Heterogeneity shapes invasion: host size and environment influence susceptibility to a nonnative pathogen.

Theoretical study of invasion dynamics has suggested that spatial heterogeneity should strongly influence the rate and extent of spreading organisms. However, empirical support for this prediction is scant, and the importance of understanding heterogeneity for real-world systems has remained ambiguous. This study quantified the influence of host and environmental heterogeneity on the dynamics of a 19-year disease invasion by the exotic and fatal pathogen, Phytophthora lateralis, within a stream population of its host tree, Port Orford cedar (Chamaecyparis lawsoniana). Using dendrochronology, we reconstructed the invasion history along a 1350-m length of infected stream, which serves as the only route of pathogen dispersal. Contrary to theoretical predictions, the temporal progression of the disease invasion was not related to a host's downstream spatial position, but instead was determined by two sources of heterogeneity: host size and proximity to the stream channel. These sources of heterogeneity influenced both the epidemic and endemic dynamics of this pathogen invasion. This analysis provides empirical support for the influence of heterogeneity on the invasion dynamics of a commercially important forest pathogen and highlights the need to incorporate such natural variability into both invasion theory and methods aimed at controlling future spread.     

Quantity and quality: unifying food web and ecosystem perspectives on the role of resource subsidies in freshwaters

Although the study of resource subsidies has emerged as a key topic in both ecosystem and food web ecology, the dialogue over their role has been limited by separate approaches that emphasize either subsidy quantity or quality. Considering quantity and quality together may provide a simple, but previously unexplored, framework for identifying the mechanisms that govern the importance of subsidies for recipient food webs and ecosystems. Using a literature review of > 90 studies of open-water metabolism in lakes and streams, we show that high-flux, low-quality subsidies can drive freshwater ecosystem dynamics. Because most of these ecosystems are net heterotrophic, allochthonous inputs must subsidize respiration. Second, using a literature review of subsidy quality and use, we demonstrate that animals select for high-quality food resources in proportions greater than would be predicted based on food quantity, and regardless of allochthonous or autochthonous origin. This finding suggests that low-flux, high-quality subsidies may be selected for by animals, and in turn may disproportionately affect food web and ecosystem processes (e.g., animal production, trophic energy or organic matter flow, trophic cascades). We then synthesize and review approaches that evaluate the role of subsidies and explicitly merge ecosystem and food web perspectives by placing food web measurements in the context of ecosystem budgets, by comparing trophic and ecosystem production and fluxes, and by constructing flow food webs. These tools can and should be used to address future questions about subsidies, such as the relative importance of subsidies to different trophic levels and how subsidies may maintain or disrupt ecosystem stability and food web interactions.     

Responses of a bacterial pathogen to phosphorus limitation of its aquatic invertebrate host

Host nutrition is thought to affect the establishment, persistence, and severity of pathogenic infections. Nutrient-deficient foods possibly benefit pathogens by constraining host immune function or benefit hosts by limiting parasite growth and reproduction. However, the effects of poor elemental food quality on a host's susceptibility to infection and disease have received little study. Here we show that the bacterial microparasite Pasteuria ramosa is affected by the elemental nutrition of its aquatic invertebrate host, Daphnia magna. We found that high food carbon:phosphorus (C:P) ratios significantly reduced infection rates of Pasteuria in Daphnia and led to lower within-host pathogen multiplication. In addition, greater virulent effects of bacterial infection on host reproduction were found in Daphnia-consuming P-deficient food. Poor Daphnia elemental nutrition thus reduced the growth and reproduction of its bacterial parasite, Pasteuria. The effects of poor host nutrition on the pathogen were further evidenced by Pasteuria's greater inhibition of reproduction in P-limited Daphnia. Our results provide strong evidence that elemental food quality can significantly influence the incidence and intensity of infectious disease in invertebrate hosts.     

Monopolization in a resource queue: water striders competing for food and mates

The monopolization of resources plays an important theoretical role in the literature on competition for food and mates. We used 12 groups of male water striders (Aquariusremigis) to: (1) test the general prediction that monopolization of both food and mates decreases as the temporal clumping of resources increases, (2) compare the efficiency of two indices of resource monopolization, coefficient of variation and Q (Ruzzante et al. 1996), and (3) quantitatively assess the resource queue model of Blanckenhorn and Caraco (1992). Each group of six males competed for both food items and mates released from the upstream end of a laboratory stream. The mean inter-arrival time for resource units (food or females) was 10 min, with four levels of temporal clumping (variance in inter-arrival time: 0, 25, 50 or 320 min²). As predicted, the monopolization of both food and mates decreased as the temporal clumping of resource arrival increased, although monopolization was greater for food than for mates. Q detected the difference in monopolization of food and mates, whereas the coefficient of variation did not, because Q is independent of mean resource abundance. The resource queue model successfully predicted monopolization of both resource types, explaining 89% and 76% of variation in the proportion of food and mates acquired by the six males. The success of the model suggests that the scaling of handling time to the variance in resource inter-arrival time should play an important role in any general theory of resource monopolization. Received: 28 February 1997 / Accepted after revision: 26 September 1997     

Mother really knows best: host choice of adult phytophagous insect females reflects a within-host variation in suitability as larval food

Non-random distribution patterns of specialized phytophagous insects on their hosts may depend on intraspecific differences in plant tissue quality, including nutrients and secondary compounds. Secondary compounds are involved in plant resistance, but are also important for the recognition and acceptability of plants as resources by specialized insects. If individuals within a plant species vary in their content of such secondary substances, there may also be qualitative differences between them. In such cases, natural selection will favor insects with the ability to distinguish and prefer the more suitable plants. In Sweden, the leaf beetle Gonioctena linnaeana Schrank (Coleoptera, Chrysomelidae) is highly specialized on one host, the native willow Salix triandra L (Salicaceae). Field observations reveal that some host plants in a population harbor many feeding larvae, causing severe defoliation, whereas neighboring plants may have few or no feeding larvae. Our hypothesis is that the distribution pattern of G. linnaeana larvae in this population results from qualitative differences between individual host plants in combination with the ability of G. linnaeana females to distinguish between plants that are suitable and not suitable for offspring performance. We examine whether larval survival differs depending on diet and whether the content of secondary chemical compounds explains female preference. Based on the higher survival rate of larvae reared on leaves from preferred hosts, we conclude that G. linnaeana females have evolved a behavior that maximizes offspring performance and thus positively affects female fitness. A chemical survey of the plants indicates that luteolin-7-glucoside and an unidentified flavonoid are important for separating the preferred from the non-preferred plants.     

History of titanium dioxide regulation as a food additive: a review

Environmental Chemistry Letters - Since its discovery in the late eighteenth century and mass production in the early twentieth century, titanium dioxide has been used in a wide range of...     

Nutrient enrichment reduces constraints on material flows in a detritus-based food web

Most aquatic and terrestrial ecosystems are experiencing increased nutrient availability, which is affecting their structure and function. By altering community composition and productivity of consumers, enrichment can indirectly cause changes in the pathways and magnitude of material flows in food webs. These changes, in turn, have major consequences for material storage and cycling in the ecosystem. Understanding mechanisms and predicting consequences of nutrient-induced changes in material flows requires a quantitative food web approach that combines information on consumer energetics and consumer-resource stoichiometry. We examined effects of a whole-system experimental nutrient enrichment on the trophic basis of production and the magnitude and pathways of carbon (C), nitrogen (N), and phosphorus (P) flows in a detritus-based stream food web. We compared the response of the treated stream to an adjacent reference stream throughout the study. Dietary composition and elemental flows varied considerably among invertebrate functional feeding groups. During nutrient enrichment, increased flows of leaf litter and amorphous detritus to shredders and gatherers accounted for most of the altered flows of C from basal resources to consumers. Nutrient enrichment had little effect on patterns of material flows but had large positive effects on the magnitude of C, N, and P flows to consumers (mean increase of 97% for all elements). Nutrient-specific food webs revealed similar flows of N and P to multiple functional groups despite an order of magnitude difference among groups in consumption of C. Secondary production was more strongly related to consumption of nutrients than C, and increased material flows were positively related to the degree of consumer-resource C:P and C:N imbalances. Nutrient enrichment resulted in an increased proportion of detrital C inputs consumed by primary consumers (from -15% to 35%) and a decreased proportion of invertebrate prey consumed by predators (from -80% to 55%). Our results demonstrate that nutrient enrichment of detritus-based systems may reduce stoichiometric constraints on material flows, increase the contribution of consumers to C, N, and P cycling, alter the proportion of C inputs metabolized by consumers, and potentially lead to reduced ecosystem-level storage of C.     

The fear of being eaten reduces energy transfer in a simple food chain

Food chain length is an important property of ecosystems, but the mechanisms maintaining it remain elusive. Classical views suggest that energetic inefficiencies (the "energy-flow hypothesis") limit food chain length, but others have argued that better explanations reside in more complex scenarios that consider the stability of food webs or the combined effects of productivity and ecosystem size. We argue that abandonment of the energy-flow hypothesis is premature. For a simple tritrophic rocky intertidal food chain, we show that the efficiency of energy transfer is strongly influenced by predation risk and consumer density. Effects tied to predation risk were particularly strong, explaining 32% of the variation in growth efficiency (compared to 15% for density effects) and reducing it by 44-76%. Hence, the stress (fear of being eaten) that predators impose on prey may be instrumental in limiting energy transfer up the food chain and thus contribute to the shortening of food chains.     

The changing status of water as a natural resource

This paper considers the way freshwater resources are perceived in an era of climate change. It is suggested that in many parts of the world water is moving from being a renewable resource (or continuous natural resource) to a potentially renewable resource (PRR). In some areas water is moving from being a PRR to a nonrenewable resource. Selected water problems from three continents are investigated in the context of this theoretical construct. Recent experience in the United Kingdom is then also investigated as a case study of these changes before brief conclusions are drawn.     

Kelp as a trophic resource for marine suspension feeders: a review of isotope-based evidence

Kelp forests are enormously productive, and they and adjacent habitats support large populations of suspension feeders. What do these suspension feeders eat? Intuitively, we might expect that kelp primary production is a key form of trophic support for these animals. Indeed, a large and growing number of studies using carbon stable isotope data, typically collected over short time periods, have asserted that detritus from kelps is an important, and in some cases the main, food source for coastal benthic suspension feeders. This view has been incorporated into several textbooks and review papers covering kelp forest ecosystems, and loss of trophic support for benthic suspension feeders is now often invoked as an ecosystem consequence of top-down or other impacts on kelp forests. More direct evidence, however, suggests that these animals mainly eat phytoplankton and, in some cases, bacteria or zooplankton. Because isotope values of pure coastal phytoplankton, uncontaminated with detritus, are difficult to obtain, present studies have largely relied on single measurements from offshore environments or from the literature, which typically reflects offshore values. We review the evidence showing that phytoplankton isotope values can, and are expected to, vary widely in coastal waters and that inshore phytoplankton may often be enriched in ¹³C compared to offshore phytoplankton. This unaccounted-for variation may have systematically biased the results of such trophic studies toward finding large contributions of kelp detritus to suspension-feeder diets. We review some key stable isotope studies and put forth evidence for alternative explanations of the isotope patterns presented. Finally, we make recommendations for future isotope studies and describe several approaches for progress in this area. New techniques, particularly flow cytometry and compound-specific stable isotope analysis, provide ways to shed light on this interesting and important ecological issue.     

Larval growth and settlement of the European oyster (Ostrea edulis) as a function of food quality measured as fatty acid composition

Larval growth rate and settlement of the European flat oyster Ostrea edulis were experimentally studied as a function of the composition of dietary fatty acids. Diets differing in fatty acid composition were composed by mixtures of the microalgae Isochrysis galbana, Pavlova lutheri and Chaetoceros calcitrans. Fatty acid content in the tissue of the feeding larvae, analyzed by gas chromatography and mass spectrometry, reflected the composition in the diet. Larval growth rate was significantly correlated to the three omega-3 polyunsaturated fatty acids (PUFA) C18:3, C18:4 and C22:6, with minor differences for neutral and polar lipids. No relation between growth rate and the omega-3 PUFA C20:5 was detected, a PUFA often implied as essential for bivalves. It is suggested that naturally occurring variability in fatty acid composition may constrain larval growth. In settlement experiments in both still water and flume flow little substrate selectivity was found for some contrasting substrates. It is concluded that differences in dietary fatty acids may explain as much of settlement success as the variability of substrates. Received: 12 October 1998 / Accepted: 6 April 1999     

Cascading effects from predator removal depend on resource availability in a benthic food web

We tested joint effects of predator loss and increased resource availability on the grazers’ trophic level and the propagation of trophic interactions in a benthic food web by excluding larger predatory fish from cages and manipulating nutrients in the coastal zone of the Baltic Sea. The combination of nutrient enrichment and excluding larger predators induced an increase in medium-sized predatory fish (three-spined stickleback). The meso-predator fish in turn did not change the total abundance of the invertebrate herbivores, but did cause a substantial shift in their community composition towards the dominance of gastropods by reducing amphipods by 40–60%, while gastropods were left unchanged. The shift in grazer composition generated a 23 times higher producer biomass, but only under nutrient enrichment. Our results show that top-predator declines can substantially shift the species composition at the grazers’ level, but that cascading effects on producers by a trophic cascade strongly depend on resource availability.     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

Benthic harpacticoids as a food source for fish

The trophic relationships between meiobenthic harpacticoids and juvenile predatory fish were investigated in a shallow lagoon on Bermuda. Harpacticoids represent an essential food source for the juveniles of several demersal fish species. Juveniles up to 3 cm length of one fish species feed exclusively on harpacticoids. Meiobenthic harpacticoids are therefore an important component of food chains leading to higher trophic levels. However, the feeding pressure exerted by the fish on the harpacticoid populations is negligible.     

On detritus as a food source for pelagic filter-feeders

Data on the amount of organic detritus within the particle size fraction 1 to 150 are presented for the Western Kiel Bight. Grouped into a mixed surface layer and a stratified lower layer in accordance with the main hydrographic features of the Western Baltic Sea, the rounded-off values show a seasonal variation between 100 and 600 mg m^-3 expressed as dry weight of organic matter. The overall average for both layers is about 200 mg m^-3. Organic detritus thus comprises more than 40% of total organic matter in the above size class, which is the class most easily accessible to the relatively small filter-feeders in this area. Attempting to trace the origin of organic detritus, a positive correlation to phytoplankton standing stock was found in some cases, suggesting the predominance of autochthonous detritus. Proceeding from the assumption that pelagic filter-feeders select their food mainly by size and not by taste, it is concluded that organic detritus plays an important role as a supplementary food source, being ingested together with phytoplankton and small nonmotile heterotrophs. The nutritive value of detritus is increased by the adsorption of dissolved organic matter and above all through the subsequent colonization by bacteria, which utilize the dissolved substances. Detritus particles serving as a substratum for bacteria thus form a means whereby dissolved organic substances reenter the food chain. The ingestion of detritus by filter-feeders is, therefore, thought to be instrumental in increasing the effectivity of energy transfer from the primary to the secondary food-chain level.     

Food selection by copepods: discrimination on the basis of food quality

The copepod Acartia tonsa displayed nearly two-fold higher ingestion rates on faster-growing cells of the diatom Thalassiosira weissflogii compared to ingestion rates on slower-growing cells of that species at the same cell concentration. Ingestion rates on slow-growing cells were also enhanced by the addition of cell-free aliquots of algal exudate to the experimental feeding chambers. In addition, the faster-growing algal cells were selectively ingested by the copepod when the two cell types were mixed together in different proportions, indicating that physiological differences between growing cells are a critical factor in the food detection/selection process of zooplankton. Consideration of cell carbon, nitrogen, and protein composition suggests that the copepods are maximizing nitrogenous ingestion (total protein and/or nitrogen). Selectivity for cells with higher protein content results in a higher daily protein ration, even if the selection process results in a decreased rate of ingestion in mixtures of cell types.     

Meta-analysis: trophic level, habitat, and productivity shape the food web effects of resource subsidies

Studies of the effects of cross-habitat resource subsidies have been a feature of food web ecology over the past decade. To date, most studies have focused on demonstrating the magnitude of a subsidy or documenting its effect in the recipient habitat. Ecologists have yet to develop a satisfactory framework for predicting the magnitude of these effects. We used 115 data sets from 32 studies to compare consumer responses to resource subsidies across recipient habitat type, trophic level, and functional group. Changes in consumer density or biomass in response to subsidies were inconsistent across habitats, trophic, and functional groups. Responses in stream cobble bar and coastline habitats were larger than in other habitats. Contrary to expectation, the magnitude of consumer response was not affected by recipient habitat productivity or the ratio of productivity between donor and recipient habitats. However, consumer response was significantly related to the ratio of subsidy resources to equivalent resources in the recipient habitat. Broad contrasts in productivity are modified by subsidy type, vector, and the physical and biotic characteristics of both donor and recipient habitats. For this reason, the ratio of subsidy to equivalent resources is a more useful tool for predicting the possible effect of a subsidy than coarser contrasts of in situ productivity. The commonness of subsidy effects suggests that many ecosystems need to be studied as open systems.     

Adaptive introgression as a resource for management and genetic conservation in a changing climate

Current rates of climate change require organisms to respond through migration, phenotypic plasticity, or genetic changes via adaptation. We focused on questions regarding species’ and populations’ ability to respond to climate change through adaptation. Specifically, the role adaptive introgression, movement of genetic material from the genome of 1 species into the genome of another through repeated interbreeding, may play in increasing species’ ability to respond to a changing climate. Such interspecific gene flow may mediate extinction risk or consequences of limited adaptive potential that result from standing genetic variation and mutation alone, enabling a quicker demographic recovery in response to changing environments. Despite the near dismissal of the potential benefits of hybridization by conservation practitioners, we examined a number of case studies across different taxa that suggest gene flow between sympatric or parapatric sister species or within species that exhibit strong ecotypic differentiation may represent an underutilized management option to conserve evolutionary potential in a changing environment. This will be particularly true where advanced‐generation hybrids exhibit adaptive traits outside the parental phenotypic range, a phenomenon known as transgressive segregation. The ideas presented in this essay are meant to provoke discussion regarding how we maintain evolutionary potential, the conservation value of natural hybrid zones, and consideration of their important role in adaptation to climate.