Resistance and re-organization of an ecosystem in response to biological invasion: Some hypotheses

Using a dynamic model of Lake Chozas developed by Marchi et al. (2011), we tested three hypotheses about recovery of the indigenous community and water quality after radical changes caused by introduction of an invasive allochthonous crayfish, Procambarus clarkii:

1.

Can the lake resist the pressure of an invasive species, like P. clarkii, by adaptation?

2.

Can the ecosystem recover when all the crayfish are removed and low phosphorus concentrations persist in inflow water?

3.

Does the simulated recovery of submerged vegetation occur at a total phosphorus concentration below 100 mg TP m⁻³, as estimated by Scheffer et al. (1993), Scheffer (1997), Jeppesen et al. (1998) and Zhang et al. (2003)?

We obtained the following answers:

1.

Lake Chozas can at least partly resist by adaptation. A combination of possible parameter changes could lead to a significant increase in eco-exergy.

2.

Removal of the phosphorus represented by crayfish (by harvesting) implies complete recovery of the lake and its eco-exergy, albeit not necessarily with the same organisms having the same properties.

3.

The expected hysteresis created by introduction and harvesting of crayfish is observed under the following conditions: phytoplankton dominance at total phosphorus ≥ about 200-250 mg TP m⁻³ and submerged vegetation returns at total phosphorus < 100 mg TP m⁻³.

     

The free energy and information embodied in the amino acid chains of organisms

It is generally accepted as a useful and workable hypothesis that when an ecosystem receives an inflow of exergy (energy that can do work) it will utilize this flow of exergy to move as far away from thermodynamic equilibrium as possible after the exergy (energy) for maintenance has been covered. If more combinations of system components including organisms are offered, the combination of components and processes that will bring the system most away from thermodynamic equilibrium will win.The amino acid sequences of the proteins e.g. enzymes determine and control the life processes of the organisms and may be viewed as information sensu lato. The free energy of oxidation of the amino acids and the peptide bonds of the cell enzymes expresses therefore the exergy content, eco-exergy or work capacity that the information contributes to “moving further away from thermodynamic equilibrium”. In this paper eco-exergy is calculated and plotted versus the β-values (a measure of the information contained in the genome) for different organisms. The eco-exergy density was previously (see [J?rgensen et al., 1995] and [J?rgensen et al., 2005]) proposed to be calculated as the summation of the product of the β-values representing the information of the genome multiplied by the concentrations of the respective ecosystem components. This analysis shows a strong correlation between the β-values and free energy released when oxidizing the enzymes. The β-values can therefore be assumed to represent the free energy that the organisms have invested in genetic information.     

The role of conjugation in the gene-individual-population relationships in increasing eco-exergy

The genotypic and phenotypic processes were incorporated into one system in the gene-individual-population relationships under the framework of Individual based models (IBMs). The gene types addressing different degrees of metabolic efficiency and toxin susceptibility were provided as attributes in the individuals. Subsequently ecological processes such as food competition and movement were allowed concurrently on the 2-D space to determine the suitable species adapted to the system. The integrative gene-individual-population model accordingly responded to gene exchanges between the neighboring individuals through conjugation. At a substantially low level of gene exchange, system heterogeneity increased to produce high levels of eco-exergy, being presented by species diversity and total population size in the system. The issues related to genetic and ecological effects in the integrative gene-individual-population relationships were further discussed.     

Dynamic model of Lake Chozas (León, NW Spain)—Decrease in eco-exergy from clear to turbid phase due to introduction of exotic crayfish

We developed a dynamic model of the phosphorus cycle in Lake Chozas, a small shallow water body in León (NW Spain). The calibrated model simulated seasonal dynamics of phosphorus concentrations in major components of the lake's ecological network before and after 1997, the year when an invasive allochthonous crustacean, the Louisiana red swamp crayfish (Procambarus clarkii), was introduced into the lake. The shift from clean to turbid phase, due to grazing by crayfish on submerged vegetation, caused a gradual decrease in eco-exergy, reflecting an increase in entropy, related to breakdown of ecosystem internal equilibria. This case study verifies the hypothesis of Marchi et al. (2010) that, after an initial relatively stable state, the allochthonous species may cause an increase in entropy indicating perturbation of the ecosystem.     

Ecosystem services as a counterpart of emergy flows to ecosystems

A generic input-state-output scheme has been used to represent ecosystem dynamics. Systemic approaches to ecosystems use functions that are based either on inputs, state or outputs of the system. Some examples of approaches that use a combination of functions have been recently proposed. For example the use of eco-exergy to emergy flow can be seen as a mixed input-state approach; more recently, to connect the state to the output of the ecosystem, the relation of eco-exergy and ecosystems services has been proposed. This paper studies the link between the useful output of an ecosystems and its input through the relation between ecosystem services and emergy flow, in a kind of grey/black box scheme (i.e., without considering the state and the structure of the ecosystem). No direct connection between the two concepts can be determined, but identifying and quantifying the emergy flows feeding an ecosystem and the services to humans coming from them facilitate the sustainable conservation of Nature and its functions. Furthermore, this input-output relation can be established in general by calculating the ratio of the value of the ecosystem services to the emergy flow that supports the system. In particular, the ratio of the world ecosystem services to the emergy flow supporting the entire biosphere has been calculated showing that, at least at the global level, Nature is more efficacious in producing “money” (in form of ecosystem services) than economic systems (e.g., national economies and their GDP).     

Network calculations and ascendency based on eco-exergy

Ascendency is an index of activity and organization in living systems calculated in terms of flows. The concern here is with how that quantity behaves when the flows in question are measured in terms of eco-exergy. The storage of eco-exergy has served as a goal function in assessing parameter values for structurally dynamic models, but network magnitudes and topologies can change in response to significant changes in the forcing functions. As storages are relatively insensitive to such changes, it is advisable in such cases to explore how changes in a flow variable, like ascendency, might capture network adaptations. It happens that changes in ascendency calculated in terms of flows of simple energy are small in comparison to corresponding variations in the storages of eco-exergy. But when ascendency is reckoned in terms of flows of eco-exergy, its changes in response to network changes are more comparable to those in the storages. Ascendency seems to be more sensitive to changes in flow topology, however, so that a combination of eco-exergy storage and eco-exergy ascendency would probably be most appropriate for situations where changes in flow topology are significant.     

Application of eco-exergy for assessment of ecosystem health and development of structurally dynamic models

Eco-exergy has been widely used in the assessment of ecosystem health, parameter estimations, calibrations, validations and prognoses. It offers insights into the understanding of ecosystem dynamics and disturbance-driven changes. Particularly, structurally dynamic models (SDMs), which are developed using eco-exergy as the goal function, have been applied in explaining and exploring ecosystem properties and changes in community structure driven by biotic and abiotic factors. In this paper, we review the application of eco-exergy for the assessment of ecosystem health and development of structurally dynamic models (SDMs). The limitations and possible future applications of the approach are also addressed.