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.     

The effect of burning and sheep grazing on soil water composition in a blanket bog: evidence for soil structural changes?

This study considers the impact of managed rotational burning of vegetation and sheep grazing upon the composition of soil waters within an upland peat soil. The study has considered soil water compositions from a complete factorial design of treatment plots where three different burning treatments were considered in replication with grazing and no grazing. All plots were sampled across a complete year with three dipwells in each plot. The study included aluminium (Al), iron, calcium, sodium (Na), magnesium (Mg), potassium, sulphate, chloride (Cl^?), bromide, fluoride, phosphate (PO )and nitrate; and in order to clarify the nature of the results, the pH, conductivity and dissolved organic carbon were also considered, but the major results for these are reported elsewhere. The study finds: (1) Ca, Na, Mg and PO concentrations are significantly lower on all burnt plots, with only Al concentration being significantly higher on burnt plots. (2) Only Cl^? showed any significant changes (a decrease) with the presence of sheep grazing, and then only when plots were also burnt. (3) A principal component analysis shows that the composition of most soil waters can be described by rainwater and soil water components, but in unburnt plots a base‐rich, high ionic strength water is sometimes present. The study suggests that burning, but not grazing, caused significant changes in soil water composition leading to increased interaction between incoming rainwaters and the peat soil but led to loss of interaction with deeper waters. However, no evidence was found for structural change in the soils even after long term (50 years) grazing and burning management. Copyright © 2007 John Wiley & Sons, Ltd.     

放牧对浑善达克沙地丘间低地植被群落及土壤的影响

为探究放牧对丘间低地植被群落及土壤的影响,分析了浑善达克沙地丘间低地在重度和轻度放牧下植被的地上生物量、物种重要值、物种丰富度、Shannon-Wiener多样性指数、Pielou均匀度指数、Simpson生态优势度指数及0~10 cm土壤中黏粒、粉粒、砂粒、有机碳和全氮含量。结果表明：（1）植被地上生物量、土壤黏粒、粉粒、砂粒、土壤有机碳和全氮含量在不同放牧程度下存在极显著差异（P<0.01）,物种丰富度、Shannon-Wiener多样性指数和Pielou均匀度指数在不同放牧程度下差异不显著（P>0.05）,Simpson生态优势度指数在不同放牧程度下差异显著（P<0.05）。（2）放牧区植被以禾本科为主,禾本科植被地上生物量占整个植被群落的54.88%~57.76%;重度放牧区植被地上生物量为63.59 g·m^-2,比轻度放牧区低26.61%。（3）羊草（Leymus chinensis）和狗尾草（Setaria viridis）是放牧区的优势种,其重要值依次为14.16%（重度放牧区）和19.10%（轻度放牧区）、13.40%（重度放牧区）和15.42%（轻度放牧区）,重度放牧下雾滨藜（Bassia dasyphylla）、克氏针茅（Stipa krylovii）和猪毛蒿（Artemisia scoparia）的重要值较显著高于轻度放牧区,而羊草则低于轻度放牧区。（4）重度放牧加剧了土壤沙漠化,使砂粒含量增加,草地生产力下降,土壤有机碳和全氮含量降低。     

景电灌区移民对祁连山植被恢复的相对生态价值

近些年来,生态系统的服务价值研究已成为生态学以及生态经济学领域中的一个热点问题。祁连山被称为伸进西北干旱区的一座湿岛,在我国"一带一路"建设中占有重要地位。景电灌区兴建以来从祁连山移出了大量农牧民。那么,景电灌区移民对祁连山植被恢复的生态价值如何呢?以景电灌区移民涉及到的祁连山东端景泰、古浪、天祝3县山区为研究区,用价值工程方法对从祁连山区向景电灌区移民退耕退牧还林还草的生态价值做了分析,并与模型因子当量法的计算结果进行了比较。结果表明:祁连山向景电灌区移民退耕退牧还林还草总的生态价值为37.458 1×10⁸元。其中,退耕还林还草的生态价值为37.438 6×10⁸元,退牧后草场植被盖度增加的生态价值为194.79×10⁴元。计算结果为用COSTANZA和谢高地模型因子当量法计算得的祁连山向景电灌区移民退耕退牧还林还草总的生态服务价值40.054 0×10⁸元的93.52%。两种方法计算结果祁连山向景电灌区移民退耕退牧还林还草总的生态价值为景电工程年总成本1.45×10⁸元的...     

Soil and vegetation seasonal changes in the grazing Andean Mountain grasslands

Andean grasslands ecosystems are fragile environments with rigorous climatologic conditions and low and variable food for the grazing. The Apolobamba area is located in the Bolivian Andean Mountains. Its high grasslands provide a natural habitat for wild and domestic camelids such as vicuna（Vicugna vicugna） and alpaca（Lama pacos）. The botanical diversity plays an essential role in maintaining vital ecosystem functions. The objectives of this research were to determine the seasonal changes in soil properties, to study the vegetation changes during the wet and dry seasons and the influence of soil properties and camelid densities on the vegetation in the Apolobamba grasslands. Four zones with different vicuna populations were selected to be studied. The following soil parameters were determined： total organic carbon, total nitrogen, available phosphorous, cation exchange capacity, exchangeable cations, pH and texture. The vegetation season changes were studied through botanical identification, above-ground biomass, plant cover and species richness. Results showed that some soil properties such as C/N ratio, CEC, silt and clay percentages kept stable against the seasonal changes. Generally, soil nutrients were relatively higher during the dry season in the surface and subsurface. The results did not point out the predominant vegetation growth during the wet season. The seasonal vegetation growth depended on each species. Thegood soil fertility corresponded to the highest plant cover. Soil fertility presented no influence on the above-ground biomass of the collected species. The negative influence of camelid grazing on soil properties could not be assessed. However, overgrazing could affect some plant species. Therefore, protection is needed in order to preserve the biodiversity in the Andean mountain grasslands.     

Preliminary study on different nutrient pools supplies for the phytoplankton growth in the Jiaozhou Bay in China in the fall of 2004

The source and significance of two nutrients, nitrogen and phosphorous, were investigated by a modified dilution method performed on seawater samples from the Jiaozhou Bay, in autumn 2004. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the internal and external nutrient pools, as well as nutrient supplied through remineralization by microzooplankton. The results indicated that the phytoplankton net growth rate increased in turn from inside the bay, to outside the bay, to in the Xiaogang Harbor. The phytoplankton maximum growth rates and microzooplankton grazing mortality rates were 1.14 and 0.92 d-1 outside the bay, 0.42 and 0.32 d-1 inside the bay and 0.98 and 0.62 d-1 in the harbor respectively. Outside the bay, the remineralized nitrogen (Kr=24.49) had heavy influence on the growth of the phytoplankton. Inside the bay, the remineralized phosphorus(Kr=3.49) strongly affected the phytoplankton growth. In the harbor, the remineralized phosphorus (Kr=3.73) was in larger demand by phytoplankton growth. The results demonstrated that the different nutrients pools supplied for phytoplankton growth were greatly in accordance with the phytoplankton community structure, microzooplankton grazing mortality rates and environmental conditions. It is revealed that nutrient remineralization is much more important for the phytoplankton growth in the Jiaozhou Bay than previously believed.     

Effects of grazing and climate change on species diversity in sandy grassland, Inner Mongolia, China

To understand the effects of animal grazing activities and climate change on sandy grassland vegetation in northern China, a field grazing and protected enclosure experiment was conducted from 1992 through 2006 in Horqin Sand Land, Inner Mongolia. The results showed that (1) the grazing was primary responsible for changes of the vegetation richness and diversity in the grazing grassland and that changing climate was the main reason for changes in the species richness and diversity in the grassland protected from grazing; (2) light and moderate grazing can promote restoration of the richness and the diversity in the degraded grassland, and heavy grazing could result in a decrease of the richness and diversity; (3) heavy grazing can result in significant decrease of the perennial diversity, and moderate and light grazing promotes increase of the perennial diversity; the grazing, whether heavy or moderate and light grazing, was beneficial to increase of the annual diversity; (4) heavy grazing was not beneficial to diversity of Graminean and Chenopodiaceae, and moderate and light grazing was favorable the diversity of Compositae and Chenopodiaceae; (5) the warm-humid climate was favorable to increase of the richness and the diversity, and the warm-drought climate could result in decease of the richness and the diversity; (6) increased precipitation was favorable to perennial diversity and the diversity of Graminean, Leguminosae, and Compositae, and decreased precipitation had few effects on the annual diversity and Chenopodiaceae diversity.     

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.     

Evidence for the grazing hypothesis: Grazing reduces phytoplankton responses of the HNLC ecosystem to iron enrichment in the western subarctic pacific (SEEDS II)

A mesoscale iron-enrichment study (SEEDS II) was carried out in the western subarctic Pacific in the summer of 2004. The iron patch was traced for 26 days, which included observations of the development and the decline of the bloom by mapping with sulfur hexafluoride. The experiment was conducted at almost the same location and the same season as SEEDS (previous iron-enrichment experiment). However, the results were very different between SEEDS and SEEDS II. A high accumulation of phytoplankton biomass (∼18 mg chl m⁻³) was characteristic of SEEDS. In contrast, in SEEDS II, the surface chlorophyll-a accumulation was lower, 0.8 to 2.48 mg m⁻³, with no prominent diatom bloom. Photosynthetic competence in terms of F _v/F _m for the total phytoplankton community in the surface waters increased after the iron enrichments and returned to the ambient level by day 20. These results suggest that the photosynthetic physiology of the phytoplankton assemblage was improved by the iron enrichments and returned to an iron-stressed condition during the declining phase of the bloom. Pico-phytoplankton (<2 μm) became dominant in the chlorophyll-a size distribution after the bloom. We observed a nitrate drawdown of 3.8 μM in the patch (day 21), but there was no difference in silicic acid concentration between inside and outside the patch. Mesozooplankton (copepod) biomass was three to five times higher during the bloom-development phase in SEEDS II than in SEEDS. The copepod biomass increased exponentially. The grazing rate estimation indicates that the copepod grazing prevented the formation of an extensive diatom bloom, which was observed in SEEDS, and led to the change to a pico-phytoplankton dominated community towards the end of the experiment.