Effects of food-level acclimation on digestive enzyme activities and feeding behavior of Calanus pacificus

A recent hypothesis in the zooplankton literature states that zooplankton acclimate to ambient food concentrations such that higher digestive enzyme activities and, consequently, higher maximum ingestion rates are achieved at higher food levels. To test this hypothesis, adult female Calanus pacificus, collected from the main basin of Puget Sound, Washington, USA, in August 1979 and May 1982, were conditioned for 2 wk at different concentrations of the diatom Thalassiosira weissflogii (=fluviatilis). Ingestion rates and the activity of the digestive enzymes laminarinase, maltase, and cellobiase were measured periodically during acclimation and in a block-designed feeding experiment at the end of acclimation. Consistent with the hypothesis, maximum ingestion rate and digestive enzyme activity were positively correlated. However, in contrast to the hypothesized mechanism, this result arose because both maximum ingestion rate and digestive enzyme activity were negatively correlated with food concentration during acclimation. The enhanced ingestion of copepods following long-term (12 to 14 d) acclimation to low food is similar to that previously described for short-term (e.g. 1 d) starvation. It might be energetically optimal for copepods experiencing a patchy food environment to maintain higher levels of digestive enzymes at low food concentrations in order to exploit high concentrations of food when encountered.     

Modeling high-frequency position data of large herbivores with a phase-state model

Understanding the rules and factors that drive the foraging behavior of large herbivores is important to describe their interaction with the landscape at various spatial scales. Some unresolved questions refer to landscape-behavioral interactions that result in oriented or random search in seasonally changing landscapes. Remotely sensed position data indicate that herbivores select local patches of heterogeneous landscapes depending on a complex host of dynamically varying animal and environmental conditions. Since foraging paths consist in successions of relatively short steps, increasing the frequency at which position information is acquired would contribute to entangle the mechanisms resulting in herbivores’ foraging paths. We addressed the question whether herbivores would obtain information at a patch scale that would modify their distribution at a landscape scale based on directed movement or navigation ability. We considered a set of 100,000 high-frequency (1 min intervals) position data of several free-ranging sheep (Ovis aries) at a seasonal-varying range (Patagonian Monte, Argentina) and observed their movements at landscape and at single vegetation patch scales. At a landscape scale, we inspected the spatial co-variation of seasonally varying forage offer and ewes’ movement speeds. At a patch scale, we developed a phase-state (P-S) model of movement cycles based on the occurrence of behavioral phases along foraging paths, and fitted it to the observed daily time series of ewes’ movement speeds. Ewes were preferentially distributed in areas with high forage offer during periods of low forage availability and the reverse occurred during the season of high forage availability. Parameters of the model of activity cycles amenable to control by ewes (duration of speed phases, time elapsed between speed cycles) did not covariate with forage offer, but varied significantly among ewes. The shape (kurtosis) parameter of the model of movement cycles, one which is unlikely under ewes’ control, co-varied significantly with spatial forage offer but did not differ among ewes. We conclude that ewes allocated foraging time along a series of similar movement efforts irrespective of forage availability at small patches. Average forage scarcity at multi-patch level increases the ratio of searching to feeding time. This results in apparent selective time allocation to richer forage areas but does not imply evidence for oriented movement at a landscape scale. We advance a behavioral-based definition of forage patches and discuss its implications in developing foraging theory and models. The P-S model applied to high-frequency position data of large herbivores substantially improves the interpretation of the factors controlling their time allocation in space with respect to previous models of herbivore spatial behavior by discriminating among behavioral-based and environmentally induced components of their movements.     

A simple plankton model for the Oregon upwelling ecosystem: Sensitivity and validation against time-series ocean data

Simple plankton models serve as useful platforms for testing our understanding of the mechanisms underlying ecosystem dynamics. A simple, one-dimensional plankton model was developed to describe the dynamics of nitrate, ammonium, two phytoplankton size-classes, meso-zooplankton, and detritus in the Oregon upwelling ecosystem. Computational simplicity was maintained by linking the biological model to a one-dimensional, cross-shelf physical model driven by the daily coastal upwelling index. The model sacrificed resolution of regional-scale and along-shore (north to south) processes and assumed that seasonal productivity is primarily driven by local cross-shelf Ekman transport of surface waters and upwelling of nutrient-rich water from depth.Our goals were to see how well a simple plankton model could capture the general temporal and spatial dynamics of the system, test system sensitivity to alternate parameter set values, and observe system response to the effective scale of potential retention mechanisms. Model performance across the central Oregon shelf was evaluated against two years (2000-2001) of chlorophyll and copepod time-series observations. While the modeled meso-zooplankton biomass was close in scale to the observed copepod biomass, phytoplankton was overestimated relative to that inferred from the observed surface chlorophyll concentration. Inshore, the system was most sensitive to the nutrient uptake kinetics of diatom-size phytoplankton and to the functional grazing response of meso-zooplankton. Meso-zooplankton was more sensitive to alternate parameter values than was phytoplankton. Reduction of meso-zooplankton cross-shelf advection rates (crudely representing behavioral retention mechanisms) reduced the scale of model error relative to the observed seasonal mean inshore copepod biomass but had little effect of the modeled meso-zooplankton biomass offshore nor upon phytoplankton biomass across the entire shelf.     

A mathematical model of PCB bioaccumulation in plankton

In batch experiments exposing individual plankton constituents to Aroclor 1254 PCB, the rate at which the organism approaches partitioning equilibrium appears to be partly size-dependent while the extent of PCB accumulation is species-specific. The sorptive desorptive kinetics of PCB in these experiments can be described mathematically by a first-order expression. Employing this expression in a model plankton food web permits examination of the role of feeding and sorptive processes in determining PCB body burden under various environmental conditions. When ingestion rates exceed desorption and excretion rates, a consuming organism accumulates PCB above levels predicted by equilibrium partitioning relationships. Feeding-induced oscillations in PCB body burden could thus obscure the reduction of soluble PCB concentration which determine a “baseline” PCB body burden. Unless referenced to a specific set of biological and environmental conditions, the importance of direct partitioning from water vs. food uptake appears to be a moot topic.     

Simple predator–prey interactions control dynamics in a plankton food web model

A plankton food web model is analysed using interaction parameter values appropriate to the upper mixed layer of the high latitude oceans. The dynamics of this four-variable system are analysed in terms of the dynamics of much simpler two-variable predator–prey subsystems. Thus, the food web's robust, periodic, four-dimensional dynamics are explained by means of two-dimensional spirals and limit cycles. These dynamical subsystems are coupled by means of an omnivore that transfers control of the dynamics between the two predator–prey subsystems. The food web may substantially decouple the predator–prey subsystems so that the oscillating phytoplankton/zooplankton blooms exhibit population collapses when bacterial ‘breathers’ briefly dominate after growing dramatically from low background levels. This regular bloom/breather behaviour becomes benignly chaotic when the system is mildly forced by the annual cycle of the sun's irradiance.     

Study of the nutrient and plankton dynamics in Lake Tanganyika using a reduced-gravity model

An eco-hydrodynamic (ECOH) model is proposed for Lake Tanganyika to study the plankton productivity. The hydrodynamic sub-model solves the non-linear, reduced-gravity equations in which wind is the dominant forcing. The ecological sub-model for the epilimnion comprises nutrients, primary production, phytoplankton biomass and zooplankton biomass. In the absence of significant terrestrial input of nutrients, the nutrient loss is compensated for by seasonal, wind-driven, turbulent entrainment of nutrient-rich hypolimnion water into the epilimnion, which gives rise to high plankton productivity twice in the year, during the transition between two seasons. Model simulations predict well the seasonal contrasts of the measured physical and ecological parameters. Numerical tests indicate that the half saturation constant for grazing by zooplankton and the fish predation rate on zooplankton affect the zooplankton biomass measurably more than that of phytoplankton biomass. This work has implications for the application of this model to predict the climatological biological productivity of Lake Tanganyika.     

Investigations into a plankton population model: Mortality and its importance in climate change scenarios

The potential for marine plankton ecosystems to influence climate by the production of dimethylsulphide (DMS) has been an important topic of recent research into climate change. Several General Circulation Models, used to predict climate change, have or are being modified to include interactions of ecosystems with climate. Climate change necessitates that parameters within ecosystem models must change during long-term simulations, especially mortality parameters that increase as organisms are pushed toward the boundaries of their thermal tolerance. Changing mortality parameters can have profound influences on ecosystem model dynamics. There is therefore a pressing need to understand the influence of varying mortality parameters on the long-term behaviour of ecosystem models. This work examines the sensitivity of a model of DMS production by marine ecosystems to variations in three linear mortality coefficients. Significant differences in behaviour are observed, and we note the importance of these results in formulating ecosystem models for application in simulations of climate change.     

Interactions of marine plankton with transuranic elements

In a series of laboratory experiments, the biokinetics of ²⁴¹Am, an important transuranium element, was studied in Meganyctiphanes norvegica, a euphausiid common in the northwestern Mediterranean. The euphausiids accumulated Am from water by passive adsorption onto exoskeletons, achieving wet weight concentration factors on the order of 10² after 1 wk exposure; concentration factors varied inversely with the size of the euphausiids and linearly with their surface area:wet weight ratios. Essentially all (96±10%) of the Am taken up from water was associated with the exoskeleton, so that negligible Am was retained by the euphausiids after molting. The retention half-time of Am in molts was 2.9 d. Euphausiids could also concentrate Am from feeding suspensions by ingesting Am-labelled diatom cells, although there was negligible Am assimilation (3±2% after 4 d feeding); after passage through the gut, virtually all (99%) of the ingested Am was defecated within 1 wk. The retention half-time of Am in fecal pellets was 41 and 51 d at 13° and 5°C, respectively. In oceanic waters, where the preponderance of ²⁴¹Am is in the dissolved phase, uptake of Am from water by euphausiids would be the dominant route of bioaccumulation. The results underscore the importance of sinking biogenic debris from zooplankters in mediating the vertical transport of Am in the sea. Given their retention half-times for ²⁴¹Am and their rapid sinking rates, fecal pellets and discarded molts have the potential to deliver most of their Am to the sediments.     

Adding complex trophic interactions to a size-spectral plankton model: Emergent diversity patterns and limits on predictability

A new model in the NPZ (nutrient-phytoplankton-zooplankton) style is presented, mechanistically simple but with 40 size classes each of phytoplankton (1-20 μm) and small zooplankton (2.1-460 μm), in order to resolve one level of trophic interactions in detail. General, empirical allometric relationships are used to parameterize both the optimal prey size and size selectivity for each grazer class, as is rarely done. This inclusion of complex predator-prey linkages and realistic prey preferences yields a system with an emergent pattern of phytoplankton diversity consistent with global ocean observations, i.e., a parabolic relationship between diversity (as measured by the Shannon evenness) and biomass. It also yields significant long-term time evolution, which places limits on the extent to which the community response to nutrient forcing can be predicted from forcing in a pragmatic sense. When a simple annual cycle in nutrient supply is repeated exactly for many years, transient fluctuations up to a factor of two in spring bloom magnitude persist for 10-20 years before a stable seasonal biomass cycle is achieved. When the amplitude of the nutrient-supply annual cycle is given a random interannual modulation, these long-lived transients add significant noise to a 100-year correlation between annual-mean nutrient supply and annual-mean biomass. This noise is 20% of total interannual variance in the model base case, and ranges from 0% to 40% depending on the grazer size selectivity. In general, unpredictability on the bloom timescale is damped when food-web complexity is increased by making grazers less selective, while unpredictability on the interannual scale shows the opposite pattern, increasing with increasing food-web complexity up to a high threshhold, past which community structure and biomass time evolution both suddenly simplify. These results suggests a new strategy for ensemble ecosystem forecasting and uncertainty estimation, analogous to methods common in circulation and climate modeling, in which internal variability (predator-prey interactions in the biological case; eddies and climate-system oscillations in the physical case) are resolved and quantified, rather than suppressed.     

Fitness related diet-mixing by intraspecific host-plant-switching of specialist insect herbivores

Generalist insect herbivores may profit by feeding on a mixture of plant species that differ in nutritional quality. Herbivore performance can also be affected by intraspecific host plant variation. However, it is unknown whether conspecific plant individuals differ sufficiently to promote diet-mixing behavior in specialist herbivores. We experimentally tested this "specialist diet-mixing hypothesis" for specialist caterpillars (Chrysopsyche imparilis, Lasiocampidae) in a West African savanna. The caterpillars switched regularly between host tree individuals (Combretum fragrans, Combretaceae). To examine whether switching benefited caterpillar performance via diet-mixing, the caterpillars were reared either on leaves from several plant individuals (mixed diet) or on leaves from a single plant. The strongest effect of diet-mixing was found for fecundity, with females reared on a mixed diet laying significantly more eggs than sisters receiving a single-plant diet. In addition, a mixed diet decreased variability in egg size and increased the growth of second-instar caterpillars. Supplementary food choice experiments were conducted to assess a potential influence of lowered host quality (induced by herbivory) on caterpillar behavior; no such effect was found. By linking intraspecific host-switching behavior and herbivore performance, this study provides new information on the relevance of intraspecific plant variation for herbivorous insects.     

A complex social structure with fission–fusion properties can emerge from a simple foraging model

Precisely how ecological factors influence animal social structure is far from clear. We explore this question using an agent-based model inspired by the fission–fusion society of spider monkeys (Ateles spp). Our model introduces a realistic, complex foraging environment composed of many resource patches with size varying as an inverse power law frequency distribution with exponent β. Foragers do not interact among them and start from random initial locations. They have either a complete or a partial knowledge of the environment and maximize the ratio between the size of the next visited patch and the distance traveled to it, ignoring previously visited patches. At intermediate values of β, when large patches are neither too scarce nor too abundant, foragers form groups (coincide at the same patch) with a similar size frequency distribution as the spider monkey’s subgroups. Fission–fusion events create a network of associations that contains weak bonds among foragers that meet only rarely and strong bonds among those that repeat associations more frequently than would be expected by chance. The latter form subnetworks with the highest number of bonds and a high clustering coefficient at intermediate values of β. The weak bonds enable the whole social network to percolate. Some of our results are similar to those found in long-term field studies of spider monkeys and other fission–fusion species. We conclude that hypotheses about the ecological causes of fission–fusion and the origin of complex social structures should consider the heterogeneity and complexity of the environment in which social animals live.     

Growth of plankton algae as a function of N-concentration,measured by means of batch technique

Employing a special batch technique which involves extremely low concentrations of the alga Selenastrum capricornutum, it is shown that a growth rate of ca. 75% of optimum growth occurs at a concentration of 0.02 g-at N(NH₄ ⁺) l^-1. The low concentrations of this N-source in the oligotrophic parts of the oceans (ca. 0.06 g-at l^-1) thus give rise to a borderline deficiency. As the half-saturation constant (K _S) for growth was recorded at 0.05 g-at N(NO₃ ^-), this ion can hardly be of much importance at the low concentrations present in these parts of the oceans. It is wrong to assert that N is the primary limiting nutrient. Phosphate also is found at concentrations giving rise to a borderline deficiency.     

A simple population theory for mutualism by the use of lattice gas model

The population dynamics of species interactions provides valuable information for life sciences. Lotka-Volterra equations (LVEs) are known to be the most popular model, and they are mainly applied to the systems of predation and competition. However, LVEs often fail to catch the population dynamics of mutualism; the population sizes of species increase infinitely under certain condition (divergence problem). Furthermore, LVEs never predicts the Allee effect in the systems of obligate mutualism. Instead of LVEs, several models have been presented for mutualism; unfortunately, they are rather complicated. It is, therefore, necessary to introduce a simpler theory for mutualism. In the present paper, we apply the lattice gas model which corresponds to the mean-field theory of the usual lattice model. The derived equations are cubic and contain only essential features for mutualism. In the case of obligate mutualism, the dynamics exhibits the Allee effect, and it is almost the same as in the male-female systems. In our model, the population sizes never increase infinitely, because our model contains not only intra- but also interspecific competitions. If the density of one species increases disproportionately in respect of its mutual partners, then this might imply downward pressure on the population abundance of the mutual partner species and such feedback would eventually act as a controlling influence on the population abundance of either species. We discuss several assumptions in our model; in particular, if both species can occupy in each cell simultaneously, then the interspecific competition disappears.     

Turbulence and phytoplankton diversity: A general model of the “paradox of plankton”

The principle of “competitive exclusion” predicts that no two species can occupy the same ecological niche at the same time and place (Hardin, 1960). Hutchinson (1953, 1961) suggested that the vast diversity of phytoplankton observed in many aquatic environments presents an apparent contradiction to this principle. Since all phytoplankton compete for the same basic resources, and since the euphotic zones of most natural waters are relatively homogeneous, such coexisting plankters appear to be simultaneously occupying the same niche. In this paper we present simulation results from a mathematical model wherein we examine the hypothesis that physical turbulence in an aquatic system can mollify interactive pressures between plankton populations and allow coexistence of species competing for the same resources. Using Bella's (1972) highly simplified model as a point of departure, we develop a new model, explicitly incorporating gross physiological mechanisms, to investigate the effects of both advective and turbulent components of water movement on the growth of three competing phytoplankton species. We observed that, in the absence of water motion, no two species were able to coexist, while under the hypothetical conditions of advection without turbulence (laminar flow), just two species were able to occur contemporaneously. Coexistence of all three species was achieved only with the addition of a random turbulent component to the model's hydrodynamic function. Moreover, this general coexistence was observed only when the major turbulent frequency approached the turnover rate of phytoplankton populations. We suggest that there is a limited region of periodicities and magnitudes for hydrodynamic energy in which phytoplankton can coexist, and that most natural aquatic environments fall within this region. We further speculate that, in general, the coupling of physical and biological processes in nature may be influenced by the relative frequency characteristics of those processes.     

Implications of a simple competition model for the stability of an intercropping system

It is a common view that intercropping systems of agricultural crops produce more stable yields than do systems in which the same crops are grown in monoculture. This paper discusses a modelling approach which has been used to support the notion that whether or not intercropping is more stable than monoculture depends on the mode of interaction among crops, i.e. whether two different crops suppress or enhance each other. It is shown here that this notion is not supported by the model used. We conclude that the relative merits of the two cropping systems depend on the proportion of land allocated to each crop rather than on the mode of interaction. The model suggests that if the optimum allocation of land is considered, both systems will be equally stable.     

Managing plant population spread: prediction and analysis using a simple model

Models can be used to direct the management of population spread for the control of invasives or to encourage species of conservation value. Analytical models are attractive because of their theoretical basis and limited data requirements, but there is concern that their simplicity may limit their practical utility. We address the applied use of simple models in a study of a declining annual herb, Rhinanthus minor. We parameterized a population-spread model using field data on demography and dispersal for four management systems: grazed only (GR), hay-cut once (H1), hay-cut twice (H2), and hay-cut with autumn grazing (HG). Within a replicated experiment we measured spread rates of introduced R. minor populations over eight years. The modeled and measured spread rates were very similar in terms of both patterns of management effects and absolute values, so that in both cases HG > H2, H1 > GR. The treatments affected both dispersal and demography (establishment and survival) and so we used decomposition approaches to analyze the major causes of differences in population spread. Increased dispersal under hay-cutting was more important than demographic changes and accounted for approximately 70% of the differences in spread rate between the hay-cut and grazed-only treatments. Furthermore, management effects on the tail of the dispersal curve were by far the most critical in governing spread. This study suggests that simple models can be used to inform practical conservation management, and we demonstrate straightforward uses of our model to predict the impacts of different management strategies. While simple models can give accurate projections, we emphasize that they must be parameterized with high-quality data gathered at the appropriate spatial scale.     

A simple spatially explicit ideal-free distribution: a model and an experiment

This investigation presents a simple spatially explicit analysis of the ideal-free distribution. The traditional ideal-free distribution assumes discrete sites with definite boundaries, and predicts how many individuals should occupy each site. In contrast, the present analysis assumes that a forager’s gains gradually decline with distance from a site, and asks where in space individuals ought to be. Although many interesting situations may arise, the analysis asks how individuals should position themselves as the distance between two identical sources increases. Nash equilibrium positions should follow a pitchfork pattern as the distance between sites is increased; that is, an individual should maintain a position between two sources when they are close together but should move nearer one of the sources when they are far apart. In addition, the text describes an experimental study that parallels the theoretical analysis. The experiment supports the predicted pitchfork pattern, and provides somewhat weaker support for the predicted differences in ”individual” and ”paired” pitchforks. Received: 14 June 2000 / Revised: 20 September 2000 / Accepted: 7 October 2000     

Negative effects of vertebrate herbivores on invertebrates in a coastal dune community

Although competition has been a major focus in ecology for the past century, most empirical and theoretical studies in this area have emphasized interactions between closely related species. However, there is growing evidence that negative interactions among distantly related taxa also occur and may be far more important than previously thought. In this study, we took advantage of an 11-year-old replicated vertebrate-exclosure experiment in a coastal dune community in northern California, USA, to examine the effects of the two most common vertebrate herbivores (jackrabbits and black-tailed deer) on the abundance of the three most visible invertebrate herbivores (two snail, a moth, and a grasshopper species). Our results indicate that four of the six possible pairwise interactions were significantly negative for the invertebrates. Jackrabbits reduced the abundances of snails by 44-75%, tiger moth caterpillars by 36%, and grasshoppers by 62%. Deer reduced the abundances of snails by 32%, increased the abundances of caterpillars by 31%, and had no measurable effect on grasshopper abundance. Our data also revealed that jackrabbits significantly decreased the volume of forbs and common shrubs and the flowering by grasses in our study plots. We were unable to detect an effect of deer on these measures of vegetation. These results suggest that by changing vegetation, jackrabbits may reduce invertebrate populations that are limited by food, protective structures, or microclimate provided by plants. Of these three mechanisms, only shade was strongly supported as limiting snail numbers in smaller-scale manipulations. In most systems, as in this one, the number of pairs of distantly related herbivores far exceeds the number of pairs of congeners. Since interactions among distantly related herbivores may be common in many cases, these interactions are likely to be important and should receive far more attention from ecologists.