Why allometric scaling enhances stability in food web models

It has recently been shown that the incorporation of allometric scaling into the dynamic equations of food web models enhances network stability if predators are assigned a higher body mass than their prey. We investigate the underlying mechanisms leading to this stability increase. The dynamic equations can be written such that allometric scaling influences these equations at three places: the time scales of predator and prey dynamics become separated, the energy outflow to the predators is decreased, and intraspecific competition is increased relative to metabolic rates. For five food web topologies and various network sizes (i.e., species richness), we study the effect of each of these modifications on the percentage of surviving species separately and find that the decreased interaction strengths and the increased intraspecific competition are responsible for the enhanced stability. We also investigate the range of parameter values for which an enhanced stability is observed.     

Replicability of physicochemistry and macroinvertebrate assemblages in stream mesocosms: implications for experimental research

1. The replicability of mesocosms is assessed in a case study of artificial streams (4 m²) flanking a lowland chalk river in southern U.K. Among‐mesocosm comparisons of physicochemistry and macroinvertebrate assemblage composition (in drift and benthos) were made during a 2‐year survey of 12 outdoor once‐through linear channels. 2. Mesocosm physicochemistry was highly replicable, with statistically significant congruence in temporal variation across the mesocosm series, low spatial variation and no significant blocking or positional effects arising from mesocosm layout. Parallel physicochemistry was attributed to the outdoor stream‐side location and short water residence times. 3. Mesocosms were colonized by 127 macroinvertebrate taxa from 15 taxonomic orders. Both benthic and drifting assemblages were comparable among mesocosms, with no significant effect of mesocosm layout. Small differences in taxonomic composition were, however, evident among particular mesocosms, with higher (e.g. Tinodes, Limnius and Elmis) or lower (e.g. Pisidium and Valvata) abundances observed for a minority (5%) of taxa. We conclude that large (4 m²) outdoor flow – through mesocosms can be replicable when located near to the source system and allowed to establish naturally.     

Mycorrhizal respiration: implications for global scaling relationships

Most plant species form mycorrhizas, yet these are neglected by plant physiologists. One consequence of this neglect is reduced ability to predict plant respiration, because respiration rate (R) in mycorrhizal roots might be higher than in non-mycorrhizal roots owing to increased substrate availability associated with enhanced nutrient uptake, coupled with increased respiratory product demand. Other predictions include that mycorrhizal colonization will affect scaling of R with tissue nitrogen concentrations; that mycorrhizal and non-mycorrhizal root R differ in their response to nutrient supply; and that the impact of colonization on R is related to fungal biomass. Failure to examine properly the role of colonization in determining root R means that current interpretations of root and soil respiration data might be flawed.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Allometric scaling law in a simple oxygen exchanging network: possible implications on the biological allometric scaling laws

A simple model of an oxygen exchanging network is presented and studied. This network's task is to transfer a given oxygen rate from a source to an oxygen consuming system. It consists of a pipeline, that interconnects the oxygen consuming system and the reservoir and of a fluid, the active oxygen transporting element, moving through the pipeline. The network optimal design (total pipeline surface) and dynamics (volumetric flow of the oxygen transporting fluid), which minimize the energy rate expended in moving the fluid, are calculated in terms of the oxygen exchange rate, the pipeline length, and the pipeline cross-section. After the oxygen exchanging network is optimized, the energy converting system is shown to satisfy a 3/4-like allometric scaling law, based upon the assumption that its performance regime is scale invariant as well as on some feasible geometric scaling assumptions. Finally, the possible implications of this result on the allometric scaling properties observed elsewhere in living beings are discussed.     

Food web complexity and stability across habitat connectivity gradients

The effects of habitat connectivity on food webs have been studied both empirically and theoretically, yet the question of whether empirical results support theoretical predictions for any food web metric other than species richness has received little attention. Our synthesis brings together theory and empirical evidence for how habitat connectivity affects both food web stability and complexity. Food web stability is often predicted to be greatest at intermediate levels of connectivity, representing a compromise between the stabilizing effects of dispersal via rescue effects and prey switching, and the destabilizing effects of dispersal via regional synchronization of population dynamics. Empirical studies of food web stability generally support both this pattern and underlying mechanisms. Food chain length has been predicted to have both increasing and unimodal relationships with connectivity as a result of predators being constrained by the patch occupancy of their prey. Although both patterns have been documented empirically, the underlying mechanisms may differ from those predicted by models. In terms of other measures of food web complexity, habitat connectivity has been empirically found to generally increase link density but either reduce or have no effect on connectance, whereas a unimodal relationship is expected. In general, there is growing concordance between empirical patterns and theoretical predictions for some effects of habitat connectivity on food webs, but many predictions remain to be tested over a full connectivity gradient, and empirical metrics of complexity are rarely modeled. Closing these gaps will allow a deeper understanding of how natural and anthropogenic changes in connectivity can affect real food webs.     

Biomechanics and allometric scaling in primate locomotion and morphology

Body size has a dominant influence on locomotor performance and the morphology of the locomotor apparatus. In locomotion under the influence of gravity, body mass acts as weight force and is a mechanical variable. Accordingly, the application of biomechanical principles and methods allows a functional understanding of scaling effects in locomotion. This is demonstrated here using leaping primates as an example. With increasing body size, the decreasing ratio of muscle force available for acceleration during takeoff to the body mass that has to be accelerated dictates both the movement pattern and the proportions of the hindlimbs. In an arm-swinging movement, the long, heavy arms of the large-bodied leapers are effectively used to gain additional momentum. A new perspective on decreasing size identifies the absolutely small acceleration distance and time available for propulsion as factors limiting leaping distance and extensively determining locomotor behavior and body proportions. As the mechanical constraints differ according to body size for a given mode of locomotion, a typological approach to morphology in relation to locomotor category is ruled out. Across locomotor categories, dynamic similarity (sensu Alexander) can be expected if the propulsive mechanisms as well as the selective pressures acting upon locomotion are the same.     

Ecological relationships between stream communities and spatial scale: implications for designing catchment-level monitoring programmes

1. Stream communities are structured by factors acting over multiple spatial and temporal scales. Identifying what factors are driving spatial patterns in stream communities is a central aim of ecology.
2. Here we used two large European data sets of fish, invertebrates, macrophytes, benthic diatoms and environmental data in two stream groups (lowland and mountain) to determine the importance of variables at different spatial scales (geographical, regional, local) on community structure.
3. Both geographical position and ecoregion were selected first in canonical correspondence analysis (CCA), clearly showing the broad spatial gradients covered in the data set. Secondary predictors (after accounting for spatial and/or ecoregion effects) were similar between stream groups and among the four organism groups. In particular, conductivity and N concentration were strong predictors reflecting catchment land use.
4. Using partial CCA, we assessed the individual importance of the three spatial scales on the community structure of the four organism groups in the two stream groups. The majority of among-site variability (22–29%) was accounted for by local scale variables (e.g. water chemistry and substratum type), with regional and spatial variables accounting 11–13% and 5–6%, respectively. Our findings indicate that the four organism groups are responding similarly to the different levels of spatial scale, implying much redundancy which should be consider when implementing studies of bioassessment.     

Trophic relationships in a tropical stream food web assessed by stable isotope analysis

1. Stable isotope analysis, coupled with dietary data from the literature, was used to investigate trophic patterns of freshwater fauna in a tropical stream food web (Guadeloupe, French West Indies).
2. Primary producers (biofilm, algae and plant detritus of terrestrial origin) showed distinct δ¹³C signatures, which allowed for a powerful discrimination of carbon sources. Both autochthonous (¹³C-enriched signatures) and allochthonous (¹³C-depleted signatures) resources enter the food web. The migrating behaviour of fishes and shrimps between marine and freshwater during their life cycles can be followed by carbon isotopes. Here, shrimp δ¹³C signatures were shown to shift from −16‰ (for juveniles under marine influence) to −24.7‰ (for adults in freshwater habitats). For resident species, δ¹³C values partly reflected the species' habitat preferences along the river continuum : species living in river mouths were ¹³C-enriched in comparison with those collected upstream.
3. Nitrogen isotopic ratios were also discriminating and defined three main trophic guilds among consumers. The δ¹⁵N values of herbivores/detritivores were 5.0–8.4‰, omnivores 8.8–10.2‰ and carnivores 11–12.7‰.
4. Mixing model equations were employed to calculate the possible range of contribution made by respective food sources to the diet of each species. The results revealed the importance of omnivorous species and the dependence of riverine biota on terrestrial subsidies, such as leaf detritus and fruits. Finally, the abundance of shrimps and their feeding habits placed in relief their key role in tropical freshwater food webs. Isotopic analysis provides a useful tool for assessing animal feeding patterns.     

Disturbance, biological legacies and community development in stream mesocosms

Disturbances reduce the biota in stream ecosystems, and leave biological legacies, including remnant species, which potentially influence post-disturbance community development but are poorly understood. We investigated whether three remnant species, the snail Radix peregra, the mayfly Serratella ignita and the freshwater shrimp Gammarus pulex, affected community development in mesocosms that mimicked disturbed habitat patches in streams. Following 21 days of colonisation, we found that the occurrence of legacy effects depended on the identity of the remnant species. Radix had the strongest effect. By bulldozing epilithon, the snails acted as ecological engineers that promoted settlement of filter feeders (Simuliidae) and invertebrate predators (especially Pentaneura and Aphelocheirus) and strongly deterred settlement of non-predatory chironomids (e.g. Heterotrissocladius and Microtendipes). Gammarus increased in density (by 665%) where remnant, probably through rapid reproduction. Baetis and Pentaneura were scarce, and Asellus absent, in remnant Gammarus treatments, as a consequence of interference and/or predation by the amphipods. In contrast, Serratella tolerated the colonisation of immigrant species and did not affect the structure of the developing benthic community. Despite the observed effects on the presence and abundance of benthos, remnant fauna had no significant effect on assemblage taxon richness, or that of any specific trophic group. The contrasting effects of remnant species on immigrant colonisation echoed differences in their life-history traits and foraging behaviours. Our results indicate that biota can generate spatial patchiness of epilithon and benthic invertebrates in stream ecosystems.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

Resting metabolism of helodermatid lizards: allometric and ecological relationships

We measured metabolic rates at 15 and 25°C in 42 helodermatid lizards ranging in mass from 26 to 1616 g. No consistent repeatable daily rhythms of metabolism were detected. There were no significant differences in metabolic rates between the two species of Heloderma. The temperature coefficient for metabolism (Q ₁₀) was 3.0 between 15 and 25°C. The mass exponent for helodermatids (0.69) differed significantly from the among-species mass exponent of 0.80 for all squamates combined. However, adult Heloderma had a mass exponent of 0.80. Rates of metabolism of adult helodermatids were lower than those of other squamate reptiles, and at 15°C periods of apnea contributed to a further reduction in metabolic rate. Our finding that helodermatids have low SMRs supports the hypothesis that ecology is important in influencing metabolic rate, and that “reclusive” squamates have lower rates of metabolism than do nonreclusive species.     

Algal-periphyton population and community changes from zinc stress in stream mesocosms

Three treatments of zinc (0.05, 0.5, 1.0 mg Zn l⁻¹) and a control could be identified by different algal communities in outdoor, flow-through, stream mesocosms. Established communities were continuously exposed to Zn, and samples were collected on days 0, 2, 5, 10, 20 and 30 after treatment began. Experiments were conducted in spring, summer, and fall 1984. Control stream mesocosms could be identified by diatoms in all seasons. The 0.05 mg Zn l⁻¹ treatment could be identified by certain diatom taxa being more abundant than in the control in all seasons and by a filamentous green alga in summer and fall. The 0.5 mg Zn l⁻¹ treatment could be identified by a filamentous green alga in fall. The 1.0 mg Zn l⁻¹ treatment was dominated by unicellular green algae in all seasons and by a filamentous blue-green alga in summer. A similarity index (SIMI) indicated that Zn-stressed samples generally became less similar to control samples as Zn concentration increased from 0.05 to 1.0 mg Zn l⁻¹. Total biovolume-density of all taxa responded slower than individual taxa in spring and failed to distinguish between Zn treatments in summer and fall. Zinc bound to periphyton was much better than total Zn in water for identifying Zn treatments. Zinc treatments as low as 0.05 mg Zn l⁻¹ changed algal species composition despite 0.047 mg Zn l⁻¹ being the Criterion of the US Environmental Protection Agency for the 24-h average of total recoverable Zn.     

A statistical model for the genetic origin of allometric scaling laws in biology

Many biological processes, from cellular metabolism to population dynamics, are characterized by particular allometric scaling (power-law) relationships between size and rate. Although such allometric relationships may be under genetic determination, their precise genetic mechanisms have not been clearly understood due to a lack of a statistical analytical method. In this paper, we present a basic statistical framework for mapping quantitative genes (or quantitative trait loci, QTL) responsible for universal quarter-power scaling laws of organic structure and function with the entire body size. Our model framework allows the testing of whether a single QTL affects the allometric relationship of two traits or whether more than one linked QTL is segregating. Like traditional multi-trait mapping, this new model can increase the power to detect the underlying QTL and the precision of its localization on the genome. Beyond the traditional method, this model is integrated with pervasive scaling laws to take advantage of the mechanistic relationships of biological structures and processes. Simulation studies indicate that the estimation precision of the QTL position and effect can be improved when the scaling relationship of the two traits is considered. The application of our model in a real example from forest trees leads to successful detection of a QTL governing the allometric relationship of third-year stem height with third-year stem biomass. The model proposed here has implications for genetic, evolutionary, biomedicinal and breeding research.     

Resource translocation within seagrass clones: allometric scaling to plant size and productivity

The allometric scaling of resource demand and translocation within seagrass clones to plant size (i.e. shoot mass and rhizome diameter), shoot production and leaf turnover was examined in situ in eight seagrass species (Cymodocea nodosa, Cymodocea serrulata, Halophila stipulacea, Halodule uninervis, Posidonia oceanica, Thalassodendron ciliatum, Thalassia hemprichii and Zostera noltii), encompassing most of the size range present in seagrass flora. One fully developed shoot on each experimental rhizome was incubated for 2–3 h with a pulse of NaH¹³CO₃ (235 μmol) and ¹⁵NH₄Cl (40 μmol). The mobilisation of incorporated tracers across the clone was examined 4 days later. Carbon and nitrogen demand for shoot production across seagrass species scaled at half of the shoot mass, whereas seagrass leaves incorporated tracers (¹³C and ¹⁵N) at rates proportional to the shoot mass. The shoots of all seagrass species shared resources with neighbours, particularly with younger ones. The time scales of physiological integration and the absolute amount of resources shared by seagrass ramets scaled at 2.5 power of the rhizome diameter. Hence, the ramets of larger species were physiologically connected for longer time scales and share larger absolute amounts of resources with neighbours than those of smaller species. The different pattern of resource translocation exhibited by seagrasses helps explain the ecological role displayed by these species and the success of large seagrasses colonising nutrient-poor coastal areas, where they often dominate.     

Species-specific allometric scaling under self-thinning: evidence from long-term plots in forest stands

Experimental plots covering a 120 years' observation period in unthinned, even-aged pure stands of common beech (Fagus sylvatica), Norway spruce (Picea abies), Scots pine (Pinus sylvestris), and common oak (Quercus Petraea) are used to scrutinize Reineke's (1933) empirically derived stand density rule [see text], N=tree number per unit area, [see text]=mean stem diameter), Yoda's (1963) self-thinning law based on Euclidian geometry ([see text] [see text]=mean biomass per tree), and basic assumptions of West, Brown and Enquist's (1997, 1999) fractal scaling rules ([see text] [see text] w=biomass per tree, d=stem diameter). RMA and OLS regression provides observed allometric exponents, which are tested against the exponents, expected by the considered rules. Hope for a consistent scaling law fades away, as observed exponents significantly correspond with the considered rules only in a minority of cases: (1) exponent r of [see text] varies around Reineke's constant -1.605, but is significantly different from r=-2, supposed by Euclidian or fractal scaling, (2) Exponent c of the self-thinning line [see text] roams roughly about the Euclidian scaling constant -3/2, (3) Exponent a of [see text] tends to follow fractal scaling 8/3. The unique dataset's evaluation displays that (4) scaling exponents and their oscillation are species-specific, (5) Euclidian scaling of one relation and fractal scaling of another are coupled, depending on species. Ecological implications of the results in respect to self-tolerance (common oak>Norway spruce>Scots pine>common beech) and efficiency of space occupation (common beech>Scots pine>Norway spruce>common oak) are stressed and severe consequences for assessing, regulating and scheduling stand density are discussed.     

Stem and wood allometric relationships in Cacteae (Cactaceae)

Allometric relationships in organisms are considered a universal phenomenon. A positive scaling has been reported between stem size and cellular size of tracheary elements in wood of different vascular plants, but few studies have been carried out in slow-growing succulent plants. The aim of this study was to evaluate if a relationship exists between size, growth form and wood cell size among individual species of Cacteae. Forty-four species belonging to 16 genera of the tribe Cacteae with differing growth forms and sizes were studied. When analyzing plant size, we found a positive allometric scaling and the larger-sized species showing a higher percentage of succulent tissue and less accumulation of wood tissue. The positive scaling found between plant size (height and diameter) and vessel elements and fiber length support the universality of the allometric relationship proposed for other vascular plants with non-succulent stems. Notably, wide-band tracheids do not scale with plant size or growth form. Succulence associated with narrow vessel elements with distinctive helical secondary walls and wide-band tracheids suggest they are the key adaptations to tolerate drought and provide support to the stems of most taxa in Cacteae. Fibers do not have the primary role of giving mechanical support; therefore, we assume the scarce fibers in clusters represent reaction wood that, along with the fundamental tissue, maintains the vertical position and shape of those species growing in rocky cracks. Our results with species having short succulent stems support the universal theory of positive allometric scaling of vascular plants.     

Soil acidity, ecological stoichiometry and allometric scaling in grassland food webs

The factors regulating the structure of food webs are a central focus of community and ecosystem ecology, as trophic interactions among species have important impacts on nutrient storage and cycling in many ecosystems. For soil invertebrates in grassland ecosystems in the Netherlands, the site-specific slopes of the faunal biomass to organism body mass relationships reflected basic biochemical and biogeochemical processes associated with soil acidity and soil C : N : P stoichiometry. That is, the higher the phosphorus availability in the soil, the higher, on average, the slope of the faunal biomass size spectrum (i.e., the higher the biomass of large-bodied invertebrates relative to the biomass of small invertebrates). While other factors may also be involved, these results are consistent with the growth rate hypothesis from biological stoichiometry that relates phosphorus demands to ribosomal RNA and protein production. Thus our data represent the first time that ecosystem phosphorus availability has been associated with allometry in soil food webs (supporting information available online). Our results have broad implications, as soil invertebrates of different size have different effects on soil processes.     

Pitfalls and challenges of estimating population growth rate from empirical data: consequences for allometric scaling relations

The intrinsic rate of increase is a fundamental concept in population ecology, and a variety of problems require that estimates of population growth rate be obtained from empirical data. However, depending on the extent and type of data available (e.g. time series, life tables, life history traits), several alternative empirical estimators of population growth rate are possible. Because these estimators make different assumptions about the nature of age‐dependent mortality and density‐dependence of population dynamics, among other factors, these quantities capture fundamentally different aspects of population growth and are not interchangeable. Nevertheless, they have been routinely commingled in recent ecoinformatic analyses relating to allometry and conservation biology. Here we clarify some of the confusion regarding the empirical estimation of population growth rate and present separate analyses of the frequency distributions and allometric scaling of three alternative, non‐interchangeable measures of population growth. Studies of allometric scaling of population growth rate with body size are additionally sensitive to the statistical line fitting approach used, and we find that different approaches yield different allometric scaling slopes. Across the mix of population growth estimators and line fitting techniques, we find scattered and limited support for the key allometric prediction from the metabolic theory of ecology, namely that log₁₀(population growth rate) should scale as ?0.25 power of log₁₀(body mass). More importantly, we conclude that the question of allometric scaling of population growth rate with body size is highly sensitive to previously unexamined assumptions regarding both the appropriate population growth parameter to be compared and the line fitting approach used to examine the data. Finally, we suggest that the ultimate test of allometric scaling of maximum population growth rates with body size has not been done and, moreover, may require data that are not currently available.     

Nitrogen in insects: implications for trophic complexity and species diversification

Disparities in nutrient content (nitrogen and phosphorus) between herbivores and their plant resources have lately proven to have major consequences for herbivore success, consumer-driven nutrient cycling, and the fate of primary production in ecosystems. Here we extend these findings by examining patterns of nutrient content between animals at higher trophic levels, specifically between insect herbivores and predators. Using a recently compiled database on insect nutrient content, we found that predators exhibit on average 15% greater nitrogen content than herbivores. This difference persists after accounting for variation from phylogeny and allometry. Among herbivorous insects, we also found evidence that recently derived lineages (e.g., herbivorous Diptera and Lepidoptera) have, on a relative basis, 15%-25% less body nitrogen than more ancient herbivore lineages (e.g., herbivorous Orthoptera and Hemiptera). We elaborate several testable hypotheses for the origin of differences in nitrogen content between trophic levels and among phylogenetic lineages. For example, interspecific variation in insect nitrogen content may be directly traceable to differences in dietary nitrogen (including dilution by gut contents), selected for directly in response to the differential scarcity of dietary nitrogen, or an indirect consequence of adaptation to different feeding habits. From some functional perspectives, the magnitude rather than the source of the interspecific differences in nitrogen content may be most critical. We conclude by discussing the implications of the observed patterns for both the trophic complexity of food webs and the evolutionary radiation of herbivorous insects.     

Theoretical and empirical derivation of cardiovascular allometric relationships in children.

Basic fluid dynamic principles were used to derive a theoretical model of optimum cardiovascular allometry, the relationship between somatic and cardiovascular growth. The validity of the predicted models was then tested against the size of 22 cardiovascular structures measured echocardiographically in 496 normal children aged 1 day to 20 yr, including valves, pulmonary arteries, aorta and aortic branches, pulmonary veins, and left ventricular volume. Body surface area (BSA) was found to be a more important determinant of the size of each of the cardiovascular structures than age, height, or weight alone. The observed vascular and valvar dimensions were in agreement with values predicted from the theoretical models. Vascular and valve diameters related linearly to the square root of BSA, whereas valve and vascular areas related to BSA. The relationship between left ventricular volume and body size fit a complex model predicted by the nonlinear decrease of heart rate with growth. Overall, the relationship between cardiac output and body size is the fundamental driving factor in cardiovascular allometry.