An ecosystem model is an abstract, usually mathematical, representation of an ecological system (ranging in scale from an individual population, to an ecological community, or even an entire biome), which is studied to better understand the real system. [2]
Using data gathered from the field, ecological relationships—such as the relation of sunlight and water availability to photosynthetic rate, or that between predator and prey populations—are derived, and these are combined to form ecosystem models. These model systems are then studied in order to make predictions about the dynamics of the real system. Often, the study of inaccuracies in the model (when compared to empirical observations) will lead to the generation of hypotheses about possible ecological relations that are not yet known or well understood. Models enable researchers to simulate large-scale experiments that would be too costly or unethical to perform on a real ecosystem. They also enable the simulation of ecological processes over very long periods of time (i.e. simulating a process that takes centuries in reality, can be done in a matter of minutes in a computer model). [3]
Ecosystem models have applications in a wide variety of disciplines, such as natural resource management, [4] ecotoxicology and environmental health, [5] [6] agriculture, [7] and wildlife conservation. [8] Ecological modelling has even been applied to archaeology with varying degrees of success, for example, combining with archaeological models to explain the diversity and mobility of stone tools. [9]
There are two major types of ecological models, which are generally applied to different types of problems: (1) analytic models and (2) simulation / computational models. Analytic models are typically relatively simple (often linear) systems, that can be accurately described by a set of mathematical equations whose behavior is well-known. Simulation models on the other hand, use numerical techniques to solve problems for which analytic solutions are impractical or impossible. Simulation models tend to be more widely used, and are generally considered more ecologically realistic, while analytic models are valued for their mathematical elegance and explanatory power. [10] [11] [12] Ecopath is a powerful software system which uses simulation and computational methods to model marine ecosystems. It is widely used by marine and fisheries scientists as a tool for modelling and visualising the complex relationships that exist in real world marine ecosystems. [13] [14] [15] [16] [17] [18] [19]
The process of model design begins with a specification of the problem to be solved, and the objectives for the model. [21]
Ecological systems are composed of an enormous number of biotic and abiotic factors that interact with each other in ways that are often unpredictable, or so complex as to be impossible to incorporate into a computable model. Because of this complexity, ecosystem models typically simplify the systems they are studying to a limited number of components that are well understood, and deemed relevant to the problem that the model is intended to solve. [22] [23]
The process of simplification typically reduces an ecosystem to a small number of state variables and mathematical functions that describe the nature of the relationships between them. [24] The number of ecosystem components that are incorporated into the model is limited by aggregating similar processes and entities into functional groups that are treated as a unit. [25] [26]
After establishing the components to be modeled and the relationships between them, another important factor in ecosystem model structure is the representation of space used. Historically, models have often ignored the confounding issue of space. However, for many ecological problems spatial dynamics are an important part of the problem, with different spatial environments leading to very different outcomes. Spatially explicit models (also called "spatially distributed" or "landscape" models) attempt to incorporate a heterogeneous spatial environment into the model. [27] [28] [29] A spatial model is one that has one or more state variables that are a function of space, or can be related to other spatial variables. [30]
After construction, models are validated to ensure that the results are acceptably accurate or realistic. One method is to test the model with multiple sets of data that are independent of the actual system being studied. This is important since certain inputs can cause a faulty model to output correct results. Another method of validation is to compare the model's output with data collected from field observations. Researchers frequently specify beforehand how much of a disparity they are willing to accept between parameters output by a model and those computed from field data. [31] [32] [33] [34] [35]
One of the earliest, [36] and most well-known, ecological models is the predator-prey model of Alfred J. Lotka (1925) [37] and Vito Volterra (1926). [38] This model takes the form of a pair of ordinary differential equations, one representing a prey species, the other its predator.
where,
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Volterra originally devised the model to explain fluctuations in fish and shark populations observed in the Adriatic Sea after the First World War (when fishing was curtailed). However, the equations have subsequently been applied more generally. [39] Although simple, they illustrate some of the salient features of ecological models: modelled biological populations experience growth, interact with other populations (as either predators, prey or competitors) and suffer mortality.[ citation needed ]
A credible, simple alternative to the Lotka-Volterra predator-prey model and its common prey dependent generalizations is the ratio dependent or Arditi-Ginzburg model. [40] The two are the extremes of the spectrum of predator interference models. According to the authors of the alternative view, the data show that true interactions in nature are so far from the Lotka-Volterra extreme on the interference spectrum that the model can simply be discounted as wrong. They are much closer to the ratio dependent extreme, so if a simple model is needed one can use the Arditi-Ginzburg model as the first approximation. [41]
The theoretical ecologist Robert Ulanowicz has used information theory tools to describe the structure of ecosystems, emphasizing mutual information (correlations) in studied systems. Drawing on this methodology and prior observations of complex ecosystems, Ulanowicz depicts approaches to determining the stress levels on ecosystems and predicting system reactions to defined types of alteration in their settings (such as increased or reduced energy flow, and eutrophication. [42]
Conway's Game of Life and its variations model ecosystems where proximity of the members of a population are factors in population growth.
Theoretical ecology is the scientific discipline devoted to the study of ecological systems using theoretical methods such as simple conceptual models, mathematical models, computational simulations, and advanced data analysis. Effective models improve understanding of the natural world by revealing how the dynamics of species populations are often based on fundamental biological conditions and processes. Further, the field aims to unify a diverse range of empirical observations by assuming that common, mechanistic processes generate observable phenomena across species and ecological environments. Based on biologically realistic assumptions, theoretical ecologists are able to uncover novel, non-intuitive insights about natural processes. Theoretical results are often verified by empirical and observational studies, revealing the power of theoretical methods in both predicting and understanding the noisy, diverse biological world.
The Lotka–Volterra equations, also known as the Lotka–Volterra predator–prey model, are a pair of first-order nonlinear differential equations, frequently used to describe the dynamics of biological systems in which two species interact, one as a predator and the other as prey. The populations change through time according to the pair of equations:
Alfred James Lotka was an American mathematician, physical chemist, and statistician, famous for his work in population dynamics and energetics. A biophysicist, Lotka is best known for his proposal of the predator–prey model, developed simultaneously but independently of Vito Volterra. The Lotka–Volterra model is still the basis of many models used in the analysis of population dynamics in ecology.
Population dynamics is the type of mathematics used to model and study the size and age composition of populations as dynamical systems.
Population ecology is a sub-field of ecology that deals with the dynamics of species populations and how these populations interact with the environment, such as birth and death rates, and by immigration and emigration.
A metapopulation consists of a group of spatially separated populations of the same species which interact at some level. The term metapopulation was coined by Richard Levins in 1969 to describe a model of population dynamics of insect pests in agricultural fields, but the idea has been most broadly applied to species in naturally or artificially fragmented habitats. In Levins' own words, it consists of "a population of populations".
The paradox of enrichment is a term from population ecology coined by Michael Rosenzweig in 1971. He described an effect in six predator–prey models where increasing the food available to the prey caused the predator's population to destabilize. A common example is that if the food supply of a prey such as a rabbit is overabundant, its population will grow unbounded and cause the predator population to grow unsustainably large. That may result in a crash in the population of the predators and possibly lead to local eradication or even species extinction.
A functional response in ecology is the intake rate of a consumer as a function of food density. It is associated with the numerical response, which is the reproduction rate of a consumer as a function of food density. Following C. S. Holling, functional responses are generally classified into three types, which are called Holling's type I, II, and III.
In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological community, or life assemblage. The term community has a variety of uses. In its simplest form it refers to groups of organisms in a specific place or time, for example, "the fish community of Lake Ontario before industrialization".
A population model is a type of mathematical model that is applied to the study of population dynamics.
The green chromide is a species of cichlid fish that is native to fresh and brackish water habitats in some parts in India such as Kerala, Goa, Chilika Lake in Odisha and Sri Lanka. The species was first described by Marcus Elieser Bloch in 1790. This species and other members of the genus Etroplus are relatively closely related to the Paretroplus cichlids from Madagascar.
A fishery is an area with an associated fish or aquatic population which is harvested for its commercial or recreational value. Fisheries can be wild or farmed. Population dynamics describes the ways in which a given population grows and shrinks over time, as controlled by birth, death, and migration. It is the basis for understanding changing fishery patterns and issues such as habitat destruction, predation and optimal harvesting rates. The population dynamics of fisheries is used by fisheries scientists to determine sustainable yields.
Ecopath with Ecosim (EwE) is a free and open source ecosystem modelling software suite, initially started at NOAA by Jeffrey Polovina, but has since primarily been developed at the formerly UBC Fisheries Centre of the University of British Columbia. In 2007, it was named as one of the ten biggest scientific breakthroughs in NOAA's 200-year history. The NOAA citation states that Ecopath "revolutionized scientists' ability worldwide to understand complex marine ecosystems". Behind this lie more than three decades of development work in association with a thriving network of fisheries scientists such as Villy Christensen, Carl Walters and Daniel Pauly, and software engineers around the world. EwE is funded through projects, user contributions, user support, training courses and co-development collaborations. Per November 2021 there are an estimated 8000+ users across academia, non-government organizations, industry and governments in 150+ countries.
The paradox of the pesticides is a paradox that states that applying pesticide to a pest may end up increasing the abundance of the pest if the pesticide upsets natural predator–prey dynamics in the ecosystem.
The generalized Lotka–Volterra equations are a set of equations which are more general than either the competitive or predator–prey examples of Lotka–Volterra types. They can be used to model direct competition and trophic relationships between an arbitrary number of species. Their dynamics can be analysed analytically to some extent. This makes them useful as a theoretical tool for modeling food webs. However, they lack features of other ecological models such as predator preference and nonlinear functional responses, and they cannot be used to model mutualism without allowing indefinite population growth.
A trophic function was first introduced in the differential equations of the Kolmogorov predator–prey model. It generalizes the linear case of predator–prey interaction firstly described by Volterra and Lotka in the Lotka–Volterra equation. A trophic function represents the consumption of prey assuming a given number of predators. The trophic function was widely applied in chemical kinetics, biophysics, mathematical physics and economics. In economics, "predator" and "prey" become various economic parameters such as prices and outputs of goods in various linked sectors such as processing and supply. These relationships, in turn, were found to behave similarly to the magnitudes in chemical kinetics, where the molecular analogues of predators and prey react chemically with each other.
Lev R. Ginzburg is a mathematical ecologist and the president of the firm Applied Biomathematics.
The Arditi–Ginzburg equations describes ratio dependent predator–prey dynamics. Where N is the population of a prey species and P that of a predator, the population dynamics are described by the following two equations:
Philosophy of ecology is a concept under the philosophy of science, which is a subfield of philosophy. Its main concerns centre on the practice and application of ecology, its moral issues, and the intersectionality between the position of humans and other entities. This topic also overlaps with metaphysics, ontology, and epistemology, for example, as it attempts to answer metaphysical, epistemic and moral issues surrounding environmental ethics and public policy.
Eco-evolutionary dynamics refers to the reciprocal effects that ecology and evolution have on each other. The effects of ecology on evolutionary processes are commonly observed in studies, but the realization that evolutionary changes can be rapid led to the emergence of eco-evolutionary dynamics. The idea that evolutionary processes can occur quickly and on one timescale with ecological processes led scientists to begin studying the influence evolution has on ecology along with the affects ecology has on evolution. Recent studies have documented eco-evolutionary dynamics and feedback, which is the cyclic interaction between evolution and ecology, in natural and laboratory systems at different levels of biological organization, such as populations, communities, and ecosystems.