Carbon cycling and POC turnover in the mesopelagic zone of the ocean: Insights from a simple model

Carbon budgets of the mesopelagic zone are poorly constrained, highlighting our lack of understanding of the biota that inhabit this environment and their role in the cycling and sequestering of carbon in the deep ocean. A simple food web model of the mesopelagic zone is presented that traces the turnover of particulate organic carbon (POC), supplied as sinking detritus, through to its respiration by the biota via three pathways: colonization and solubilization of detritus by attached bacteria, production of free-living bacteria following losses of solubilization products during particle degradation, and consumption by detritivorous zooplankton. The relative consumption of detritus by attached bacteria was initially specified as 76%, with the remaining 24% by detritivores. Highlighting an asymmetry between consumption and respiration, the resulting predicted share of total respiration due to bacteria was 84.7%, with detritivores accounting for just 6.6% (with 6.5% and 2.2% by bacterivores and higher zooplankton, respectively). Bacteria thus dominated respiration and thereby acted as the principal sink for POC supplied to the mesopelagic zone, whereas zooplankton mainly recycled carbon back to the base of the food web as detritus or dissolved organic carbon rather than respiring it to CO₂. Estimates of respiration are therefore not necessarily a reliable indicator of the relative roles of bacteria and zooplankton in consuming and processing POC in the mesopelagic zone of the ocean. The work highlighted a number of major unknowns, including how little we know in general about the dynamics and metabolic budgets of bacteria and zooplankton that inhabit the mesopelagic zone and, specifically, the degree to which the solubilized products of enzymatic hydrolysis of POC by attached bacteria are lost to the surrounding water, the magnitude and factors responsible for bacterial growth efficiency, the role of microbes in the nutrition of detritivores, and the recycling processes by which zooplankton return what they consume to the food web as detritus and dissolved organic matter.     

国际海洋浮游动物研究进展

综述了国际上有关海洋浮游动物种群、群落结构、多样性及浮游动物对全球气候变化响应等方面研究进展。海洋浮游动物种类繁多, 数量丰富, 分布广泛, 是海洋生态系统中最重要的生物类群。在海洋食物网中, 浮游动物通过摄食浮游植物控制初级生产力, 同时, 又被更高营养阶层的动物(鱼、虾、鲸、海鸟等)捕食, 充当次级生产者的角色, 其群落结构、种群动态和物种多样性影响鱼类和其他海洋动物资源量, 浮游动物是海洋食物网中关键环节。海洋生态系统动力学过程的关键环节是浮游生物的生物学和生态学过程, 多项国际研究计划以生物多样性和年际变化趋势为研究重点并联系全球变化及响应, 因此, 浮游动物的研究已成为海洋生态研究的核心内容之一。国际上对浮游动物的研究主要集中在以下6个方向:(1)浮游动物生境、种群的分布和扩散动力学研究;(2)浮游动物的群落结构和多样性;(3)浮游动物的实验生态和现场受控生态研究;(4)浮游动物对全球气候变化的响应;(5)深海、南北极、低氧区等极端生境的浮游动物生态学研究;(6)浮游动物研究新技术和方法。     

Trophic efficiency of the planktonic food web in a coastal ecosystem dominated by Phaeocystis colonies

The trophic efficiency of the planktonic food web in the Phaeocystis-dominated ecosystem of the Belgian coastal waters was inferred from the analysis of the carbon flow network of the planktonic system subdivided into its different trophodynamic groups. A carbon budget was constructed on the basis of process-level field experiments conducted during the spring bloom period of 1998. Biomass and major metabolic activities of auto- and heterotrophic planktonic communities (primary production, bacterial production, nanoproto-, micro- and mesozooplankton feeding activities) were determined in nine field assemblages collected during spring at reference station 330. In 1998, the phytoplankton spring flowering was characterised by a moderate diatom bloom followed by a massive Phaeocystis colony bloom. Phaeocystis colonies, contributing 70% to the net primary production, escaped the linear food chain while the early spring diatom production supplied 74% of the mesozooplankton carbon uptake. The rest of mesozooplankton food requirement was, at the time of the Phaeocystis colony bloom, partially fulfilled by microzooplankton. Only one-third of the microzooplankton production, however, was controlled by mesozooplankton grazing pressure. Ungrazed Phaeocystis colonies were stimulating the establishment of a very active microbial network. On the one hand, the release of free-living cells from ungrazed colonies has been shown to stimulate the growth of microzooplankton, which was controlling 97% of the nanophytoplankton production. On the other hand, the disruption of ungrazed Phaeocystis colonies supplied the water column with large amounts of dissolved organic matter available for planktonic bacteria. The budget calculation suggests that ungrazed colonies contributed up to 60% to the bacterial carbon demand, while alternative sources (exudation, zooplankton egestion and lysis of other organisms) provided some 30% of bacterial carbon requirements. This suggests that the spring carbon demand of planktonic bacteria was satisfied largely by autogenic production. The trophic efficiency was defined as the ratio between mesozooplankton grazing on a given source and food production. In spite of its major contribution to mesozooplankton feeding, the trophic efficiency of the linear food chain, restricted to the grazing on diatoms, represented only 5.6% of the available net primary production. The trophic efficiency of the microbial food chain, the ratio between mesozooplankton grazing on microzooplankton and the resource inflow (the bacterial carbon demand plus the nanophytoplankton production) amounted to only 1.6%. These low trophic efficiencies together with the potential contribution of ungrazed Phaeocystis-derived production to the bacterial carbon demand suggest that during spring 1998 most of the Phaeocystis-derived production in the Belgian coastal area was remineralised in the water column.     

Box model analysis on phytoplankton production and grazing pressure in a eutrophic estuary

To determine the quantitative relationship between phytoplankton production and zooplankton grazing pressure in Atsumi Bay, a eutrophic and partially-mixed estuary, a series of investigations, including measurements of hydrographic conditions, dissolved oxygen, dissolved total nitrogen, particulate organic nitrogen, and phyto- and zooplankton biomass were conducted 13 times at intervals of 2–7 days in June and July 1984. Continuous measurements of water flow and salinity were also carried out to examine transverse flow and horizontal diffusivity. The supply of freshwater and nitrogen was estimated from given data. The changes of hydrographic condition, net photosynthetic rate and community primary production were calculated by a two-layered box model analysis. The grazing rate on phytoplankton obtained as the difference between net photosynthetic rate and community primary production was compared to the one estimated from zooplankton biomass and sardine,Sardinops melanosticta, biomass. The agreement between the data was remarkable in the upper layer, showing the grazing pressure on phytoplankton followed phytoplankton production, suggesting that a large part of produced phytoplankton was immediately grazed by zooplankton. Consequently, the community primary production was depressed to a fairly lower level. An important role of nutrient supply and water circulation, to limit phytoplankton production, was also confirmed. Dynamic response observed between the calculated grazing pressure and the biomass of phytoplankton and protozoa was also analyzed.     

Grazing experiments and model simulations of the role of zooplankton in Phaeocystis food webs

A combined empirical and modelling study was conducted to further examine the potential importance of grazing by zooplankton in pelagic food webs in which Phaeocystis is a significant or dominant component. Laboratory experiments were designed to measure ingestion of Phaeocystis and other potential prey items which co-occur with Phaeocystis. Grazers included copepods and ciliates, and prey included Phaeocystis colonies and solitary cells, diatoms, ciliates, bacteria, and detritus. These data were expressed in the model currency of nitrogen units, and fit to hyperbolic tangent equations which included minimum prey thresholds. These equations and literature data were used to constrain a food web model whose purpose was to investigate trophic interactions rather than to mimic actual events. Nevertheless, the model output was similar to the general pattern and magnitude of development of Phaeocystis–diatom communities in some environments where they occur, e.g. north Norwegian waters. The model included three forms of nitrogen, three phytoplankton groups, bacteria, two zooplankton groups, and detritus, with detailed flows between compartments. An important component of the model was inclusion of variable prey preferences for zooplankton. The experiments and model simulations suggest several salient conclusions. Phaeocystis globosa colonies were eaten by a medium-sized copepod species, but ingestion appeared to be strongly dependent upon a proper size match between grazer and prey. If not, colonies were eaten little if at all. Phaeocystis solitary cells were ingested rapidly by ciliate microzooplankton, in agreement with prior literature observations. In contrast, detritus was eaten comparatively slowly by both ciliates and copepods. Both types of zooplankton exhibited apparent minimum prey thresholds below which grazing did not occur or was inconsequential. Model simulations implied that transitions between life cycle stages of Phaeocystis may potentially be important to phytoplankton–zooplankton interactions, and that relative rates of ingestion of Phaeocystis by various zooplankton may have significant impacts upon material fluxes through and out of Phaeocystis–diatom ecosystems. Indirect effects of trophic interactions appear to be equally significant as direct effects.     

Growth and succession patterns of major phylogenetic groups of marine bacteria during a mesocosm diatom bloom

Our objective was to track microbial processes associated with serial degradation of organic matter derived from algal blooms. To do this, we analyzed population fluctuations and growth responses of major phylogenetic groups of free-living marine bacteria. We used bromodeoxyuridine immunocytochemistry–fluorescence in situ hybridization methodology to examine marine bacterial community development during and after a diatom bloom in a mesocosm. We revealed that the Roseobacter/Rhodobacter, SAR11, Alteromonas, and Bacteroidetes groups were clearly major phylotypes responsible for most free-living bacterial biomass and production throughout the experiment. The clearest bacterial response was a proliferation of the Alteromonas group (cells with large volumes) during development of the bloom (up to 30?% of actively growing cells). Populations of these bacteria declined sharply thereafter, likely due to grazing. Alteromonas group responses suggest that these bacteria strongly influenced the flux of organic matter at an early bloom stage. The growth potential of Bacteroidetes was relatively large as the bloom peaked; this early development probably contributed to the initial stage of bloom decomposition. In contrast, the contribution of Roseobacter/Rhodobacter to total bacterial production increased at a late stage of decomposing of the bloom. The contributions of Betaproteobacteria, SAR11, and SAR86 groups to total bacterial abundance and production were relatively minor throughout the experiment. These results imply that the ability to utilize organic matter derived from diatoms varies among bacterial phylotypes, and, frequently, less abundant but ecological specialist taxa such as Alteromonas may play major roles in the flux of organic matter during diatom blooms.     

Microheterotrophic pathways in the southern Benguela upwelling system

During the 1980s, there was a major shift in the understanding of the structure of marine foodwebs. As a result, the microbial loop has been incorporated into the classical concept of the planktonic food chain. Heterotrophic bacteria and Protozoa have been shown to be important components of the plankton biomass in many parts of the world's oceans, and their role in the trophic dynamics of pelagic foodwebs has been studied intensively. In the southern Benguela, field, laboratory and modelling studies have been combined to calculate carbon and nitrogen fluxes through the microheterotrophic portion of the pelagic foodweb. A size-based simulation model incorporating recent hypotheses on the structure and functioning of the pelagic foodweb after upwelling predicts rapid growth of a phytoplankton community dominated by netphytoplankton and chain-forming nanophytoplankton cells. After nitrate-depletion the bloom declines, to be followed by a bloom of single nanophytoplankton cells dependent upon regenerated nitrogen. During conditions of high nitrate availability and a netphytoplankton-dominated community, mesozooplankton ingest 44 per cent of the total primary production through herbivory and 1,4 per cent through carnivory. During periods of low nitrate availability and a nanophytoplankton-dominated community, mesozooplankton ingest 0,6 per cent of the total primary production through herbivory and 5 per cent through carnivory. Food chains are longer and microheterotrophs are an important link between primary producers and the larger heterotrophs. Simulation results show that microheterotrophs are an important component of the pelagic foodweb, primarily as regenerators of nitrogen, which sustains phytoplankton growth, and as a food source for larger heterotrophs of the metazoan foodweb during the nanophytoplankton-dominated bloom.     

Assessing food web dynamics and relative importance of organic matter sources for fish species in two Portuguese estuaries: a stable isotope approach

Stable carbon and nitrogen isotopes (δ13C, δ1?N) were used to analyse food web dynamics of two of the main estuaries of the Portuguese coast: Tejo and Mira. The ultimate sources of organic matter supporting production of some of the most abundant and commercially important fish species were determined; and seasonal, inter- and intra- estuarine differences in the trophic relations among producers and consumers were identified. Stable isotope analysis was performed in different producers, primary consumers (main prey items for fish) and fish species (Solea solea, Solea senegalensis, Pomatoschistus microps, Dicentrarchus labrax, Liza ramada, Diplodus vulgaris and Atherina presbyter) of two areas in each estuary, in July and October 2009. Model calculations showed that the main prey for the fish species in the Tejo estuary used mostly salt marsh-derived organic matter as nutritional sources, with no marked differences between the sampled months. Trophic levels of fish species from the same estuary differed at multiple scales: inter-species, seasonally and spatially (both between and within estuaries). Significant differences in isotopic composition of fish species were more pronounced spatially (between the two sampled areas in the estuary) than seasonally (between sampled months). Trophic relationships in both estuaries demonstrated that organic matter is transferred to higher trophic positions mainly through benthic pathways. This shows the flexibility of these species to share resources and to exploit temporary peaks in prey populations. The present results showed that extensive disturbance in intertidal habitats from both estuaries may potentially change the balance of organic matter in the base of these complex food webs.     

Linking bacterial community structure to carbon fluxes in marine environments

Microbial oceanography is undergoing a dramatic revolution thanks to the rapid development of novel techniques that allow the examination of microbial diversity and functions via molecular methods, including genomic and metagenomic analyses. During the past decade, studies have revealed previously unknown and surprisingly diverse bacterial communities in marine waters. These studies have radically changed our understanding of spatiotemporal patterns in marine bacterial community composition and the distribution of specific genes. However, our knowledge of the role of individual bacterial subgroups in oceanic food webs and biogeochemical cycles remains limited. To embed the internal dynamics of bacterial communities into marine biogeochemistry models, the characteristic parameters of individual bacterial subgroups (i.e., growth, mortality, and utilization of dissolved organic matter) must be determined. Here, we survey the approaches used to assess variation in and factors controlling bacterial communities in marine environments, emphasizing the importance of quantitative studies that examine growth and grazing parameters of bacterial subgroups.     

The biogeochemical fate and toxicity of mercury in Controlled Experimental Ecosystems

The affinity of mercury for organic matter was the most important parameter governing its chemical speciation, transport and toxicity in Controlled Experimental Ecosystems (CEEs) of the Controlled Ecosystem Pollution Experiment (CEPEX) deployed in Saanich Inlet, B.C., Canada, Particulate, colloidal and high molecular weight dissolved forms of mercury accounted for about 90% of the total mercury present in the water column. Bioassays indicated that organic matter in the above size classes had an important influence on the toxicity of mercury to phytoplankton.Sorption of mercury to particulate organic matter in the CEEs could be described by either Freundlich or linear adsorption isotherms. An observed depth-dependent increase in mercury content of particulate matter was attributed to selective biological degradation of non-mercury-binding organic components of the particles during settling.The high affinity of mercury for settling particulate organic matter led to rapid removal of mercury from CEEs. Removal rates could be described as first order and were a function of primary production rates. Deviations of measured half-removal times from those predicted by an empirically formulated relationship between primary production and mercury half-removal times could be related successfully to the influence of other occasionally significant biological parameters such as zooplankton grazing activity.     

Balance Between Autotrophic and Heterotrophic Components and Processes in Microbenthic Communities of Sandy Sediments: A Field Study

The microscopic community of a microtidal sandy sediment on the Swedish west coast was studiedin situat two depths (0·5 and 4 m) on four occasions (January, April, August and October). Biomass of microalgae, bacteria, ciliates and meiofauna, as well as primary and bacterial productivity, were quantified. Meiofaunal grazing on algae and bacteria was measured simultaneously by radiolabelling intact sediment cores. Autotrophic biomass dominated the microbial community at both depths and on all sampling occasions, accounting for 47–87% of the microbial biomass. Meiofauna contributed 10–47%, while bacteria and ciliates together made up less than 6%. The microflora was dominated by attached (epipsammic) diatoms, but occasional ‘ blooms ’ of motile species occurred. Vital cells of planktonic diatoms contributed to benthic algal biomass in spring. Primary productivity exceeded bacterial productivity in April and August at both depths, while the balance was reversed in October and January. Meiofauna grazed between 2 and 12% of the algal biomass per day, and between 0·3 and 37% of the bacterial biomass. Almost an order of magnitude more algal (17–138 mg C m⁻²) than bacterial (0·1–33 mg C m⁻²) carbon was grazed daily. At the shallow site, primary productivity always exceeded grazing rates on algae, whereas at the deeper site, grazing exceeded primary productivity in October and January. Bacterial productivity exceeded grazing at both depths on all four occasions. Thus, meiofaunal grazing seasonally controlled microalgal, but not bacterial, biomass. These results suggest that, during summer, only a minor fraction (<10%) of the daily microbenthic primary production appears to enter the ‘ small food web ’ through meiofauna. During spring and autumn, however, a much larger fraction (≈30–60%) of primary production may pass through meiofauna. During winter, meiofaunal grazing is a less important link in the shallow zone, but at sublittoral depths, algal productivity may be limiting, and meiofauna depend on other food sources, such as bacteria and detritus.     

Sources and transfer of organic matter in food webs of a Mediterranean coastal environment: Evidence for spatial variability

The spatial variability in the food web structure of a Mediterranean semi-enclosed coastal environment (Stagnone di Marsala, Italy) was investigated using stable carbon and nitrogen isotopes. Organic matter sources and consumers were sampled in two locations with different environmental features (e.g. hydrodynamic regime, open-sea influence, vegetal coverage). Overall more ¹³C-enriched and ¹⁵N-depleted values were found in the central location than in the southern for organic matter sources and consumers. Pelagic consumers (zooplankton and juveniles of transient fish) showed slight spatial differences and in both locations seemed to depend on phytoplankton as the ultimate energy source. In contrast, benthic consumers (epifauna and resident fish) exhibited remarkable differences between locations. Spatial differences in organic matter sources were smaller than in benthic consumers and thus consumers presumably exploited different ultimate organic matter sources in the two locations. Sedimentary organic matter and epiphytes appeared to be the main primary producers transferred within the food web in both locations, and seagrasses seemed to play a non-negligible trophic role in the central location. The results of this paper corroborate the finding food webs are characterised by high spatial variability even on a small spatial scale and environmental heterogeneity more than primary production that seems to influence the trophic role of autotrophs.     

Variation of reactivity of particulate and sedimentary organic matter along the Zhujiang River Estuary

To investigate organic matter source and reactivity in the Zhujiang River (Pearl River)Estuary and its adjacent areas, particulate organic carbon (POC), particulate hydrolysable amino acids (PHAA), and Chl a during two cruises in July 1999 and July 2000 were measured. The highest POC and PHAA concentration was observed in the waters with maximum Chl a. The spectra distribution,relative content (dry weight in milligram per gram), PHAA-C% POC and other indicators such as the ratios of amino acids vs. amino sugars (AA/AS) and glucosamine vs. galactosamine (Glum/Gal) suggested that particulate amino acids in the water column and sediments in the Zhujiang River Estuary were mainly derived from biogenic processes rather than transported from terrestrial erosion. In inner estuary where high turbidity was often observable, organic matter was mainly contributed by re-suspension of bottom sediments with revealed zooplankton, microbial reworked characteristics, which suggest that these organic matters were relatively “old“. In the estuarine brackish region, organic matter in water column is mainly contributed by relatively fresh, easily degradable phytoplankton derived organic matter.During physical - biological processes within the eastuary, organic matter derived from phytoplankton was subjected to alteration by zooplankton grazing and bacterial reworking.     

Determining how the pelagic ecosystem over the continental shelf of the Bay of Biscay (NE Atlantic) functions: An approach using mesozooplankton enzyme activities as descriptors

A fisheries research cruise conducted in 2000 offered a first opportunity to take simultaneous measurements of the activities of three enzymes in mesozooplankton samples collected at a regional scale over the continental shelf of the Bay of Biscay in the NE Atlantic, with the aim of characterizing main aspects of the functioning of the biotic environment of small pelagic fish populations. The activity of the digestive endopeptidase trypsin was selected to characterize the assimilation rate of proteins, whereas pyruvate kinase (PK) was chosen as an indicator of carbohydrate assimilation and aspartate transcarbamylase (ATC) provided an overall assessment of mesozooplankton productivity. The Bay of Biscay region is subject to various strong physical driving forces that directly affect the primary structure of the pelagic food web. On our cruise, the phytoplankton biomass distribution reflected these different physical influences: diatoms dominated the nutrient-enriched coastal water; picoplankton dominated the northern-central part where nutrients were depleted; and nanoplankton were abundant at the shelf break where internal waves provided an input of nutrients. These and other results (on bacteria, particulate organic carbon distribution, among others) illustrate the differences that exist in the microbial food webs of different sectors of the bay. The living matter produced was characterized by the quality and quantity of the smallest prey items that were available to higher trophic levels. Variations in mesozooplankton enzyme activities may agree well not only with classically expected results, but also present unexpected special features: high ATC specific activities were measured around the mouth of the Gironde, in the nutrient-rich desalted water of the plume, but surprisingly not in front of the Loire river. PK specific activities reflected preponderantly the balance between phytoplankton cells sizes and the related bacterial abundance resulting from nutrient limitation (mainly P), that induces varying carbohydrates production potential. Trypsin specific activities were moderately variable, except in a restricted area where a highly abundant protein content characterized the particulate matter and in the plume of water flowing out of the Gironde. It is concluded that the presented approach of the metabolism of mesozooplankton communities may provide novel views on crucial processes occurring at the mesoscale, which fits in generally well with the scales of ecological factors mostly influential on small pelagic fish populations.     

Production and degradation of fluorescent dissolved organic matter derived from bacteria

Dynamics of fluorescent dissolved organic matter (FDOM) in ocean environments has received attention over the past few decades. Although it has appeared that in situ production of oceanic FDOM is mainly due to bacteria, the production and bio- and photodegradation processes of bacterial FDOM have not been elucidated. In this study, a culture experiment with bacteria was carried out to assess the production and biodegradation processes of bacterial FDOM. Photodegradation of bacterial FDOM and dissolved organic carbon (DOC) was also examined by exposure to a solar simulator. Bacterial FDOM consists of six components which were determined by parallel factor analysis (PARAFAC). Fluorescence intensities of protein-like FDOM increased with the bacterial biomass, but the increases of humic-like FDOM lagged behind the protein-like FDOM by 5–10 days. Exposure to simulated sunlight caused significant decreases in fluorescence intensities of all components; 52–94% of the initial intensities were lost during 24 h. While, the DOC concentration exhibited a small decrease through the experiment (1.9–11.1%). These results showed that photodegradability of bacteria derived DOC was much less than the fluorescence, indicating that the lifetime of bacteria-derived DOC is much longer than the length estimated by the fluorescence. The role of photobleached FDOM derived from bacteria may be significant in the biogeochemical cycle at the surface layer.     

Food web flows through a sub-arctic deep-sea benthic community

The benthic food web of the deep Faroe–Shetland Channel (FSC) was modelled by using the linear inverse modelling methodology. The reconstruction of carbon pathways by inverse analysis was based on benthic oxygen uptake rates, biomass data and transfer of labile carbon through the food web as revealed by a pulse-chase experiment. Carbon deposition was estimated at 2.2 mmol C m⁻² d⁻¹. Approximately 69% of the deposited carbon was respired by the benthic community with bacteria being responsible for 70% of the total respiration. The major fraction of the labile detritus flux was recycled within the microbial loop leaving merely 2% of the deposited labile phytodetritus available for metazoan consumption. Bacteria assimilated carbon at high efficiency (0.55) but only 24% of bacterial production was grazed by metazoans; the remaining returned to the dissolved organic matter pool due to viral lysis. Refractory detritus was the basal food resource for nematodes covering ∼99% of their carbon requirements. On the contrary, macrofauna seemed to obtain the major part of their metabolic needs from bacteria (49% of macrofaunal consumption). Labile detritus transfer was well-constrained, based on the data from the pulse-chase experiment, but appeared to be of limited importance to the diet of the examined benthic organisms (<1% and 5% of carbon requirements of nematodes and macrofauna respectively). Predation on nematodes was generally low with the exception of sub-surface deposit-feeding polychaetes that obtained 35% of their energy requirements from nematode ingestion. Carnivorous polychaetes also covered 35% of their carbon demand through predation although the preferred prey, in this case, was other macrofaunal animals rather than nematodes. Bacteria and detritus contributed 53% and 12% to the total carbon ingestion of carnivorous polychaetes suggesting a high degree of omnivory among higher consumers in the FSC benthic food web. Overall, this study provided a unique insight into the functioning of a deep-sea benthic community and demonstrated how conventional data can be exploited further when combined with state-of-the-art modelling approaches.     

Microzooplankton feeding impact in a coastal upwelling system on the NW Iberian margin: The Ría de Vigo

The dilution technique, combined with identification and enumeration of pico-, nano- and micro-plankton by microscopy, was used to estimate microzooplankton impact on the microbial community in surface waters of a coastal embayment on the NW Iberian upwelling system. Microzooplankton were important consumers of autotrophic and heterotrophic plankton in this system, feeding up to 93% of standing stock and more than 100% of production of several groups. Heterotrophic bacteria and heterotrophic picoflagellates experienced the highest and constant impact, with 75–84% of their standing stocks and 85–102% of their production being channelled through the microbial food web. Pico- and nano-phytoplankton were also consumed, although maximum grazing occurred on diatoms during upwelling events, coinciding with highest primary production. Predation on pico-nano-heterotrophs was especially relevant under downwelling conditions, when consumption of total carbon and particularly autotrophic carbon was considerably lower than during upwelling. The results suggest that the existence of a multivorous food web, extending from the microbial loop to the herbivorous food web, could be a major feature in this coastal upwelling system. The microbial loop, which occurs as a permanent background in the system, would contribute to sustain the microbial food web during downwelling, whereas the herbivorous food web could coexist with a microbial food web based on large diatoms during upwelling. The multivorous food web would partially divert diatoms from sinking and hence favour the retention of organic matter in the water column. This could enhance the energy transfer to higher pelagic trophic levels in coastal upwelling systems.     

基于碳、氮稳定同位素的雅浦海沟底栖生物食物来源和营养级初探

为了分析雅浦海沟中底栖生物群落的食物来源和营养级，本研究分析了雅浦海沟真光层中浮游植物和浮游动物、海底沉积物和巨型底栖生物（海绵、海参、海蛇尾、海星、海葵和钩虾）中的碳、氮稳定同位素组成。研究发现雅浦海沟真光层中的浮游植物和浮游动物δ13C值[（-22.8±0.4）‰和（-21.8±0.8）‰]和δ15N值[（5.4±0.4）‰和（6.8±0.2）‰]与巨型底栖生物的δ13C值（-20.1‰~-16.8‰）和δ15N值（11.9‰~17.9‰）的差异超过了一个营养级，表明作为底栖生物的初始食物来源的浮游植物和浮游动物在向下输送的过程中经历了食物链传递和细菌的降解。巨型底栖生物的δ15N和δ13C值之间无显著的相关性，此外不同物种之间营养级也存在明显差异，表现为海绵的营养级相对较高（3.4~4.7），海参（3.3~3.6）、海蛇尾（3.4~3.5）和海星（3.2~3.7）的营养级较为接近，钩虾（2.9~3.3）和海葵（3.1）的营养级则相对略低，反映了底栖生物不同物种之间食物来源的多样化。     

Spatial distribution of zooplankton in the intertidal marsh creeks of the Yangtze River Estuary,China

Zooplankton are important grazers of primary production within intertidal marshes and are the optimal prey of higher trophic consumers; however, the patterns of their spatial distribution in marsh creeks are rarely reported. The zooplankton in the intertidal creeks with different salinities at Dongtan marshes of the Yangtze River Estuary was surveyed. The mean zooplankton densities in the intertidal creeks were 53,638 ind. m⁻³ in April and 132,916 ind. m⁻³ in July, respectively, which were as high as in the near-shore subtidal waters of the Yangtze River Estuary. This high abundance implied the important roles of zooplankton in the matter flux between marshes and near-shore waters through complex intertidal creek systems. Zooplankton total densities changed significantly from northern to southern creeks. ANOSIM and CCA analyses revealed that the zooplankton community structure were significantly different among the northern, eastern and southern creeks, and between two sampling seasons. Salinity accounted for most of the spatial variation of zooplankton community, whereas water temperature, chlorophyll a concentration, and pH were the main reasons of the temporal variation observed. Copepods were the most abundant zooplankton group. A total of 24 copepod species, belonging to 15 families and 20 genera, were recorded. Planktonic copepods preferred the northern and eastern creeks, with higher densities in July than in April, while benthic copepods predominated only in the northern creeks in April. Since the role of benthic and planktonic copepods may differ in transporting nutrients in the intertidal creeks, it is suggested that the variations in their distribution may influence the ecological functions of zooplankton in the estuarine matter fluxes both spatially and temporally.