions from all the AMT cruises
rather than clustering the earlier AMTs versus the recent ones
the phytoplankton community composition is shaped by the strong gradients of the environmental conditions
Figure 9 Principal Component Analysis performed on the phytoplankton taxonomic over the environmental data
The colour code represents the dominant phytoplankton taxon by carbon biomass
This may be related to the fact that dust deposition was occasionally slightly higher during AMT29 compared to AMT28
as indicated by in-situ measurements of Mn concentrations (pmol m-3) measured from dust samples collected in situ during the cruises (data not shown
The increase of Autotrophic dinoflagellates carbon biomass in AMT29
It can be due to the natural community seasonal evolution
with a higher abundance of dinoflagellates
or it could be an indicator of an effective shift in the community occurring independent of the seasonality
when we compare the overall median values of dinoflagellates in AMT25
this aspect should be investigated in future AMTs
as more data are needed to evaluate whether there is a consistent shift or not
specifying that dinoflagellates can be major grazers of diatoms
where the maximum cell number (3.5 x 104 cells L-1) was observed in NADR
It should be noted that the dinoflagellates satellite product relies on the assumption that peridinin is the diagnostic pigment for this taxonomic class
and although peridinin is only present in dinoflagellate species
whereas the abundance of nano- and microplankton fractions is strongly dependent on the nutrient availability
Variability of phytoplankton species is an inherent and well-known characteristic of marine ecosystems, however assessing phytoplankton variability patterns and the way communities respond to changes, in terms of shift between taxa, cell size, or trophic strategy, is still a challenging research subject. As pointed out by Behrenfeld and Boss (2014)
factors controlling phytoplankton blooms remain controversial despite a century of investigation
results showed that blooms were mainly produced by diatom species
and that blooms of 105 or 106 cells L-1 were only observed in earlier AMTs
suggesting conditions which would favour diversity
from the total of 20 diatom species producing a bloom identified for the seven AMT cruises
only three taxa were observed in the recent ones (2015-2019)
belonging to the pennate-colonial species type
constituting as well a decrease in shape diversity in relation to the earlier AMT cruises species)
who stated that the toxic species Prorocentrum cordatum has been reported to become more widespread in recent years; hence these results constitute an alert for future studies in the English Channel
Concerning filamentous Cyanobacteria, the abundances of Trichodesmium spp seem to be remarkably similar. The maxima values of Trichodesmium spp were 960 filaments in AMT 28 (11.45oN; -27.71oW), and 720 in AMT29 (11.85oN, 27.96oW). Tyrrell et al. (2003), in AMTs 1,3, 5 and 7 reported high filament abundances in the region between 0° and ~15oN, with a maximum value of 1040 filaments of Trichodesmium spp. for AMT 5. In AMT25, Brotas et al. (2022)
and the presence of filamentous Cyanobacteria from 23°N to 2oN
not only the presence but also the abundance levels of filamentous Cyanobacteria seem to be a common feature in this area
as well as in the earlier Boreal Autumn AMTs (1
This work provides observations on the abundance and diversity of phytoplankton taxa on a gradient of trophic conditions in the North and South Atlantic
Our data contribute to the main goal of the AMT program
regarding the regular and long-term acquisition of a series of observations
crucial to understand the response of biological communities to ongoing climate change
whilst providing data for the development and testing of more accurate prediction models for future climatic scenarios
The results obtained showed a common pattern from both cruises of phytoplankton biomass distribution and relative dominance of groups
across the Atlantic: the biomass values for the different biogeographic oceanic provinces were similar in both cruises
Interannual major differences were found in high latitudes
where blooms of diatoms and dinoflagellates appeared
This work explored complementary approaches to study phytoplankton size classes
namely in situ determination of size classes by chemotaxonomy
in situ enumeration of cell abundance of all taxonomic groups
and remote sensing of the relative contribution of the three size classes to Chla
We found a good relationship between satellite determination of phytoplankton size classes with in situ size classes estimated as carbon biomass
albeit with a relatively small number of available matchups
An increase in the picoplankton fraction was detected in southern Atlantic, along AMT25, 28 and 29, consistent with earlier reports (Racault et al., 2014; Agirbas et al., 2015)
In spite of the differences in route tracks between the different AMTs
the analysis of the more abundant microplankton species of diatoms
dinoflagellates and filamentous Cyanobacteria between AMT cruises from the 1990s and more recent expeditions
which might be considered for future studies
namely i) Trichodesmium blooms are a common feature in earlier and recent AMTs
ii) the small Athecate Dinoflagellate category is also a common taxa throughout the different AMTs
iii) there is a possible indication for a shift in most abundant diatom species
where the nutrient concentrations are high
iv) the Prorocentrum cordatum dinoflagellate blooms in recent cruises may be in line with the widespread of toxic species
Additionally, we produced a database of cell carbon content per species, and its abundance in the sampled Atlantic regions, extending the database from the late 1990s, published by Sal et al. (2013)
leaving a record for future research on biodiversity
The original contributions presented in the study are included in the article/Supplementary Material
Further inquiries can be directed to the corresponding author
All authors contributed to the conceptual development of the paper
Authors contributed differently to the following aspects: cruise participation (GT
microscope identification and enumeration (VV)
All authors contributed to manuscript revision
AT benefited a grant from European Union’s (EU) Horizon 2020 CERTO (Copernicus Evolution: Research for harmonised Transitional water Observation) project (No
We thank the officers and crew of the British Antarctic Survey vessel RRS James Clark Ross and RRS Discovery
We are indebted to NEODAAS for processing the satellite imagery
We thank Carolina Sá and Giulia Sent for their help in AMT29
We thank the three reviewers for valuable suggestions
The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest
All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations
Any product that may be evaluated in this article
or claim that may be made by its manufacturer
is not guaranteed or endorsed by the publisher
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmars.2023.1229692/full#supplementary-material
Supplementary Table 1 | Supplementary material
the cell biovolume (µm3) and cell carbon content (pg Carbon Cell-1)
Supplementary Table 2 | Supplementary material
Supplementary Table 3 | Supplementary material
Temporal changes in total and size-fractioned chlorophyll-a in surface waters of three provinces in the Atlantic Ocean (September to November) between 2003 and 2010
A synthesis of the environmental response of the North and South Atlantic Sub-Tropical Gyres during two decades of AMT
On the roles of cell size and trophic strategy in North Atlantic diatom and dinoflagellate communities Limnol
GO-SHIP repeat hydrography nutrient manual: the precise and accurate determination of dissolved inorganic nutrients in seawater
Resurrecting the ecological underpinnings of ocean plankton blooms
doi: 10.1146/annurev-marine-052913-021325
CrossRef Full Text | Google Scholar
Obtaining phytoplankton diversity from ocean color: A scientific roadmap for future development
An ordination of the upland forest communities of Southern Wisconsin
Google Scholar
The automatic determination of nitrate in seawater
CrossRef Full Text | Google Scholar
Uncertainty in ocean-colour estimates of chlorophyll for phytoplankton groups
CrossRef Full Text | Google Scholar
The influence of the Indian Ocean Dipole on interannual variations in phytoplankton size structure as revealed by Earth Observation
A three-component model of phytoplankton size class for the Atlantic Ocean
Complementary approaches to assess phytoplankton groups and size classes on a long transect in the Atlantic Ocean
Phytoplankton species counts from Atlantic Ocean surface water samples collected during JR15001 (AMT25) in September-November 2015
doi: 10.5285/eb752e17-cd14-3889-e053-6c86abc0ec59
CrossRef Full Text | Google Scholar
MAREDAT: towards a world atlas of MARine Ecosystem DATa
CrossRef Full Text | Google Scholar
Cermeño P.
The role of nutricile depth in regulating the ocean carbon cycle
CrossRef Full Text | Google Scholar
Non-parametric multivariate analyses of changes in community structure
CrossRef Full Text | Google Scholar
Testing of null hypotheses in exploratory community analyses: similarity profiles and biota-environment linkage
CrossRef Full Text | Google Scholar
PubMed Abstract | CrossRef Full Text | Google Scholar
Primary production of the biosphere: integrating terrestrial and oceanic components
A practical strategy for analysing multispecies distribution patterns
Google Scholar
and pumps to microbial multitasking: Evolving concepts of marine microbial ecology
The current configuration of the OSTIA system for operational production of foundation sea surface temperature and ice concentration analyses
Methods of Sea-water Analysis (Weiheim: Verlag Chemie)
Google Scholar
Response of coccolithophore communities to oceanographic and atmospheric processes across the North- and Equatorial Atlantic
Synoptic relationships between surface Chlorophyll-a and diagnostic pigments specific to phytoplankton functional types
“Phytoplankton functional types from space,” in Reports of the International Ocean Color Coordinating Group No
Google Scholar
Are phytoplankton blooms occurring earlier in the Arctic
doi: 10.1111/j.1365-2486.2010.02312.x
CrossRef Full Text | Google Scholar
Khanaychenko A
Bloom-forming potentially toxic dinoflagellates Prorocentrum cordatum in marine plankton food webs
CrossRef Full Text | Google Scholar
Simultaneous determination of selected nutrients in seawater
Google Scholar
Le Quéré C.
Ecosystem dynamics based on plankton functional types for global ocean biogeochemistry models
Menden-Deuer S.
Carbon to volume relationships for dinoflagellates
CrossRef Full Text | Google Scholar
HPLC determination of microalgae pigments: comparing resolution and sensitivity of a C18 and a C8 method
CrossRef Full Text | Google Scholar
Large-scale distribution of Atlantic nitrogen fixation controlled by iron availability
Remote sensing of phytoplankton functional types
Google Scholar
Carbon sequestration in the deep Atlantic enhanced by Saharan dust
CrossRef Full Text | Google Scholar
Google Scholar
Phongphattarawat S.
Ecophysiological basis of spatiotemporal patterns in picophytoplankton pigments in the global ocean
Coccolithophore ecology in the tropical and subtropical Atlantic Ocean: New perspectives from the Atlantic meridional transect (AMT)
Plankton indicators and ocean observing systems: support to the marine ecosystem state assessment
The atlantic meridional transect programme
Ecology of Phytoplankton (Cambridge: Cambridge University Press)
Google Scholar
Towards a functional classification of the freshwater phytoplankton
CrossRef Full Text | Google Scholar
CrossRef Full Text | Google Scholar
Sathyendranath S.
An ocean-colour time series for use in climate studies: the experience of the ocean-colour climate change initiative (OC-CCI)
Sathyendranath S.
Copernicus marine service ocean state report
CrossRef Full Text | Google Scholar
Sathyendranath S.
Reconciling models of primary production and photoacclimation
Pelagic ecosystem structure: heterotrophic compartments of the plankton and their relationship to plankton size fractions
Google Scholar
Determining Atlantic Ocean province contrasts and variations
CrossRef Full Text | Google Scholar
Coupling ecological concepts with an ocean-colour model: Phytoplankton size structure
Latitudinal changes in the standing stocks of nano- and picoeukaryotic phytoplankton in the Atlantic Ocean
Microbial community structure and standing stocks in the NE Atlantic in June and July of 1996
Large-scale latitudinal distribution of Trichodesmium spp
Vertical distribution of phytoplankton communities in open ocean: An assessment based on surface chlorophyll
CrossRef Full Text | Google Scholar
Utermöhl H
Zur Vervolkommung der quantitativen Phytoplankton: methodik
Mitteilungen Internationale Vereinigung f̈r Theoretische und Angewandte
Google Scholar
Phytoplankton pigment distribution in relation to upper thermocline in the eastern Mediterranean Sea during winter
Rethinking the marine carbon cycle: factoring in the multifarious lifestyles of microbes
Satellite monitoring of surface phytoplankton functional types in the Atlantic Ocean over 20 years (2002–2021)
Separation of chlorophylls and carotenoids from marine phytoplankton: a new HPLC method using a reversed phase C8 column and pyridine-containing mobile phases
Clewley D and Groom SB (2023) Assessing phytoplankton community composition in the Atlantic Ocean from in situ and satellite observations
Received: 26 May 2023; Accepted: 16 October 2023;Published: 03 November 2023
Copyright © 2023 Brotas, Ferreira, Veloso, Tracana, Guerreiro, Tarran, Woodward, Ribeiro, Netting, Clewley and Groom. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY)
distribution or reproduction in other forums is permitted
provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited
in accordance with accepted academic practice
distribution or reproduction is permitted which does not comply with these terms
*Correspondence: Vanda Brotas, dmJyb3Rhc0BmYy51bC5wdA==
Disclaimer: All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations
Any product that may be evaluated in this article or claim that may be made by its manufacturer is not guaranteed or endorsed by the publisher
94% of researchers rate our articles as excellent or goodLearn more about the work of our research integrity team to safeguard the quality of each article we publish
DSM is buying Amyris Brasil from its long-term partner Amyris for $58 million
The acquisition includes the Brotas 1 production plant in Sao Paulo
and intellectual property related to farnesene
a bio-based key intermediate for many applications
The deal also includes an additional value share arrangement over a three-year period amounting to $37.5 million
which is subject to the usual closing conditions
is expected to complete in the next few months
Brotas 1 produces about 24,000 t/y of farnesene and provides DSM with an abundant availability of sugar cane-based raw materials for its pipeline of bio-based products. DSM said it plans to optimize the site’s operational performance.
The Dutch life sciences group added that it would honor existing supply agreements to both Amyris and other parties, including the supply of specialty compounds to Amyris until it starts up a second facility, Brotas 2, which will produce specialty, high-performance ingredients.
Amyris said the sale will allow it to accelerate the construction of Brotas 2, as well as complete its plant in Sao Martinho to focus on sweeteners. “The combination of these actions provides us the manufacturing footprint to meet our current demand through the next three to five years and to manage this within our funding constraints,” said John Melo, president and CEO of Amyris.
CHEManager Spotlight is an exclusive event tailored for practitioners and decision-makers in the chemical industry. This part of our event series delves into the latest trends and innovations in logistics to streamline your operations and drive efficiency.
CHEManager Innovation Pitch supports innovation in the chemistry and life sciences start-up scene. The platform allows founders, young entrepreneurs, and start-ups to present their companies to the industry.
(Oslo/Florianópolis, 12 September 2023) Statkraft, Europe’s largest generator of renewable energy has purchased 18.69% of the shares in its Brazilian subsidiary Statkraft Energias Renováveis (SKER) from Fundação dos Economiários Federais (Funcef), the third largest pension fund in Brazil. As of today, 100% of SKER is owned by the Statkraft Group.
“With this acquisition, we will be able to bring even more agility to our strategic plan by increasing our ownership in Brazil. I would also like to thank Funcef for the long successful partnership we had along this journey together," says Ingeborg Dårflot, Executive Vice President International in Statkraft.
SKER is in the final stages of constructing the Santa Eugênia Wind Complex, located in Bahia, the Northeast region of the country. The renewable energy production of the complex is expected to reach 2,300 Gigawatt-hours (GWh) per year, enough to power 1.17 million Brazilian households. The complex will comprise 14 wind farms, totaling 91 wind turbines with a capacity of 5.7 megawatts each.
In the state of Bahia, Statkraft is also constructing the greenfield wind project Morro do Cruzeiro, an expansion of the operational Brotas de Macaúbas Wind Complex. The expansion consists of 14 turbines with an installed capacity of 79.8 MW. Given the excellent wind conditions in the region, the project will generate 386 GWh of renewable energy per year, enough to power more than 190,000 households. Both ongoing projects are financed with resources from the Banco do Nordeste do Brasil (BNB).
The company conducts market operations from its trading offices in Rio de Janeiro and Sao Paolo and is a pioneer in offering traceable renewable energy with proof of origin in Brazil, certifying its energy production's sustainability through the I-REC (International Renewable Energy Certificate) program.
Last month, Statkraft announced the signing of an agreement with EDPR to purchase two wind farms in Brazil. This transaction, together with the acquisition of FUNCEF's shares today, is an important part of Statkraft's global growth. Based on projects under construction, in acquisition and operation, Statkraft will soon reach 1.3 GW of installed capacity in Brazil, strengthening its position as an international leader in renewable energy supply.
Recently, Statkraft also obtained approval from Aneel for seven solar projects with a total installed capacity of 228 MW in Bahia. The solar projects are extensions of the wind farms Brotas de Macaúbas Complex (Sol de Brotas 1 plant), Morro do Cruzeiro Complex (Sol de Brotas 2 plant) and Santa Eugênia Winds Complex (Sol de Brotas 3, 4, 5, 6, and 7 plants).
The project complies with Aneel's regulations for hybrid plants and, in Statkraft's case, will leverage the complementary nature of its wind and solar power generating facilities. With the received authorization, the company will begin the implementation strategy for its first solar project developed by the company in Brazil.
About StatkraftStatkraft is a leading company in hydropower internationally and Europe’s largest generator of renewable energy. The Group produces hydropower, wind power, solar power, gas-fired power and supplies district heating. Statkraft is a global company in energy market operations. Statkraft has 5,700 employees in 21 countries.
For more information, please contact: Mariana Aoad, Communication Manager, Statkraft Brazil Mob: +55 21 999647681 E-mail: mariana.aoad@statkraft.com Torbjørn Steen, Vice President External Communications, Statkraft AS Mob: +47 911 66 888 E-post: torbjorn.steen@statkraft.com
ACS’s Basic Package keeps you connected with C&EN and ACS.
$80 Regular Members & Society Affiliates
ACS’s Standard Package lets you stay up to date with C&EN, stay active in ACS, and save.
ACS’s Premium Package gives you full access to C&EN and everything the ACS Community has to offer.
Biobased chemicals maker Amyris has agreed to sell its Brazilian farnesene fermentation plant to Dutch specialties firm DSM for $96 million. The deal includes intellectual property for producing farnesene, a key intermediate chemical made from sugar.
The announcement came three days after Amyris disclosed its earnings for the third quarter. The company booked $24.2 million in revenues from product sales and collaborations, down from $26.5 million in the year-ago quarter. And its quarterly loss ballooned to almost $34 million from $19.7 million.
Founded in 2003 to commercialize science from the lab of University of California, Berkeley, chemical engineer Jay D. Keasling, Amyris bet big on the potential of its fermentation technology. But investors have recently grown inpatient. Its stock price has fallen from a high of close to $80.00 per share in 2013 to less than $4.00 today.
The purchase is the second cash infusion from DSM. In May, the two firms began working together to develop nutritional ingredients such as microbially produced vitamin A. As part of the agreement, DSM invested $50 million in Amyris.
Amyris has operated the farnesene plant in Brotas, Brazil, since 2013. Equipped with six 200,000-L fermenters, the facility was designed to serve large markets. In a conference call with analysts, CEO John Melo said Amyris requires new capacity to manufacture the smaller batches of specialty chemicals desired by its other customers in health, personal care, and flavors and fragrance markets.
Meanwhile, DSM plans to ramp up its own biobased production. “Our know-how in fermentation, downstream process development, and large-scale manufacturing will allow us to further improve the operational performance of the facility,” said Chris Goppelsroeder, president of DSM Nutritional Products.
Sign up for C&EN's must-read weekly newsletter
Cowen & Co. stock analyst Christopher Souther agrees that DSM can likely improve the plant’s operations. “We see the sale as strong evidence of the value of Amyris’s platform and view it as a positive given the company’s strong need for liquidity,” he wrote in a note to investors.
Melo said Amyris is on track to hit $130 million in revenues this year, or double last year’s results. The company plans to manufacture a low-calorie sweetener in 2018 and build a second facility without taking on more debt.
This article has been sent to the following recipient:
Copyright © 2025 American Chemical Society. All Rights Reserved.
WindStatkraft to host WEG 7MW Brazilian wind turbine as pair eye global opportunitiesAs interest grows in diversifying pool of OEMs, WEG also sees potential for harnessing Brazilian and Indian supply chains
Volume 8 - 2021 | https://doi.org/10.3389/fmars.2021.682621
has been assessed at synoptic temporal and spatial scales with satellite remote sensing (RS) for over two decades
RS algorithms to monitor relative size classes abundance are widely used; however
as well as the assessment of phytoplankton structure
the main motivation of this work it to discuss the links between size classes and phytoplankton groups
in order to foster the capability of assessing phytoplankton community structure and phytoplankton size fractionated carbon budgets
photosynthetic pigments concentration and cell numbers per taxa) collected in surface samples along a transect on the Atlantic Ocean
during the 25th Atlantic Meridional Transect cruise (AMT25) between 50° N and 50° S
from nutrient-rich high latitudes to the oligotrophic gyres
We compared phytoplankton size classes from two methodological approaches: (i) using the concentration of diagnostic photosynthetic pigments
and assessing the abundance of the three size classes
and (ii) identifying and enumerating phytoplankton taxa by microscopy or by flow cytometry
and dividing the community into five size classes
The distribution of phytoplankton community in the different oceanographic regions is presented in terms of size classes
and discussed in relation to the environmental oceanographic conditions
The distribution of seven functional types along the transect showed the dominance of picoautotrophs in the Atlantic gyres and high biomass of diatoms and autotrophic dinoflagellates (ADinos) in higher northern and southern latitudes
where larger cells constituted the major component of the biomass
Total carbon ranged from 65 to 4 mg carbon m–3
at latitudes 45° S and 27° N
The pigment and cell carbon approaches gave good consistency for picoplankton and microplankton size classes
but nanoplankton size class was overestimated by the pigment-based approach
The limitation of enumerating methods to accurately resolve cells between 5 and 10 μm might be cause of this mismatch
was fitted to the Chlorophyll a (Chla) data and
to extract the biomass of three size classes of phytoplankton
The general pattern of the model fitted to the carbon data was in accordance with the fits to Chla data
The ratio of the parameter representing the asymptotic maximum biomass gave reasonable values for Carbon:Chla ratios
but with higher values for picoplankton (170) than for combined pico-nanoplankton (36)
The approach may be useful for inferring size-fractionated carbon from Earth Observation
improved understanding of biogeochemical cycles
The CO2 cycle is affected greatly by primary production by plants on land and algae in the oceans
Chlorophyll a (Chla) has traditionally been used as a proxy for phytoplankton biomass and can be assessed by Earth Observation (EO) on synoptic spatial scales and over an increasing timespan (currently
and their occurrence is strongly linked with oceanographic conditions
requiring a better understanding of the correspondence of PSC and PFT
and the possibility of assessment of phytoplankton size structure in terms of Carbon by EO
The present work assesses phytoplankton community size structure using both pigments and carbon concentration (converted from cell enumeration) and applies the Brewin et al. (2010) model to both datasets
making this work the first attempt to apply the model to carbon data: this provides the basis to infer size fractionated carbon from EO data
The main objectives of this study were the following: (i) to analyse and compare PSCs with two methodological approaches: the pigment approach and cell enumeration converted into carbon; (ii) to apply the abundance size class model of Brewin et al. (2010) to a size class abundance based on carbon
(iii) to study the biomass and composition of phytoplankton communities along distinct environmental conditions of the Atlantic Ocean
discussing the links between phytoplankton taxonomic groups
The ultimate goal aims to contribute to strengthening the capability of assessing phytoplankton community structure and phytoplankton carbon budgets by satellite EO
This work was conducted on board the Royal Research Ship James Clark Ross within the AMT programme. The cruise took place from September 15th to November 3rd, 2015, along a track spanning from 50.033° N and 4.375° W to 49.777° S and 54.509° W. Figure 1 plots the cruise track over images of SST (Sea Surface Temperature) and Chla obtained for the whole cruise period
Figure 1. Composite images of Sea Surface Temperature (SST), left, and Chla, right, obtained from September 11, 2015 until November 4, 2015, showing the cruise track with the sampling stations. Chla image was obtained from the OC-CCI portal (www.oceancolour.org)
Seawater samples were taken from 24 × 20 L OTE (Ocean Test Equipment) CTD bottles mounted on a stainless steel rosette frame and a Seabird CTD system
For microscopic and flow cytometric enumeration of phytoplankton
taken from a water depth between 2 and 5 m were analysed
The samples were analysed on the ship as soon as possible after sampling and were not stored or preserved
Micro-molar nutrient analysis was carried out on board using a four channel SEAL analytical AAIII segmented flow nutrient auto-analyser. The colorimetric analysis methods used were: Nitrate (Brewer and Riley, 1965, modified), Nitrite (Grasshoff, 1976), and Phosphate and Silicate (Kirkwood, 1989). Sample handling and protocols were carried out where possible according to GO-SHIP protocols (Becker et al., 2020)
Mixed Layer Depth (MLD) was determined following the temperature criterion (Levitus, 1982)
which defines the mixed layer as the depth at which temperature change from the surface value is 0.5°C
The distinction between autotrophic and heterotrophic dinoflagellate species followed the same database
only coccolithophore species >10 μm were identified and enumerated (due to coccolith degradation in Lugol’s-fixed samples)
Species identifications were checked against species records of AMTs 1
in common with the 25th Atlantic Meridional Transect (AMT25) cruise
Flow cytometer targets the smaller component of phytoplankton cells
from equivalent spherical diameter (ESD) of <1 μm up to ca 10 μm
whereas microscopy may identify cells with ESD ≥10 μm
In order to compare all components of the phytoplankton community
cell abundance of each taxa was converted into carbon content (mg carbon m–3)
It should be noted that the estimation of carbon content through biovolume implies a certain bias in the data
as in larger cells the carbon content is probably overestimated in comparison to smaller cells
where the cellular content is more tightly packed
the total number of cells was estimated by measuring the cell dimension
biovolume and carbon conversion factors were applied
For cells measured by flow cytometry, cell numbers of Prochlorococcus and Synechococcus were converted to cell carbon by applying conversion factors of 32 and 110 fg carbon cell–1, respectively, following Tarran et al. (2001). Cell carbon content factors for PEUK, NEUK, CRYPTO, and COCCOS, were 0.44, 3.53, 23.66, and 33.37 pg carbon cell–1 respectively (Tarran et al., 2006)
Due to a malfunction with the flow cytometer during the first week of the cruise
and COCCOS were counted only from south of 37° N
A volume of 1–4 L of seawater was vacuum-filtered through 25 mm diameter GF/F filters
Filters were folded into 2 mL cryovials and stored immediately in the −80°C freezer
Phytoplankton pigments were determined later in the laboratory using HPLC, following the method of Zapata et al. (2000) with adaptations described in (Brewin et al., 2017a)
Pigments were extracted from the sample filters into 2 mL 90% acetone using an ultrasonic probe (50 W) for 35 s
Extracts were clarified by filtration and analysed by reversed-phase HPLC using a Thermo Accela Series HPLC system with chilled autosampler (4°C) and photodiode array detector
The column used was a Waters C8 Symmetry; 150 × 2.1 mm; 3.5 μm particle size
with a flow rate of 200 μL min–1
A quality control filter was applied to all pigment data, following Aiken et al. (2009)
which uses the relationship of accessory pigments (i.e.
all carotenoids plus chlorophylls b and c) and total Chla (the sum of monovinyl chlorophyll-a
divinyl chlorophyll-a and chlorophyllide a) to accept or eliminate particular samples
after investigating the influence of ambient light on a vast dataset
The fractions were then multiplied by the in situ total chlorophyll-a concentration (TChla) to derive size-specific chlorophyll-a concentrations
refining the factor WfucoPfuco,n with coefficients obtained from a wide pigments database; this factor is subtracted in the microplankton equation and added in the nanoplankton one
This approach was followed in the present work
We used the three size classes defined above for the pigment approach
we divided the whole community into five size classes
in order to account for the large diversity of cell morphology and characteristics of all nanoplankton components
as well as for the smaller diatoms and dinoflagellates
This resulted in splitting the nanoplankton size class into three subcategories
we will identify the size classes as pigment approach size classes or carbon approach size classes
Each species/taxon was allocated to one class
in accordance with the value of carbon content per cell
The five classes were as follows: class 1: <2 pg carbon cell–1; class 2: 2–10 pg carbon cell–1; class 3: 10–50 pg carbon cell–1; class 4: 50–180 pg carbon cell–1; and class 5: >180 pg carbon cell–1
List of the five plankton classes defined according to: Carbon per cell content
and size as equivalent spherical diameter (ESD)
Relation between taxonomic group or entity
and phytoplankton functional groups (PFTs)
The upper limit for picoplankton is not agreed in the literature, varying between 2 μm (Jeffrey et al., 2011) and 3 μm (Buitenhuis et al., 2012), who note that the size range is a source of ambiguity for estimating carbon budget. By establishing 2.7 μm we are near the value indicated by Buitenhuis et al. (2012)
with ESD from 10.3 to 17.5 μm is relevant to estimate the contribution of smaller diatoms and dinoflagellates
Cp,nm and Cpm are the asymptotic maximum values of Chla for the two size classes (<20 μm and <2 μm
The Chla concentration of nanophytoplankton (Cn) and micro-phytoplankton (Cm) are simply calculated as Cn = Cp,n−Cp and Cm = C−Cp,n
Dp,n and Dp reflect the fraction contributed by each size class to TChla as TChla tends to zero
and they should take values in the range between 0 and 1 to ensure size-fractionated Chla does not increase faster than TChla
as the fraction of cells <2 μm must obviously be smaller than the fraction of cells <20 μm
we also fitted the same set of equations to the carbon data
and Dp) are with respect to carbon biomass
The concentrations of nitrate + nitrite, phosphate and silicate, from surface to 200 m depth, as well as the depth of the mixed layer is illustrated in Figure 2
Concentration of nitrate + nitrite and phosphate in the surface were below quantification limits in most stations
MLD was from 50 to 11 m in the Northern hemisphere
reaching values higher than 90 m in the southern hemisphere (from latitudes 30 to 38° S)
with a maximum value of 187 m at 36° S
with the Mixed Layer Depth plotted along the transect
High nutrient values throughout the water column up to the surface were present only south of 40° S
The low concentration of nutrients down to 200 m in the North Atlantic Gyre (NAG) and South Atlantic Gyre (SATL) is clearly visible
Nitrate + nitrite values within the MLD were above quantification limit only in three distinct areas
Values for silicate within the MLD were always above quantification limit
whereas for phosphate values were below quantification limit in NAG but not in SATL
Using the cell carbon content allowed comparison of the biomass contribution of all phytoplankton classes/divisions in the Atlantic transect
considering both the cell counts done by microscopy and by FC
taxa from the following groups were enumerated: bacillaryophyceae (diatoms
coccolithophores > 10 μm (25 taxa)
coccolithophores <10 μm (COCCOS)
poorly known; its biomass attained values above 5 mg carbon m–3 only in a few stations of high northern and southern latitudes
as well as in the equatorial upwelling region
with values of 2–4 mg carbon m–3
Relative abundance of major phytoplankton groups
enumerated by FC or microscopy and estimated in mg carbon m–3
The correspondence between groups and size classes is given by the colour code: Picoplankton
Latitudinal distribution of the carbon biomass (mg carbon m–3) of the main phytoplankton groups
(A) Total carbon of all groups and Prochlorococcus spp.; (B) Synechococcus spp
and picoeukaryotes (PEUK); (C) autotrophic and heterotrophic dinoflagellates and diatoms; (D) coccolithophores
other nanoplankton and filamentous cyanobacteria
Diatoms were present at the majority of stations
High biomass values were found in the English Channel and southern latitudes
reaching a maximum value at most southern station (49° S)
where the genus Pseudonitzschia and the large species Proboscia alata were dominant
Diatom biomass also increased between 1.7 and 8° S
with Hemiaulus hauckii as the more abundant species
attained significant concentrations at latitude 15° N
but not coincident with the Trichodesmium peak
Autotrophic and heterotrophic dinoflagellates are presented separately
as each group has very different ecological requirements
Autotrophic small unchained species were present at low abundance along the transect
From latitudes 45.6° to 41.5° N
the small toxic species Prorocentrum cordatum was abundant (from 0.8 to 2.4 × 103 cells L–1) In the South Atlantic
a sharp increase in ADinos occurred at 42.7° S
corresponding to 28.6 mg carbon m–3
Heterotrophic dinoflagellates had very low abundance throughout the transect
with median and maximum values of 0.11 and 1.92 mg carbon m–3
The maximum value was recorded at latitude 47.52° N
with the species Protoperidinium divergens
The category “NEUK” (Figure 4D) includes cells counted under the microscope
namely Prasinophyceae and the genus Phaeocystis (haptophyta) as well as cells enumerated using the flow cytometer such as cryptophytes and other nanoeukaryotes
It should be underlined that cryptophytes and nanoeukaryotes enumerated by FC accounted
for 99% of total carbon of the total “NEUK” category
This discrepancy is due to the difficulty in identifying small (<10 μm) flagellated cells by microscopy
as well as to the different order of magnitude of multiplying factors used in microscopy
versus 1250–2000 for the 500–800 μL of a sample analysed by FC
to a final result for both with cell abundance expressed per litre
Due to malfunctioning of the flow cytometer during the first week of the cruise
and COCCOS <10 μm diameter were not recorded in the northern latitudes
the plot only displays “NEUK” from 37° N southwards
showing a biomass increase south of the equator and in the Southern Ocean
Coccolithophores include cells identified and counted by microscopy (>10 μm diameter) and cells enumerated by FC (<10 μm diameter)
The percentage of species with >10 μm diameter in relation to the total is higher than 20% in the majority of the transect
attaining a maximum of 93% at latitude 19.3° N
due to the species Umbilicosphaera cf sibogae
with cell diameters of 20–30 μm; the total coccolithophore biomass also increased in the equatorial upwelling and southern latitudes (>30° S)
due to high cell numbers of the same species
The species Emiliania huxleyi was present throughout the transect
and principally Trichodesmium were present between 30° N and 4° N
attaining values >1 mg carbon m–3 between 15 and17° N
where MLD was shallower and surface nitrate + nitrite values negligible
whereas the regions of the northern and southern gyres registered a maximum of 45 and 48
Normalised carbon abundance distribution (mg carbon m–3) of phytoplankton functional types (PFTs) along the transect
and diazotrophs; (B) autotrophic dinoflagellates (ADinos)
Positive and negative anomalies observed for each PFT and normalised to its maximum value
Phytoplankton functional types contributing to biomass increase along the transect
The dynamics of picoautotrophs, depicted in Figure 5, can be complemented with results from Figure 4: Prochlorococcus were the dominant taxa in both gyres
a shift from Prochlorococcus to Synechococcus occurred near more nutrient-rich or colder regions
PEUK responded positively to nutrient increase
the nutrients concentration in surface waters
were very low (often below quantification limit)
The exception was the concentration of silicate
where a significant correlation was found with diatom carbon cell abundance (r2 = 0.49
The temperature along the transect (Figure 1) spanned from 5°C in the two southern stations
being higher than 20°C in the large region from 40° N to 26° S
It should be highlighted that whilst in the northern hemisphere this study corresponds to autumn
in the southern hemisphere represents spring
A further analysis of the different species dominance in these regions is beyond the scope of this work
Figure 6 illustrates the distribution of three size classes (pico-, nano-, and microplankton) according to both the pigment approach and the cell/carbon approach. Figure 6A shows the value of mg Chla m–3 allocated to each size class
as well as the TChla concentration along the transect
The TChla median value is 0.13 mg m–3
with higher values in the northern and southern stations
higher values of TChla were due to higher Chla values estimated for all the 3 size classes
The sharp increase of nanoplankton Chla occurring north of latitude 44° N
and microplankton obtained by pigment (determined by pigments’ concentration) and carbon approaches (determined by cell enumeration)
(A) Chla concentration (mg Chla m–3) observed along the transect using the pigment approach for the pico-
(B) Total Chla and total carbon (mg m–3); (C) mg carbon m–3 for 3 size classes pico-
Note that total carbon values in (B) and nano-values in (C) north of 37° N are underestimated due to the lack of NEUK FC data
Figure 6B shows total carbon (mg carbon m–3) and TChla (mg Chla m–3) along the transect; although there is a general agreement of the two
there are notable differences in higher latitudes
this mismatch is undoubtedly due to the lack of flow cytometer cell values for NEUK
and COCCOS <10 μm diameter from 49 to 37° N
implying that these taxa must be an important component of the phytoplankton community in this region
cell counts by microscopy registered high values for prasinophytes
coccolithophores and for the genus Phaeocystis in this region
justifying the assumption that FC counts for the above groups would also have showed high biomass levels
with ESD from 30 to 100 μm and by ADinos
The linear increase in diameter corresponds to an exponential increase in cell volume and consequently in cell carbon content
which may be biased by the volume-carbon equations used
as it would be expected that larger cells have a less dense cell content
The carbon nano class, corresponding to organisms with ESD between 2.7 and 17.5 μm, had low biomass throughout the latitudinal transect (Figure 6C)
the contribution of three intermediate carbon classes was compared (data not shown): class 2
with a median value of 80% of the total carbon nano class
Chla-microplankton was matched with carbon size class 5 (cells with >180 pg carbon cell–1) and also with the sum of carbon classes 4 and 5
corresponding to cells with >50 pg carbon cell–1 and ESD >10 μm
so as to account for the strong shape variability of diatoms and dinoflagellates
Both correlations gave the same equation coefficients and r2 value
Chla-nanoplankton was matched with carbon size classes 2 (cells 2–10 pg carbon cell–1)
the sum of classes 2 and 3 (cells 2–50 pg carbon cell–1) and the sum of classes 2
corresponding to cells from 2 to 180 pg carbon cell–1
with a range of ESD from 2.7 to 17.5 μm
A significant correlation was obtained for the smallest cells and for the sum of the three carbon size classes defined within nanoplankton
due to the methodological issues regarding identification and enumeration of cells within an ESD range of 10–17.5 μm
it reflects the dominance of small species in nanoplankton size class
Results of linear regression Type II of log10-transformed carbon and log10-transformed Chla for the total data set
as well as for the matching size classes estimated by the pigment or carbon approaches
The three-component size class model of Brewin et al. (2010) was applied to TChla and, for the first time, also to TCarbon (Figure 7)
Fourth row: size-specific fractional contributions to Fm
Total chlorophyll-a is plotted in the top six graphs (blue markers)
and TCarbon on the six bottom graphs (red markers)
The general pattern of the model fitted to TCarbon is in accordance with the one obtained with TChla
With the exception of the graph showing the fraction of nanoplankton as a function of TChla (third graph second row) or TCarbon (third graph fourth row)
which is due to the methodological issues regarding the estimation of nanoplankton cells mentioned above
The ratio of the parameters (Cp,nm and Cpm) obtained for TChla and TCarbon give a value for Carbon:Chla of 170 for picoplankton and 36 for nano plus picoplankton classes
as picoplankton cells have higher carbon content per μm3 cell volume
The overall median Carbon:Chla ratio estimated from the in situ data was 112
it should be noted that Cp,nm and Cpm parameters have a high uncertainty as the phytoplankton biomass concentrations along the AMT are low (TChla maximum was 0.7 mg m–3 and TCarbon maximum was 70 mg m–3)
The Dp,n and Dp parameters are slightly higher for TChla than for TCarbon
but both highlight the dominance of picoplankton at low total biomass
This work has showed that both approaches give a similar general picture in terms of biomass size distribution and composition in the Atlantic
although with poorer results for the nanoplankton fraction
hence the lack of correlation between carbon values and Chla (Fdino)
This observation aims to provide an alert to the diversity of pigment composition found in some taxonomic groups
which might be considered flaws in the pigment approach for some regions/occasions
disadvantages of microscopy should also be remembered
the stronger dependence of the operator expertise for species identification
as well as the longer time consuming of microscopy in comparison with pigments analysis by HPLC
Our results gave significant correlations between both HF and BF with coccolithophore carbon biomass
suggesting the existence of small Prymnesiophyceae and Pelagophyceae in PEUK
whereas phytoplankton cell size displays a continuum
But it is also important to underline the common conclusion of the above three studies
reinforcing the need for complementary methodological approaches
to better characterize the taxa within each size class
in particular regarding the nanoplankton fraction
From the estimation of Fdiat, and the diatom carbon biomass, the average Carbon:Chla value of 60 was obtained, close to the value found by Sathyendranath et al. (2009) for diatoms in northwest Atlantic; however
a wide variation of this ratio was observed
which is related to the wide biovolume and shape variability of diatom cells
The results obtained in our work show a generally good agreement between the use of pigments and use of species/entities identification and enumeration converted into carbon biomass
this work can contribute to the capability of assessing phytoplankton carbon budgets by satellite remote sensing in future studies
the classical phytoplankton species microscopy identification data presented in this article are the first from AMT after a 15 years gap
The cell carbon approach allowed for the comparison of all groups, counted by microscopy or FC. The pattern of spatial distribution of total biomass of the phytoplankton community shows a strong relation with nutrient availability within the MLD, in accordance with the observations of Marañón et al. (2012) that resource availability and not temperature is the key factor explaining the relative success and abundance of different algal size classes
whereas our results indicate percentages >50% in most of north and southern Atlantic stations
reaching 94% at latitude of 49.78° S; we think that this difference is due by the higher detail of our measurements
Carbon concentration of each size class versus total carbon concentration (determined by FCor microscopy)
3 and 4 (>2 <180 pgC cell–1) and microplankton carbon (Micro > 180 Carbon
The correlation between each size class and TCarbon was significant (p > 0.01)
justifying it with a selective advantage for diazotrophic organisms in inorganic nitrogen depleted zones; moreover
these authors showed that activity of diazotrophs can in turn be controlled by the availability of other potentially limiting nutrients
Currently, PFT products are available through the CMEMS, using the approach of Xi et al. (2020)
These authors used nine reflectance bands and a large and widespread diagnostic pigment database
and Synechococcus-like cyanobacteria (SLC)
The relative dominance of PFTs they estimated for October (their Figure 12) can be compared with our results
The dominance of SLC in the gyres and WTRA and the dominance of Prochlorococcus does not correspond to our observations
agreeing with their own conclusion of a low performance for these two PFTs
The dominance of diatoms off the Patagonian coast is in agreement with our results
but not the dominance of haptophytes in temperate regions
we would argue that there is still a great need for ground-truth validation of EO-based PFT retrievals
The roadmap for a future successful discrimination of PFTs, lies in a multi-disciplinary approach which includes in situ measurements of phytoplankton groups (with an array of existing methodologies) and corresponding optical properties, observable by satellites (Bracher et al., 2017)
by providing results for the whole phytoplankton community in the Atlantic
This work showed a good agreement between complementary assessments of phytoplankton structure
the widespread chemotaxonomic approach and the less used
collected along a transect in the Atlantic Ocean
covering a wide range of oceanic provinces
allowed us to quantify phytoplankton taxonomic groups
and discuss their classification within a size class and functional types framework
We applied, for the first time, the Brewin et al. (2010) model to in situ carbon data to assess phytoplankton size structure
The results obtained encourage the application of this model to cell carbon data
establishing the link between PSC assessment by EO
this study presents a comprehensive assessment of all phytoplankton groups in the Atlantic ocean
contributing to a better knowledge of phytoplankton community
providing biomass values for the various regions
namely the composition of the picoeukaryote and nanoplankton fractions
Relevant aspects to be considered in the future have also been pointed out by this work: notably regarding the nanoplankton fraction
which is overestimated by the pigment approach
and poorly evaluated by the combination of microscopy and FC
the correspondence of nanoplankton measured by FC only and the nanoplankton in the pigment approach
validates the use of FC to quantify nanoplankton cells
Our results also evidence the importance of enumerating coccolithophores with ESD >10 μm
as the percentage of species with this dimension were at least 40% of the total coccolithophores concentration
The original contributions presented in the study are included in the article/Supplementary Material
further inquiries can be directed to the corresponding author
All authors contributed to the conceptual development of the manuscript
and CB contributed to cruise participation
VV contributed to microscope identification and enumeration
VB received a sabbatical grant from FCT (SFRH/BSAB/142981/2018)
This work received funding from the European Union’s Horizon 2020 Research and Innovation Programme under grant agreement n° 810139 Project Portugal Twinning for Innovation and Excellence in Marine Science and Earth Observation – PORTWIMS
It also received support by national funds through FCT – Fundação IP
This work was also a contribution to ESA project Ocean Colour Climate Change Initiative
and Interreg Project Innovation in the Framework of the Atlantic Deep Ocean (iFADO)
RB was supported by a UKRI Future Leader Fellowship (MR/V022792/1)
and we also acknowledge John Stephens for providing us the MLD data for the AMT25
We are also indebted to the three reviewers who improved the manuscript
The Supplementary Material for this article can be found online at: https://www.frontiersin.org/articles/10.3389/fmars.2021.682621/full#supplementary-material
Supplementary Table 1 | List of species found and their abundances per sampling station (cell abundances determined by microscopy, and expressed in carbon). The codes of column A are the same as indicated by Sal et al. (2013)
A synthesis of the environmental response of the North and South Atlantic sub-tropical gyres during two decades of AMT
Phytoplankton pigments and functional types in the Atlantic Ocean: a decadal assessment
Obtaining phytoplankton diversity from ocean color: a scientific roadmap for future development
CrossRef Full Text | Google Scholar
Modelling size-fractionated primary production in the Atlantic Ocean from remote sensing
Influence of light in the mixed-layer on the parameters of a three-component model of phytoplankton size class
Comparison of two methods to derive the size-structure of natural populations of phytoplankton
Sensing the ocean biological carbon pump from space: a review of capabilities
Natural variability of phytoplanktonic absorption in oceanic waters: influence of the size structure of algal populations
Understanding bio-optical properties of phytoplankton in the western Iberian coast: application of theoretical models
Deriving phytoplankton size classes from satellite data: validation along a trophic gradient in the Eastern Atlantic
Picophytoplankton biomass distribution in the global ocean
Species Richness and cell-size distribution: size structure and phytoplankton communities
CrossRef Full Text | Google Scholar
Evaluation of diagnostic pigments to estimate phytoplankton size classes
“Phytoplankton size,” in Primary Productivity and Biogeochemical Cycles in the Sea
Corredor-Acosta
Phytoplankton size structure in association with mesoscale eddies off central-southern chile: the satellite application of a phytoplankton size-class model
A three component classification of phytoplankton absorption spectra: application to ocean-colour data
Dimensions of marine phytoplankton diversity
Stress of life at the ocean’s surface: latitudinal patterns of UV sunscreens in plankton across the Atlantic
Analytical phytoplankton carbon measurements spanning diverse ecosystems
Google Scholar
Biovolume calculation for pelagic and benthic microalgae
An absorption model to determine phytoplankton size classes from satellite ocean colour
“Phytoplankton functional types from space,” in Reports of the International Ocean Color Coordinating Group No
Google Scholar
An exact solution for modeling photoacclimation of the carbon-to-chlorophyll ratio in phytoplankton
CrossRef Full Text | Google Scholar
““Microalgal classes and their signature pigments
chemotaxonomy and applications in oceanography,” in Phytoplankton Pigments: Characterization and Applications in Oceanography
Johnsen (Cambridge: Cambridge University Press)
Simultaneous Determination of Selected Nutrients in Seawater
Google Scholar
Carbon-based phytoplankton size classes retrieved via ocean color estimates of the particle size distribution
Seasonal variation in remotely-sensed phytoplankton size structure around southern Africa
CrossRef Full Text | Google Scholar
Le Quéré
Google Scholar
Google Scholar
Spatial and temporal variations of satellite-derived phytoplankton size classes using a three-component model bridged with temperature in Marginal Seas of the Western Pacific Ocean
Synergistic exploitation of hyper- and multispectral precursor Sentinel measurements to determine phytoplankton functional types at best spatial and temporal resolution (SynSenPFT)
Marañón
Resource supply overrides temperature as a controlling factor of marine phytoplankton growth
Marañón
and phytoplankton size structure in the ocean
Marañón
Scaling of phytoplankton photosynthesis and cell size in the ocean
CrossRef Full Text | Google Scholar
Biovolumes and size-classes of phytoplankton in the Baltic Sea
Google Scholar
Ocean’s least productive waters are expanding
CrossRef Full Text | Google Scholar
Coccolithophore ecology in the tropical and subtropical Atlantic Ocean: new perspectives from the Atlantic meridional transect (AMT)
20 years of the Atlantic meridional transect
Phytoplankton Pigments: Characterization and Applications in Oceanography
CrossRef Full Text | Google Scholar
Size-partitioned phytoplankton carbon and carbon-to-chlorophyll ratio from ocean colour by an absorption-based bio-optical algorithm
CrossRef Full Text | Google Scholar
Carbon-to-chlorophyll ratio and growth rate of phytoplankton in the sea
Seasonal ITCZ migration dynamically controls the location of the (sub-)tropical Atlantic biogeochemical divide
Heterotrophic dinoflagellates: a significant component of microzooplankton biomass and major grazers of diatoms in the sea
CrossRef Full Text | Google Scholar
Geometric models for calculating cell biovolume and surface area for phytoplankton
primary production and potential export production in the Atlantic Ocean
Vertical distribution of phytoplankton communities in open ocean: an assessment based on surface chlorophyll
Utermöhl
Zur vervolkommung der quantitativen phytoplankton: methodik
mitteilungen internationale vereinigung für theoretische und angewandte
CrossRef Full Text | Google Scholar
“Ecology and diversity of picoeukaryotes,” in Microbial Ecology of the Oceans
Global retrieval of phytoplankton functional types based on empirical orthogonal functions using CMEMS GlobColour merged products and further extension to OLCI data
A bridge between ocean optics and microbial ecology
CrossRef Full Text | Google Scholar
Ferreira A and Groom SB (2022) Complementary Approaches to Assess Phytoplankton Groups and Size Classes on a Long Transect in the Atlantic Ocean
Copyright © 2022 Brotas, Tarran, Veloso, Brewin, Woodward, Airs, Beltran, Ferreira and Groom. This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY)
and although our state doesn't have such well-defined seasons
the warm weather is inviting you to hit the road and visit cities that are worth a visit
the municipality preserves its tropeira culture
present in its cuisine with dishes such as arroz-de-carreteiro and feijão-de-tropeiro
as well as offering cozy inns for all styles
is the largest in Latin America and a highlight of the municipality
Other attractions include the Manoel Jorge Forest
with the largest karting track in the world
the Festival of Nations brings together culture and solidarity
making it an unmissable destination for all ages
Limeira has attractions such as the Horto Florestal
The city also preserves historic coffee plantations and important events
ideal for hiking and for getting in touch with the rich biodiversity of the Atlantic Forest
the Municipal Spa of Águas de Lindóia not only offers baths for treating ailments
its tourist attractions include Morro do Araçoiaba
where the first mines in the region and the first smelting ovens were discovered
One example is the well-known natural pools with clear waters and white sandy bottoms
You can also go hang gliding and abseiling
Its mountainous terrain is ideal for those looking to get in touch with nature
The city offers many options and is known for rafting
Parece que a página que você está procurando não está disponível