Maurice Levasseur

Synthesis of iron fertilization experiments: From the Iron Age in the Age of Enlightenment

Comparison of eight iron experiments shows that maximum Chl a, the maximum DIC removal, and the overall DIC/Fe efficiency all scale inversely with depth of the wind mixed layer (WML) defining the light environment. Moreover, lateral patch dilution, sea surface irradiance, temperature, and grazing play additional roles. The Southern Ocean experiments were most influenced by very deep WMLs. In contrast, light conditions were most favorable during SEEDS and SERIES as well as during IronEx-2. The two extreme experiments, EisenEx and SEEDS, can be linked via EisenEx bottle incubations with shallower simulated WML depth. Large diatoms always benefit the most from Fe addition, where a remarkably small group of thriving diatom species is dominated by universal response of Pseudo-nitzschia spp. Significant response of these moderate (10–30 μm), medium (30–60 μm), and large (>60 μm) diatoms is consistent with growth physiology determined for single species in natural seawater. The minimum level of “dissolved” Fe (filtrate < 0.2 μm) maintained during an experiment determines the dominant diatom size class. However, this is further complicated by continuous transfer of original truly dissolved reduced Fe(II) into the colloidal pool, which may constitute some 75% of the “dissolved” pool. Depth integration of carbon inventory changes partly compensates the adverse effects of a deep WML due to its greater integration depths, decreasing the differences in responses between the eight experiments. About half of depth-integrated overall primary productivity is reflected in a decrease of DIC. The overall C/Fe efficiency of DIC uptake is DIC/Fe ∼ 5600 for all eight experiments. The increase of particulate organic carbon is about a quarter of the primary production, suggesting food web losses for the other three quarters. Replenishment of DIC by air/sea exchange tends to be a minor few percent of primary CO2 fixation but will continue well after observations have stopped. Export of carbon into deeper waters is difficult to assess and is until now firmly proven and quite modest in only two experiments.

A global database of sea surface dimethylsulfide (DMS) measurements and a procedure to predict sea surface DMS as a function of latitude, longitude, and month

A database of 15,617 point measurements of dimethylsulfide (DMS) in surface waters along with lesser amounts of data for aqueous and particulate dimethylsulfoniopropionate concentration, chlorophyll concentration, sea surface salinity and temperature, and wind speed has been assembled. The database was processed to create a series of climatological annual and monthly 1°×1° latitude-longitude squares of data. The results were compared to published fields of geophysical and biological parameters. No significant correlation was found between DMS and these parameters, and no simple algorithm could be found to create monthly fields of sea surface DMS concentration based on these parameters. Instead, an annual map of sea surface DMS was produced using an algorithm similar to that employed by Conkright et al. [1994]. In this approach, a first-guess field of DMS sea surface concentration measurements is created and then a correction to this field is generated based on actual measurements. Monthly sea surface grids of DMS were obtained using a similar scheme, but the sparsity of DMS measurements made the method difficult to implement. A scheme was used which projected actual data into months of the year where no data were otherwise present.

PHYSIOLOGICAL ACCLIMATION OF MARINE PHYTOPLANKTON TO DIFFERENT NITROGEN SOURCES1

We examined the energetic dependency of the biochemical and physiological responses of Thalassiosira pseudonana Hasle and Heimdal. Chaetoceros gracilis Schütt, Dunaliella tertiolecta Butcher, and Gymnodinium sanguineum Hirasaka to NH4+, NO3−, and urea by growing them at subsaturating and saturating photon flux (PF). At subsaturating PF, when energy was limiting, NO3− and NH4+ grown cells had similar growth rates and C and X quotas. Therefore, NO3− grown cells used up to 48% more energy than NH4+ grown cells to assimilate carbon and nitrogen. Based on our measurements of pigments, chlorophyll‐a‐specific in vivo absorption cross‐section, and fluorescence‐chlorophyll a−1, we suggest that NO3−, grown cells do not compensate for the greater energy requirements of NO3− reduction by trapping more light energy. At saturating PF, when energy is not limiting, the utilization of NO3−, compared to NH4+ resulted in lower growth rates and N quotas in Thalassiosira pseudonana and lower N quotas in Chaetoceros gracilis, suggesting enzymatic rather than energetic limitations to growth. The utilization of urea compared to Nh4+ resulted in lower growth rates in Chaetoceros gracilis and Gymnodinium sanguineum (saturating PF) and in lower N quotas in all species tested at both subsaturating and saturating PF. The high C:N ratios observed in all urea‐grown species suggest that nitrogen assimilation may be limited by urea uptake or deamination and that symptoms of N limitation in microalgae may be induced by the nature of the N source in addition to the N supply rate. Our results provide new eridence that the maximum growth rates of microalgae may be limited by enzymatic processes associated with the assimilation of NO3−, or urea.

Carbon budget of sea‐ice algae in spring: Evidence of a significant transfer to zooplankton grazers

The fate of ice-bottom algae, before and after release from the first-year sea ice into the water column, was assessed during the period of ice-algal growth and decline in Resolute Passage (Canadian Arctic). During spring 1992 (from April to June), algae in the bottom ice layer and those suspended and sinking in the upper water column (top 15 m) were sampled approximately every 4 days. Ice-bottom chlorophyll a reached a maximum concentration of 160 mg m−2 in mid-May, after which it decreased to lower values. In the water column, chlorophyll a concentrations were low until the period of ice-algal decline (∼0.1 mg m−3), with most biomass in the 65% of total export) and that ice-algae were ingested by under-ice grazers after release from the ice. These results stress the importance of ice algae for pelagic consumers during the early stages of ice melt and show that the transfer of ice algae to higher trophic levels extends beyond the period of maximum algal production in the ice bottom.

A new source of dimethylsulfide (DMS) for the arctic atmosphere: ice diatoms

We report the first evidence that pennate diatoms growing within the bottom layer of first-year ice in the Arctic produce significant amounts of particulate dimethylsulfoniopropionate (DMSPp) and dissolved DMSP+DMS. In 1992 in Resolute Passage, a tributary of Barrow Strait, DMSPp concentrations within the bottom layer of ice reached 1055 mg S m-3 at the end of the vernal bloom, a value one order of magnitude higher than the maximum value reported in antarctic ice. Bottom-ice concentrations in DMSPp and DMSPd+DMS were significantly correlated with the abundance of the dominant pennate diatom Nitzschia frigida. Intracellular concentration in DMSP of ice algae was very low (0.001 pg cell-1) at the end of April when algae were light-limited and reached 1.17 pg cell-1 in mid-May following an increase in light and algal growth. We calculate that the rapid release of the dissolved DMSP+DMS from the ice into surface waters following the ice break-up will generate a sea-to-air DMS flux of 0.7 mg S m-2 d-1, a pulse ten times higher than the mean arctic summer flux. We estimate that this 1-d pulse represents up to 5% of the annual DMS emission in the Arctic.

Production of DMSP and DMS during a mesocosm study of an Emiliania huxleyi bloom: influence of bacteria and Calanus finmarchicus grazing

We investigated the influence of bacteria and metazooplankton on the production of dimethylsulfoniopropionate (DMSP) and dimethylsulfide (DMS) during blooms of Emiliania huxleyi (Lohmann) Hay and Mohler in seawater mesocosms. The phytoplankton succession was marked by the rapid collapse of an initial Skeletonema costatum (Greville) Cleve bloom followed by a small E. huxleyi bloom. The collapse of the diatom bloom was accompanied by an increase in concentrations of dissolved DMSP (DMSPd) and bacterial abundance and activity (as determined by the thymidine incorporation technique). The increase in bacterial activity was followed by a rapid decrease in DMSPd concentrations which remained low for the rest of the experiment, even during the subsequent collapse of the E. huxleyi blooms. The absence of DMSPd and DMS peaks during the declining phase of the E. huxleyi blooms was attributed to the high bacterial activity prevailing at that time. The influence of metazooplankton grazing on DMSP and DMS production was investigated by adding moderate (24 mg dry weight m-3) and high (520 mg dry weight m-3) concentrations of Copepodite Stage V and adults of Calanus finmarchicus to two of four filtered (200 μm mesh net) enclosures during the E. huxleyi blooms. The addition of C. finmarchicus, even in high concentrations, had no apparent effect on the dynamics of E. huxleyi, suggesting that the copepods were not grazing significantly on nanophytoplankton. The addition of copepods in high concentrations favored an accumulation of chlorophyll a and particulate DMSP. These results suggest that copepods were preying on the herbivorous microzooplankton which, in turn, was controlling the biomass of nanophytoplankton. DMS production was also enhanced in the enclosure with maximum metazooplankton biomass, suggesting that the grazing of C. finmarchicus on microzooplankton containing DMSP may contribute to DMS production. These results provide strong support to the emerging idea that bacteria and metazooplankton grazing play a dominant role in determining the timing and magnitude of DMS pulses following phytoplankton blooms.

Flux estimation of oceanic dimethyl sulfide around North America

Simultaneous measurements of atmospheric and surface water dimethyl sulfide (DMS) concentrations were taken aboard the icebreaker USCGC Polar Sea from July to October 1994, as part of a joint Canada/United States circumnavigation of North America, an expedition with a unique Arctic Ocean transect. Atmospheric DMS concentrations around North America varied between 0.25 and 50 nmol m−3 (mean = 5.1 nmol m−3, σ = 8.5 nmol m−3, n = 89) with highest values occurring near (south of) the Arctic ice edge. Surface water DMS concentrations ranged between 0.1 and 12.6 nmol L−1 (mean = 2.2 nmol L−1, σ = 2.7 nmol L−1, n = 46) with highest values in the western Arctic Ocean and off the U.S. east coast, near the Sargasso sea. In the Arctic Ocean, maximum concentrations in air and water were found along the ice edge in the Chukchi Sea region. Atmospheric DMS decay rates of 68% per day and 38% per day were deduced from observations between 70° and 76°N (continental shelf and slope of Chukchi Sea) on the west side and between 80° and 90°N (central Arctic Ocean) for the east side of the Arctic Ocean, respectively. Ocean to atmosphere flux estimates of DMS were determined using the Liss-Merlivat empirical dependence of exchange coefficient on wind speed, DMS air concentrations, Henrys law constants, and DMS water concentrations. DMS fluxes varied between 0.0017 and 30 μmol m−2 d−1, respectively, with higher fluxes in regions with open water. The Arctic Ocean contributed 0.063 Tg S (DMS) (0.4% of DMS from the world oceans) during the summer of 1994. A simple one-dimensional (1-D) photochemical box model, applied to six case studies, showed that the atmospheric lifetime of DMS in the high Arctic was 2.5 to 8 days, whereas at 16°–33°N it was 1 to 2 days. Modeled DMS decay rates for these regions, using the 1-D model, accounted for only 33% of the measured decay rate. This result also suggests that halogen chemistry, reactions with Br/BrO, may be an important sink for DMS in the Arctic atmosphere.

CONTROL OF GERMINATION OF ALEXANDRIUM TAMARENSE (DINOPHYCEAE) CYSTS FROM THE LOWER ST. LAWRENCE ESTUARY (CANADA)

Cysts of the toxic dinoflagellate Alexandrium tamarense (Lebour) Balech 1992 from the lower St. Lawrence estuary were used in a test of the following hypotheses: (1) cyst germination is triggered by a change in temperature, and (2) germination rate varies throughout the year and is controlled by a circannual internal biological clock. Results show that cyst germination was not affected significantly by temperature of incubation over the range 1°–16° C, and light showed no significant stimulation of germination. This is supported by the lack of effect of cyst incubation conditions during evaluation of the seasonal changes in germination rate (two temperatures: 4° and 15° C, and two light conditions: darkness and 150 μmol photons·m−2·s−1). Thus, direct environmental control through short‐term increases in temperature and exposure to light has no effect on the germination of the cysts tested. The rate of germination, observed monthly over a 16‐month period, showed low germination (<20%) over most of the period tested, except for a maximum reaching more than 50% germination in August to October of the second year of the experiment. This pattern was observed for cysts both from monthly field collections and from laboratory‐stored cysts kept under constant environmental conditions (4° C, in the dark). The peak in germination observed under constant environmental conditions (in the laboratory), the almost coincidental increase in cyst germination observed for the field‐collected cysts, and the absence of effects of temperature and light during incubation could be explained either by a temperature‐controlled cyst maturation period (the time‐temperature hypothesis of Huber and Nipkow 1923) or by the presence of an internal biological clock. However, the large decline in the rate of germination 2 months after the maximum provides strong support for the biological clock hypothesis. The ca. 12‐month maturation (dormancy) period observed for the laboratory‐stored cysts is the longest reported for this species to our knowledge; this might be related to the low storage temperature (4° C), which is close to bottom temperatures generally encountered in this environment (0° to 6° C). Similar field and laboratory storage temperatures could explain the coincidental increase in germination rate in the fall of the second year if cyst maturation is controlled by temperature. A fraction of the laboratory‐stored cysts did not follow a rhythmic pattern: A rather constant germination rate of about 20% was observed throughout the year. This continuous germination of likely mature cysts may supplement the local blooms of this toxic dinoflagellate, as these often occur earlier than peak germination observed in late summer. It seems that two cyst germination strategies are present in the St. Lawrence: continuous germination after cyst maturation, with temperature controlling the length of the maturation period, and germination controlled by a circannual internal rhythm.

Influence of transport and ocean ice extent on biogenic aerosol sulfur in the Arctic atmosphere

The recent decline in sea ice cover in the Arctic Ocean could affect the regional radiative forcing via changes in sea ice-atmosphere exchange of dimethyl sulfide (DMS) and biogenic aerosols formed from its atmospheric oxidation, such as methanesulfonic acid (MSA). This study examines relationships between changes in total sea ice extent north of 70 degrees N and atmospheric MSA measurement at Alert, Nunavut, during 1980-2009; at Barrow, Alaska, during 1997-2008; and at Ny-Alesund, Svalbard, for 1991-2004. During the 1980-1989 and 1990-1997 periods, summer (July-August) and June MSA concentrations at Alert decreased. In general, MSA concentrations increased at all locations since 2000 with respect to 1990 values, specifically during June and summer at Alert and in summer at Barrow and Ny-Alesund. Our results show variability in MSA at all sites is related to changes in the source strengths of DMS, possibly linked to changes in sea ice extent as well as to changes in atmospheric transport patterns. Since 2000, a late spring increase in atmospheric MSA at the three sites coincides with the northward migration of the marginal ice edge zone where high DMS emissions from ocean to atmosphere have previously been reported. Significant negative correlations are found between sea ice extent and MSA concentrations at the three sites during the spring and June. These results suggest that a decrease in seasonal ice cover influencing other mechanisms of DMS production could lead to higher atmospheric MSA concentrations.

A first appraisal of prognostic ocean DMS models and prospects for their use in climate models

Ocean dimethylsulfide (DMS) produced by marine biota is the largest natural source of atmospheric sulfur, playing a major role in the formation and evolution of aerosols, and consequently affecting climate. Several dynamic process-based DMS models have been developed over the last decade, and work is progressing integrating them into climate models. Here we report on the first international comparison exercise of both 1D and 3D prognostic ocean DMS models. Four global 3D models were compared to global sea surface chlorophyll and DMS concentrations. Three local 1D models were compared to three different oceanic stations (BATS, DYFAMED, OSP) where available time series data offer seasonal coverage of chlorophyll and DMS variability. Two other 1D models were run at one site only. The major point of divergence among models, both within 3D and 1D models, relates to their ability to reproduce the summer peak in surface DMS concentrations usually observed at low to mid- latitudes. This significantly affects estimates of global DMS emissions predicted by the models. The inability of most models to capture this summer DMS maximum appears to be constrained by the basic structure of prognostic DMS models: dynamics of DMS and dimethylsulfoniopropionate (DMSP), the precursor of DMS, are slaved to the parent ecosystem models. Only the models which include environmental effects on DMS fluxes independently of ecological dynamics can reproduce this summer mismatch between chlorophyll and DMS. A major conclusion of this exercise is that prognostic DMS models need to give more weight to the direct impact of environmental forcing (e.g., irradiance) on DMS dynamics to decouple them from ecological processes.