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Anderson, J. T., Hood, R. R., & Zhang, X. (2003). Quantification of Pfiesteria piscicida growth and encystment parameters using a numerical model. Marine Ecology-Progress Series, 246, 105–113.
Abstract: In the past decade, there has been growing interest in understanding the physiological ecology and life cycle of toxic forms of Pfiesteria piscicida. However, transformations among non-inducible (NON-IND; formerly described as nontoxic) stages have received less attention despite the fact that NON-IND stages are found in nature and may be ecologically important as prey and predators. NON-IND stages are also mixotrophic and have the ability to retain and utilize prey chloroplasts in a process termed 'kleptoplastidic mixotrophy'. Quantifying growth, grazing and encystment rates from P. piscicida laboratory experiments is confounded by the interrelationship between mixotrophy and life stage transformations. By fitting a numerical model to a laboratory experiment on NON-IND P. piscicida, we were able to isolate the potential mechanisms that cause encystment and speculate on the interrelationship between adverse conditions (i.e. low light and limiting prey) and life stage transformations. The structure of the laboratory experiment allowed for the estimation of several growth and encystment parameters including grazing rates, gross growth and assimilation efficiencies, as well as the retention time of chloroplasts. Model results suggest a link between encystment and mixotrophic ability. Furthermore, the model results suggest that encystment rates and gross growth and assimilation efficiencies calculated from the model are lower than expected.
Keywords: Pfiesteria piscicida; harmful algal blooms; numerical models;; kleptochloroplastidy; mixotrophy; life history transformations; resting; stages; ambush-predator dinoflagellate; toxic dinoflagellate; life-cycle;; estuarine dinoflagellate; gyrodinium-uncatenum; fish kills;; dinophyceae; mixotrophy; behavior; chloroplasts
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Roman, M., Zhang, X., McGilliard, C., & Boicourt, W. (2005). Seasonal and annual variability in the spatial patterns of plankton biomass in Chesapeake Bay. Limnology and Oceanography, 50(2), 480–492.
Abstract: We conducted high-resolution, underway sampling in April, July, and October for 6 yr (1995-2000) in the large estuary, Chesapeake Bay. This period included climatological extremes in freshwater inputs that strongly influenced both the overall stocks and spatial distribution of phytoplankton and zooplankton. Higher biomass of both phytoplankton and zooplankton occurred in springs, when freshwater input into Chesapeake Bay was above the average discharge. While whole-Bay productivity appears to be influenced by freshwater flow variability, mesoscale patterns in plankton biomass are driven by freshwater inputs, circulation, and bathymetry. Persistent maxima in plankton biomass occurred in areas of physical and topographic discontinuities such as the upper-Bay salt front, plume fronts, the hydraulic control region, tidal fronts, and near a topographically induced eddy. Although the contribution of these hot spots to the whole-Bay standing stock of plankton may vary due to changes in the background levels of plankton, controlled in part by freshwater discharge, they nevertheless represent predictable areas of higher forage for planktivorous fish. Enhanced trophic coupling between plankton and fish at these physical discontinuities may be one reason why estuaries have higher fisheries yields in relation to their primary production than lakes and other marine systems.
Keywords: estuarine turbidity maximum; long-term trends; white perch;; zooplankton; fish; phytoplankton; abundance; counter; chlorophyll;; nutrients
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Roman, M. R., Adolf, H. A., Landry, M. R., Madin, L. P., Steinberg, D. K., & Zhang, X. (2002). Estimates of oceanic mesozooplankton production: a comparison using the Bermuda and Hawaii time-series data. Deep-Sea Research Part Ii-Topical Studies in Oceanography, 49(1-3), 175–192.
Abstract: Mesozooplankton growth rates were estimated for the Hawaiian (HOT) and Bermuda (BATS) ocean time-series stations using the empirical model of Hirst and Lampitt (Marine Biology 132 (1998) 247), which predicts copepod growth rate from temperature and body size. Using this approach we derived seasonal and annual estimates of mesozooplankton production as well as rates of mesozooplankton ingestion and egestion using assumed growth and assimilation efficiencies for the period 1994-1997. Annual mesozooplankton production estimates at HOT (average 0.79 Mol C m(-2) yr(-1)) were higher than production estimates at BATS (average 0.3 3 mol Cm-2 yr(-1)) due to both higher mesozooplankton biomass and higher estimated mesozooplankton individual growth rates. Annual primary production at the two sites was similar (average 14.92 mol Cm-2 yr(-1) at HOT and 13.43 mol Cm-2 yr(-1) at BATS). Thus, mesozooplankton production was a greater fraction of primary production at HOT (0.05) as compared to BATS (0.02). Mesozooplankton potentially contributed more to the gravitational flux of carbon at HOT, where the ratio of the average annual estimate of mesozooplankton fecal pellet carbon production/annual estimate of carbon flux at the base of the euphotic zone was 1.03 compared to the same ratio of 0.39 at BATS. Mortality estimates were similar to estimates of mesozooplankton production when compared over the entire study period. The higher mesozooplankton biomass and derived rate parameters at HOT compared to BATS may be due to the more episodic nature of nutrient inputs at BATS, which could result in mismatches between increases in phytoplankton production and the grazing/production response by mesozooplankton. In addition, there is evidence to suggest that there are periodic blooms of gelatinous macrozooplankton (salps) at BATS that may not be captured sufficiently by the monthly sampling program. Thus the gelatinous zooplankton would add to the overall grazing impact on the phytoplankton at BATS as well as the contribution of zooplankton to the gravitational flux of biogenic material via fecal pellet production. (C) 2001 Elsevier Science Ltd. All rights reserved.
Keywords: marine planktonic copepods; northwestern sargasso sea; zooplankton; growth-rates; weight-specific growth; acartia-tonsa; egg-production;; southern-california; mesoscale eddies; food limitation; particle-flux
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Roman, M. R., Dam, H. G., Le Borgne, R., & Zhang, X. (2002). Latitudinal comparisons of equatorial Pacific zooplankton. Deep-Sea Research Part Ii-Topical Studies in Oceanography, 49(13-14), 2695–2711.
Abstract: Zooplankton biomass and rates of ingestion, egestion and production in the equatorial Pacific Ocean along 140degreesW and 180degrees exhibit maximum values in the High-Nutrient Low-Chlorophyll (HNLC) zone associated with equatorial upwelling (5degreesS-5degreesN) as compared to the more oligotrophic regions to the north and south. Zooplankton biomass and rates are not usually highest on the equator, but increase “downstream” of the upwelling center as the zooplankton populations exhibit a delayed response to enhanced phytoplankton production. The vertical distribution of zooplankton biomass in the equatorial HNLC area tends to be concentrated in surface waters and is more uniform with depth in oligotrophic regions to the north and south of the equatorial upwelling zone. In general, the amount of mesozooplankton (> 200 mum) carbon biomass is approximately 25% of estimated phytoplankton biomass and 30% of bacterial biomass in the HNLC area of the equatorial Pacific Ocean. Zooplankton grazing on phytoplankton is low in the equatorial Pacific Ocean, generally <5% of the total chlorophyll-a standing stock grazed per day. Based on estimates of metabolic demand, it is apparent that zooplankton in the equatorial Pacific Ocean are omnivores, consuming primarily microzooplankton and detritus. Estimated zooplankton growth rates in the warm waters of the HNLC equatorial Pacific Ocean are high, ranging from 0.58 d(-1) for 64-200 mum zooplankton to 0.08 d(-1) for 1000-2000 mum zooplankton. Thus, the numerical and functional response of equatorial zooplankton to increases in phytoplankton production are more rapid than normally occurs in sub-tropical and temperate waters. Potential zooplankton fecal pellet production, estimated from metabolic demand, is approximately 1.6 times the estimated gravitational carbon flux at 150 m in the zone of equatorial upwelling (5degreesS-5degreesN) and 1.1 times the export flux in the more oligotrophic regions to the north and south. The active flux of carbon by diel migrant zooplankton in the HNLC zone is a minor fraction of the gravitational flux (2% at 140degreesW, 4% at 180degrees) but increases in the more oligotrophic regions to the north and south where there is a deeper mixed layer and a greater relative proportion of diel migrant zooplankton. (C) 2002 Elsevier Science Ltd. All rights reserved.
Keywords: marine planktonic copepods; weight-specific growth; phytoplankton; growth; standing stocks; organic-carbon; fecal pellets; export flux;; nitrogen; mesozooplankton; diel
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Zhang, X., Dam, H. G., White, J. R., & Roman, M. R. (1995). Latitudinal Variations in Mesozooplankton Grazing and Metabolism in the Central Tropical Pacific During the Us Jgofs Eqpac Study. Deep-Sea Research Part Ii-Topical Studies in Oceanography, 42(2-3), 695–714.
Abstract: Pigment ingestion rates by three size classes of mesozooplankton (200-500 mu m, 500-1000 mu m and 1000-2000 mu m) within the euphotic zone were measured during the Survey 1 (February/March) and Survey 2 (August/September) cruises of the 1992 United States Joint Global Ocean Flux Study (U.S. JGOFS) in the central equatorial Pacific (EqPac). Survey 1 was characterized by El Nino conditions while Survey 2 was characterized by typical climatological conditions, The small animals (200-500 mu m) contributed more than 50% (range: 34-80%) to the total mesozooplankton grazing reported here. Mesozooplankton grazing was higher within the equatorial region (5 degrees S-5 degrees N) than at higher latitudes (5 degrees S-12 degrees S, 5 degrees N-12 degrees N). The carbon-specific ingestion rates of both the large and small animals tended to be higher during Survey 1 than during Survey 2. In contrast to the carbon-specific ingestion rates, the mesozooplankton biomass during Survey 1 was lower than that during Survey 2. Thus, the higher ingestion rates during Survey 1 were offset by lower biomass and the mesozooplankton grazing was quite similar during Surveys 1 and 2. Mesozooplankton removal of chi a was higher within the high-phytoplankton-biomass equatorial region than at higher latitudes during Survey 1, but regional differences were not significant during Survey 2. Mesozooplankton community grazing was equivalent to an average daily removal of 3.0% (range: 0.5-7.7%) and 2.2% (range: 0.8-3.5%) of the total chi a standing stock within the euphotic zone during Surveys 1 and 2, respectively. However, mesozooplankton grazing was equivalent to a daily removal of 47% (range: 15-91%) and 36% (range: 23-66%) of the > 5 mu m chi a standing stock within the euphotic zone during Surveys 1 and 2, respectively. Assuming a carbon-to-chi a ratio of 58, we estimate that mesozooplankton grazing removed an average of 6% (range: 2-12%) of C-14 primary production during Survey 1, and 5% (range: 2-10%) during Survey 2. Removal rates were not significantly different between the equatorial region and the higher latitudes. Rates of respiration and ammonium excretion were estimated from empirical models based on animal weight and water temperature. Phytoplankton ingestion could not satisfy the estimated daily maintenance-carbon demands of mesozooplankton. The shortages were more pronounced in the large size fraction than in the small size fraction. The estimated ammonium excretion by the mesozooplankton could support 4-15% and 3-17% of C-14 primary production during Surveys 1 and 2, respectively.
Keywords: santa-monica-basin; equatorial pacific; digestive acclimation;; pseudocalanus-minutus; planktonic copepods; calanus-pacificus;; acartia-tonsa; chlorophyll; food; size
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Zhang, X., Roman, M., Sanford, A., Adolf, H., Lascara, C., & Burgett, R. (2000). Can an optical plankton counter produce reasonable estimates of zooplankton abundance and biovolume in water with high detritus? Journal of Plankton Research, 22(1), 137–150.
Abstract: The Optical Plankton Counter (OPC) has been used in oceanic and fresh waters to estimate zooplankton abundance and biovolume. However, it is not clear whether the OPC can produce accurate estimates of zooplankton abundance and biovolume in waters with high detritus. In order to test the capability of the OPC to estimate zooplankton abundance and biovolume in Chesapeake Bay, two sets of laboratory experiments were conducted using water with high detritus concentrations collected from the upper Choptank estuary of Chesapeake Bay and laboratory cultured Artemia. Our results suggest that the OPC is able to produce accurate estimates of zooplankton biovolume after correcting for the influence of background detritus in all the detritus concentrations used, but accurate estimates of zooplankton abundance only in water with background detritus <100 particles l(-1). The relationship between light attenuation and OPC background particle concentrations provides a useful way to estimate OPC background particle concentrations when direct OPC background particle measurements are not available. Light attenuation corrected OPC particle abundance and particle volume gave accurate estimates of zooplankton abundance and biovolume. However, the accuracy of the corrected OPC measurements by the estimated background particle concentrations was not as high as the corrected OPC measurements by the direct background particle measurements.
Keywords: size distribution; calibration; ocean
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Elmore, A. J., Shi, X., Gorence, N. J., Li, X., Jin, H., Wang, F., et al. (2008). Spatial distribution of agricultural residue from rice for potential biofuel production in China. Biomass Bioenerg., 32(1), 22–27.
Abstract: In China, agricultural residues (particularly from rice) are widely used for energy and other applications, albeit on a localized scale and often at poor rates of efficiency. if some portion of this biomass were to be reallocated and transported to central biomass energy facilities, an initial component of the design process would be to gain an understanding of the spatial distribution of biomass production. In this paper, we present a method that utilizes China-wide data sets of net primary production (NPP) from the moderate-resolution imaging spectrometer (MODIS) and detailed land cover maps produced from Landsat-enhanced thematic mapper plus (ETM+) data to calculate the spatial distribution of rice straw for the period 2000-2004. Through a comparison with census statistics, we show that remote measures of rice straw can reasonably predict census results at the provincial scale. Remote sensing results have the added benefits of being a quick and inexpensive solution for providing spatially detailed information. Therefore, these data can be used for applications such as the spatial optimization of energy production infrastructure. In an error analysis including climate and land use variables, we found that data on sown rice area is the largest source of error. Therefore, the most important improvement to this method would be more accurate and more frequently updated maps of agricultural land use. (c) 2007 Elsevier Ltd. All rights reserved.
Keywords: China; agricultural residues; rice; net primary productivity; remote sensing; MODIS; GIS
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Hood, R. R., Zhang, X., Glibert, P. A., Roman, M. R., Stoecker, D. K., AF Hood, R. R., et al. (2006). Modeling the influence of nutrients, turbulence and grazing on Pfiesteria population dynamics. Harmful Algae, 5(4), 459–479.
Abstract: A semi-idealized marine ecosystem model, designed as a heuristic tool for exploring the population dynamics of non-inducible versus toxic forms of Pfiesteria is described. The model is based on empirical evidence suggesting that these differing functional types of Pfiesteria also differ substantially in terms of what they eat and how they utilize it to optimize their growth. Non-inducible strains are similar to other mixotrophic dinoflagellates, whereas toxic strains may consume organic matter and detritus, produce toxins and attack fish. In our model formulation we represent these differences in a simplified way: the non-inducible strain is kleptochloroplastidic and it can take up DIN, but it cannot utilize DON, whereas the toxic strain is heterotrophic, it cannot utilize DIN, but it can utilize DON directly. These differences give rise to very different impacts on prey and nutrient concentrations in our model. Under high DIN/DON ratio conditions, the non-inducible cells grew much faster and were therefore more likely to bloom, but this advantage is substantially mitigated when the DIN/DON ratio is low. A turbulence parameterization was also incorporated into our model. The effect of this was to reduce the grazing rate of Pfiesteria when turbulence levels are high. According to our model, increased turbulence is more detrimental to the toxic functional type because it grows more slowly. The further imposition of microzooplankton grazing in the model showed that top-down control effects can be very significant, which is consistent with both laboratory and field studies and the general idea that plankton blooms can only happen in the absence of substantial grazing control. In general, our model results suggest that non-toxic blooms are more likely to occur in more turbulent inorganic-nutrient rich conditions, which are often found in more open coastal and estuarine waters that are subject to high inorganic loading. In contrast, toxic blooms are more likely to occur in calm, organic-nutrient rich conditions, which are often found in shallow, protected tributaries that are subject to high organic nutrient loading. Our model results also support the idea that the absence of strong grazing pressure is a prerequisite to bloom formation for both non-inducible and toxic strains of Pfiesteria. These results are generally consistent with observed patterns of toxic Pfiesteria blooms in Chesapeake Bay, the Neuse River of North Carolina and many other coastal and estuarine environments. (c) 2006 Elsevier B.V. All rights reserved.
Keywords: Pfiesteria; ecosystem model; zooplankton grazing; turbulence; nutrients; DIN; DON PHYTOPLANKTON; GROWTH; PREY
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Roman, M. R., Pierson, J. J., Kimmel, D. G., Boicourt, W. C., & Zhang, X. (2012). Impacts of Hypoxia on Zooplankton Spatial Distributions in the Northern Gulf of Mexico. Estuaries Coasts, 35(5), 1261–1269.
Abstract: The northern Gulf of Mexico (NGOMEX) was surveyed to examine the broad-scale spatial patterns and inter-relationships between hypoxia (< 2 mg L-1 dissolved oxygen) and zooplankton biovolume. We used an undulating towed body equipped with sensors for conductivity, temperature, depth, oxygen, fluorescence, and an optical plankton counter to sample water column structure, oxygen, and zooplankton at high spatial resolution (1 m-vertical; 0.25-1 km-horizontal). We contrast the distribution of zooplankton during summer surveys with different freshwater input, stratification, and horizontal and vertical extent of bottom-water hypoxia. Bottom-water hypoxia did not appear to influence the total amount of zooplankton biomass present in the water column or the areal integration of zooplankton standing stock in the NGOMEX region surveyed. However, where there were hypoxic bottom waters, zooplankton shifted their vertical distribution to the upper water column during the day where they normally would reside in deeper and darker waters. When bottom waters were normoxic (> 2 mg L-1 dissolved oxygen), the daytime median depth of the water column zooplankton was on average 7 m deeper than the median depth of zooplankton in water columns with hypoxic bottom waters. A reduction in larger zooplankton when there were hypoxic bottom waters suggests that if zooplankton cannot migrate to deeper, darker water under hypoxic conditions, they may be more susceptible to size-selective predation by visual predators. Thus, habitat compression in the northern Gulf of Mexico due to hypoxic bottom water may have implications for trophic transfer by increasing the contact between predators and prey.
Keywords: Gulf of Mexico; Hypoxia; Spatial distributions; Zooplankton
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Zhang, X., Roman, M., Kimmel, D., McGilliard, C., & Boicourt, W. (2006). Spatial variability in plankton biomass and hydrographic variables along an axial transect in Chesapeake Bay. J. Geophys. Res.-Oceans, 111(C5 AR C05S11 DI ARTN C05S11).
Abstract: [ 1] High-resolution, axial sampling surveys were conducted in Chesapeake Bay during April, July, and October from 1996 to 2000 using a towed sampling device equipped with sensors for depth, temperature, conductivity, oxygen, fluorescence, and an optical plankton counter (OPC). The results suggest that the axial distribution and variability of hydrographic and biological parameters in Chesapeake Bay were primarily influenced by the source and magnitude of freshwater input. Bay-wide spatial trends in the water column-averaged values of salinity were linear functions of distance from the main source of freshwater, the Susquehanna River, at the head of the bay. However, spatial trends in the water column-averaged values of temperature, dissolved oxygen, chlorophyll-a and zooplankton biomass were nonlinear along the axis of the bay. Autocorrelation analysis and the residuals of linear and quadratic regressions between each variable and latitude were used to quantify the patch sizes for each axial transect. The patch sizes of each variable depended on whether the data were detrended, and the detrending techniques applied. However, the patch size of each variable was generally larger using the original data compared to the detrended data. The patch sizes of salinity were larger than those for dissolved oxygen, chlorophyll-a and zooplankton biomass, suggesting that more localized processes influence the production and consumption of plankton. This high-resolution quantification of the zooplankton spatial variability and patch size can be used for more realistic assessments of the zooplankton forage base for larval fish species.
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