Population-level compensation impedes biological control of an invasive forb and indirect release of a native grass

The intentional introduction of specialist insect herbivores for biological control of exotic weeds provides ideal but understudied systems for evaluating important ecological concepts related to top-down control, plant compensatory responses, indirect effects, and the influence of environmental context on these processes. Centaurea stoebe (spotted knapweed) is a notorious rangeland weed that exhibited regional declines in the early 2000s, attributed to drought by some and to successful biocontrol by others. We initiated an experiment to quantify the effects of the biocontrol agent, Cyphocleonus achates, on Ce. stoebe and its interaction with a dominant native grass competitor, Pseudoroegneria spicata, under contrasting precipitation conditions. Plots containing monocultures of each plant species or equal mixtures of the two received factorial combinations of Cy. achates herbivory (exclusion or addition) and precipitation (May-June drought or "normal," defined by the 50-year average) for three years. Cy. achates herbivory reduced survival of adult Ce. stoebe plants by 9% overall, but this effect was stronger under normal precipitation compared to drought conditions, and stronger in mixed-species plots compared to monocultures. Herbivory had no effect on Ce. stoebe per capita seed production or on recruitment of seedlings or juveniles. In normal-precipitation plots of mixed composition, greater adult mortality due to Cy. achates herbivory resulted in increased recruitment of new adult Ce. stoebe. Due to this compensatory response to adult mortality, final Ce. stoebe densities did not differ between herbivory treatments regardless of context. Experimental drought reduced adult Ce. stoebe survival in mixed-species plots but did not impede recruitment of new adults or reduce final Ce. stoebe densities, perhaps due to the limited duration of the treatment. Ce. stoebe strongly depressed P. spicata reproduction and recruitment, but these impacts were not substantively alleviated by herbivory on Ce. stoebe. Population-level compensation by dominant plants may be an important factor inhibiting top-down effects in herbivore-driven and predator-driven cascades.     

Salinity and substratum effects on condition index of the bivalve Rangia cuneata

The condition index (CI) of the clam Rangia cuneata was measured monthly at 8 stations in 2 substrate types and 3 salinity regimes in the James River estuary, Virginia, USA. CI was higher in clams from sand bottoms than from mud, and decreased in both substrata in fresher water. Values were lowest in early spring and at a maximum in early autumn, with a pronounced autumn peak in the sand substratum. This pattern coincided with the annual salinity cycle. A field experiment showed that some factor associated with the water overlying a sand or mud substratum, rather than the nature of the substratum itself, was important in determining CI. The hypothesis is presented that this factor may be the higher suspended solids concentrations measured immediately over mud bottoms.Virginia Institute of Marine Science Contribution No. 782.     

Population and leaf-level variation of iridoid glycosides in the invasive weed Verbascum thapsus L. (common mullein): implications for herbivory by generalist insects

Plant–insect interactions, which are strongly mediated by chemical defenses, have the potential to shape invasion dynamics. Despite this, few studies have quantified natural variation in key defensive compounds of invasive plant populations, or how those defenses relate to levels of herbivory. Here, we evaluated variation in the iridoid glycosides aucubin and catalpol in rosette plants of naturally occurring, introduced populations of the North American invader, Verbascum thapsus L. (common mullein; Scrophulariaceae). We examined two scales that are likely to structure interactions with insect herbivores—among populations and within plant tissues (i.e., between young and old leaves). We additionally estimated the severity of damage incurred at these scales due to insect chewing herbivores (predominantly grasshoppers and caterpillars), and evaluated the relationship between iridoid glycoside content and leaf damage. We found significant variation in iridoid glycoside concentrations among populations and between young and old leaves, with levels of herbivory strongly tracking leaf-level investment in defense. Specifically, across populations, young leaves were highly defended by iridoids (averaging 6.5× the concentration present in old leaves, and containing higher proportions of the potentially more toxic iridoid, catalpol) and suffered only minimal damage from generalist herbivores. In contrast, old leaves were significantly less defended and accordingly more substantially utilized. These findings reveal that quantitative variation in iridoid glycosides is a key feature explaining patterns of herbivory in an introduced plant. In particular, these data support the hypothesis that defenses limit the ability of generalists to feed on mullein’s well-defended young leaves, resulting in minimal losses of high-quality tissue, and increasing performance of this introduced species.     

Condition-dependent control of paternity by female purple martins: implications for coloniality

Proposed causal links between extra-pair copulation (EPC) and colony formation in socially monogamous birds hinge on the question of which sex controls fertilizations. We examined in colonial purple martins Progne subis (1) whether EPCs were forced or accepted by females, and (2) the degree to which apparently receptive females were able to obtain EPCs against their mates’ paternity defenses. Paternity analyses of multilocus DNA fingerprinting confirmed previous findings of a marked relationship between age class and extra-pair fertilizations (EPFs), with young males losing paternity of 43% (n = 53) of their putative offspring compared to 4% (n = 85) by old males. All assignable extra-pair offspring were sired by old males, with one male obtaining most EPFs each year. Contrary to the hypothesis that EPCs are forced, EPF frequency within age class did not increase with seasonal increases in the number of males per fertile female. Whereas the male control hypothesis predicted that the male age class that mate-guarded more would be cuckolded less, the reverse was true: young males guarded significantly more intensely. The male age class difference in cuckoldry could not be explained by the possibility that young and inexperienced females (which are usually paired to young males) were more vulnerable to forced copulation because EPFs were unrelated to female age. These findings suggest that females (1) pair with old males and avoid EPCs, or (2) pursue a mixed mating strategy of pairing with young males and accepting EPCs from old males. The receptivity to EPCs by females paired to young males put them in conflict with their mates. Two factors determined the paternity achieved by young males: (1) the relative size of the male to the female, with young males achieving much higher paternity when they were larger than their mates, and (2) the intensity of mate-guarding. Both variables together explained 77% of the variance in paternity and are each aspects of male-female conflict. Given female receptivity to EPCs, mate-guarding can be viewed as male interference with female mating strategies. We conclude that EPCs are rarely or never forced, but the opportunity for females paired to young males to obtain EPCs is relative to the ability of their mates to prevent them from encountering other males. Evidence of mixed mating strategies by females, combined with other features of the martin mating system, is consistent with the female-driven “hidden lek hypothesis” of colony formation which predicts that males are drawn to colonies when females seek extra-pair copulations. Received: 23 March 1995/Accepted after revision: 14 January 1996     

Carbon isotope effects during precipitation of barium carbonate: implications for environmental studies

Here we demonstrate that precipitation of dissolved inorganic carbon (DIC) to at pH 10.26, 11.38 and 12.11 does not cause measurable carbon isotope fractionation. However, NaOH solution prepared under standard laboratory procedure and used to increase pH value for precipitation of BaCO₃ can be contaminated by atmospheric CO₂. In our experiment, δ¹³C value in the contaminant DIC was −24.30‰, which resulted in decrease in the δ¹³C(DIC) value by 0.03, 0.07 and 0.22 at pH 10.26, 11.38 and 12.11, respectively.     

Herbivore vs. nutrient control of marine primary producers: context-dependent effects

Pervasive overharvesting of consumers and anthropogenic nutrient loading are changing the strengths of top-down and bottom-up forces in ecosystems worldwide. Thus, identifying the relative and synergistic roles of these forces and how they differ across habitats, ecosystems, or primary-producer types is increasingly important for understanding how communities are structured. We used factorial meta-analysis of 54 field experiments that orthogonally manipulated herbivore pressure and nutrient loading to quantify consumer and nutrient effects on primary producers in benthic marine habitats. Across all experiments and producer types, herbivory and nutrient enrichment both significantly affected primary-producer abundance. They also interacted to create greater nutrient enrichment effects in the absence of herbivores, suggesting that loss of herbivores produces more dramatic effects of nutrient loading. Herbivores consistently had stronger effects than did nutrient enrichment for both tropical macroalgae and seagrasses. The strong effects of herbivory but limited effects of nutrient enrichment on tropical macroalgae suggest that suppression of herbivore populations has played a larger role than eutrophication in driving the phase shift from coral- to macroalgal-dominated reefs in many areas, especially the Caribbean. For temperate macroalgae and benthic microalgae, the effects of top-down and bottom-up forces varied as a function of the inherent productivity of the ecosystem. For these algal groups, nutrient enrichment appeared to have stronger effects in high- vs. low-productivity systems, while herbivores exerted a stronger top-down effect in low-productivity systems. Effects of herbivores vs. nutrients also varied among algal functional groups (crustose algae, upright macroalgae, and filamentous algae), within a functional group between temperate and tropical systems, and according to the metric used to measure producer abundance. These analyses suggest that human alteration of food webs and nutrient availability have significant effects on primary producers but that the effects vary among latitudes and primary producers, and with the inherent productivity of ecosystems.     

Variation in secondary metabolite and aragonite concentrations in the tropical green seaweed Neomeris annulata: effects on herbivory by fishes

The tropical alga Neomeris annulata (Dickie) (Dasycladaceae: Chlorophyta) produces brominated sesquiterpenes and deposits aragonite throughout the thallus. This study, conducted throughout 1990–1991, showed that the fleshy, apical portions of the thalli (tips) were high in secondary metabolite concentrations (1.5%, mean combined secondary metabolites based on dry mass) and relatively low in calcium carbonate (aragonite form) (65.2% ash). The basal portions were lower in combined secondary metabolites (0.2% dry mass), and higher in aragonite (90.0% ash). The crude organic extract of the alga deterred fish feeding in the field at concentrations of 5, 10 and 15% dry mass, but not at a lower concentration of 1.5%. Natural concentrations of crude organic extract ranged from 1.5 to 15.3% in whole individuals and averaged 5.1% based on dry mass. Two brominated sesquiterpenes were isolated as major metabolites from the crude extract, but only one deterred feeding at natural concentrations. Ash concentrations in N. annulata were 60% dry mass in both the tips and bases. Aragonite strongly deterred feeding at concentrations of 65 and 90% dry mass. When a naturally occurring combination of organic extract and aragonite in the tips (10% crude extract and 65% aragonite) was compared with that of the bases (0.8% crude extract and 90% aragonite), no significant difference in grazing was observed. Combinations of secondary metabolites and aragonite were also tested against one or the other single defense. The combination of defenses proved a more effective deterrent than either secondary metabolites or aragonite alone.     

Evidence for regional heterogeneous atmospheric particulate matter distribution in China: implications for air pollution control

Environmental Chemistry Letters - China has suffered from severe nationwide air quality degradation for decades. PM2.5, the atmospheric particulate matter with an aerodynamic equivalent diameter of...     

The effects of direct and indirect constraints on biological communities

Human activities are expected to result in a diversity of directional or stochastic constraints that affect species either directly or by indirectly impacting their resources. However, there is no theoretical framework to predict the complex and various effects of these constraints on ecological communities. We developed a dynamic model that mimics the use of different resource types by a community of competing species. We investigated the effects of different environmental constraints (affecting either directly the growth rate of species or having indirect effects on their resources) on several biodiversity indicators. Our results indicate that (i) in realistic community models (assuming uneven resource requirements among species) the effects of perturbations are strongly buffered compared to neutral models; (ii) the species richness of communities can be maximized for intermediate levels of direct constraints (unimodal response), even in the absence of trade-off between competitive ability and tolerance to constraints; (iii) no such unimodal response occurs with indirect constraints; (iv) an increase in the environmental (e.g., climatic) variance may have different effects on community biomass and species richness.     

Modeling seed dispersal distances: implications for transgenic Pinus taeda.

Predicting forest-tree seed dispersal across a landscape is useful for estimating gene flow from genetically engineered (GE) or transgenic trees. The question of biocontainment has yet to be resolved, although field-trial permits for transgenic forest trees are on the rise. Most current field trials in the United States occur in the Southeast where Pinus taeda L., an indigenous species, is the major timber commodity. Seed dispersal distances were simulated using a model where the major determinants were: (1) forest canopy height at seed release, (2) terminal velocity of the seeds, (3) absolute seed release, and (4) turbulent-flow statistics, all of which were measured or determined within a P. taeda plantation established from seeds collected from wild forest-tree stands at the Duke Forest near Durham, North Carolina, USA. In plantations aged 16 and 25 years our model results showed that most of the seeds fell within local-neighborhood dispersal distances, with estimates ranging from 0.05 to 0.14 km from the source. A fraction of seeds was uplifted above the forest canopy and moved via the long-distance dispersal (LDD) process as far as 11.9-33.7 km. Out of 10(5) seeds produced per hectare per year, roughly 440 seeds were predicted to be uplifted by vertical eddies above the forest canopy and transported via LDD. Of these, 70 seeds/ha traveled distances in excess of 1 km from the source, a distance too great to serve as a biocontainment zone. The probability of LDD occurrence of transgenic conifer seeds at distances exceeding 1 km approached 100%.     

Understanding abundance patterns of a declining seabird: implications for monitoring.

The Kittlitz's Murrelet (Brachyramphus brevirostris) is a rare, non-colonial seabird often associated with tidewater glaciers and a recent candidate for listing under the Endangered Species Act. We estimated abundance of Kittlitz's Murrelets across space and time from at-sea surveys along the coast of Alaska (USA) and then used these data to develop spatial models to describe abundance patterns and identify environmental factors affecting abundance. Over a five-week period in the summer of 2005, we recorded 794 Kittlitz's Murrelets, 16 Marbled Murrelets (B. marmoratus), and 70 unidentified murrelets. The overall population estimate (N, mean +/- SE) during the peak period (3-9 July) was 1317 +/- 294 birds, decreasing to 68 +/- 37 by the last survey period (31 July-6 August). Density of Kittlitz's Murrelets was highest in pelagic waters of Taan Fjord (18.6 +/- 7.8 birds/km2, mean +/- SE) during 10-16 July. Spatial models identified consistent "hotspots" of Kittlitz's Murrelets, including several small areas where high densities of murrelets were found throughout the survey period. Of the explanatory variables that we evaluated, tidal current strength influenced murrelet abundance most consistently, with higher abundance associated with strong tidal currents. Simulations based on the empirically derived estimates of variation demonstrated that spatial variation strongly influenced power to detect trend, although power changed little across the threefold difference in the coefficient of variation on detection probability. We include recommendations for monitoring Kittlitz's Murrelets (or other marine species) when there is a high degree of uncertainty about factors affecting abundance, especially spatial variability.     

Weather effects on avian breeding performance and implications of climate change

The influence of recent climate change on the world's biota has manifested broadly, resulting in latitudinal range shifts, advancing dates of arrival of migrants and onset of breeding, and altered community relationships. Climate change elevates conservation concerns worldwide because it will likely exacerbate a broad range of identified threats to animal populations. In the past few decades, grassland birds have declined faster than other North American avifauna, largely due to habitat threats such as the intensification of agriculture. We examine the effects of local climatic variations on the breeding performance of a bird endemic to the shortgrass prairie, the Lark Bunting (Calamospiza melanocorys) and discuss the implications of our findings relative to future climate predictions. Clutch size, nest survival, and productivity all positively covaried with seasonal precipitation; yet relatively intense daily precipitation events temporarily depressed daily survival of nests. Nest survival was positively related to average temperatures during the breeding season. Declining summer precipitation may reduce the likelihood that Lark Buntings can maintain stable breeding populations in eastern Colorado although average temperature increases of up to 3 degrees C (within the range of this study) may ameliorate declines in survival expected with drier conditions. Historic climate variability in the Great Plains selects for a degree of vagility and opportunism rather than strong site fidelity and specific adaptation to local environments. These traits may lead to northerly shifts in distribution if climatic and habitat conditions become less favorable in the drying southern regions of the Great Plains. Distributional shifts in Lark Buntings could be constrained by future changes in land use, agricultural practices, or vegetative communities that result in further loss of shortgrass prairie habitats.     

Physical and biological disturbances interact differently with productivity: effects on floral and faunal richness

Physical and biological disturbances are ecological processes affecting patterns in biodiversity at a range of scales in a variety of terrestrial and aquatic systems. Theoretical and empirical evidence suggest that effects of disturbance on diversity differ qualitatively and quantitatively, depending on levels of productivity (e.g., the dynamic equilibrium model). In this study we contrasted the interactive effects between physical disturbance and productivity to those between biological disturbance and productivity. Furthermore, to evaluate how these effects varied among different components of marine hard-substratum assemblages, analyses were done separately on algal and invertebrate richness, as well as richness of the whole assemblage. Physical disturbance (wave action) was simulated at five distinct frequencies, while biological disturbance (grazing periwinkles) was manipulated as present or absent, and productivity was manipulated as high or ambient. Uni- and multivariate analyses both showed significant effects of physical disturbance and interactive effects between biological disturbance and productivity on the composition of assemblages and total species richness. Algal richness was significantly affected by productivity and biological disturbance, whereas invertebrate richness was affected by physical disturbance only. Thus, we show, for the first time, that biological disturbance and physical disturbance interact differently with productivity, because these two types of disturbances affect different components of assemblages. These patterns might be explained by differences in the distribution (i.e., press vs. pulse) and degree of selectivity between disturbances. Because different types of disturbance can affect different components of assemblages, general ecological models will benefit from using natural diverse communities, and studies concerned with particular subsets of assemblages may be misleading. In conclusion, this study shows that the outcome of experiments on effects of disturbance and productivity on diversity is greatly influenced by the composition of the assemblage under study, as well as on the type of disturbance that is used as an experimental treatment.     

Role of herbivory in controlling phytoplankton abundance: annual pigment budget for a temperate marine fjord

An annual pigment budget was constructed for Dabob Bay, Washington (USA) by comparing the downward vertical loss of phytoplankton pigments (chlorophyll and pheopigments) to the production of chlorophyll within the euphotic zone. The vertical flux of pigments was measured with sediment traps deployed at intervals of 1 to 6 wk over a 2.5 yr period yielding 763 d of trap exposure (28 November 1978–16 June 1981). The production rate of chlorophyll was calculated from measurements of algal specific growth rates, chlorophyll (chl) crops, primary production (as carbon) and appropriate C: chl ratios. Sixty one to 77% of the annual chlorophyll production was accounted for by the vertical flux of pheopigments resulting from herbivorous zooplankton grazing (macrozooplankton). Algal sinking, represented by downward chlorophyll flux, accounted for only 5 to 6% of the annual chlorophyll production. The remaining fraction of chlorophyll production was estimated to be consumed by small herbivores (microzooplankton), whose fecal material contributes to the suspended pool of pheopigments found in the euphotic zone. The suspended pheopigments are continuously removed by photodegradation. In Dabob Bay, the major process controlling phytoplankton abundance is zooplankton grazing and it appears that the ultimate fate of most phytoplankton is to be consumed by herbivores.     

Antibacterial effects of Anodonta cygnea fluids on Escherichia coli and enterococci multi-drug-resistant strains: environmental implications

The persistence of bacteria in Anodonta cygnea largely depends on the hemolymph bactericidal activity against aquatic micro-organisms.

The aim of this study was to assess the in vitro bactericidal activity of whole hemolymph against multi-drug-resistant Escherichia coli and enterococci strains. Four mussels were injected with multi-resistant strains of E. coli in order to determine their in vivo bactericidal reactivity. In vitro experiments showed that the hemocytes viability decreased almost 70% in 4 h, the same happened in the control hemolymph. Enterococci tested in in vitro experiments were more susceptible than E. coli to whole hemolymph and hemocytes fractions. None of the enterococci bacteria was detected in the hemolymph fractions from 24 to 72 h, while E. coli was still detectable 72 h after inoculation both in the control and respective plasma fractions. The microbial inhibition on both hemolymph and cell pellets suggest that hemocytes may be the main responsibles for that process.

In vivo experiments showed that hemolymph inhibited the E. coli load injected and it was correlated with a high tendency for the increase in hemocytes counts, after 72 h. High adherence of E. coli and an intense pseudopods manifestation, mainly to the granulocytes, suggest a clear phagocytosis process.     

The policy implications of non-convex environmental damages: A smog control case study

Nitrogen oxides and hydrocarbons react to form atmospheric ozone, but, for given concentrations of either one, increasing amounts of the other give rise to declining increments of ozone. Thus, the damage functions from hydrocarbon and nitrogen oxide emissions are non-convex. A case study of control strategies in the New York metropolitan regions show that, because control costs rise so steeply for both pollutants, a cost-effective approach is to control emissions of both, despite the non-convexities. However, the loss function, for deviations from the least-cost strategy, lies well within the range of scientific uncertainty regarding precursor-ozone relationships, and the least-cost strategy choice is highly sensitive to small changes in the relative control costs. Therefore, decisions regarding controls on these emissions, especially nitrogen oxide emissions, should be based also on other considerations, such as the reduction in acid deposition.     

Tissue type matters: selective herbivory on different life history stages of an isomorphic alga

Selective grazing by herbivores can have large effects on the population dynamics and community structure of primary producers. However, the ecological impacts of within-species herbivore preference for tissues of different phases (e.g., ploidy levels) or reproductive status remain relatively poorly known, especially among algae and other species with free-living haploid (gametophyte) and diploid (sporophyte) phases. We tested for herbivore selectivity among tissue types of the isomorphic (identical haploid and diploid free-living stages) red alga Mazzaella flaccida. Laboratory feeding assays demonstrated that the snail Tegula funebralis exhibited more than a threefold preference for gametophyte reproductive tissue over other tissue types, due to morphological differences. In contrast, the urchin Strongylocentrotus purpuratus did not distinguish as clearly between gametophytes and sporophytes; but it did prefer sporophyte reproductive to nonreproductive tissue, due to differences in water-soluble chemicals. Field surveys of grazer damage on M. flaccida blades were consistent with these laboratory preferences, with more damage found on gametophytes than sporophytes and reproductive than nonreproductive tissues. Differential fecundity can contribute to a skew in relative frequencies of phases in the field, and our results suggest that differential grazing by snails may contribute to this pattern and thus play a role in algal population biology.     

Temperature effects on accumulation and retention of radionuclides in the sea star,Asterias forbesi: implications for contaminated northern waters

Radioactive waste disposal and nuclear testing concentrated in high latitudes in the northern hemisphere have resulted in the accumulation of radionuclides in Arctic marine ecosystems, but little is known of the consequences for marine biota in these waters. Under controlled laboratory conditions in May through September 1994, we examined the bioaccumulation in sea stars, Asterias forbesi (Desor), or the radionuclides ²⁴¹Am, ⁵⁷Co and ¹³⁷Cs, all of which are important components of disposed radioactive wastes. Experiments at 2 and 12°C determined the relative importance of food (the bivalve, Macoma balthica) and water as sources of radionuclides and assessed the influence of temperature on radionuclide influx and efflux rates. The lower temperature greatly increased the retention of radionuclides ingested with food; for instance, the biological half-life (tb _1/2) of ²⁴¹Am in the sea stars was 31 d at 12°C, but was virtually infinite at 2°C. Retention of ingested ⁵⁷Co was also increased at 2°C (tb _1/2=41 d). ¹³⁷Cs was not accumulated from food. Low temperature significantly reduced net influx rates of ¹³⁷Cs from water, but did not affect net uptake of ²⁴¹Am or ⁵⁷Co. Temperature had little effect on the retention of all three isotopes obtained from the dissolved phase. These experiments suggest that extrapolation of results of previous radioecological studies, conducted at warmer temperatures, to polar or temperate winter environments may be problematic, and that nuclear waste isotopes obtained through trophic transfer may be retained far more efficiently in high latitude marine biota than by fauna from warmer ecosystems.     

Patterns of microzooplankton growth in dilution experiments across a trophic gradient: Implications for herbivory studies

To investigate the growth and grazing patterns of microzooplankton (MZP) in environments of differing productivity, dilution experiments measuring phytoplankton growth (μ) and grazing mortality (m) rates were performed using samples from contrasting locations along the Texas coast. Samples were collected from estuaries, coastal lagoons and offshore Gulf of Mexico locations in the spring and summer of 2001. MZP growth rates were determined in each dilution treatment. Although MZP biomass changed over time in most dilution treatments, adjusting μ and m for the actual grazer gradient (represented by geometric mean MZP biomass) did not cause a significant deviation from the nominal dilution gradient. Likewise, these adjustments did not yield significant regressions where none existed before adjustment. The dynamics of MZP taxonomic groups (ciliates, dinoflagellates) and size categories differed suggesting that in some cases internal predation may lead to trophic cascades. MZP biomass was higher in productive coastal waters and included a larger proportion of dinoflagellates than in the oligotrophic, ciliate-dominated waters of the Gulf of Mexico. The MZP biomass-to-chlorophyll a ratio was lowest in the hypereutrophic Nueces River, where MZP biomass significantly increased in all dilution treatments (net growth rates up to 2 day⁻¹) suggesting a strong top–down control. In the brown-tide dominated Upper Laguna Madre and the oligotrophic seagrass-dominated Lower Laguna Madre MZP growth was decoupled from that of phytoplankton. At these sites, MZP were likely fueled by bacterial carbon and mixotrophy, respectively. Observing the growth response of MZP in dilution experiments can provide insight into trophic structure and efficiency of the microbial food web.