Temperature and phosphorus interacts in controlling the picoplankton carbon flux in the Adriatic Sea: an experimental versus field study

Temperature and phosphorus positively interacted in controlling picoplankton biomass production and its transfer towards higher trophic levels. Two complementary approaches (experimental and field study) indicated several coherent patterns: (1) the impact of temperature on heterotrophic bacteria was high at temperatures lower than 16°C and levelled off at higher temperatures, whereas this impact on autotrophic picoplankton was linear along the entire range of the investigated temperatures; (2) the addition of phosphorus increased the values of picoplankton production and grazing, but did not change the nature of their relationships with temperature substantially; (3) the picoplankton carbon flux towards higher trophic levels was larger during the warmer months (grazing by HNF dominated during the warmer period and by ciliates during the colder period) and also strengthened in conditions without phosphorus limitation; (4) the hypothesis that the available phosphorus can be better utilized at higher temperatures was confirmed for both autotrophic and heterotrophic picoplankton; (5) the hypothesis that the rise in temperature stimulates growth only in conditions of sufficient phosphorus was confirmed only for heterotrophic bacteria. Therefore, in the global warming scenario, an increase of the picoplankton carbon flux towards higher trophic levels can be expected in the Adriatic Sea, particularly under unlimited phosphorus conditions.     

Autotrophic and Heterotrophic Picoplankton in Wetlands: Differences with Lake Patterns

This study describes the occurrence, importance and seasonal patterns of picoplankton in two wetlands (TDNP and La Safor), and compares them to a system of fifteen interconnected lakes (Ruidera). In TDNP we performed a six‐year monthly study in three sites of the wetland. Bacterial abundance increased throughout time and the autotrophic picoplankton (APP) range was wide (up to 33 × 10⁶ cells/ml). The annual averaged APP contribution to total picoplankton and phytoplankton biovolumes was 0.5–22% and 0.03–6% respectively. There were large differences among sites in terms of APP absolute and relative abundance and seasonal patterns. In La Safor, the APP relative contribution to picoplankton and phytoplankton biovolumes was 0–25% and 0–40%, respectively, while in the Ruidera lakes was 0–47% and 0–5%, respectively. In the three systems there was a significant correlation between bacterial abundance and chlorophyll a but the slopes of the linear regressions were different. No significant relationships were found of APP abundance and trophic status in the wetlands, but were noted in the lake system. There was no clear relationship of APP contribution to total phytoplankton biomass to the trophic gradient in wetlands. In the lakes, the higher contribution of APP was found in those with higher trophic levels.     

Systematics and Ecology of Chlorophyte Picoplankton in German Inland Waters along a Nutrient Gradient

The community structure and succession of autotrophic picoplankton in several oligotrophic to hypertrophic German freshwater ecosystems were studied with emphasis on the occurrence and characterization of chlorophyte picoplankton. Depending on the trophic status and the time of the year, the relation of green eukaryotic picoplankton to picocyanobacteria, the contribution of the picoplankton to the total phytoplankton biomass, and the succession and dominance of picoplankton groups changed considerably. A significant correlation between the picoplankton abundances, their biomass and their biomass contribution could not be found. Although the chlorophyte picoplankton were similar with respect to their ultrastructure, phylogenetic analyses of the rbcL genes revealed that these organisms evolved independently within several green algal lineages. The most common picoplanktonic green algae in the lakes that were studied belong to the genera Choricystis and Pseudodictyosphaerium. Considering the new molecular biological findings, the systematics of picoplanktonic green algae from freshwater and marine habitats are discussed.     

Effects of UV radiation on grazing by two marine heterotrophic nanoflagellates on autotrophic picoplankton

Phytoplankton <4-3 µm in diameter, or autotrophic picoplankton,can constitute the majority of the biomass and productivityof photosynthetic organisms in marine and freshwater systems.Indirect evidence has indicated that mortality of autotrophicpicoplankton occurs principally at night in the open ocean,but continuously in coastal water. The predominant view of thefate of autotrophic picoplankton production in the ocean isthat they are consumed by heterotrophic nanoflagellates. A possiblemechanism to explain these observations is that grazing of heterotrophicnanoflagellates on autotrophic picoplankton is inhibited byultraviolet radiation (W), at least in clear open-ocean environments.A series of laboratory experiments was conducted to examinethe effects of UV radiation on the grazing impact of two heterotrophicnanoflagellates on Synechococcus spp., a commonly occurringgenus of autotrophic picoplankton. The two nanoflagellates usedwere Paraphysomonas bandaieensis and Paraphysomonas imperforata.For both nanoflagellates, there was an inverse relationshipbetween the grazing mortality of Synechococcus and UV irradiance.The grazing mortality of Synechococcus was reduced less withP.imperforata than with P.bandaiensis. In some experiments,the effect of UV on the grazing impact of the nanoflagellatepopulations was caused in part by UV-related reductions in nanoflagellatesurvival. However, UV reduced the grazing impact of nanoflagellatesprimarily by reducing the rates of consumption of Synechococcusby individual nanoflagellates, to a degree directly relatedto UV irradiance. The results suggat that UV radiation may bean important selection factor in clear open-ocean water, andthat in order to predict the effect of increasing UV radiationon marine microbial plankton communities, we must consider interactionsbetween trophic levels as well as effects on single trophiclevels.     

Picoplankton in Six New Zealand Lakes: Abundance in Relation to Season and Trophic State

The abundance of picoplankton (0.2-2 μm) was measured seasonally in the surface waters of six New Zealand lakes that represent a range of trophic states. The lakes were: Wakatipu, Te Anau, Manapouri, Hayes, Mahinerangi and Ross Creek Reservoir. Among the lakes, picoplankton abundance was associated positively with temperature; picoplankton were most abundant in summer and autumn when they attained densities of 108,000-270,000 cells/ml in the oligotrophic lakes. In these lakes, prokaryotic picoplankton was generally an order of magnitude more abundant than eukaryotic picoplankton. Consistent with the hypothesis that picoplankton are more important in oligotrophic than eutrophic ecosystems, there was a weak negative correlation between the density of prokaryotic picoplankton in the lakes and the level of chlorophyll a. The presence of large numbers of chroococcoid cyanobacteria in the guts of Ceriodaphnia dubia and Bosmina meridionalis implies that prokaryotic picoplankton are collected, but not digested, by these species.     

Is the ‘microbial loop’ an early warning indicator of anthropogenic stress?

Various components of the Microbial loop such as bacteria, heterotrophic nanoflagellates and autotrophic picoplankton were analyzed, for the first time across the Great Lakes, during a cruise in the summer of 1988. In addition, the size fractionated primary productivity using carbon-14 techniques was also determined. The statistical analysis indicated that bacteria, autotrophic picoplankton and ultraplankton/picoplankton productivity were significantly higher in Lakes Ontario and Erie than Lakes Huron and Michigan. The autotrophic picoplankton and ultraplankton/picoplankton productivity was higher in Lake Erie compared to Lake Ontario.The autotrophic picoplankton showed sensitivity to nutrients and contaminants in various types of environments. A dramatic decrease of autotrophic picoplankton in eutrophic-contaminated areas, such as Ashbridges Bay, Hamilton Harbour and western Lake Erie was observed. Conversely, in Saginaw Bay, another eutrophic environment, the autotrophic picoplankton were significantly higher than in Lake Huron. The sensitivity of autotrophic picoplankton to nutrients/contaminants might have implications to trophic interactions. Our results suggest that structural and functional characteristics of the microbial loop may be operating differently in stressed versus unstressed ecosystems. The possibility of using autotrophic picoplankton as an early warning indicator of environmental perturbation is proposed.     

The microbial food web structure of a hypertrophic warm-temperate shallow lake, as affected by contrasting zooplankton assemblages

The composition of zooplankton is known to affect the structure of the microbial trophic web. The zooplankton of the hypertrophic Laguna Chascomús (Argentina) is generally dominated by rotifers and cyclopoids copepods. An unusual dominance by small-cladocerans was observed after a massive winter fish kill in 2007. We hypothesized that small-cladocerans would increase the grazing pressure on heterotrophic flagellates (HF), reducing the degree of coupling between HF and picoplankton. The aim of this study was to investigate the microbial food web structure under two contrasting zooplankton assemblages. The lake was sampled every other week between 2007 and 2009. The abundances of heterotrophic bacteria (HB) and picocyanobacteria (Pcy) laid among the highest values reported for aquatic systems (>10⁸ and 10⁷ cells ml^?1, respectively). Pcy averaged 53% of total picoplanktonic biomass. When small-cladocerans dominated zooplankton HF reached the higher abundance (>10⁵ cells ml^?1) and picoplankton showed the opposite pattern, while the proportion of grazing resistant morphologies (i.e. microaggregates of Pcy) was higher. In contrast, when rotifers dominated, HF abundance decreased and picoplankton increased. Our data suggest that the degree of HF–HB coupling was affected by changes in zooplankton dominance. In contrast to our initial hypothesis, the present results suggest that large numbers of rotifers (>5,000 ind. l^?1) are more efficient than small-cladocerans at controlling HF populations.     

Picoplankton photosynthesis and diurnal variations in photosynthesis-irradiance relationship in a eutrophic and meso-oligotrophic lake

The abundance and relative importance of autotrophic picoplankton were investigated in two lakes of different trophic status. In the eutrophic lake, measurements of primary production were performed on water samples in situ and in a light incubator three times during the day whereas for the oligotrophic lake, only one measurement of primary production was performed on water samples in the incubator. Dark-carbon losses of phytoplankton from Lake Loosdrecht were investigated in time series. Cell numbers of autotrophic picoplankton in eutrophic Lake Loosdrecht (3.2 × 10⁴ cells ml^–1) were lower than in meso-oligotrophic Lake Maarsseveen (9.8 and 11.4 × 10⁴ cells ml^–1 at the surface and bottom respectively). In the phytoplankton of both lakes the ratio of picoplankton production increased with decreasing light intensity. In Lake Loosdrecht depth-integrated contribution of picoplankton to total photosynthesis was less than 4%. The P-I-relationship showed diurnal variations in light saturated photosynthesis, while light limited carbon uptake remained constant during the day. Dark carbon losses from short-term labelled phytoplankton during the first 12 hours of the night period accounted for 10–25% of material fixed during the preceeding light period.     

Ecophysiological and Trophic Implications of Light-Stimulated Amino Acid Utilization in Marine Picoplankton

By using microautoradiography, light-stimulated utilization of dissolved amino acids for natural marine phytoplankton assemblages was demonstrated. The <2-μm-size (diameter) picoplankton, known to be a dominant fraction of marine primary production, revealed a widespread capability for this process. Autofluorescent (chlorophyll a-containing) picoplankton and some larger phytoplankton from diverse oceanic locations, as well as isolates of the representative cyanobacterial picoplankton Synechococcus spp. (WH7803, WH8101), showed light-stimulated incorporation of amino acids at trace concentrations. Dark-mediated amino acid utilization was dominated by nonfluorescent bacterial populations. Among autofluorescent picoplankton, light-stimulated exceeded dark-mediated amino acid incorporation by 5 to 75%; light-stimulated amino acid incorporation was only partially blocked by the photosystem II inhibitor 3(3,4-dichloro-phenyl)-1,1-dimethy-lurea (2 × 10^-5 M), suggesting a photoheterotrophic incorporation mechanism. Parallel light versus dark incubations with glucose and mannitol indicated a lack of light-stimulated utilization of these nonnitrogenous compounds. Since marine primary production is frequently nitrogen limited, light-mediated auxotrophic utilization of amino acids and possibly other dissolved organic nitrogen (DON) constituents may represent exploitation of the relatively large DON pool in the face of dissolved inorganic nitrogen depletion. This process (i) increases the efficiency of DON retention at the base of oceanic food webs and (ii) may in part be responsible for relatively high rates of picoplankton production under conditions of chronic dissolved inorganic nitrogen limitation. Picoplanktonic recycling of organic matter via this process has important ramifications with respect to trophic transfer via the “microbial loop.”     

Seasonal Trophic Niche Shift and Cascading Effect of a Generalist Predator Fish

Few studies have examined how foraging niche shift of a predator over time cascade down to local prey communities. Here we examine patterns of temporal foraging niche shifts of a generalist predator (yellow catfish, Pelteobagrus fulvidraco) and the abundance of prey communities in a subtropical lake. We predicted that the nature of these interactions would have implications for patterns in diet shifts and growth of the predator. Our results show significant decreases in planktivory and benthivory from late spring to summer and autumn, whereas piscivory increased significantly from mid-summer until late autumn and also increased steadily with predator body length. The temporal dynamics in predator/prey ratios indicate that the predation pressure on zooplankton and zoobenthos decreased when the predation pressure on the prey fish and shrimps was high. Yellow catfish adjusted their foraging strategies to temporal changes in food availability, which is in agreement with optimal foraging theory. Meanwhile the decrease in planktivory and benthivory of yellow catfish enabled primary consumers, such as zooplankton and benthic invertebrates, to develop under low grazing pressure via trophic cascading effects in the local food web. Thus, yellow catfish shifts its foraging niche to intermediate consumers in the food web to benefit the energetic demand on growth and reproduction during summer, which in turn indirectly facilitate the primary consumers. In complex food webs, trophic interactions are usually expected to reduce the strength and penetrance of trophic cascades. However, our study demonstrates strong associations between foraging niche of piscivorous fish and abundance of prey. This relationship appeared to be an important factor in producing top-down effects on both benthic and planktonic food webs.     

Autotrophic picoplankton in southern Lake Baikal: abundance, growth and grazing mortality during summer

Autotrophic picoplankton were highly abundant during the thermalstratification period in late July in the pelagic area (waterdepth 500–1300 m) of southern Lake Baikal; maximum numberswere 2 x 10⁶ cells ml^–1 in the euphotic zone ({small tilde}15m). Unicellular cyanobacteria generally dominated the picoplanktoncommunity, although unidentified picoplankton that fluorescedred under blue excitation were also abundant (maximum numbers4 x 10⁵ cells ml^–1) and contributed up to {small tilde}40%of the total autotrophic picoplankton on occasions. Carbon andnitrogen biomasses of autotrophic picoplankton estimated byconversion from biovolumes were 14–84 µg C l^–1and 3.6–21 µg N l^–1. These were comparableto or exceeded the biomass of heterotrophic bacteria. Autotropicpicoplankton and bacteria accounted for as much as 33% of paniculateorganic carbon and 81% of nitrogen in the euphotic zone. Measurementsof the photosynthetic uptake of [^l4C]bicarbonate and the growthof picoplankton in diluted or size-fractionated waters revealedthat 80% of total primary production was due to picoplankton,and that much of this production was consumed by grazers inthe <20 µ.m cell-size category. These results suggestthat picoplankton-protozoan trophic coupling is important inthe pelagic food web and biogeochemical cycling of Lake Baikalduring summer.     

Predation on marine picoplankton populations examined with an 'add-in' approach

NW Mediterranean surface water was spiked with picoplanktonprey (heterotrophic bacteria or cyanobacteria) or predators(bacterivorous microflagellates or ciliates) to investigatedifferential grazing pressure on picoplankton populations. Addinga particular prey type did not yield different growth patternsfor heterotrophic bacteria and cyanobacteria. but gave eithersimilar, positive, effects on both picoplankton types or similarnegative effects. Natural populations of both predator typesincreased with additions of cyanobacteria, but not heterotrophicbacteria. Ciliate additions gave marked decreases in cyanobacteria.While individual groups of grazers may preferentially consumecyanobacteria, selective grazing is probably not responsiblefor the maintenance of apparently stable populations of differentgroups of picoplankters during the summer.     

Occurrence,Dynamics and Production of Picoplankton in Hungarian Shallow Lakes

Phototrophic picoplankton were detected in 10⁵-10⁶ cells/ml concentrations in seven water bodies of differing chemistry and trophic level. Dominant picoplankters were, at all sites, coccoid cyanobacteria of 0.8-1.2 μm dimensions, exhibiting red or yellow autofluorescence. Apart from the effects of water temperature their quantitative dynamics were significantly influenced by the nitrogen supply and the herbivorous zooplankton (Cladocera). Their maximum contribution to the total planktonic primary production was about 50%, both in mesotrophic and hypereutrophic environments. However during the bloom of filamentous nitrogen-fixing blue-greens their role became negligible. In phytoplankton communities the significance of picoplankton is overestimated several times, when based on the cell count, while it is underestimated on the basis of biomass. The most useful characteristics of phytoplankton size groups was the total surface area of their cells.     

The response of marine picoplankton to ocean acidification

Since industrialization global CO(2) emissions have increased, and as a consequence oceanic pH is predicted to drop by 0.3-0.4 units before the end of the century - a process coined 'ocean acidification'. Consequently, there is significant interest in how pH changes will affect the ocean's biota and integral processes. We investigated marine picoplankton (0.2-2?μm diameter) community response to predicted end of century CO(2) concentrations, via a 'high-CO(2) ' (～?750?ppm) large-volume (11?000?l) contained seawater mesocosm approach. We found little evidence of changes occurring in bacterial abundance or community composition due to elevated CO(2) under both phytoplankton pre-bloom/bloom and post-bloom conditions. In contrast, significant differences were observed between treatments for a number of key picoeukaryote community members. These data suggested a key outcome of ocean acidification is a more rapid exploitation of elevated CO(2) levels by photosynthetic picoeukaryotes. Thus, our study indicates the need for a more thorough understanding of picoeukaryote-mediated carbon flow within ocean acidification experiments, both in relation to picoplankton carbon sources, sinks and transfer to higher trophic levels.     

Nutrient and Grazer-Mediated Effects on Picoplankton and Size Structure in Phytoplankton Communities

This study examines the factors which contribute to the abundance of algal picoplankton in lakes. A three-year field study of a meso-eutrophic lake was compared with observations from oligotrophic and highly eutrophic lakes in the region. Trophic state alone (oligotrophic vs. eutrophic) was not a good predictor of the importance of picoplankton; smaller cells were relatively abundant when phosphorus was limiting other phytoplankters, but also when nitrogen was in surplus. Subsequent field experiments found that picoplankton growth was stimulated by N, but not by P additions. This relationship was strongly affected by light and grazer levels. Grazers apparently mediate the effects of nutrient deficiency, and favor the growth of larger algal size classes, especially nanoplankton. The flux of P within experimental enclosures was controlled by picoplankton abundance under low nutrient conditions, but was a function of total phytoplankton biomass under P surplus.     

Predation impact of ciliated and flagellated protozoa during a summer bloom of brown sulfur bacteria in a meromictic coastal lake

Anaerobic phagotrophic protozoa may play an important role in the carbon flux of chemically stratified environments, especially when phototrophic sulfur bacteria account for a high proportion of the primary production. To test this assumption, we investigated the vertical and temporal distribution of microbial heterotrophs and of autotrophic picoplankton throughout the water column of the meromictic coastal lake Faro (Sicily, Italy), in the summer of 2004, coinciding with a bloom of brown-colored green sulfur bacteria. We also assessed the grazing impact of ciliated and flagellated protozoa within the sulfur bacteria plate using a modification of the fluorescently labeled bacteria uptake approach, attempting to minimize the biases intrinsic to the technique and to preserve the in situ anoxic conditions. Significant correlations were observed between ciliate biomass and bacteriochlorophyll e concentration, and between heterotrophic nanoflagellate biomass and chlorophyll a concentration in the water column. The major predators of anaerobic picoplankton were pleuronematine ciliates and cryptomonad flagellates, with clearances of 26.6 and 9.5 nL per cell h⁻¹, respectively, and a cumulative impact on the picoplankton gross growth rate ranging between 36% and 72%. We concluded that protozoan grazing channels a large proportion of anaerobic picoplankton production to higher trophic levels without restraining photosynthetic bacteria productivity.     

互利关系和植物多样性对节肢动物群落不同营养级的影响路径和强度

食物网中的上行效应和下行效应对于群落的动态和生态系统功能有十分重要的影响,旨在探讨互利关系和植物多样性对节肢动物群落中食物网不同营养级之间的影响。通过随机裂区试验方法,分别设置了3种蚂蚁-紫胶虫互利关系处理(有互利关系、无互利关系和自然对照)以及3种植物多样性处理(单一种植、2树种混植和3树种混植),于2016年8月和9月分两次用手捡法、网扫法和震落法采集试验地寄主植物上所有的节肢动物,并按照不同营养级将其分类。利用结构方程模型分析方法对不同营养级之间的相互作用的路径和强度进行了比较,结果显示:1)互利关系对捕食者和消费者均有显著的下行作用,有互利关系处理下蚂蚁对捕食者的路径强度要强于自然对照组,互利关系对捕食者的影响要强于对消费者的影响。2)植物多样性会通过影响植物的生物量而对消费者和捕食者产生显著的上行效应影响,这种影响会随着营养级的升高而显著减小。3)消费者主要受植物多样性的上行效应影响,而捕食者主要受互利关系的下行效应影响。有互利关系的食物网结构更加复杂,营养级之间的相互作用更为显著。探讨了以蚂蚁-紫胶虫互利关系为核心作用的紫胶林生态系统中互利关系和植物多样性对节肢动物食物网中各营养级的影响,揭示了上行效应和下行效应对各营养级的作用途径和强度,其结果有一定的理论参考价值。     

Seasonal Changes in Chlorophyll-containing Picoplankton Populations of Ten Lakes in Northern England

Key features of photosynthetic picoplankton populations were compared during 1988 in ten lakes in northern England ranging from oligotrophic to slightly eutrophic; two of the three eutrophic lakes were shallow and lacked a thermocline. Measurements were made at 0.5 m depth of temperature, total chlorophyll a, chlorophyll-containing picoplankton cell density, mean picoplankton cell volume and percentage of phycoerythrin-rich cells in the total picoplankton population. All lakes showed maxima for total chlorophyll concentration and picoplankton cell density in mid- to late summer. The maximum value for picoplankton density ranged from 3.4 × 10³ (Esthwaite Water) to 1.3 × 10⁶ cells ml⁻¹ (Ennerdale Water). There was a significant negative relationship (p < 0.05) between log₁₀ of maximum picoplankton cell density and maximum total chlorophyll, the latter being taken as an indicator of lake trophic status. The ratio of maximum to minimum picoplankton density during the year in a particular lake ranged from 39 to 2360 and showed no obvious relationship to lake type. Overall, the seasonal range in picoplankton density was about one order of magnitude greater than the range in total chlorophyll a, but there were considerable differences between lakes. Phycoerythrin-rich picoplankton as a percentage of total picoplankton reached a maximum in summer in all lakes. Values were always very low (<5%) in the two shallow eutrophic lakes, but reached 97% and over in the four most oligotrophic lakes. In two of the oligotrophic lakes, Wast-water and Ennerdale Water, phycoerythrin-rich picoplankton was a major component of the summer phytoplankton biomass.     

Effect of larval ontogeny, turbulence and light on prey attack rate and swimming activity in herring larvae

The effects of ontogeny (larval size), light and turbulence on the attack rate and swimming activity (proportion of time swimming and duration of swimming bout) of herring larvae (15-28 mm TL) have been investigated. Emphasis was put on the experimental design in order to create a set-up where the turbulence intensity distribution could be accurately measured as well as controlled in the entire experimental tank.Both larval size (ontogeny) and light had a significant positive effect on prey attack rate. Likewise, an intermediate increase in turbulence had a positive effect on prey attack rate, but this effect was dependent of light intensity and larval size.At low light (1.5 μE m² s⁻¹) intermediate turbulence increased the prey attack rate significantly for larger larvae (26 and 28 mm), while at high light (18 μE m² s⁻¹) intermediate turbulence had only a significant positive effect on the attack rate of smaller larvae 20 and 23 mm.In general, our data show a dome-shaped response of turbulence on attack rate and a U-shaped response of turbulence on swimming activity.For herring larvae >20 mm, the maximum (attack rate) and minimum (swimming activity) response of turbulence were found at intermediate turbulence intensities (energy dissipation rates between 7∗10⁻⁸ and 1∗10⁻⁶ W/kg). The highest turbulence level tested (8∗10⁻⁶ W/kg) showed only negative effects, as attack rates where at the lowest and swimming activity at the highest.Swimming activity increased with larval size or light, and decreased at intermediate turbulence. Compared to turbulent intensities under natural conditions this implies that larger herring larvae at 10 m depth have to be exposed to wind speeds of more than 17 m/s before negative effects on attack rate and swimming activity occurs.     

Effects of disturbance on species diversity: a multitrophic perspective

Models of the effects of disturbance on ecological communities have largely considered communities of competing species at a single trophic level. In contrast, most real communities have multiple interacting trophic levels. I explored several versions of simple single- and multitrophic models to determine whether predictions of the intermediate disturbance hypothesis (IDH), derived from considering only a single trophic level, apply to multitrophic situations. The IDH was predicted by models of competing species at a single trophic level but did not hold in many situations with more natural trophic structure. In general, basal species in a food web tended to follow the IDH, whereas competitors at top trophic levels did not. Additional analyses indicated that outside immigration interacted with trophic structure to produce widely differing predictions about the consequences of disturbance and that density-dependent disturbance events could recapture the IDH in some multiple trophic level situations. Model predictions matched the results of empirical studies to date: the IDH has generally been supported for species competing for nondynamic basal resources but not for mobile aquatic invertebrates at higher trophic levels. The model analysis also verified basic predictions of verbal models addressing the effects of physical stress. Three different aspects of disturbance and their contributions to species coexistence were identified: changes in average mortality rates, changes in temporal variability, and changes in spatial heterogeneity. The results indicate that the IDH should be applied with caution to real multitrophic communities.