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Biological stoichiometry: a chemical bridge between ecosystem ecology and evolutionary biology.

The mission of the American Society of Naturalists is "to advance and diffuse knowledge of organic evolution and other broad biological principles so as to enhance the conceptual unification of the biological sciences." In this article, I argue that the area of biology least integrated with knowledge of organic evolution is the field of ecosystem ecology, as evidenced by a semiquantitative literature survey of use of terms in the scientific literature. I present an overview of recent theoretical developments and empirical findings in the emerging field of biological stoichiometry (the study of the balance of energy and multiple chemical elements in living systems). These developments hold some promise as a means to conceptually integrate ecosystem ecology, with its emphasis on flows and pools of energy and chemical elements, with evolutionary biology, with its emphasis on genetic fitness and the biochemical products of the genome. For example, recent evidence indicates that organismal C : P and N : P ratios have a major impact on biologically mediated flows of energy and phosphorus; in turn, variations among taxa in these ratios are connected to evolved differences in organismal growth rate because of the connection between growth rate and the need for increased allocation to P-rich ribosomal RNA. In this way, evolutionary change in growth-related traits, by altering organismal P requirements, has direct biogeochemical implications, while ecosystem conditions can constrain evolutionary acceleration of growth rates by imposing a direct P limitation on production of the needed biochemical machinery of growth. Thus, stoichiometric theory provides a broad biological principle that can interconvert the currencies and concerns of ecosystem ecology and evolutionary biology, facilitating integration of diverse fields of study and contributing to conceptual unification of the biological sciences.

Biological Evolution↗

Sequentially assembled food webs and extremum principles in ecosystem ecology.

1. Successional changes during sequential assembly of food webs were examined. This was carried out by numerical methods, drawing one species at a time from a species pool and obtaining the permanent (persistent) community emerging at each step. Interactions among species were based on some simple rules about body sizes of consumers and their prey, and community dynamics were described in terms of flows of biomass density. 2. Sequential assembly acted as a sieve on the communities, assembled communities having many properties different on average from those of feasible, stable communities taken at random from the species pools. 3. Time-series of community development were consistent with certain functions thought to go to an extremum (maximum or minimum) in ecosystem ecology, including a rapid early increase in net primary productivity and ascendency, although a clear trend in total biomass density was not evident and resilience decreased rather than increased. 4. In addition, more gradual changes in food web structure took place during succession to which the ecosystem goal functions were relatively insensitive. These changes included gradual increases in the number of species, invasion resistance, number of loops of length > 2 and number of prey species per consumer species. 5. We therefore argue that ecosystem and community dynamics can offer complementary insights into the process of ecological succession. The extremum principles of ecosystem ecology highlight some of the major properties of succession, whereas the community ecology sheds light on some more subtle changes taking place within the networks.

Animals↗

Seabirds as indicators of changes in marine ecosystems: ecological monitoring on Machias Seal Island.

Changes in marine ecosystems can be manifested in many different ways, on different temporal and spatial scales. Seabirds are top consumers in marine foodwebs and offer opportunities to detect and assess the biological effects of changes in physical parameters (sea-surface temperature [SST], salinity, depth of thermocline etc.) of the marine ecosystem. We compare six-eight years' of data on the biology (diet, and breeding success) of four species of seabird (arctic tern Sterna paradisaea and common tern S. hirundo, which feed at the sea surface; and Atlantic puffin Fratercula antica and razorbill Alca torda, which dive 30-60 m for their prey) breeding on Machias Seal Island (MSI) in the Bay of Fundy with both our own meteorological and oceanographic measurements, and with standard measurements from conventional sources. These are compared with fisheries data on changes in the main prey of all the seabirds concerned (juvenile or '0-group' herring Clupea harengus) which are the most direct link between the seabirds and the physical properties of the marine system. We explore relationships between seabird productivity and diet, and other aspects of both herring biology (larval surveys, and fat content) and oceanography (SST data from the island, and remotely sensed data from the entrance to the Bay of Fundy). Timing of laying by puffins followed SST variation at neither the local (MSI) nor regional scales, but at the scale of the North Atlantic, following the trend of populations breeding off northern Norway. The proportion of herring in the diet of terns over 6 years varied inversely with herring larval abundance the previous fall; this relationship was not statistically significant in the puffin and razorbill. A major new finding is the considerable (approximately 50%) inter-annual variation in the energy density (fat content) of juvenile herring that are the main seabird prey; breeding success of both species of tern varied in parallel with the energy density of juvenile herring in the diet until the last two years of the study, when sandlance (Ammodytes sp.) and euphausid shrimp predominated in the diet. Our long-term research approach combines traditional population monitoring (of numbers of breeding birds) with demographic, behavioural and environmental monitoring, to provide new understanding of the marine ecosystem as well as of seabirds.

Animals↗

Application of eutrophication indices for assessment of the Bulgarian Black Sea coastal ecosystem ecological quality.

The present paper is an attempt to test the applicability of the trophic state index (TRIX) for scaling the eutrophication along the Bulgarian Black Sea coastal zone in concert with a number of chemical and biological descriptors aimed at selection of relevant indicators of marine coastal area ecological quality. The following environmental parameters have been considered: t(o), salinity, nutrients--inorganic P, N and dissolved Si, dissolved oxygen and oxygen saturation, phytoplankton--taxonomic structure, abundance and biomass, chlorophyll a, zooplankton--taxonomic structure, abundance and biomass. Principal Component Analysis was applied in order to figure out and score the most relevant combination of parameters to discriminate between sites and select representative descriptors (pressure/state) of eutrophication. The following variables are defined as relevant descriptors for classification of the sites: nutrients (N, P, Si) and their molar ratios (N:P and Si:P), the capacity of the system to produce and sustain organic matter (chlorophyll a, phytoplankton biomass), phytoplankton taxonomic dominance (Bacilariophyceae:Dinophyceae biomass ratio), grazing pressure (phytoplankton:zooplankton biomass, Bacilariophyceae:Copepoda), plankton diversity index (Hb and Ha) and the trophic state index (TRIX). The investigated sites under a different anthropogenic impact are classified according to selected descriptors and their water quality state.

Animals↗

Historical overfishing and the recent collapse of coastal ecosystems.

Ecological extinction caused by overfishing precedes all other pervasive human disturbance to coastal ecosystems, including pollution, degradation of water quality, and anthropogenic climate change. Historical abundances of large consumer species were fantastically large in comparison with recent observations. Paleoecological, archaeological, and historical data show that time lags of decades to centuries occurred between the onset of overfishing and consequent changes in ecological communities, because unfished species of similar trophic level assumed the ecological roles of overfished species until they too were overfished or died of epidemic diseases related to overcrowding. Retrospective data not only help to clarify underlying causes and rates of ecological change, but they also demonstrate achievable goals for restoration and management of coastal ecosystems that could not even be contemplated based on the limited perspective of recent observations alone.

Animals↗

[Consequences of antibiotic therapy to the intestinal ecosystem].

Ecological impact of antibiotherapy results from the interaction between microorganisms in the ecosystems and antibiotics at which they are exposed. The amount of antibiotics use in the world is continuously increasing. The fraction devoted to human care is only about half the total amount. There are multiple other fields of usage, in agriculture, breeding and veterinary medicine. Bacterial ecosystems exposed at antibiotherapy in man are mainly the skin and the gastrointestinal and respiratory tracts. The gastrointestinal system is quantitatively predominant and the consequences of the bacterial imbalance induced by antibiotics are potentially severe. It is the reason why it is the most extensively studied, in the literature and in the present review. The origin of resistant bacteria will be briefly discussed.

Animals↗

Are bacterial biofilms constrained to Darwin's concept of evolution through natural selection?

Numerous antimicrobial agents have been developed which act at the molecular, cellular, and organismal levels. However, few have been developed which act at the community-level. This results largely from the failure of Darwinian selection theory to envision communities as units of proliferation and evolution. It is thus difficult to conceive of microbial communities as causative agents and to develop antimicrobials which are effective against them. Consequently, we find it necessary to consider a more comprehensive biological paradigm which envisions biofilm communities and other microbial associations (e.g. mixed infections, food spoilage, tooth decay) as units of existence, activity, ecology, proliferation, survival, and evolution. These communities exist in the same sense that organisms exist as units of ecological activity. This is a simpler, more comprehensive, and more unifying theory of ecology. It is simpler in that it no longer requires convoluted explanations of altruistic behavior in terms of individual selection. It is more comprehensive by not constraining evolution to the selection of any single level of biological organization (genes, races, lineages, or groups). It unifies in that it bridges the boundaries between microbial ecology, evolutionary ecology and ecosystem ecology. The basis for this theory lies in recognizing that life consists of various forms of information (order) which evolve not only through genetic recombination and mutation, but also through the recombination of organisms within communities (as well as other mechanisms, some of which are considered beyond the realm of biology). It also involves setting aside the concept of evolution through selection and competition, in favor of evolution through proliferation and association.

Bacteria↗

Environment, ecosystems, and ecological behavior: a dialogue toward developing nursing ecological theory.

Current epistemological foundations of nursing theory incorporate minimal ecology theory. The purpose of this article is to present a nursing ecological theory with a goal to broaden current nursing perspectives by incorporating expanded concepts of global ecosystems, communities, and interrelationships derived from ecological sciences. A theory derivation process is utilized, and a nursing ecological model is proposed. Nurses face a challenge to translate global environment concern and ecological beliefs into professional activities. Elucidating a nursing ecological theory may guide our profession toward new directions in holistic care and will be good for the care of our patients, profession, and the Earth.

Concept Formation↗

Using ecosystem engineers to restore ecological systems.

Ecosystem engineers affect other organisms by creating, modifying, maintaining or destroying habitats. Despite widespread recognition of these often important effects, the ecosystem engineering concept has yet to be widely used in ecological applications. Here, we present a conceptual framework that shows how consideration of ecosystem engineers can be used to assess the likelihood of restoration of a system to a desired state, the type of changes necessary for successful restoration and how restoration efforts can be most effectively partitioned between direct human intervention and natural ecosystem engineers.

Ecology↗

Valuing biodiversity: reality or mirage?

The objective of this paper was to consider the social value of biological diversity and explore if this value could be expressed in terms of a unidimensional metric in money. Economics distinguishes between use-values and non-use-values, which are critically evaluated for valuing biodiversity. It is shown that these utility-based valuations have severe limitations as they treat species in isolation from their ecological contexts. In contrast, ecosystem ecology regards ecosystems as an integrated non-linear and nonconvex system in which ecosystem functions can be understood as a four-component cycle; exploitation, accumulation of biomass, creative destruction and renewal. Within such a cycle, ecosystems can be seen to have two properties: stability and resilience. A good proxy for resilience is the probability of extinction of species, and social value of biodiversity can be expressed as a partial ordering with this probability as an index. This approach is consistent with decision theory, of which social choice is an important component, pioneered by Arrow.

Animals↗

Paradox of enrichment: destabilization of exploitation ecosystems in ecological time.

Six reasonable models of trophic exploitation in a two-species ecosystem whose exploiters compete only by depleting each other's resource supply are presented. In each case, increasing the supply of limiting nutrients or energy tends to destroy the steady state. Thus man must be very careful in attempting to enrich an ecosystem in order to increase its food yield. There is a real chance that such activity may result in decimation of the food species that are wanted in greater abundance.

Competitive Behavior↗

Effects of field exposure to diazinon on small mammals inhabiting a semienclosed prairie grassland ecosystem. I. Ecological and reproductive effects.

The widespread use of cholinesterase-inhibiting pesticides in the environment presents increasing concerns about their effects on human, wildlife, and ecosystem health. As a group, these pesticides are generally highly toxic and have great potential for negatively affecting nontarget organisms. Small mammals have proven to be ideal biomonitors of environmental contaminants, and were used here to test for possible effects of a widely used cholinesterase-inhibiting insecticide, diazinon, in a natural field setting. Using 12 0.1-ha terrestrial mesocosms, we examined the effects of low-level diazinon exposure on the small mammal communities inhabiting semienclosed grassland ecosystems. Our primary objective was to test the hypothesis that diazinon, applied at two different recommended label application rates, would not cause any observable adverse ecological or reproductive effects on small mammal populations and communities. Experimental small mammal communities consisting of Sigmodon hispidus, Microtus ochrogaster, Reithrodontomys fulvescens, and Mus musculus were stocked at natural densities and sex ratios inside empty mesocosms. Diazinon 4E was applied at two different maximum recommended label application rates, 0.56 kg a.i./ha (1x) and 4.5 kg a.i./ha (8x), and controls remained unsprayed, with four enclosures (replicates) per treatment. Two 30-d trials were run during peak rodent breeding seasons and enclosures were sampled on days 2, 16, and 30 of each trial. Recovery of small mammals was not significantly different among treatments, although fewer animals were recovered from the diazinon-exposed enclosures in both trials. Analysis of trapping data suggested that the normally strong competitive relationship between Sigmodon and Microtus may be altered by the pesticide, favoring Microtus in the diazinon-exposed enclosures. Incidence of reproductive condition was found to be reduced 20 to 80% and 33 to 100% in diazinon-exposed males and females, respectively. Reproductive productivity, including percentage of pregnant females and of females giving birth, was significantly reduced in diazinon-exposed animals. Percentage of pregnant females ranged from 13.6 to 43.5% in diazinon-exposed animals compared to 40 to 80% for control animals, and percentage of females giving birth ranged from 0 to 17% in diazinon-exposed animals compared to 22 to 50% for control animals. Generally, the effects found in this study suggest that diazinon was relatively persistent in the sprayed enclosures and that oral routes of exposure (consumption of dead and dying arthropods, grooming) may have been important. Ecological relationships and reproduction in both herbivorous and omnivorous mammals were negatively impacted by diazinon exposure. Overall, ecological relationships in the enclosed prairie grassland ecosystem were disrupted by diazinon, probably through a combination of sublethal effects, particularly reproductive effects, impacting individuals and their populations. This suggests that negative impacts on populations and community structure and function may persist longer than diazinon persists in the environment.

Animals↗

Autocatalysis in cultural ecology: model ecosystems and the dynamics of biocultural evolution.

Using a well-known mathematical model frequently applied in theoretical population dynamics, certain ecological mechanisms are investigated that are inherent in the organic evolution of cultural capacities in man. Culture is argued to involve ecological interactions exhibiting analogies to the interaction of chemical species in autocatalytic biomolecular reactions. In the model, biocultural evolution proceeds by more and more broadening ecological niches and, thus, releasing competitive selection pressure on the populations involved. This, in turn, facilitates the maintenance of polymorphism in these populations as well as the individual acquisition of organic traits through learning and cultural transmission. The result is that the genetic variance in phenotypic expressions decreases at an accelerated rate.

Cultural Evolution↗

Microbial ecology of the closed artificial ecosystem MELiSSA (Micro-Ecological Life Support System Alternative): reinventing and compartmentalizing the Earth's food and oxygen regeneration system for long-haul space exploration missions.

MELiSSA is a bioregenerative life support system designed by the European Space Agency (ESA) for the complete recycling of gas, liquid and solid wastes during long distance space exploration. The system uses the combined activity of different living organisms: microbial cultures in bioreactors, a plant compartment and a human crew. In this minireview, the development of a short-cut ecological system for the biotransformation of organic waste is discussed from a microorganism's perspective. The artificial ecological model--still in full development--that is inspired by Earth's own geomicrobiological ecosystem serves as an ideal study object on microbial ecology and will become an indispensable travel companion in manned space exploration.

Bacteria↗