Individual and interactive effects of ocean acidification,global warming,and UV radiation on phytoplankton

Rising carbon dioxide (CO₂) concentrations in the atmosphere result in increasing global temperatures and ocean warming (OW). Concomitantly, dissolution of anthropogenic CO₂ declines seawater pH, resulting in ocean acidification (OA) and altering marine chemical environments. The marine biological carbon pump driven by marine photosynthesis plays an important role for oceanic carbon sinks. Therefore, how ocean climate changes affect the amount of carbon fixation by primary producers is closely related to future ocean carbon uptake. OA may upregulate metabolic pathways in phytoplankton, such as upregulating ß-oxidation and the tricarboxylic acid cycle, resulting in increased accumulation of toxic phenolic compounds. Ocean warming decreases global phytoplankton productivity; however, regionally, it may stimulate primary productivity and change phytoplankton community composition, due to different physical and chemical environmental requirements of species. It is still controversial how OA and OW interactively affect marine carbon fixation by photosynthetic organisms. OA impairs the process of calcification in calcifying phytoplankton and aggravate ultraviolet (UV)-induced harms to the cells. Increasing temperatures enhance the activity of cellular repair mechanisms, which mitigates UV-induced damage. The effects of OA, warming, enhanced exposure to UV-B as well as the interactions of these environmental stress factors on phytoplankton productivity and community composition, are discussed in this review.     

Oceanic sinks for atmospheric CO2

There is approximately 50 times more inorganic carbon in the global ocean than in the atmosphere. On time scales of decades to millions of years, the interaction between these two geophysical fluids determines atmospheric CO₂ levels. During glacial periods, for example, the ocean serves as the major sink for atmospheric CO₂, while during glacial–interglacial transitions, it is a source of CO₂ to the atmosphere. The mechanisms responsible for determining the sign of the net exchange of CO₂ between the ocean and the atmosphere remain unresolved. There is evidence that during glacial periods, phytoplankton primary productivity increased, leading to an enhanced sedimentation of particulate organic carbon into the ocean interior. The stimulation of primary production in glacial episodes can be correlated with increased inputs of nutrients limiting productivity, especially aeolian iron. Iron directly enhances primary production in high nutrient (nitrate and phosphate) regions of the ocean, of which the Southern Ocean is the most important. This trace element can also enhance nitrogen fixation, and thereby indirectly stimulate primary production throughout the low nutrient regions of the central ocean basins. While the export flux of organic carbon to the ocean interior was enhanced during glacial periods, this process does not fully account for the sequestration of atmospheric CO₂. Heterotrophic oxidation of the newly formed organic carbon, forming weak acids, would have hydrolyzed CaCO₃ in the sediments, increasing thereby oceanic alkalinity which, in turn, would have promoted the drawdown of atmospheric CO₂. This latter mechanism is consistent with the stable carbon isotope pattern derived from air trapped in ice cores. The oceans have also played a major role as a sink for up to 30% of the anthropogenic CO₂ produced during the industrial revolution. In large part this is due to CO₂ solution in the surface ocean; however, some, poorly quantified fraction is a result of increased new production due to anthropogenic inputs of combined N, P and Fe. Based on ‘circulation as usual’, models predict that future anthropogenic CO₂ inputs to the atmosphere will, in part, continue to be sequestered in the ocean. Human intervention (large-scale Fe fertilization; direct CO₂ burial in the deep ocean) could increase carbon sequestration in the oceans, but could also result in unpredicted environmental perturbations. Changes in the oceanic thermohaline circulation as a result of global climate change would greatly alter the predictions of C sequestration that are possible on a ‘circulation as usual’ basis.     

127 Climate Variability and Phytoplankton Production in PNW Coast Estuaries

The west coast of North America receives strong forcing from climate modes such as El Niño-Southern Oscillation and the Pacific Decadal Oscillation. Estuaries are poised at a sensitive interface because estuarine biota and habitat will be affected by variability in properties and processes associated with the ocean, the watershed, and the local weather. Observations from the Washington coast and Willapa Bay illustrate these three arenas of influence. Variation in ocean upwelling and ocean thermocline depth associated with the 1997–98 El Niño versus the 1999 La Niña affected temperature and nutrient availability in Willapa Bay. Variation in river flow associated with the 2000–01 drought affected estuarine circulation and residence time. And, variation in prevailing wind direction and/or cloudiness was highly correlated with phytoplankton biomass. This situation is responsible for the complexity of understanding climate impacts on estuarine systems. In order to help evaluate which mechanisms, remote oceanic processes or local watershed/estuarine characteristics, most affected Willapa Bay's phytoplankton production, several phytoplankton species were used as indicators of water mass origin and compared with primary productivity data to assess whether phytoplankton blooms were dominated by endemic or imported species. Our analysis resolved that the highest primary production (and the appearance of Pseudo-nitzschia spp.) was associated with oceanic intrusions of phytoplankton biomass into Willapa Bay. This result underscores the dominant role that variation in ocean and climate play in controlling Pacific Northwest estuarine production. However, while the largest blooms were oceanic in origin, numerous medium-sized production events were from either oceanic or estuarine sources, indicating a complex picture.     

Ocean productivity and climate change

Satellite measurements and the development of new techniques have confirmed the importance of ocean biology in controlling the carbon dioxide (CO(2)) content of the atmosphere. The marine sedimentary record shows that climate change and the ocean carbon cycle are closely linked: during glacial periods, marine productivity was enhanced and atmospheric CO(2) levels were reduced. Global warming may have the opposite effect, with reduced uptake of CO(2) exacerbating the problems of climate change.     

Phytoplankton decline in the eastern North Pacific transition zone associated with atmospheric blocking

Global climate change can significantly influence oceanic phytoplankton dynamics, and thus biogeochemical cycles and marine food webs. However, associative explanations based on the correlation between chlorophyll‐a concentration (Chl‐a) and climatic indices is inadequate to describe the mechanism of the connection between climate change, large‐scale atmospheric dynamics, and phytoplankton variability. Here, by analyzing multiple satellite observations of Chl‐a and atmospheric conditions from National Center for Environmental Prediction/National Center for Atmospheric Research reanalysis datasets, we show that high‐latitude atmospheric blocking events over Alaska are the primary drivers of the recent decline of Chl‐a in the eastern North Pacific transition zone. These blocking events were associated with the persistence of large‐scale atmosphere pressure fields that decreased westerly winds and southward Ekman transport over the subarctic ocean gyre. Reduced southward Ekman transport leads to reductions in nutrient availability to phytoplankton in the transition zone. The findings describe a previously unidentified climatic factor that contributed to the recent decline of phytoplankton in this region and propose a mechanism of the top‐down teleconnection between the high‐latitude atmospheric circulation anomalies and the subtropical oceanic primary productivity. The results also highlight the importance of understanding teleconnection among atmosphere–ocean interactions as a means to anticipate future climate change impacts on oceanic primary production.     

Effects of biotic disturbances on forest carbon cycling in the United States and Canada

Forest insects and pathogens are major disturbance agents that have affected millions of hectares in North America in recent decades, implying significant impacts to the carbon (C) cycle. Here, we review and synthesize published studies of the effects of biotic disturbances on forest C cycling in the United States and Canada. Primary productivity in stands was reduced, sometimes considerably, immediately following insect or pathogen attack. After repeated growth reductions caused by some insects or pathogens or a single infestation by some bark beetle species, tree mortality occurred, altering productivity and decomposition. In the years following disturbance, primary productivity in some cases increased rapidly as a result of enhanced growth by surviving vegetation, and in other cases increased slowly because of lower forest regrowth. In the decades following tree mortality, decomposition increased as a result of the large amount of dead organic matter. Net ecosystem productivity decreased immediately following attack, with some studies reporting a switch to a C source to the atmosphere, and increased afterward as the forest regrew and dead organic matter decomposed. Large variability in C cycle responses arose from several factors, including type of insect or pathogen, time since disturbance, number of trees affected, and capacity of remaining vegetation to increase growth rates following outbreak. We identified significant knowledge gaps, including limited understanding of carbon cycle impacts among different biotic disturbance types (particularly pathogens), their impacts at landscape and regional scales, and limited capacity to predict disturbance events and their consequences for carbon cycling. We conclude that biotic disturbances can have major impacts on forest C stocks and fluxes and can be large enough to affect regional C cycling. However, additional research is needed to reduce the uncertainties associated with quantifying biotic disturbance effects on the North American C budget.     

Vegetation-atmosphere interactions and their role in global warming during the latest Cretaceous

Forest vegetation has the ability to warm Recent climate by its effects on albedo and atmospheric water vapour, but the role of vegetation in warming climates of the geologic past is poorly understood. This study evaluates the role of forest vegetation in maintaining warm climates of the Late Cretaceous by (1) reconstructing global palaeovegetation for the latest Cretaceous (Maastrichtian); (2) modelling latest Cretaceous climate under unvegetated conditions and different distributions of palaeovegetation; and (3) comparing model output with a global database of palaeoclimatic indicators. Simulation of Maastrichtian climate with the land surface coded as bare soil produces high-latitude temperatures that are too cold to explain the documented palaeogeographic distribution of forest and woodland vegetation. In contrast, simulations that include forest vegetation at high latitudes show significantly warmer temperatures that are sufficient to explain the widespread geographic distribution of high-latitude deciduous forests. These warmer temperatures result from decreased albedo and feedbacks between the land surface and adjacent oceans. Prescribing a realistic distribution of palaeovegetation in model simulations produces the best agreement between simulated climate and the geologic record of palaeoclimatic indicators. Positive feedbacks between high-latitude forests, the atmosphere, and ocean contributed significantly to high-latitude warming during the latest Cretaceous, and imply that high-latitude forest vegetation was an important source of polar warmth during other warm periods of geologic history.     

Physical conditions on the early Earth

The formation of the Earth as a planet was a large stochastic process in which the rapid assembly of asteroidal-to-Mars-sized bodies was followed by a more extended period of growth through collisions of these objects, facilitated by the gravitational perturbations associated with Jupiter. The Earth's inventory of water and organic molecules may have come from diverse sources, not more than 10% roughly from comets, the rest from asteroidal precursors to chondritic bodies and possibly objects near Earth's orbit for which no representative class of meteorites exists today in laboratory collections. The final assembly of the Earth included a catastrophic impact with a Mars-sized body, ejecting mantle and crustal material to form the Moon, and also devolatilizing part of the Earth. A magma ocean and steam atmosphere (possibly with silica vapour) existed briefly in this period, but terrestrial surface waters were below the critical point within 100 million years after Earth's formation, and liquid water existed continuously on the surface within a few hundred million years. Organic material delivered by comets and asteroids would have survived, in part, this violent early period, but frequent impacts of remaining debris probably prevented the continuous habitability of the Earth for one to several hundred million years. Planetary analogues to or records of this early time when life began include Io (heat flow), Titan (organic chemistry) and Venus (remnant early granites).     

The plankton multiplier--positive feedback in the greenhouse

The plankton multiplier is a positive feedback mechanism linkingthe greenhouse effect and biological pump (Woods.J.D., RoyalCommission on Environmental Pollution, 1990). As pollution increasesthe atmospheric concentration of carbon dioxide, the enhancedgreenhouse effect induces radiative forcing of the ocean, whichdiminishes the depth of winter convection, reducing the annualresupply of nutrients to the euphotic zone and therefore theannual primary production. That weakens the biological pump,which contributes to oceanic uptake of CO₂,. As the ocean takesup less CO₂, more remains in the atmosphere, accelerating therise in radiative forcing. We have used a mathematical modelof the upper ocean ecosystem, based on the Lagrangian Ensemblemethod, to estimate the sensitivity of the biological pump toradiative forcing, which lies at the heart of the plankton multiplier.We conclude that increasing radiative forcing by 5 W m^–(equivalent to doubling atmospheric CO₂) reduces the deep fluxof paniculate carbon by 10%. That sensitivity is sufficientto produce significant positive feedback in the greenhouse.It means that the plankton multiplier will increase the rateof climate change in the 21st century. It also suggests thatthe plankton multiplier is the mechanism linking the Milankovicheffect to the enhanced greenhouse effect that produces globalwarming at the end of ice ages.     

铁施肥促进海洋浮游植物和生物碳泵的不确定性

铁作为浮游植物所必需的微量元素,限制了全球超过三分之一海域的初级生产力,尤其是在高营养盐、低叶绿素海域(high nutrient low chlorophyll,HNLC)。长期以来海洋铁施肥被认为是一项可以降低大气二氧化碳含量的地球工程策略。然而通过13次海洋人工铁施肥(artificial ocean iron fertilization,aOIF)实验发现,铁的额外添加对海洋深层碳输出量的促进作用要显著低于预期。本文简要地总结了碳在海洋和大气中的循环过程,回顾了人工铁施肥实验对生物碳泵和碳通量等的影响,分析了从海洋铁施肥到海洋碳汇关键生物地球化学过程的影响因素。综上分析发现,科学界对生物碳泵过程及其调控机制的认识仍十分浅薄,考虑到海洋铁施肥还会对海洋生态系统带来一定的负面作用,铁施肥能否作为降低大气中CO₂的有效手段,以达到碳中和并缓解温室效应仍需进一步研究。     

采用遥感手段估算海洋初级生产力研究进展

海洋初级生产力的精确估算对渔业资源评估与管理、海洋生态系统和全球变化等研究具有重要意义.传统的现场测量与估算方法必须依赖于随船采样数据.卫星遥感具有能够获取实时的、大尺度的、动态的海洋环境参数的优点,因此卫星遥感日益成为大尺度海洋初级生产力估算的重要手段.本文从海洋水色传感器的发展历程出发,着重归纳了以叶绿素、浮游植物碳和浮游植物吸收系数为参量的海洋初级生产力的遥感估算方法,并就这3类模型的适应性和复杂程度进行了讨论.在此基础上,进一步分析评价了全球海洋初级生产力遥感估算的研究现状.鉴于当前海洋初级生产力遥感估算研究中存在的问题,今后的研究需要在4个方面进一步加强:1)对全球海洋初级生产力估算进行分区域研究;2)加深对浮游植物吸收系数的研究;3)提高海洋遥感技术水平;4)加强实地测量技术的研究.     

海洋生态系统固碳能力估算方法研究进展

气候变化受到全球关注,大气中CO2含量与气候变化息息相关。海洋是地球上最大的活跃碳库,在气候变化中扮演着举足轻重的作用。定量估算海洋中碳元素的吸收、转移、埋藏速率在全球碳循环及全球气候变化研究中有重要意义。目前,海洋固碳能力估算研究包括:利用海-气界面CO2分压差法估算海洋海-气界面CO2交换通量,根据海水中叶绿素含量建立的生态学数理模型法估算真光层浮游生物的初级生产力,234Th—238U不平衡法估算POC输出通量,210Pb定年法估算有机碳沉积通量。但迄今为止的研究工作尚有一定局限性,碳在大气—海水—沉积物3种介质间交换通量间相互影响的研究较少,海洋中碳垂直传输过程的主要影响因素和关键控制因子尚不明确,在海洋生态系统固碳能力估算方法方面国内外还没有统一的规范和标准。为进一步完善海洋生态系统固碳能力的估算方法,今后的工作应注重海洋固碳整套观测技术、分析和估算方法研究,并建立海洋碳汇估算指标体系、指标标准体系、以及评价标准体系,为我国的碳"减排"、"增汇"国家需求提供技术支持。     

The possible roles of algae in restricting the increase in atmospheric CO2 and global temperature

ABSTRACT

Anthropogenic inputs are increasing the CO₂ content of the atmosphere, and the CO₂ and total inorganic C in the surface ocean and, to a lesser degree, the deep ocean. The greenhouse effect of the increased CO₂ (and, to a lesser extent, other greenhouse gases) is very probably the major cause of present global warming. The warming increases temperature of the atmosphere and the surface ocean to a greater extent than the deep ocean, with shoaling of the thermocline, decreasing nutrient flux to the surface ocean where there is greater mean photosynthetic photon flux density. These global changes influence algae in nature. However, it is clear that algae are important, via the biological pump, in decreasing the steady state atmospheric and ocean surface CO₂, and thus decreasing radiative forcing, a reduction enhanced by algal increases in albedo. As well as these natural processes there are possibilities that algae can, with human intervention, partly offset the increase in atmospheric CO₂. One possibility is to grow algae as sources of fuel for transport, in principle providing an energy source that is close to CO₂-neutral. The other possibility is to increase the role of algae in sequestering CO₂ as organic C over periods of hundreds or more years in the deep ocean and marine sediments and/or increasing albedo and decreasing radiative forcing of temperature. There are problems, currently unresolved, in the economically viable production of algal biofuels without carbon trading subsidies. Enhanced algal CO₂ sequestration also has costs, both in resource input (phosphorus (P) from high P content rocks, a limited resource with a competing use as an agricultural fertilizer) and adverse environmental effects. For example, ocean anoxic zones producing N₂O and increased algal production of short-lived halocarbons by algae that both, through breakdown, destroy O₃ and increase UV flux to the Earth’s surface.     

Biotic and Human Vulnerability to Projected Changes in Ocean Biogeochemistry over the 21st Century

Ongoing greenhouse gas emissions can modify climate processes and induce shifts in ocean temperature, pH, oxygen concentration, and productivity, which in turn could alter biological and social systems. Here, we provide a synoptic global assessment of the simultaneous changes in future ocean biogeochemical variables over marine biota and their broader implications for people. We analyzed modern Earth System Models forced by greenhouse gas concentration pathways until 2100 and showed that the entire world''s ocean surface will be simultaneously impacted by varying intensities of ocean warming, acidification, oxygen depletion, or shortfalls in productivity. In contrast, only a small fraction of the world''s ocean surface, mostly in polar regions, will experience increased oxygenation and productivity, while almost nowhere will there be ocean cooling or pH elevation. We compiled the global distribution of 32 marine habitats and biodiversity hotspots and found that they would all experience simultaneous exposure to changes in multiple biogeochemical variables. This superposition highlights the high risk for synergistic ecosystem responses, the suite of physiological adaptations needed to cope with future climate change, and the potential for reorganization of global biodiversity patterns. If co-occurring biogeochemical changes influence the delivery of ocean goods and services, then they could also have a considerable effect on human welfare. Approximately 470 to 870 million of the poorest people in the world rely heavily on the ocean for food, jobs, and revenues and live in countries that will be most affected by simultaneous changes in ocean biogeochemistry. These results highlight the high risk of degradation of marine ecosystems and associated human hardship expected in a future following current trends in anthropogenic greenhouse gas emissions.     

Viral attack exacerbates the susceptibility of a bloom‐forming alga to ocean acidification

Both ocean acidification and viral infection bring about changes in marine phytoplankton physiological activities and community composition. However, little information is available on how the relationship between phytoplankton and viruses may be affected by ocean acidification and what impacts this might have on photosynthesis‐driven marine biological CO₂ pump. Here, we show that when the harmful bloom alga Phaeocystis globosa is infected with viruses under future ocean conditions, its photosynthetic performance further decreased and cells became more susceptible to stressful light levels, showing enhanced photoinhibition and reduced carbon fixation, up‐regulation of mitochondrial respiration and decreased virus burst size. Our results indicate that ocean acidification exacerbates the impacts of viral attack on P. globosa, which implies that, while ocean acidification directly influences marine primary producers, it may also affect them indirectly by altering their relationship with viruses. Therefore, viruses as a biotic stressor need to be invoked when considering the overall impacts of climate change on marine productivity and carbon sequestration.     

The effects of natural iron fertilisation on deep-sea ecology: the Crozet Plateau, Southern Indian Ocean

The addition of iron to high-nutrient low-chlorophyll (HNLC) oceanic waters stimulates phytoplankton, leading to greater primary production. Large-scale artificial ocean iron fertilization (OIF) has been proposed as a means of mitigating anthropogenic atmospheric CO(2), but its impacts on ocean ecosystems below the photic zone are unknown. Natural OIF, through the addition of iron leached from volcanic islands, has been shown to enhance primary productivity and carbon export and so can be used to study the effects of OIF on life in the ocean. We compared two closely-located deep-sea sites (～400 km apart and both at ～4200 m water depth) to the East (naturally iron fertilized; +Fe) and South (HNLC) of the Crozet Islands in the southern Indian Ocean. Our results suggest that long-term geo-engineering of surface oceanic waters via artificial OIF would lead to significant changes in deep-sea ecosystems. We found that the +Fe area had greater supplies of organic matter inputs to the seafloor, including polyunsaturated fatty acid and carotenoid nutrients. The +Fe site also had greater densities and biomasses of large deep-sea animals with lower levels of evenness in community structuring. The species composition was also very different, with the +Fe site showing similarities to eutrophic sites in other ocean basins. Moreover, major differences occurred in the taxa at the +Fe and HNLC sites revealing the crucial role that surface oceanic conditions play in changing and structuring deep-sea benthic communities.     

Quantifying the ocean, freshwater and human effects on year-to-year variability of one-sea-winter Atlantic salmon angled in multiple Norwegian rivers

Many Atlantic salmon, Salmo salar, populations are decreasing throughout the species' distributional range probably due to several factors acting in concert. A number of studies have documented the influence of freshwater and ocean conditions, climate variability and human impacts resulting from impoundment and aquaculture. However, most previous research has focused on analyzing single or only a few populations, and quantified isolated effects rather than handling multiple factors in conjunction. By using a multi-river mixed-effects model we estimated the effects of oceanic and river conditions, as well as human impacts, on year-to-year and between-river variability across 60 time series of recreational catch of one-sea-winter salmon (grilse) from Norwegian rivers over 29 years (1979-2007). Warm coastal temperatures at the time of smolt entrance into the sea and increased water discharge during upstream migration of mature fish were associated with higher rod catches of grilse. When hydropower stations were present in the course of the river systems the strength of the relationship with runoff was reduced. Catches of grilse in the river increased significantly following the reduction of the harvesting of this life-stage at sea. However, an average decreasing temporal trend was still detected and appeared to be stronger in the presence of salmon farms on the migration route of smolts in coastal/fjord areas. These results suggest that both ocean and freshwater conditions in conjunction with various human impacts contribute to shape interannual fluctuations and between-river variability of wild Atlantic salmon in Norwegian rivers. Current global change altering coastal temperature and water flow patterns might have implications for future grilse catches, moreover, positioning of aquaculture facilities as well as the implementation of hydropower schemes or other encroachments should be made with care when implementing management actions and searching for solutions to conserve this species.     

Assessing the distribution of sedimentary C40 carotenoids through time

A comprehensive marine biomarker record of green and purple sulfur bacteria (GSB and PSB, respectively) is required to test whether anoxygenic photosynthesis represented a greater fraction of marine primary productivity during the Precambrian than the Phanerozoic, as current models of ocean redox evolution suggest. For this purpose, we analyzed marine rock extracts and oils from the Proterozoic to the Paleogene for C₄₀ diagenetic products of carotenoid pigments using new analytical methods. Gas chromatography coupled with tandem mass spectrometry provides a new perspective on the temporal distributions of carotenoid biomarkers for phototrophic sulfur bacteria, specifically okenane, chlorobactane, and paleorenieratane. According to conventional paleoredox interpretations, this revised stratigraphic distribution of the GSB and PSB biomarkers implies that the shallow sunlit surface ocean (<24 m) became sulfidic more frequently in the geologic past than was previously thought. We reexamine whether there is evidence supporting a planktonic source of GSB and PSB pigments in marine systems or whether additional factors are required to explain the marine phototrophic sulfur bacteria record. To date, planktonic GSB and PSB and their pigments have been identified in restricted basins and lakes, but they have yet to be detected in the unrestricted, transiently sulfidic, marine systems. Based on modern observations, additional environmental factors, including basin restriction, microbial mats, or sediment transport, may be required to fully explain GSB and PSB carotenoids in the geologic record.     

Carbon pump dynamics and limited organic carbon burial during OAE1a

Oceanic Anoxic Events (OAEs) are conspicuous intervals in the geologic record that are associated with the deposition of organic carbon (OC)-rich marine sediment, linked to extreme biogeochemical perturbations, and characterized by widespread ocean deoxygenation. Mechanistic links between the marine biological carbon pump (BCP), redox conditions, and organic carbon burial during OAEs, however, remain poorly constrained. In this work we reconstructed the BCP in the western Tethys Ocean across OAE1a (~120 Mya) using sediment geochemistry and OC mass accumulation rates (OC_Acc). We find that OC_Acc were between 0.006 and 3.3 gC m⁻² yr⁻¹, with a mean value of 0.79 ± 0.78 SD gC m⁻² yr⁻¹—these rates are low and comparable to oligotrophic regions in the modern oceans. This challenges longstanding assumptions that oceanic anoxic events are intervals of strongly elevated organic carbon burial. Numerical modelling of the BCP, furthermore, reveals that such low OC fluxes are only possible with either or both low to moderate OC export fluxes from ocean surface waters, with rates similar to oligotrophic (nutrient-poor, <30 gC m⁻² yr⁻¹) and mesotrophic (moderate-nutrients, ~50–100 gC m⁻² yr⁻¹) regions in the modern ocean, and stronger than modern vertical OC attenuation. The low OC fluxes thus reflect a relatively weak BCP. Low to moderate productivity is further supported by palaeoecological and geochemical evidence and was likely maintained through nutrient limitation that developed in response to the burial and sequestration of phosphorus in association with iron minerals under ferruginous (anoxic iron-rich) ocean conditions. Without persistently high productivity, ocean deoxygenation during OAE1a was more likely driven by other physicochemical and biological factors including ocean warming, changes in marine primary producer community composition, and fundamental shifts in the efficiency of the BCP with associated effects and feedbacks.     

The role of cometary particle Coalescence in chemical evolution

Important prebiotic organic compounds might have been transported to Earth in dust or produced in vapor clouds resulting from atmospheric explosions or impacts of comets. These compounds coalesced in the upper atmosphere with particles ejected from craters formed by impacts of large objects. Coalescence during exposure to UV radiation concentrated organic monomers and enhanced formation of oligomers. Continuing coalescence added material to the growing particles and shielded prebiotic compounds from prolonged UV radiation. These particles settled into the lower atmosphere where they were scavenged by rain. Aqueous chemistry and evaporation of raindrops containing nomomers in high temperature regions near the Earth's surface also promoted continued formation of oligomers. Finally, these oligomers were deposited in the oceans where continued prebiotic evolution led to the most primitive cell. Results of our studies suggest that prebiotic chemical evolution may be an inevitable consequence of impacting comets during the late accretion of planets anywhere in the universe if oceans remained on those planetary surfaces.