Functional responses for zooplankton feeding on multiple resources: a review of assumptions and biological dynamics

Modelers often need to quantify the rates at which zooplankton consume a variety of species, size classes and trophic types. Implicit in the equations used to describe the multiple resource functional response (i.e. how nutritional intake varies with resource densities) are assumptions that are not often stated, let alone tested. This is problematic because models are sensitive to the details of these formulations. Here, we enable modelers to make more informed decisions by providing them with a new framework for considering zooplankton feeding on multiple resources. We define a new classification of multiple resource responses that is based on preference, selection and switching, and we develop a set of mathematical diagnostics that elucidate model assumptions. We use these tools to evaluate the assumptions and biological dynamics inherent in published multiple resource responses. These models are shown to simulate different resource preferences, implied single resource responses, changes in intake with changing resource densities, nutritional benefits of generalism, and nutritional costs of selection. Certain formulations are further shown to exhibit anomalous dynamics such as negative switching and sub-optimal feeding. Such varied responses can have vastly different ecological consequences for both zooplankton and their resources; inappropriate choices may incorrectly quantify biologically-mediated fluxes and predict spurious dynamics. We discuss how our classes and diagnostics can help constrain parameters, interpret behaviors, and identify limitations to a formulation's applicability for both regional (e.g. High-Nitrate-Low-Chlorophyll regions comprising large areas of the Pacific) and large-scale applications (e.g. global biogeochemical or climate change models). Strategies for assessing uncertainty and for using the mathematics to guide future experimental investigations are also discussed.     

A comparison of three marine ecosystems surrounding the Korean peninsula: Responses to climate change

This study uses a comparative approach to examine responses of marine ecosystems to climatic regime shifts. The three seas surrounding the Korean peninsula, the Japan/East Sea, the East China Sea and the Yellow Sea represent three contiguous but distinct ecosystems. Sampling has been carried out by the National Fisheries Research and Development Institute of South Korea since 1965, using the same methods in all three seas. Sampling was generally synoptic. Amplitude time series of 1st EOF modes for temperature, salinity, zooplankton biomass and concentrations of four major zooplankton taxa were used to determine whether the three marine ecosystems respond in a similar manner to climate variations. Temporal patterns of the variables were strongly similar among the three seas at decadal time scales, but very weakly similar at interannual scales. All three seas responded to a climatic regime shift that occurred in 1989. Temperature, zooplankton biomass and copepod concentrations increased in the late 1980s or early 1990s in all three seas. Concentrations of amphipods, chaetognaths and euphausiids also increased in the Japan/East Sea and the East China Sea, but not the Yellow Sea. The Yellow Sea ecosystem differs strongly from the other two seas, and water exchange between the Yellow Sea and the East China Sea is much weaker than that between the East China Sea and Japan/East Sea. Spatial patterns of zooplankton determined by the EOF analysis were closely related to currents and fronts in each of the three seas.     

Elevated biomass of mesozooplankton and enhanced fecal pellet flux in cyclonic and mode-water eddies in the Sargasso Sea

Accumulating evidence points to the importance of mesoscale eddies in supplying nutrients to surface waters in oligotrophic gyres. However, the nature of the biological response and its evolution over time has yet to be elucidated. Changes in mesozooplankton community composition due to eddy perturbation also could affect biogeochemical cycling. Over the course of two summers we sampled seven eddies in the Sargasso Sea. We focused on and followed a post-phytoplankton bloom cyclonic eddy (C1) in 2004 and a blooming mode-water anticyclonic eddy (A4) in 2005. We collected zooplankton in all eddies using a Multiple Opening and Closing Net Environmental Sampling System (MOCNESS) and quantified biomass (>0.15 mm, in five size fractions) from 0 to 700 m over nine discrete depth intervals. Zooplankton biomass (>0.5 mm) in the upper 150 m was similarly enhanced at night for the periphery of C1 and the center of A4 at 0.514 g m⁻² and 0.533 g m⁻², respectively, compared to outside (0.183 g m⁻² outside C1 and 0.197 g m⁻² outside A4). Despite minimal chlorophyll a enhancement and dominance by picoplankton in C1, zooplankton biomass increased most for the largest size class (>5 mm). Gut fluorescence for euphausiids and large copepods was also elevated on the C1 periphery. In A4, peak biomass occurred at eddy center coincident with peak primary production, but was highly variable (changing by >3-fold) over time, perhaps resulting from the dense, but patchy distribution of diatom chains in this region. Shifts in zooplankton community composition and abundance were reflected in enhancement of fecal pellet production and active transport by diel vertical migration in eddies. Inside C1 the flux of zooplankton fecal pellets at 150 m in June 2004 was 1.5-fold higher than outside the eddy, accounting for 9% of total particulate organic carbon (POC) flux. The flux of fecal pellets (mostly from copepods) increased through the summer in eddy A4, matching concurrent increases in zooplankton <2 mm in length, and accounting for up to 12% of total POC flux. Active carbon transport by vertically migrating zooplankton was 37% higher on the periphery of C1 and 74% higher at the center of A4 compared to the summer mean at the Bermuda Atlantic Time-series Study (BATS) station. Despite contrasting responses by the phytoplankton community to cyclonic and mode-water eddies, mesozooplankton biomass was similarly enhanced, possibly due to differential physical and biological aggregation mechanisms, and resulted in important zooplankton-mediated changes in mesoscale biogeochemistry.     

Zooplankton Feeding Ecology: Contents of Fecal Pellets of the Cyclopoid Copepods Oncaea venusta, Corycaeus amazonicus, Oithona plumifera, and O. simplex from the Northern Gulf of Mexico

Abstract. In situ feeding habits of the cyclopoid copepods Oncaea venusta, Corycaeus amazonicus, Oithona plumifera , and O. simplex were investigated by scanning electron microscope examination of fecal pellets, the contents of which reflected copepod gut contents upon capture. Peilet contents were compared with assemblages of phytoplankton present in surface waters at times of copepod collection. All samples were from the northern Gulf of Mexico. All four copepods fed upon phytoplankton and O. venusta also ingested other crustaceans. Dominant components of fecal pellets generally did not mirror those of available phytoplankton assemblages. In some cases, O. venusta ingested primarily larger-sized particles even when these were not most abundant, and in other cases it did not ingest large cells even when they were present in bloom concentrations. The presence of small (< 2–5 urn diameter) centric diatoms in O. venusta pellets suggests the possibility of feeding by mechanisms other than suspension or raptorial feeding. Limited observations suggest that C. amazonicus and O. plumifera may feed raptorially on larger particles even when these are not particularly abundant, and that the small O. simplex (< 500 nm total length) feeds mainly upon nanoplankton. It appears that cyclopoid feeding mechanisms are complex, and likely more so than those of many calanoids.     

The vertical distribution of zooplankton in the western Irish Sea

Zooplankton diel vertical migration is evident on the mixed isothermal side of the western Irish Sea frontal system but is often influenced by large tides and persistent geostrophic currents. On the stratified side of the front, temperature acts as a controlling factor with most of the zooplankton occurring above the thermocline and carrying out pronounced vertical migration when chlorophyll a levels are low and diffuse. At higher chlorophyll levels, when discrete chlorophyll a maxima form, zooplankton vertical movement may be greatly modified with a large number of species and stages concentrating within these maxima at all times of the diel light cycle.     

Net Zooplankton and the biological pump off Sanriku, Japan

Taxonomic composition, size composition, standing stock, and chemical composition of mesozooplankton were determined to examine the contribution of their fecal pellets to the vertical flux of organic carbon at the outside, the edge, and the center of the warm core ring. The warm core ring significantly affects not only their taxonomic composition and size composition but also their standing stock and chemical composition. The zooplankton at the center of the warm core ring was characterized by the absence of carnivores at the top of the size-trophic relation and filter feeding planktonic tunicates at the bottom. Zooplankton carbon biomass at the outside of the ring was one-third less than that at the center of the ring. The vertical flux of fecal pellets obtained from the pellet volume (12.3 mgC m⁻²d⁻¹) contributed 19 to 96% of the flux (13 to 64 mgC m⁻² d⁻¹) estimated from the body carbon and the fecal pellet production rate. The estimated flux of fecal pellets was 6 to 27% of vertical carbon flux (236 mgC m⁻²d⁻¹) determined by the sediment traps. Microscopic determination of fecal pellets and plankton in the sediment trap samples indicated high grazing activity during the sinking process. Those observations might suggest that particles other than fecal pellets contributed significantly to the vertical carbon flux and fecal pellets were settled directly without loss or being recycled within the surface mixed layer.     

华侨城湿地浮游生物调查

2014年7月,对华侨城湿地浮游生物进行调查。共鉴定浮游植物4门17种,浮游动物3门12种。叶绿素a含量值较大,以及耐污浮游生物成为优势种群,表明华侨城湿地正受较严重污染。同时,淡水藻类成为水体优势藻种,表明华侨城湿地水体正逐步淡化。     

典型河口浮游动物种类数及生物量变化趋势的研究

依据国家海洋局2004-2006年每年5月和8月的黄河口、长江口和珠江口生态监控区浮游动物的监测结果,对上述河口浮游动物和桡足类种类数及浮游动物生物量分布进行了分析,并将结果与历史资料进行同测区、同季节的对比。结果表明,与全国海洋普查时期相比,2006年黄河口浮游动物种类数下降了50．6％,桡足类种类数下降了53.3％;桡足类在浮游动物中所占比重由1985年的42.4％下降到2006年的32．6％。长江口浮游动物种类数有所上升,但是桡足类种类数由1985年的62种降低到2006年的37种;桡足类在浮游动物中的比重从1958年的54．2％降低到2006年的42．5％。珠江口桡足类在浮游动物中所占的比例由1959年的59．3％下降到54．4％,未发生明显变化。黄河口春季、夏季浮游动物生物量上升趋势不明显,夏季增幅较小;长江口自2000年以后春季浮游动物平均生物量为528．36mg／m^3,是1958年的2．4倍,2005年与2006年夏季浮游动物生物量分别是1958年的2．2和2．7倍,增加趋势十分明显;珠江口春季浮游动物生物量上升趋势最为显著,2004-2006年3年春季浮游动物平均生物量是1959年同期的5．7—8．7倍,1981年、2006年夏季生物量分别比1959年增加了3．1倍和11倍。就这3个河口生物量变化趋势看,珠江口生物量的上升趋势最为明显,长江口次之,黄河口变化幅度较小。     

Direct effects of Daphnia-grazing, not infochemicals, mediate a shift towards large inedible colonies of the gelatinous green alga Sphaerocystis schroeteri

The influence of Daphnia galeata×hyalina grazing and of infochemicals released by the daphnids on the colony size and growth rate of the colonial gelatinous green alga Sphaerocystis schroeteri (Chlorococcales) was investigated in laboratory batch experiments run for 96 h. High zooplankton grazing pressure was exerted by a final concentration of 100 daphnids L⁻¹ in the Daphnia treatments. Infochemicals were obtained by filtration (0.2 μm) of water from D. galeata×hyalina cultures (200 ind. L⁻¹ exposed for 24 h). This filtrate was added to the S. schroeteri cultures in two concentrations corresponding to 7 and 50 daphnids L⁻¹, respectively. The growth rate of S. schroeteri was neither affected significantly by direct Daphnia grazing nor by the presence of Daphnia infochemicals, in comparison to the control. However, the portion of inedible S. schroeteri colonies (diameter>50 μm) increased under direct grazing pressure, whereas the Daphnia infochemicals did not influence the colony size significantly. We conclude that the shift in colony size by direct zooplankton grazing denotes an effective defence mechanism against size selective feeding for colonial gelatinous green algae. This effective defence in combination with unchanged growth rates of the larger colonies (under non-limiting nutrient and light conditions) falsifies the assumption of a trade-off between minimising grazing losses and maximising growth by optimising the colony size.     

雷州半岛灯楼角珊瑚礁海区夏季的浮游动物

2004年8月在雷州半岛灯楼角珊瑚礁海区分别使用浅水Ⅰ型和浅水Ⅱ型浮游生物网进行了浮游动物调查,共鉴定浮游动物72种和浮游幼虫19个类群(或类型),其中桡足类的种数最多达45种。结果表明,浮游动物种数分布由近岸往远岸递增,约3/4的种类属于终身浮游生物,1/4属于阶段性浮游生物。中小型的浮游动物种类多,数量大。两种网具采集的浮游动物总种数、总密度、多样性指数、均匀度和中小型优势种的密度差异相当显著,而大型优势种的密度差异不明显。用浅水Ⅱ型网采集的浮游动物总密度平均值为5270ind/m3,是浅水Ⅰ型网的110倍。桡足类占浮游动物总密度的72.45%,其次幼虫占23.41%。浮游动物密度呈斑块状分布,最高达16257ind/m3,底质为珊瑚礁的测站的浮游动物数量一般较多。优势种主要是中小型种类和底栖动物的幼虫,如强额孔雀哲水蚤、蔓足类无节幼虫、细长腹剑水蚤、驼背隆哲水蚤、无节幼虫、双壳类面盘幼虫、桡足幼体、简长腹剑水蚤等。