Resource Edibility and the Effects of Predators and Productivity on the Outcome of Trophic Interactions

Population regulation of organisms is frequently viewed as being either by predation ("top-down" regulation) or by resource productivity ("bottom-up" regulation). However, these two types of regulation should ultimately balance, and the abundances of interacting organisms that result in any given environment should depend on the densities that can satisfy the balancing conditions. Here I use a simple model to examine interactions among consumers and two types of resources that differ in edibility. I use the model to predict the abundances of all three populations, which should result in environments that differ in consumer mortality (e.g., predation intensity) and resource productivity (e.g., nutrient levels). The model predicts that the effects of predation and productivity will depend strongly on the diet breadth of the consumers. I then test the predictions of the model in a field experiment on the plankton of an oligotrophic lake by manipulating fish predation and nutrient productivity. Finally, I compare these results with those of 46 other similar experiments and show the utility of the model in understanding the results of the experiments and the observed correlations of abundances of phytoplankton and zooplankton in natural lakes. The results of the experiments confirm that the effects of predators and nutrients on consumers (zooplankton) and resources (phytoplankton) predicted by the model differ when the consumer assemblage is dominated by a generalist herbivore (Daphnia) from those expected in the absence of such a generalist herbivore.

[1]  C. Duarte,et al.  Algal cell size and the maximum density and biomass of phytoplankton1 , 1987 .

[2]  Robert H. Peters,et al.  Empirical Prediction of Crustacean Zooplankton Biomass and Profundal Macrobenthos Biomass in Lakes , 1984 .

[3]  John T. Lehman,et al.  The effect of changes in the nutrient income on the condition of Lake Washington1 , 1981 .

[4]  Dmitriĭ Olegovich Logofet,et al.  Stability of Biological Communities , 1983 .

[5]  G. Mittelbach,et al.  Trophic Relations and Ontogenetic Niche Shifts in Aquatic Ecosystems , 1988 .

[6]  S. Fretwell,et al.  The Regulation of Plant Communities by the Food Chains Exploiting Them , 2015 .

[7]  Z. Gliwicz Effect of zooplankton grazing on photosynthetic activity and composition of phytoplankton: With 4 figures and 1 table in the text , 1975 .

[8]  R. C. Hart,et al.  Zooplankton feeding on size fractionated Microcystis colonies and Chlorella in a hypertrophic lake (Hartbeespoort Dam, South Africa): implications to resource utilization and zooplankton succession , 1987 .

[9]  C. Walters,et al.  Equilibrium Models for Seasonal Dynamics of Plankton Biomass in Four Oligotrophy Lakes , 1987 .

[10]  J. P. Nilssen,et al.  Pelagic predators and interfering algae: stabilizing factors in temperate eutrophic lakes , 1987 .

[11]  THE FERTILIZATION OF GREAT CENTRAL LAKE II. ZOOPLANKTON STANDING STOCK , 1972 .

[12]  W. Kerfoot,et al.  Competition Among Cladocerans: Nature of the Interaction Between Bosmina and Daphnia , 1982 .

[13]  Experimental studies on factors limiting colonization by Daphnia pulex Leydig of coastal montane lakes in British Columbia , 1978 .

[14]  J. Popovský,et al.  Influence of the fishstock on the phosphorus — chlorophyll ratio: With 2 figures and 2 tables in the text , 1978 .

[15]  John R. Post,et al.  Trophic Relationships in Freshwater Pelagic Ecosystems , 1986 .

[16]  D. J. Hall,et al.  AN EXPERIMENTAL APPROACH TO THE PRODUCTION DYNAMICS AND STRUCTURE OF FRESHWATER ANIMAL COMMUNITIES1 , 1970 .

[17]  T. Parsons,et al.  THE FERTILIZATION OF GREAT CENTRAL LAKE. I. EFFECT OF PRIMARY PRODUCTION , 1972 .

[18]  A. Jensen,et al.  Algal Competition for Phosphorus: The Influence of Zooplankton and Fish , 1986 .

[19]  M. Rosenzweig,et al.  Stability of enriched aquatic ecosystems. , 1972, Science.

[20]  Lennart Persson,et al.  Predator Regulation and Primary Production Along the Productivity Gradient of Temperate Lake Ecosystems , 1988 .

[21]  Edward L. Mills,et al.  Evaluation of Fish Communities Through Assessment of Zooplankton Populations and Measures of Lake Productivity , 1982 .

[22]  T. Osborn Algal cell size and the maximum density and biomass of phytoplankton , 1987 .

[23]  R. Levins,et al.  The dynamics of aquatic systems. 2. The effects of nutrient enrichment on model plankton communities1,2 , 1977 .

[24]  D. Tilman Resource competition and community structure. , 1983, Monographs in population biology.

[25]  Andrew Sih,et al.  Predation: direct and indirect impacts on aquatic communities , 1988 .

[26]  K. Hambright,et al.  Experimental study of the impacts of bluegill (Lepomis macrochirus) and largemouth bass (Micropterus salmoides) on pond community structure , 1986 .

[27]  M. Rosenzweig Paradox of Enrichment: Destabilization of Exploitation Ecosystems in Ecological Time , 1971, Science.

[28]  Edward McCauley,et al.  Empirical Relationships Between Phytoplankton and Zooplankton Biomass in Lakes , 1981 .

[29]  M. Vanni Effects of Food Availability and Fish Predation on a Zooplankton Community , 1987 .

[30]  K. Hyatt,et al.  Responses of Sockeye Salmon (Oncorhynchus nerka) to Fertilization of British Columbia Coastal Lakes , 1985 .

[31]  D. O. Hessen,et al.  Food Size Spectra and Species Replacement within Herbivorous Zooplankton , 1986 .

[32]  D. Canfield,et al.  Relationships between Zooplankton Abundance and Chlorophyll a Concentrations in Florida Lakes , 1984 .

[33]  William W. Murdoch,et al.  "Community Structure, Population Control, and Competition"-A Critique , 1966, The American Naturalist.

[34]  Stabilization of a Predator-Prey Equilibrium by the Addition of a Second "Keystone" Victim , 1980, The American Naturalist.

[35]  M. Vanni Freshwater Zooplankton Community Structure: Introduction of Large Invertebrate Predators and Large Herbivores to a Small Species Community , 1988 .

[36]  W. Lampert Inhibitory and Toxic Effects of Blue‐green Algae on Daphnia , 1981 .

[37]  C. H. Waddington Towards a theoretical biology : an IUBS symposium , 1968 .

[38]  F. Briand,et al.  Zooplankton grazing and phytoplankton species richness: Field tests of the predation hypothesis1 , 1979 .

[39]  H. Bottrell A review of some problems in zooplankton production studies , 1976 .

[40]  M. Lynch Predation, competition, and zooplankton community structure: An experimental study1,2 , 1979 .

[41]  S. Rognerud,et al.  Relationships between phytoplankton and zooplankton biomass in large lakes: With 4 figures in the text , 1984 .

[42]  L. Oksanen,et al.  Exploitation Ecosystems in Gradients of Primary Productivity , 1981, The American Naturalist.

[43]  W. Murdoch,et al.  Cyclic and Stable Populations: Plankton as Paradigm , 1987, The American Naturalist.

[44]  J. Schindler Food Quality and Zooplankton Nutrition , 1971 .

[45]  S. Heaney,et al.  Some observations on the use of the in vivo fluorescence technique to determine chlorophyll‐a in natural populations and cultures of freshwater phytoplankton , 1978 .

[46]  F. Briand,et al.  Cybernetic mechanisms in lake plankton systems: how to control undersirable algae , 1978, Nature.

[47]  B. Moss,et al.  The Role of Predation in Causing Major Changes in the Limnology of a Hyper‐Eutrophic Lake , 1980 .

[48]  Paul R. Ehrlich,et al.  The "Balance of Nature" and "Population Control" , 1967, The American Naturalist.

[49]  R. Sterner Herbivores' Direct and Indirect Effects on Algal Populations , 1986, Science.

[50]  Ray W. Drenner,et al.  Experimental Analysis of the Direct and indirect Effects of an Omnivorous Filter-Feeding Clupeid on Plankton Community Structure , 1986 .

[51]  W. T. Edmondson,et al.  Daphnia in Lake Washington1 , 1982 .

[52]  S. Threlkeld Estimating cladoceran birth rates: The importance of egg mortality and the egg age distribution1 , 1979 .

[53]  S. Carpenter,et al.  Plankton Community Structure and Limnetic Primary Production , 1984, The American Naturalist.

[54]  L. Slobodkin,et al.  Community Structure, Population Control, and Competition , 1960, The American Naturalist.

[55]  R. Dorazio,et al.  Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan , 1987 .

[56]  B. C. Patten,et al.  The Plankton Community. , 1964, Science.

[57]  Stephen R. Carpenter,et al.  Complex Interactions in Lake Communities , 2011, Springer New York.

[58]  A. Infante,et al.  Differences between two species of Daphnia in the use of 10 species of algae in Lake Washington , 1985 .

[59]  D. J. Hall,et al.  An Experimental Test of the Effects of Predation Risk on Habitat Use in Fish , 1983 .

[60]  G. Mittelbach Foraging Efficiency and Body Size: A Study of Optimal Diet and Habitat Use by Bluegills , 1981 .

[61]  C. Burns THE RELATIONSHIP BETWEEN BODY SIZE OF FILTER‐FEEDING CLADOCERA AND THE MAXIMUM SIZE OF PARTICLE INGESTED , 1968 .

[62]  E. Ranta,et al.  A field manipulation of trophic interactions in rock-pool plankton , 1987 .

[63]  R. Vance,et al.  Predation and Resource Partitioning in One Predator -- Two Prey Model Communities , 1978, The American Naturalist.

[64]  D. O. Hessen,et al.  From phytoplankton to detritus and bacteria: effects of short-term nutrient and fish perturbations in a eutrophic lake , 1986 .

[65]  A. Mazumder,et al.  Effects of fertilization and planktivorous fish (yellow perch) predation on size distribution of particulate phosphorus and assimilated phosphate: Large enclosure experiments1 , 1988 .

[66]  M. Vanni Competition in zooplankton communities: Suppression of small species by Daphnia pulex1 , 1986 .

[67]  J. Post,et al.  The impact of planktivorous flsh on the structure of a plankton community , 1987 .

[68]  R. Drenner,et al.  Particle-grazing and plankton community impact of an omnivorous cichlid , 1984 .

[69]  W. Kerfoot,et al.  Ability of Daphnia to buffer trout lakes against periodic nutrient inputs: With 2 figures in the text , 1985 .

[70]  R. Carlson,et al.  Direct and indirect effects of zooplankton grazing on phytoplankton in a hypereutrophic lake , 1984 .

[71]  O. Phillips,et al.  THE EQUILIBRIUM AND STABILITY OF SIMPLE MARINE BIOLOGICAL SYSTEMS. II. HERBIVORES , 1974 .

[72]  K. Porter A METHOD FOR THE IN SITU STUDY OF ZOOPLANKTON GRAZING EFFECTS ON ALGAL SPECIES COMPOSITION AND STANDING CROP1 , 1972 .

[73]  N. Yan Empirical Prediction of Crustacean Zooplankton Biomass in Nutrient-Poor Canadian Shield Lakes , 1986 .

[74]  M. Lynch,et al.  Predation, enrichment, and phytoplankton community structure1 , 1981 .

[75]  J. Shapiro The Importance of Trophic-Level Interactions to the Abundance and Species Composition of Algae in Lakes , 1980 .

[76]  D. J. Hall,et al.  Seasonal distribution and abundance of fishes in the littoral zone of a Michigan lake , 1977 .

[77]  D. Wollkind Exploitation in Three Trophic Levels: An Extension Allowing Intraspecies Carnivore Interaction , 1976, The American Naturalist.

[78]  L. Slobodkin,et al.  Regulation in Terrestrial Ecosystems, and the Implied Balance of Nature , 1967, The American Naturalist.

[79]  H. Paerl,et al.  Toxic and inhibitory effects of the blue-green alga Microcystis aeruginosa on herbivorous zooplankton , 1987 .

[80]  Biosciences Colloquium,et al.  Analysis of ecological systems , 1980 .

[81]  R. Guillard,et al.  YELLOW‐GREEN ALGAE WITH CHLOROPHYLLIDE C 1, 2 , 1972 .

[82]  K. Webster,et al.  Some size-dependent inhibitions of larger cladoceran filterers in filamentous suspensions , 1978 .

[83]  J. E. Cohen,et al.  Environmental correlates of food chain length. , 1987, Science.

[84]  S. Carpenter,et al.  Regulation of Lake Primary Productivity by Food Web Structure. , 1987, Ecology.