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.     

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.     

Carbon dioxide release from the North Pacific abyss during the last deglaciation

Atmospheric carbon dioxide concentrations were significantly lower during glacial periods than during intervening interglacial periods, but the mechanisms responsible for this difference remain uncertain. Many recent explanations call on greater carbon storage in a poorly ventilated deep ocean during glacial periods, but direct evidence regarding the ventilation and respired carbon content of the glacial deep ocean is sparse and often equivocal. Here we present sedimentary geochemical records from sites spanning the deep subarctic Pacific that--together with previously published results--show that a poorly ventilated water mass containing a high concentration of respired carbon dioxide occupied the North Pacific abyss during the Last Glacial Maximum. Despite an inferred increase in deep Southern Ocean ventilation during the first step of the deglaciation (18,000-15,000 years ago), we find no evidence for improved ventilation in the abyssal subarctic Pacific until a rapid transition approximately 14,600 years ago: this change was accompanied by an acceleration of export production from the surface waters above but only a small increase in atmospheric carbon dioxide concentration. We speculate that these changes were mechanistically linked to a roughly coeval increase in deep water formation in the North Atlantic, which flushed respired carbon dioxide from northern abyssal waters, but also increased the supply of nutrients to the upper ocean, leading to greater carbon dioxide sequestration at mid-depths and stalling the rise of atmospheric carbon dioxide concentrations. Our findings are qualitatively consistent with hypotheses invoking a deglacial flushing of respired carbon dioxide from an isolated, deep ocean reservoir, but suggest that the reservoir may have been released in stages, as vigorous deep water ventilation switched between North Atlantic and Southern Ocean source regions.     

Long-term decline in krill stock and increase in salps within the Southern Ocean

Antarctic krill (Euphausia superba) and salps (mainly Salpa thompsoni) are major grazers in the Southern Ocean, and krill support commercial fisheries. Their density distributions have been described in the period 1926-51, while recent localized studies suggest short-term changes. To examine spatial and temporal changes over larger scales, we have combined all available scientific net sampling data from 1926 to 2003. This database shows that the productive southwest Atlantic sector contains >50% of Southern Ocean krill stocks, but here their density has declined since the 1970s. Spatially, within their habitat, summer krill density correlates positively with chlorophyll concentrations. Temporally, within the southwest Atlantic, summer krill densities correlate positively with sea-ice extent the previous winter. Summer food and the extent of winter sea ice are thus key factors in the high krill densities observed in the southwest Atlantic Ocean. Krill need the summer phytoplankton blooms of this sector, where winters of extensive sea ice mean plentiful winter food from ice algae, promoting larval recruitment and replenishing the stock. Salps, by contrast, occupy the extensive lower-productivity regions of the Southern Ocean and tolerate warmer water than krill. As krill densities decreased last century, salps appear to have increased in the southern part of their range. These changes have had profound effects within the Southern Ocean food web.     

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.     

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.     

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.     

Export and remineralization of POM in the Southern Ocean and the South China Sea estimated from ~(210)Po/~(210)Pb disequilibria

Disequilibria between 210Po and 210Pb in the upper water and their potential applications as a proxy of particle export and remineralization were examined in the Southern Ocean (station IV3) and the South China Sea (NS44). 210Po was deficit in surface waters but excessive relative to 210Pb in subsurface waters. Good positive correlation between 210Po and particulate organic carbon (POC) indicated deficits and excess of 210Po resulted from particulate organic matter (POM) export and remineralization respecti...     

The Southern Ocean biogeochemical divide

Modelling studies have demonstrated that the nutrient and carbon cycles in the Southern Ocean play a central role in setting the air-sea balance of CO(2) and global biological production. Box model studies first pointed out that an increase in nutrient utilization in the high latitudes results in a strong decrease in the atmospheric carbon dioxide partial pressure (pCO2). This early research led to two important ideas: high latitude regions are more important in determining atmospheric pCO2 than low latitudes, despite their much smaller area, and nutrient utilization and atmospheric pCO2 are tightly linked. Subsequent general circulation model simulations show that the Southern Ocean is the most important high latitude region in controlling pre-industrial atmospheric CO(2) because it serves as a lid to a larger volume of the deep ocean. Other studies point out the crucial role of the Southern Ocean in the uptake and storage of anthropogenic carbon dioxide and in controlling global biological production. Here we probe the system to determine whether certain regions of the Southern Ocean are more critical than others for air-sea CO(2) balance and the biological export production, by increasing surface nutrient drawdown in an ocean general circulation model. We demonstrate that atmospheric CO(2) and global biological export production are controlled by different regions of the Southern Ocean. The air-sea balance of carbon dioxide is controlled mainly by the biological pump and circulation in the Antarctic deep-water formation region, whereas global export production is controlled mainly by the biological pump and circulation in the Subantarctic intermediate and mode water formation region. The existence of this biogeochemical divide separating the Antarctic from the Subantarctic suggests that it may be possible for climate change or human intervention to modify one of these without greatly altering the other.     

Holocene Southern Ocean surface temperature variability west of the Antarctic Peninsula

The disintegration of ice shelves, reduced sea-ice and glacier extent, and shifting ecological zones observed around Antarctica highlight the impact of recent atmospheric and oceanic warming on the cryosphere. Observations and models suggest that oceanic and atmospheric temperature variations at Antarctica's margins affect global cryosphere stability, ocean circulation, sea levels and carbon cycling. In particular, recent climate changes on the Antarctic Peninsula have been dramatic, yet the Holocene climate variability of this region is largely unknown, limiting our ability to evaluate ongoing changes within the context of historical variability and underlying forcing mechanisms. Here we show that surface ocean temperatures at the continental margin of the western Antarctic Peninsula cooled by 3-4 °C over the past 12,000 years, tracking the Holocene decline of local (65° S) spring insolation. Our results, based on TEX(86) sea surface temperature (SST) proxy evidence from a marine sediment core, indicate the importance of regional summer duration as a driver of Antarctic seasonal sea-ice fluctuations. On millennial timescales, abrupt SST fluctuations of 2-4 °C coincide with globally recognized climate variability. Similarities between our SSTs, Southern Hemisphere westerly wind reconstructions and El Ni?o/Southern Oscillation variability indicate that present climate teleconnections between the tropical Pacific Ocean and the western Antarctic Peninsula strengthened late in the Holocene epoch. We conclude that during the Holocene, Southern Ocean temperatures at the western Antarctic Peninsula margin were tied to changes in the position of the westerlies, which have a critical role in global carbon cycling.     

Intensified deep Pacific inflow and ventilation in Pleistocene glacial times

The production of cold, deep waters in the Southern Ocean is an important factor in the Earth's heat budget. The supply of deep water to the Pacific Ocean is presently dominated by a single source, the deep western boundary current east of New Zealand. Here we use sediment records deposited under the influence of this deep western boundary current to reconstruct deep-water properties and speed changes during the Pleistocene epoch. In physical and isotope proxies we find evidence for intensified deep Pacific Ocean inflow and ventilation during the glacial periods of the past 1.2 million years. The changes in throughflow may be directly related to an increased production of Antarctic Bottom Water during glacial times. Possible causes for such an increased bottom-water production include increasing wind strengths in the Southern Ocean or an increase in annual sea-ice formation, leaving dense water after brine rejection and thereby enhancing deep convection. We infer also that the global thermohaline circulation was perturbed significantly during the mid-Pleistocene climate transition between 0.86 and 0.45 million years ago.     

Importance of stirring in the development of an iron-fertilized phytoplankton bloom

The growth of populations is known to be influenced by dispersal, which has often been described as purely diffusive. In the open ocean, however, the tendrils and filaments of phytoplankton populations provide evidence for dispersal by stirring. Despite the apparent importance of horizontal stirring for plankton ecology, this process remains poorly characterized. Here we investigate the development of a discrete phytoplankton bloom, which was initiated by the iron fertilization of a patch of water (7 km in diameter) in the Southern Ocean. Satellite images show a striking, 150-km-long bloom near the experimental site, six weeks after the initial fertilization. We argue that the ribbon-like bloom was produced from the fertilized patch through stirring, growth and diffusion, and we derive an estimate of the stirring rate. In this case, stirring acts as an important control on bloom development, mixing phytoplankton and iron out of the patch, but also entraining silicate. This may have prevented the onset of silicate limitation, and so allowed the bloom to continue for as long as there was sufficient iron. Stirring in the ocean is likely to be variable, so blooms that are initially similar may develop very differently.     

Ocean nutrient ratios governed by plankton biogeography

The major nutrients nitrate and phosphate have one of the strongest correlations in the sea, with a slope similar to the average nitrogen (N) to phosphorus (P) content of plankton biomass (N/P = 16:1). The processes through which this global relationship emerges despite the wide range of N/P ratios at the organism level are not known. Here we use an ocean circulation model and observed nutrient distributions to show that the N/P ratio of biological nutrient removal varies across latitude in Southern Ocean surface waters, from 12:1 in the polar ocean to 20:1 in the sub-Antarctic zone. These variations are governed by regional differences in the species composition of the plankton community. The covariation of dissolved nitrate and phosphate is maintained by ocean circulation, which mixes the shallow subsurface nutrients between distinct biogeographic provinces. Climate-driven shifts in these marine biomes may alter the mean N/P ratio and the associated carbon export by Southern Ocean ecosystems.     

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.     

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.     

Southern Ocean dust-climate coupling over the past four million years

Dust has the potential to modify global climate by influencing the radiative balance of the atmosphere and by supplying iron and other essential limiting micronutrients to the ocean. Indeed, dust supply to the Southern Ocean increases during ice ages, and 'iron fertilization' of the subantarctic zone may have contributed up to 40?parts per million by volume (p.p.m.v.) of the decrease (80-100 p.p.m.v.) in atmospheric carbon dioxide observed during late Pleistocene glacial cycles. So far, however, the magnitude of Southern Ocean dust deposition in earlier times and its role in the development and evolution of Pleistocene glacial cycles have remained unclear. Here we report a high-resolution record of dust and iron supply to the Southern Ocean over the past four million years, derived from the analysis of marine sediments from ODP Site 1090, located in the Atlantic sector of the subantarctic zone. The close correspondence of our dust and iron deposition records with Antarctic ice core reconstructions of dust flux covering the past 800,000 years (refs 8, 9) indicates that both of these archives record large-scale deposition changes that should apply to most of the Southern Ocean, validating previous interpretations of the ice core data. The extension of the record beyond the interval covered by the Antarctic ice cores reveals that, in contrast to the relatively gradual intensification of glacial cycles over the past three million years, Southern Ocean dust and iron flux rose sharply at the Mid-Pleistocene climatic transition around 1.25 million years ago. This finding complements previous observations over late Pleistocene glacial cycles, providing new evidence of a tight connection between high dust input to the Southern Ocean and the emergence of the deep glaciations that characterize the past one million years of Earth history.     

Oceanic Cd/P ratio and nutrient utilization in the glacial Southern Ocean

During glacial periods, low atmospheric carbon dioxide concentration has been associated with increased oceanic carbon uptake, particularly in the southern oceans. The mechanism involved remains unclear. Because ocean productivity is strongly influenced by nutrient levels, palaeo-oceanographic proxies have been applied to investigate nutrient utilization in surface water across glacial transitions. Here we show that present-day cadmium and phosphorus concentrations in the global oceans can be explained by a chemical fractionation during particle formation, whereby uptake of cadmium occurs in preference to uptake of phosphorus. This allows the reconstruction of past surface water phosphate concentrations from the cadmium/calcium ratio of planktonic foraminifera. Results from the Last Glacial Maximum show similar phosphate utilization in the subantarctic to that of today, but much smaller utilization in the polar Southern Ocean, in a model that is consistent with the expansion of glacial sea ice and which can reconcile all proxy records of polar nutrient utilization. By restricting communication between the ocean and atmosphere, sea ice expansion also provides a mechanism for reduced CO2 release by the Southern Ocean and lower glacial atmospheric CO2.     

城市湿地生态系统服务动态演化及其权衡关系

以杭州西溪湿地为例，基于1984—2018年的高分辨率遥感数据，利用InVEST模型和当量因子法核算了该地区生态系统服务的动态变化，分析了生态系统服务间的权衡关系．结果表明:1）除固碳服务外，研究期内西溪湿地生态系统服务整体呈退化趋势；2）国家湿地公园的建设，使公园内外固碳服务与生物多样性服务呈现不同的变化趋势，这2种服务类型在公园内呈现增加、而公园外受城市化过程持续影响呈现降低趋势；3）受降水量增加、不透水地面扩大等因素影响，公园内外的产水量与营养物输出量等均呈现增加趋势，旅游开发和生态保护之间的矛盾并没有完全缓解；4）生态系统服务之间存在显著的权衡关系，主要是产水服务与固碳服务、生物多样性保护服务、娱乐和文化服务之间，空间上在城市不透水地面周围更为显著．      

Evidence for deep-water production in the North Pacific Ocean during the early Cenozoic warm interval

The deep-ocean circulation is responsible for a significant component of global heat transport. In the present mode of circulation, deep waters form in the North Atlantic and Southern oceans where surface water becomes sufficiently cold and dense to sink. Polar temperatures during the warmest climatic interval of the Cenozoic era (approximately 65 to 40 million years (Myr) ago) were significantly warmer than today, and this may have been a consequence of enhanced oceanic heat transport. However, understanding the relationship between deep-ocean circulation and ancient climate is complicated by differences in oceanic gateways, which affect where deep waters form and how they circulate. Here I report records of neodymium isotopes from two cores in the Pacific Ocean that indicate a shift in deep-water production from the Southern Ocean to the North Pacific approximately 65 Myr ago. The source of deep waters reverted back to the Southern Ocean 40 Myr ago. The relative timing of changes in the neodymium and oxygen isotope records indicates that changes in Cenozoic deep-water circulation patterns were the consequence, not the cause, of extreme Cenozoic warmth.     

Biological productivity and carbon cycling in the Arctic Ocean

Primary production, bacterial production, particulate organic carbon fluxes and organic carbon burial rates were quantified during the summer period of 1999 in the Arctic Ocean via 14C uptake, 3H uptake, 234Th/238U disequilibrium and 210Pbex dating, respectively. The integrated primary production in the water column was as high as 197 mmolC/(m2@d) in the Chukchi shelf and was 3.8 mmolC/(m2@d) in the Canada Basin. These rates are higher than those reported previously. The ratios of bacterial production to primary production in the study region were higher than 0.5, indicating that microbial activity is not depressed but important in cold Arctic waters. 234Th/238U disequilibria were evident at the station in the Canada Basin. The presence of significant 234Th deficiency suggested that scavenging and removal processes are also important to biogeochemical cycles of trace elements in the Arctic Ocean. Particulate organic carbon export flux was estimated to be 1.0 mmolC/(m2@d). Measurements of sediment excess 210Pb profile in the Chukchi shelf allowed us to estimate the amount of organic carbon buried in the bottom sediment, which ranged from 25 to 35 mmolC/(m2@d) and represented about 59%-82% of the mean primary production in the euphotic zone. Overall, our results indicated that the Arctic Ocean has active carbon cycling and is not a biological desert as previously believed. Therefore, the Arctic Ocean may play an important role in the global carbon cycle and climate change.