Bloom Maximum Research Articles

Global and local factors driving the phenology of Alexandrium minutum (Halim) blooms and its toxicity

The dinoflagellate Alexandrium minutum is a toxic bloom-forming species distributed worldwide. The mechanisms driving and promoting the species blooms and their toxicity are studied and presented here. Most previously published work focuses on local and/or short-term scales. In this study, a broad temporal and spatial approach is addressed using time series covering several sites over several years and combining environmental variables and A. minutum abundances from the French English Channel − Atlantic coasts. Data were explored by means of phenology and threshold analysis.The A. minutum bloom characteristics are defined. Only one bloom per year is measured and it may reach more than a million of cells L−1. Bloom period extends from April to October and the bloom length ranges from two weeks to six months. In the ecosystems studied, water temperature and river flow, as regional and local factors respectively, are the main environmental drivers influencing the magnitude, growth rate and length of the blooms. Bloom toxicity is linked to the bloom maximum abundance and river flow. This work provides new knowledge for further managing tools for A. minutum blooms in the ecosystems studied.

Climate to fish: Synthesizing field work, data and models in a 39-year retrospective analysis of seasonal processes on the eastern Bering Sea shelf and slope

We combined field data and the output from a climate-to-fish coupled biophysical model to calculate weekly climatologies and 1971–2009 time series of physical and biological drivers for 16 distinct regions of the eastern Bering Sea shelf and slope. We focus on spatial trends and physical-biological interactions as a framework to compare model output to localized or season-specific observations. Data on pollock (≥8cm) diet were used to evaluate energy flows and zooplankton dynamics predicted by the model. Model validation shows good agreement to sea-ice cover albeit with a one month delay in ice retreat. Likewise, the timing of spring phytoplankton blooms in the model were delayed approximately one month in the south and extend further into summer, but the relative timing between the spring and fall bloom peaks was consistent with observations. Ice-related primary producers may shift the timing of the spring bloom maximum biomass earlier in years when sea ice was still present after mid-March in the southern regions. Including the effects of explicit, dynamic fish predation on zooplankton in the model shifts the seasonal spring peak and distribution of zooplankton later in the year relative to simulations with implicit predation dependent only on zooplankton biomass and temperature; the former capturing the dynamic demand on zooplankton prey by fish. Pollock diets based on stomach samples collected in late fall and winter from 1982–2013 show overwintering euphausiids and small pollock as key prey items in the outer and southern Bering Sea shelf; a characteristic not currently present in the model.The model captured two large-scale gradients, supported by field data, characterizing the overall dynamics: 1) inshore to off-shelf physical and biological differences with a gradient in inter-annual variability from higher frequency inshore to lower frequency offshore; and 2) latitudinal gradients in the timing of events. The combined effects of length of day, bathymetry, and tides, which are consistent from year to year, and the two large-scale gradients, characterize the environment on which regional differences were based and restrict their inter-annual and seasonal variability. Thus, the relative timing and sequence of events remained consistent within regions. The combination of model outputs and observational data revealed specific ecosystem processes: (1) The spatial progression in the timing, peaks and sequence of events over the shelf is driven by wind, sea ice, and stratification and creates a seasonal expansion and contraction of the warmer pelagic and bottom habitat suitable to pollock. (2) The seasonal warming of air temperature and the spring-summer expansion of the warm pelagic and bottom habitats influence the ice retreat and the associated ice edge and open water spring blooms, as well as subsequent production/abundance of copepods and euphausiids. (3) These warmer conditions favor pelagic energy flows to pollock (≥10cm) and allow their distribution to expand shoreward and northward along the shelf break. (4) The fall-winter expansion of the seasonal ice cover drives the contraction of warmer waters towards the outer and southwest shelf and favors benthic energy flows over most of the shelf. There, fall blooms allow for additional lipid storage by large copepods and euphausiids that sink close to the bottom where they either go into diapause or have a restricted diel migration over winter. (5) During these cold months, the preferred pollock habitat shifts and contracts towards the outer and southwest shelf where their increased density and reduced prey availability leads to winter pollock cannibalism and consumption of overwintering euphausiids. Our project highlights the benefits of linking continuous and long-term field work with the development and implementation of highly complex models. In the face of uncertainty, simulations such as these, tightly coupled to field programs, will be instrumental as testbeds for process exploration and management evaluation, increasing their relevance for future fisheries and ecosystem management and strategic planning.

Are phytoplankton blooms occurring earlier in the Arctic?

Time series of satellite-derived surface chlorophyll-a concentration (Chl) in 1997–2009 were used to examine for trends in the timing of the annual phytoplankton bloom maximum. Significant trends towards earlier phytoplankton blooms were detected in about 11% of the area of the Arctic Ocean with valid Chl data, e.g. in the Hudson Bay, Foxe Basin, Baffin Sea, off the coasts of Greenland, in the Kara Sea and around Novaya Zemlya. These areas roughly coincide with areas where ice concentration has decreased in early summer (June), thus making the earlier blooms possible. In the selected areas, the annual phytoplankton bloom maximum has advanced by up to 50 days which may have consequences for the Arctic food chain and carbon cycling. Outside the Arctic, the annual Chl maximum has become earlier in boreal North Pacific but later in the North Atlantic.

Dinoflagellate blooms in upwelling systems: Seeding, variability, and contrasts with diatom bloom behaviour

The influence of diatom bloom behaviour, dinoflagellate life cycles, propagule type and upwelling bloom cycles on the seeding of dinoflagellate blooms in eastern boundary current upwelling systems is evaluated. Winter–spring diatom bloom behaviour is contrasted with upwelling bloom behaviour because their phenology impacts dinoflagellate blooms. The winter–spring diatom bloom is usually sustained, whereas the classical upwelling diatom bloom occurs as a series of separate, recurrent mini-blooms intercalated by upwelling-relaxation periods, during which dinoflagellates often bloom. Four sequential wind-regulated phases characterize upwelling cycles, with each phase having different effects on diatom and dinoflagellate bloom behaviour: bloom “spin up”, bloom maximum, bloom “spin down”, and upwelling relaxation. The spin up – bloom maximum is the period of heightened diatom growth; the spin down – upwelling-relaxation phases are the periods when dinoflagellates often bloom. The duration, intensity and ratio of the upwelling and relaxation periods making up upwelling cycles determine the potential for dinoflagellate blooms to develop within a given upwelling cycle and prior to the subsequent “spin up” of upwelling that favours diatom blooms. Upwelling diatoms and meroplanktonic dinoflagellates have three types of propagules available to seed blooms: vegetative cells, resting cells and resting cysts. However, most upwelling dinoflagellates are holoplanktonic, which indicates that the capacity to form resting cysts is not an absolute requirement for growth and survival in upwelling systems. The long-term (decadal) gaps in bloom behaviour of Gymnodinium catenatum and Lingulodinium polyedrum, and the unpredictable bloom behaviour of dinoflagellates generally, are examined from the perspective of seeding strategies. Mismatches between observed and expected in situ bloom behaviour and resting cyst dynamics are common among upwelling dinoflagellates. This disassociation suggests unrecognized upwelling system factors that fall within the physical–chemical–biological domain are more important than life cycle in selecting dinoflagellates species having the survival-seeding strategies and ecophysiological adaptations required for growth in physically robust upwelling systems. It is conjectured that diatom life cycles, as a group, are geared towards exploiting seeding opportunities, whereas dinoflagellates have evolved life-cycle behaviour more attuned to survival. The role of ecological dormancy and ecological release from bloom inhibition underlying dinoflagellate bloom irregularity is considered. The expectation that the dinoflagellate species selected to bloom from among the common upwelling flora would be the same in all eastern boundary upwelling systems is not realized.

A new method of describing phytoplankton blooms: Examples from Helgoland Roads

Phytoplankton blooms, in their pivotal position in pelagic seasonal succession require precise classification criteria in order to evaluate such parameters as bloom start, bloom timing, bloom maximum and growth rates. Such bloom parameters are linked directly to species and bloom specific features. Currently the phytoplankton bloom concept, though intuitively clear, lacks operational criteria allowing the precise definition of bloom parameters. We present a semi-quantitative method of classification of marine phytoplankton blooms based on an algorithmic estimation of several bloom descriptors computed from densely recorded phytoplankton data, like the Helgoland Roads long-term time series. Combining these descriptors we propose a novel classification scheme which may serve useful in the discussion of species fitness, competition and succession of marine algae. Special emphasis is put on the detection of the bloom start, because of its crucial importance for many current research topics, including trigger mechanisms and climate-induced temporal shifts in the context of the match/mismatch hypothesis. Visual examination of scatter plots of these parameters leads us to propose three types of blooming algae.

First detection of Pectenotoxin-11 and confirmation of OA-D8 diol-ester in Dinophysis acuta from European waters by LC–MS/MS

During November 2005, a dense bloom of Dinophysis spp. dominated (>97%) by Dinophysis acuta in the Galician Rías Baixas (NW Spain), provided a unique opportunity to describe the full toxin profile – including toxins that represent a low percentage and escape detection in analyses of single-cell isolates – in plankton concentrates rich in this species. Detection and identification of toxins were carried out by liquid chromatography coupled to mass spectrometry (LC–MS/MS), a method that is based on their retention times (RT) and the fragmentation patterns of their mass spectra. OA-D8 diol-ester and PTX11 were detected in co-occurrence with okadaic acid analogues (OA, DTX2) and PTX2 in plankton concentrates dominated by D. acuta. The presence of a PTX11-isomer, which was suggested to be PTX13 or a novel PTX11-isomer, released in the sea water, was also confirmed in samples obtained after deployment of passive samplers (SPATT) in situ at the time of the D. acuta bloom maximum. The amount of PTX11 per cell of D. acuta, estimated as PTX2 equivalents, ranged between not detected and 2.2 pg cell −1, and represented a maximum of 2.9% of the total toxin content. The variation in PTX11 content per cell of D. acuta, during a daily cycle, followed the same pattern than that of PTX2, with maxima at 21:00 and 03:00 h (dark hours), but the amounts per cell were one order of magnitude lower. This is the first report of PTX11, together with the confirmation of OA-D8 diol-ester in D. acuta populations from Europe.

Short‐term variations in development of a recurrent toxic Alexandrium minutum–dominated dinoflagellate bloom induced by meteorological conditions1

Development of an Alexandrium minutum Halim bloom affecting a Mediterranean harbor was monitored in detail using a multidisciplinary approach. A. minutum was by far the most abundant species at and near the bloom maximum, but always coexisted with members of three additional dinoflagellate genera and prasinophytes. Bloom initiation (early February) occurred during prolonged influences of sunny weather conditions, when day length exceeded 10.5 h and water temperatures reached 10.2°C. Subsequent development toward its maximum (end of March) also relied on good weather conditions, with specific wind directions favoring accumulation of cells. Arrival of rainy weather, associated with frontal boundaries of large‐scale low‐atmospheric‐pressure systems and characterized by reduced solar irradiance (heavy cloud coverage), opposite wind directions, and enhanced wind speeds, always caused temporal declines of the bloom. These declines were attributed to dispersal or displacement of algae, but a vertical migration of A. minutum cells toward the sediment was not excluded. Delayed inflows of excess terrestrial rainwater along the inner harbor wall strongly reduced salinity and prolonged a temporal decline far beyond influences of bad weather. The associated nutrient supply favored development of the phytoplankton population but reduced the toxin production of A. minutum cells. The HPLC‐determined Gonyautoxin (GTX) 1 + 4/GTX 2 + 3 ratio strongly increased toward the bloom maximum. This ratio was influenced by nutrient status and cell density and has a potential value for monitoring developmental stages of blooms. Prolonged bad weather conditions eventually hindered continuation of bloom development, and subsequent declines of algal biomass were attributed to grazing.

Impact of the river Liffey discharge on nutrient and chlorophyll concentrations in the Liffey estuary and Dublin Bay (Irish Sea)

Temperature, salinity, nutrients (total oxidised nitrogen (TON), ammonium (NH 4) and orthophosphate (PO 4)) and chlorophyll a were monitored in the Liffey estuary and Dublin Bay from June 2000 to June 2003. Four groups of sites were defined comprising the upper estuary (Gp. I), the outer estuary (Gp. III) with a small set (Gp. II) of sites between Groups I and III heavily influenced by the sewage treatment works outflow, and the Bay proper (Gp. IV). Riverine inputs of TON and PO 4 were calculated at an average of 826 t N y −1 and 31 t P y −1, respectively, and were largely controlled by flow rate. The sewage treatment works were identified as a major source of PO 4 and NH 4 to the system. Mixing in the upper estuary of nutrient limited saline waters with hypernutrified river water regularly (i.e. annually) produced relatively high concentrations of chlorophyll a (>10 mg chl a m −3), and also sporadic blooms with extremely high chlorophyll a values (max. 121.6 mg chl a m −3). These latter phytoplankton blooms occurred in high salinity waters and were due to mixing of nutrient limited saline waters and nutrient rich river waters. The mean annual flux of phytoplankton carbon from the river Liffey was calculated at 23.5 t C y −1, of which half was accumulated or remineralised in the estuary and did not enter the Bay. In the Bay proper (Gp. IV) summer nutrient concentrations dropped below detection limits, and chlorophyll a concentrations followed the classic pattern with a spring bloom maximum of 5.5 mg chl a m −3. This pattern in nutrients and chlorophyll a came from the advection of waters into the Bay from an offshore source. Overall while there was considerable evidence for eutrophication in the estuary, the bay itself showed little biological response to nutrient loading.

Lipid classes and fatty acids in plankton and settling matter during the spring bloom in a cold ocean coastal environment

Lipid concentration and composition were measured in net-tow and sediment trap samples during the spring diatom bloom in Conception Bay, Newfoundland, to determine the quantity and quality of material settling from the upper mixed layer into the benthic food web. An experiment was also conducted to estimate trap accuracy and precision, in which trap poisoning was found to reduce bacterially mediated dissolution, but to increase variability in particulate lipid data. Total lipid concentrations in plankton reached a maximum of 6.3 ± 2.5% dry weight (mean ± SD) at the height of the bloom in April. Bacterial fatty acid markers were at their lowest during the bloom period (2.0 to 2.8% of total fatty acids), while polyunsaturated fatty acids (PUFA) were at their highest from the beginning of the bloom onwards (44 to 53%). The maximum fluxes of total lipids (75 ± 0.7 mg m–2 d–1) in non-poisoned traps at 80 m depth occurred at the bloom maximum while those at 220 m (35 ± 1.7 mg m–2 d–1) occurred 16 d later. The lipid and PUFA supply to the benthos was over 10 times that accumulated by important amphipod and mysid populations in Conception Bay. In the deep traps, PUFA were maintained at high levels (33 to 35%) throughout the bloom period while bacterial markers were at their lowest (2.3 to 2.9%). A comparison of fluxes showed that over half the PUFA survived transit from 80 to 220 m during the bloom period. These data indicate a rapid transfer of highly nutritious material to the benthos during the spring bloom in Conception Bay.

Biogeochemical impacts due to mesoscale eddy activity in the Sargasso Sea as measured at the Bermuda Atlantic Time-series Study (BATS)

Abstract A comparison of monthly biogeochemical measurements made from 1993 to 1995, combined with hydrography and satellite altimetry, was used to assess the impacts of nine eddy events on primary productivity and particle flux in the Sargasso Sea. Measurements of primary production, thorium-234 flux, nitrate+nitrite, and photosynthetic pigments made at the US JGOFS Bermuda Atlantic Time-series Study (BATS) site were used. During the 3 years of this study, four out of six high thorium-234 flux events occurred during the passage of an eddy. Primary production nearly as high as the spring bloom maximum was observed in two mode-water eddies (May 1993 and July 1995). The 1994 spring bloom at BATS was suppressed by the passage of an anticyclone. Distinct phytoplankton community shifts were observed in mode-water eddies, which had an increased percentage of diatoms and dinoflagellates, and in cyclones, which had an increased percentage of Synechococcus . These variations in species composition within mode-water eddies and cyclones may be associated with the ages of the sampled eddies, and/or differences in physical, chemical, and biological factors in these two distinct eddy types. In general, eddies that were 1–2 months old elicited a large biological response; eddies that were 3 months old may show a biological response and were accompanied by high thorium flux; eddies that were 4 months old or older did not show a biological response or high thorium flux. A conceptual model depicting temporal changes during eddy upwelling, maturation, and decay can explain the observations in all seven upwelling eddies present in the time-series investigated herein.

Primary productivity and its regulation in the Arabian Sea during 1995

Abstract The annual cycle of monsoon-driven variability in primary productivity was studied in 1995 during the Arabian Sea Expedition as part of the United States Joint Global Ocean Flux Studies (US JGOFS). This paper describes the seasonal progression of productivity and its regulation on a section which ran from the coast of Oman to about 1000 km offshore in the central Arabian Sea at 65°E. During the SW Monsoon (June–mid-September), the coolest water and highest nutrient concentrations were close to the coast, although they extended offshore to about 800 km; during the January NE Monsoon, deep convective mixing provided nutrients to the mixed layer in the region 400 – 1000 km offshore. As expected, the SW Monsoon was the most productive season (123±9 mmol C m−2 d−1) along the southern US JGOFS section from the coast to 1000 km offshore, but productivity in the NE Monsoon was surprisingly high (112±7 mmol C m−2 d−1). There was no onshore/offshore gradient in primary productivity from 150 to 1000 km off the Omani coast in 1995, and there was no evidence of light limitation of either primary productivity or photosynthetic performance (PoptB) from deep convective mixing during the NE Monsoon, deep wind mixing during the SW Monsoon or offshore Ekman downwelling during the SW Monsoon. Productivity during the Spring Intermonsoon (86±6 mmol C m−2 d−1) was much higher than in oligotrophic regions such as the tropical Pacific Ocean (29±2 mmol C m−2 d−1) or the North Pacific gyre region (32±8 mmol C m−2 d−1). The 1995 annual mean productivity (111±11 mmol C m−2 d−1) along this section from the Omani coast to the central Arabian Sea was about equal to the spring bloom maximum (107±23 mmol C m−2 d−1) during the 1989 North Atlantic Bloom Experiment (NABE) and the equatorial, 1°N–1°S wave guide maximum (95±6 mmol C m−2 d−1) in the Pacific Ocean during the 1992 EqPac study. The 1995 SW Monsoon primary productivity was similar to the mean value observed in the same region in 1994 by the Arabesque Expedition (127±14 mmol C m−2 d−1) and in 1964 by the ANTON BRUUN Expedition (115±27 mmol C m−2 d−1). During the 1995 SW Monsoon, strong, narrow and meandering current filaments extended from the region of coastal upwelling to about 700 km offshore; these filaments had levels of biomass, primary productivity, chlorophyll-specific productivity and diatom abundance that were elevated relative to other locations during the SW Monsoon. The SW Monsoon was the most productive period, but SW Monsoon primary productivity values were lower than predicted because efficient grazing by mesozooplankton kept diatoms from accumulating the biomass necessary for achieving the high levels of primary productivity characteristic of other coastal upwelling regions. The high rates of chlorophyll-specific productivity (PoptB>10 mmol C mg Chl−1 d−1) observed in the 1995 SW Monsoon, together with the observed dust flux and iron concentrations, indicate that the Arabian Sea was more iron replete than the equatorial Pacific Ocean or the Southern Ocean.

Phytoplankton growth rates in the Ross Sea, Antarctica, determined by independent methods: temporal variations

The development of the seasonal phytoplankton bloom in the Ross Sea was studied during two cruises. The first, conducted in November-December 1994, investigated the initiation and rapid growth of the bloom, whereas the second (December 1995-January 1996) concentrated on the bloom's maximum biomass period and the subsequent decline in biomass. Central to the under- standing of the controls of growth and the summer decline of the bloom is a quantitative assessment of the growth rate of phytoplankton. Growth rates were estimated over two time scales with different methods. The first estimated daily growth rates from isotopic incorporation under simulated in situ conditions, including 14 C, 15 N and 32 Si uptake measurements combined with estimates of standing stocks of particulate organic carbon, nitrogen and biogenic silica. The second method used daily to weekly changes in biomass at selected locations, with net growth rates being estimated from changes in standing stocks of phytoplankton. In addition, growth rates were estimated in large-volume experi- ments under optimal irradiances. Growth rates showed distinct temporal patterns. Early in the growing season, short-term estimates suggested that growth rates of in situ assemblages were less than maximum (relative to the temperature-limited maximum) and were likely reduced due to low irradi- ance regimes encountered under the ice. Growth rates increased thereafter and appeared to reach their maximum as biomass approached the seasonal peak, but decreased markedly in late December. Differences between the two major taxonomic groups present were also noted, especially from the isotopic tracer experiments. The haptophyte Phaeocystis antarctica was dominant in 1994 throughout the growing season, and it exhibited the greatest growth rates (mean 0.41 day-1) during spring. Diatom standing stocks were low early in the growing season, and growth rates averaged 0.10 day-1. In summer, diatoms were more abundant, but their growth rates remained much lower (mean of 0.08 day-1) than the potential maximum. Understanding growth rate controls is essential to the develop- ment of predictive models of the carbon cycle and food webs in Antarctic waters.

Possible food chain relationships between bacterioplankton, protozoans, and cladocerans in a reservoir

AbstractIngestion of fluorescent particles by natural protozoan assemblage was studied in the Řimov Reservoir (Southern Bohemia) from April to October, 1987. Attached and free‐living bacterial abundance, proportion of active bacteria, density of suspended particles and biomass of cladocerans were also monitored.Heterotrophic nanoflagellates (HNF; 5–12.8 102ml−1) were the dominant bacterial micrograzers during the spring period and consumed 3 to 9% of the total bacteria per day. After the spring phytoplankton bloom maximum densities of suspended particles and attached bacteria (up to 28% of the total counts) were found. Development of cladocerans in May sharply decreased the proportion of attached bacteria and kept them below 5% of the total counts.All the studied components of plankton except Cladocera decreased during the clearwater phase. The most significant drop was observed in the numbers of protozoans, and they were negligible for bacterial elimination. Bacterial losses during that time apparently were due to cladoceran grazing.During the summer period, ciliates (15–142 ml−1) were mostly dominant micrograzers, and protozoan community grazing increased up to 21% of bacterial standing stock per day. The proportion of active bacteria was strongly correlated with protozoan grazing (r=0.83).