Effect of natural iron fertilization on carbon sequestration in the Southern Ocean

The availability of iron limits primary productivity and the associated uptake of carbon over large areas of the ocean. Iron thus plays an important role in the carbon cycle, and changes in its supply to the surface ocean may have had a significant effect on atmospheric carbon dioxide concentrations over glacial-interglacial cycles. To date, the role of iron in carbon cycling has largely been assessed using short-term iron-addition experiments. It is difficult, however, to reliably assess the magnitude of carbon export to the ocean interior using such methods, and the short observational periods preclude extrapolation of the results to longer timescales. Here we report observations of a phytoplankton bloom induced by natural iron fertilization--an approach that offers the opportunity to overcome some of the limitations of short-term experiments. We found that a large phytoplankton bloom over the Kerguelen plateau in the Southern Ocean was sustained by the supply of iron and major nutrients to surface waters from iron-rich deep water below. The efficiency of fertilization, defined as the ratio of the carbon export to the amount of iron supplied, was at least ten times higher than previous estimates from short-term blooms induced by iron-addition experiments. This result sheds new light on the effect of long-term fertilization by iron and macronutrients on carbon sequestration, suggesting that changes in iron supply from below--as invoked in some palaeoclimatic and future climate change scenarios--may have a more significant effect on atmospheric carbon dioxide concentrations than previously thought.     

Deep carbon export from a Southern Ocean iron-fertilized diatom bloom

Fertilization of the ocean by adding iron compounds has induced diatom-dominated phytoplankton blooms accompanied by considerable carbon dioxide drawdown in the ocean surface layer. However, because the fate of bloom biomass could not be adequately resolved in these experiments, the timescales of carbon sequestration from the atmosphere are uncertain. Here we report the results of a five-week experiment carried out in the closed core of a vertically coherent, mesoscale eddy of the Antarctic Circumpolar Current, during which we tracked sinking particles from the surface to the deep-sea floor. A large diatom bloom peaked in the fourth week after fertilization. This was followed by mass mortality of several diatom species that formed rapidly sinking, mucilaginous aggregates of entangled cells and chains. Taken together, multiple lines of evidence-although each with important uncertainties-lead us to conclude that at least half the bloom biomass sank far below a depth of 1,000 metres and that a substantial portion is likely to have reached the sea floor. Thus, iron-fertilized diatom blooms may sequester carbon for timescales of centuries in ocean bottom water and for longer in the sediments.     

Diatom carbon export enhanced by silicate upwelling in the northeast Atlantic

Diatoms are unicellular or chain-forming phytoplankton that use silicon (Si) in cell wall construction. Their survival during periods of apparent nutrient exhaustion enhances carbon sequestration in frontal regions of the northern North Atlantic. These regions may therefore have a more important role in the 'biological pump' than they have previously been attributed, but how this is achieved is unknown. Diatom growth depends on silicate availability, in addition to nitrate and phosphate, but northern Atlantic waters are richer in nitrate than silicate. Following the spring stratification, diatoms are the first phytoplankton to bloom. Once silicate is exhausted, diatom blooms subside in a major export event. Here we show that, with nitrate still available for new production, the diatom bloom is prolonged where there is a periodic supply of new silicate: specifically, diatoms thrive by 'mining' deep-water silicate brought to the surface by an unstable ocean front. The mechanism we present here is not limited to silicate fertilization; similar mechanisms could support nitrate-, phosphate- or iron-limited frontal regions in oceans elsewhere.     

A mesoscale phytoplankton bloom in the polar Southern Ocean stimulated by iron fertilization

Changes in iron supply to oceanic plankton are thought to have a significant effect on concentrations of atmospheric carbon dioxide by altering rates of carbon sequestration, a theory known as the 'iron hypothesis'. For this reason, it is important to understand the response of pelagic biota to increased iron supply. Here we report the results of a mesoscale iron fertilization experiment in the polar Southern Ocean, where the potential to sequester iron-elevated algal carbon is probably greatest. Increased iron supply led to elevated phytoplankton biomass and rates of photosynthesis in surface waters, causing a large drawdown of carbon dioxide and macronutrients, and elevated dimethyl sulphide levels after 13 days. This drawdown was mostly due to the proliferation of diatom stocks. But downward export of biogenic carbon was not increased. Moreover, satellite observations of this massive bloom 30 days later, suggest that a sufficient proportion of the added iron was retained in surface waters. Our findings demonstrate that iron supply controls phytoplankton growth and community composition during summer in these polar Southern Ocean waters, but the fate of algal carbon remains unknown and depends on the interplay between the processes controlling export, remineralisation and timescales of water mass subduction.     

Controls on tropical Pacific Ocean productivity revealed through nutrient stress diagnostics

In situ enrichment experiments have shown that the growth of bloom-forming diatoms in the major high-nitrate low-chlorophyll (HNLC) regions of the world's oceans is limited by the availability of iron. Yet even the largest of these manipulative experiments represents only a small fraction of an ocean basin, and the responses observed are strongly influenced by the proliferation of rare species rather than the growth of naturally dominant populations. Here we link unique fluorescence attributes of phytoplankton to specific physiological responses to nutrient stress, and use these relationships to evaluate the factors that constrain phytoplankton growth in the tropical Pacific Ocean on an unprecedented spatial scale. On the basis of fluorescence measurements taken over 12 years, we delineate three major ecophysiological regimes in this region. We find that iron has a key function in regulating phytoplankton growth in both HNLC and oligotrophic waters near the Equator and further south, whereas nitrogen and zooplankton grazing are the primary factors that regulate biomass production in the north. Application of our findings to the interpretation of satellite chlorophyll fields shows that productivity in the tropical Pacific basin may be 1.2-2.5 Pg C yr(-1) lower than previous estimates have suggested, a difference that is comparable to the global change in ocean production that accompanied the largest El Ni?o to La Ni?a transition on record.     

Phosphorus limitation of nitrogen fixation by Trichodesmium in the central Atlantic Ocean

Marine fixation of atmospheric nitrogen is believed to be an important source of biologically useful nitrogen to ocean surface waters, stimulating productivity of phytoplankton and so influencing the global carbon cycle. The majority of nitrogen fixation in tropical waters is carried out by the marine cyanobacterium Trichodesmium, which supplies more than half of the new nitrogen used for primary production. Although the factors controlling marine nitrogen fixation remain poorly understood, it has been thought that nitrogen fixation is limited by iron availability in the ocean. This was inferred from the high iron requirement estimated for growth of nitrogen fixing organisms and the higher apparent densities of Trichodesmium where aeolian iron inputs are plentiful. Here we report that nitrogen fixation rates in the central Atlantic appear to be independent of both dissolved iron levels in sea water and iron content in Trichodesmium colonies. Nitrogen fixation was, instead, highly correlated to the phosphorus content of Trichodesmium and was enhanced at higher irradiance. Furthermore, our calculations suggest that the structural iron requirement for the growth of nitrogen-fixing organisms is much lower than previously calculated. Although iron deficiency could still potentially limit growth of nitrogen-fixing organisms in regions of low iron availability-for example, in the subtropical North Pacific Ocean-our observations suggest that marine nitrogen fixation is not solely regulated by iron supply.     

A massive phytoplankton bloom induced by an ecosystem-scale iron fertilization experiment in the equatorial Pacific Ocean

The seeding of an expanse of surface waters in the equatorial Pacific Ocean with low concentrations of dissolved iron triggered a massive phytoplankton bloom which consumed large quantities of carbon dioxide and nitrate that these microscopic plants cannot fully utilize under natural conditions. These and other observations provide unequivocal support for the hypothesis that phytoplankton growth in this oceanic region is limited by iron bioavailability.     

Dust storm in Asia continent and its bio-environmental effects in the North Pacific： A case study of the strongest dust event in April, 2001 in central Asia

The iron hypothesis, first proposed by John Martin in 1990[1], suggests that some surface oceans such as North and Equatorial Pacific Oceans have high nutrient but low chlorophyll (HNLC). Thus iron coming from terrestrial dusts is the primary factor limit…     

Agricultural runoff fuels large phytoplankton blooms in vulnerable areas of the ocean

Biological productivity in most of the world's oceans is controlled by the supply of nutrients to surface waters. The relative balance between supply and removal of nutrients--including nitrogen, iron and phosphorus--determines which nutrient limits phytoplankton growth. Although nitrogen limits productivity in much of the ocean, large portions of the tropics and subtropics are defined by extreme nitrogen depletion. In these regions, microbial denitrification removes biologically available forms of nitrogen from the water column, producing substantial deficits relative to other nutrients. Here we demonstrate that nitrogen-deficient areas of the tropical and subtropical oceans are acutely vulnerable to nitrogen pollution. Despite naturally high nutrient concentrations and productivity, nitrogen-rich agricultural runoff fuels large (54-577 km2) phytoplankton blooms in the Gulf of California. Runoff exerts a strong and consistent influence on biological processes, in 80% of cases stimulating blooms within days of fertilization and irrigation of agricultural fields. We project that by the year 2050, 27-59% of all nitrogen fertilizer will be applied in developing regions located upstream of nitrogen-deficient marine ecosystems. Our findings highlight the present and future vulnerability of these ecosystems to agricultural runoff.     

Effect of iron supply on Southern Ocean CO2 uptake and implications for glacial atmospheric CO2

Photosynthesis by marine phytoplankton in the Southern Ocean, and the associated uptake of carbon, is thought to be currently limited by the availability of iron. One implication of this limitation is that a larger iron supply to the region in glacial times could have stimulated algal photosynthesis, leading to lower concentrations of atmospheric CO2. Similarly, it has been proposed that artificial iron fertilization of the oceans might increase future carbon sequestration. Here we report data from a whole-ecosystem test of the iron-limitation hypothesis in the Southern Ocean, which show that surface uptake of atmospheric CO2 and uptake ratios of silica to carbon by phytoplankton were strongly influenced by nanomolar increases of iron concentration. We use these results to inform a model of global carbon and ocean nutrients, forced with atmospheric iron fluxes to the region derived from the Vostok ice-core dust record. During glacial periods, predicted magnitudes and timings of atmospheric CO2 changes match ice-core records well. At glacial terminations, the model suggests that forcing of Southern Ocean biota by iron caused the initial approximately 40 p.p.m. of glacial-interglacial CO2 change, but other mechanisms must have accounted for the remaining 40 p.p.m. increase. The experiment also confirms that modest sequestration of atmospheric CO2 by artificial additions of iron to the Southern Ocean is in principle possible, although the period and geographical extent over which sequestration would be effective remain poorly known.     

Polysaccharide aggregation as a potential sink of marine dissolved organic carbon

The formation and sinking of biogenic particles mediate vertical mass fluxes and drive elemental cycling in the ocean. Whereas marine sciences have focused primarily on particle production by phytoplankton growth, particle formation by the assembly of organic macromolecules has almost been neglected. Here we show, by means of a combined experimental and modelling study, that the formation of polysaccharide particles is an important pathway to convert dissolved into particulate organic carbon during phytoplankton blooms, and can be described in terms of aggregation kinetics. Our findings suggest that aggregation processes in the ocean cascade from the molecular scale up to the size of fast-settling particles, and give new insights into the cycling and export of biogeochemical key elements such as carbon, iron and thorium.     

Evidence against dust-mediated control of glacial-interglacial changes in atmospheric CO2

The low concentration of atmospheric CO2 inferred to have been present during glacial periods is thought to have been partly caused by an increased supply of iron-bearing dust to the ocean surface. This is supported by a recent model that attributes half of the CO2 reduction during past glacial stages to iron-stimulated uptake of CO2 by phytoplankton in the Southern Ocean. But atmospheric dust fluxes to the Southern Ocean, even in glacial periods, are thought to be relatively low and therefore it has been proposed that Southern Ocean productivity might be influenced by iron deposited elsewhere-for example, in the Northern Hemisphere-which is then transported south via ocean circulation (similar to the distal supply of iron to the equatorial Pacific Ocean). Here we examine the timing of dust fluxes to the North Atlantic Ocean, in relation to climate records from the Vostok ice core in Antarctica around the time of the penultimate deglaciation (about 130 kyr ago). Two main dust peaks occurred 155 kyr and 130 kyr ago, but neither was associated with the CO2 rise recorded in the Vostok ice core. This mismatch, together with the low dust flux supplied to the Southern Ocean, suggests that dust-mediated iron fertilization of the Southern Ocean did not significantly influence atmospheric CO2 at the termination of the penultimate glaciation.     

Effect of trace metal availability on coccolithophorid calcification

The deposition of atmospheric dust into the ocean has varied considerably over geological time. Because some of the trace metals contained in dust are essential plant nutrients which can limit phytoplankton growth in parts of the ocean, it has been suggested that variations in dust supply to the surface ocean might influence primary production. Whereas the role of trace metal availability in photosynthetic carbon fixation has received considerable attention, its effect on biogenic calcification is virtually unknown. The production of both particulate organic carbon and calcium carbonate (CaCO3) drives the ocean's biological carbon pump. The ratio of particulate organic carbon to CaCO3 export, the so-called rain ratio, is one of the factors determining CO2 sequestration in the deep ocean. Here we investigate the influence of the essential trace metals iron and zinc on the prominent CaCO3-producing microalga Emiliania huxleyi. We show that whereas at low iron concentrations growth and calcification are equally reduced, low zinc concentrations result in a de-coupling of the two processes. Despite the reduced growth rate of zinc-limited cells, CaCO3 production rates per cell remain unaffected, thus leading to highly calcified cells. These results suggest that changes in dust deposition can affect biogenic calcification in oceanic regions characterized by trace metal limitation, with possible consequences for CO2 partitioning between the atmosphere and the ocean.     

The biogeochemical behavior of dissolved aluminum in the southern Yellow Sea: Influence of the spring phytoplankton bloom

Many recent studies have investigated the nutrient-type profiles of dissolved aluminum(Al) in the ocean.Significant scavenging of dissolved Al can occur during phytoplankton blooms,but the mechanism remains unclear.The distribution of dissolved Al in the southern Yellow Sea(SYS) was investigated in winter and spring 2009.Following measurements at grid stations during the spring sampling cruise,two drifting anchor surveys of more than 100 h were conducted to trace the variation of dissolved Al concentration during the spring phytoplankton bloom(SPB).The concentration of dissolved Al in the SYS decreased from 40 nmol/L in February to 30 nmol/L in March and 10-20 nmol/L in April,while the concentration of Chl a increased from < 2 μg/L in March to > 4 μg/L in April.The concentration of dissolved Al in the SYS decreased significantly with the development of the phytoplankton bloom,which indicated biological scavenging of dissolved Al from water column.The proportion of dissolved Al scavenged from water column by different phytoplankton species differed at the two drifting stations,with greater removal efficiency demonstrated by diatoms than dinoflagellates.Phytoplankton samples collected from the Chl a maximum layer were washed with trace metal clean reagent(oxalate-EDTA-citrate,abbreviate as oxalate solution,Tovar-Sanchez et al.,2003) to enable the surface-scavenged(extracellular) and intracellular Al pools associated with phytoplankton to be differentiated.Thirty-nine to ninetysix percent of the total Al was found to be existed in the interior pools,which indicated that biological absorption was the important way to scavenge dissolved Al during phytoplankton blooms in the SYS.     

Ocean currents mediate evolution in island lizards

Islands are considered to be natural laboratories in which to examine evolution because of the implicit assumption that limited gene flow allows tests of evolutionary processes in isolated replicates. Here we show that this well-accepted idea requires re-examination. Island inundation during hurricanes can have devastating effects on lizard populations in the Bahamas. After severe storms, islands may be recolonized by over-water dispersal of lizards from neighbouring islands. High levels of gene flow may homogenize genes responsible for divergence, and are widely viewed as a constraining force on evolution. Ultimately, the magnitude of gene flow determines the extent to which populations diverge from one another, and whether or not they eventually form new species. We show that patterns of gene flow among island populations of Anolis lizards are best explained by prevailing ocean currents, and that over-water dispersal has evolutionary consequences. Across islands, divergence in fitness-related morphology decreases with increasing gene flow. Results suggest that over-water dispersal after hurricanes constrains adaptive diversification in Anolis lizards, and that it may have an important but previously undocumented role in this classical example of adaptive radiation.     

Copepod hatching success in marine ecosystems with high diatom concentrations

Diatoms dominate spring bloom phytoplankton assemblages in temperate waters and coastal upwelling regions of the global ocean. Copepods usually dominate the zooplankton in these regions and are the prey of many larval fish species. Recent laboratory studies suggest that diatoms may have a deleterious effect on the success of copepod egg hatching. These findings challenge the classical view of marine food-web energy flow from diatoms to fish by means of copepods. Egg mortality is an important factor in copepod population dynamics, thus, if diatoms have a deleterious in situ effect, paradoxically, high diatom abundance could limit secondary production. Therefore, the current understanding of energy transfer from primary production to fisheries in some of the most productive and economically important marine ecosystems may be seriously flawed. Here we present in situ estimates of copepod egg hatching success from twelve globally distributed areas, where diatoms dominate the phytoplankton assemblage. We did not observe a negative relationship between copepod egg hatching success and either diatom biomass or dominance in the microplankton in any of these regions. The classical model for diatom-dominated system remains valid.     

Climate-driven trends in contemporary ocean productivity

Contributing roughly half of the biosphere's net primary production (NPP), photosynthesis by oceanic phytoplankton is a vital link in the cycling of carbon between living and inorganic stocks. Each day, more than a hundred million tons of carbon in the form of CO2 are fixed into organic material by these ubiquitous, microscopic plants of the upper ocean, and each day a similar amount of organic carbon is transferred into marine ecosystems by sinking and grazing. The distribution of phytoplankton biomass and NPP is defined by the availability of light and nutrients (nitrogen, phosphate, iron). These growth-limiting factors are in turn regulated by physical processes of ocean circulation, mixed-layer dynamics, upwelling, atmospheric dust deposition, and the solar cycle. Satellite measurements of ocean colour provide a means of quantifying ocean productivity on a global scale and linking its variability to environmental factors. Here we describe global ocean NPP changes detected from space over the past decade. The period is dominated by an initial increase in NPP of 1,930 teragrams of carbon a year (Tg C yr(-1)), followed by a prolonged decrease averaging 190 Tg C yr(-1). These trends are driven by changes occurring in the expansive stratified low-latitude oceans and are tightly coupled to coincident climate variability. This link between the physical environment and ocean biology functions through changes in upper-ocean temperature and stratification, which influence the availability of nutrients for phytoplankton growth. The observed reductions in ocean productivity during the recent post-1999 warming period provide insight on how future climate change can alter marine food webs.     

环剥和针刺对李子生长发育的影响

于盛花期和花后１５ｄ对李子进行主干环剥，可有效地抑制新梢生长，提高坐果率和干物重，促进果实生长，两次环剥处理以盛花期剥效果较好。剥后叶内钙、铁含量降低，锌含量变化不明显。此外，针刺对李子的营养生长也有显著抑制作用。     

The decline and fate of an iron-induced subarctic phytoplankton bloom

Iron supply has a key role in stimulating phytoplankton blooms in high-nitrate low-chlorophyll oceanic waters. However, the fate of the carbon fixed by these blooms, and how efficiently it is exported into the ocean's interior, remains largely unknown. Here we report on the decline and fate of an iron-stimulated diatom bloom in the Gulf of Alaska. The bloom terminated on day 18, following the depletion of iron and then silicic acid, after which mixed-layer particulate organic carbon (POC) concentrations declined over six days. Increased particulate silica export via sinking diatoms was recorded in sediment traps at depths between 50 and 125 m from day 21, yet increased POC export was not evident until day 24. Only a small proportion of the mixed-layer POC was intercepted by the traps, with more than half of the mixed-layer POC deficit attributable to bacterial remineralization and mesozooplankton grazing. The depletion of silicic acid and the inefficient transfer of iron-increased POC below the permanent thermocline have major implications both for the biogeochemical interpretation of times of greater iron supply in the geological past, and also for proposed geo-engineering schemes to increase oceanic carbon sequestration.     

Spatial synchronization of vole population dynamics by predatory birds

Northern vole populations exhibit large-scale, spatially synchronous population dynamics. Such cases of population synchrony provide excellent opportunities for distinguishing between local intrinsic and regional extrinsic mechanisms of population regulation. Analyses of large-scale survey data and theoretical modelling have indicated several plausible synchronizing mechanisms. It is difficult, however, to determine the most important one without detailed data on local demographic processes. Here we combine results from two field studies in southeastern Norway--one identifies local demographic mechanisms and landscape-level annual synchrony among 28 enclosed experimental populations and the other examines region-level multi-annual synchrony in open natural populations. Despite fences eliminating predatory mammals and vole dispersal, the growth rates of the experimental populations were synchronized and moreover, perfectly linked with vole abundance in the region. The fates of 481 radio-marked voles showed that bird predation was the synchronizing mechanism. A higher frequency of risky dispersal movements in slowly growing populations appeared to accelerate predation rate. Thus, dispersal may induce a feedback-loop between predation and population growth that enhances synchrony.