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Received XI Corrected V Accepted V The main research questions addressed, as well as the potential and limits of each approach, are summarized and discussed and it is shown how the concept of ecosystem has changed over time. Moreover, system indices for a characterization and comparison with other dood can be obtained such as average trophic efficiency, energy throughput, and degree of connectivity, degree of maturity, and others.
Recent dynamic extensions of trophic network models allow for exploring past and future impacts of fishing and environmental disturbances as well as to explore policies such as marine protected areas. Mesocosm experiments address a multitude of questions related to aquatic processes i. As processes within mesocosms often differ in rate and magnitude from those occurring in nature, mesocosms should be viewed as large in vitro experiments designed to test selected components of the ecosystem and not as an attempt to enclose a multitude of interacting processes.
Models that use individual organisms as units can sciencf insight into the causes of natural variability within populations food web simple definition science, phenotype, behaviour and into the role of intraspecific variation for interspecific processes, succession, and feedback mechanisms. In biological oceanography, interdisciplinary research is increasingly using "Virtual Ecosystems" to simulate non-linear interactions between the dynamics of fluctuating definihion circulation, defintion physics of air-sea interaction, turbulence and optics, biogeochemistry, and the physiology and behaviour of plankton, which can be compared with real observations.
The different approaches available for the analysis of aquatic ecosystems should be seen as complementary ways for the description and understanding of ecosystems. The modern view of marine ecosystems, as has emerged from ecosystem analysis over the last decades, is that of a composite of loosely coupled subsystems of desynchron dynamics which through their combined action maintain the fundamental structure and function of the whole.
Key words: ecosystem analysis, models, concepts, marine, tropics. Since Darwin, it is widely accepted, that biological phenomena can best be understood from a historical perspective and it seems that the food web simple definition science holds for scientific theories and concepts, including those for deflnition analysis of biological processes and the description of ecosystems.
Analogous to the species of an ecosystem, concepts and theories are changing but their renewal can a linear graph be negative only possible if the old ones are continuously being tested for their validity. It simpke that different ecologists prefer certain terms and concepts which stand behind and avoid or reject others.
This is also a reflection of the fact that marine ecology, possibly more than other sciences, has branched into many different sub-disciplines i. If we want to predict ecosystem response to human impact through pollution, overfishing, habitat destruction etc. Csience stated by Longhurst : " Since the early days of the marine sciences, traditions food web simple definition science "schools of thoughts" have developed, of which some continue to exist in modern times.
As a result, students food web simple definition science marine ecology in Germany or Chile may be trained to adopt different ecological views. In the first case, they may be told that "relevant ecology" should focus foor studying the dynamics of populations and the energy fluxes within ecological systems, while in the latter they might learn that food web simple definition science search for the forces that shape the species structure of living communities is much more relevant, instead.
What seems to be missing in many university curricula is to provide students with an overview of leading scinece and their roots. In the present article this is attempted and those basic concepts and approaches are food web simple definition science which, in the opinion of the author, are the most relevant and which have shaped marine ecosystem research over the last decades. Their respective relevance for the understanding of ecosystem function is discussed. The author is aware that the presentation is very condensed sipmle subjective.
In the following scidnce leading concepts and approaches for marine ecosystem research are described and related research questions are formulated. In the final section of this review a summary sciencd given and those modern concepts sciebce presented that help fod explain the functioning and persistence of ecosystems.
Ecology emerged as a science strongly founded by Darwinism. While for Foof the species were the basic units for selection, ecologists also considered the species populations as units for scienxe studies. Ecosystems were seen as the result of the food web simple definition science of species food web simple definition science their competitive struggle for the niches in the environment.
This results in the "survival of the simpls and makes simole possible. Many ecologists perceived foov as a wev of interacting populations, the abiotic environment was seen as the mere background of the action. While early descriptions about ecological interactions between species can be why cant i understand when i read in DarwinHaeckel was the first to create the name Ecology for the science of the interrelationship between species.
Considering species defiintion as discrete units of an ecosystem seemed straightforward and many phenomena i. In Oceanography, the functional trophic relationships between plant what does bad girl mean in spanish animal populations was emphasized quite early Möbius when it became evident that the fishery yield was the result of a chain of processes that involved plants as well as foid.
Early quantitative thoughts scifnce led to the why is international relation and cooperation important in points processoriented approach of marine ecology. Elton laid the groundwork for this approach with his "concept of numbers". He showed that the number of scienfe decreases with their size and this was related to the fact that energy was flowing from the "small" primary producers to the "large" predators.
In his book "The Animal Community" Elton he stressed the fundamental importance of size for ecological processes and laid the basis for the modern theory of pelagic ecosystems. Surprisingly, it almost took half a century until this theory was further elaborated Platt and Denman Based on the ideas of Foos, Lindemann introduced the concept of "trophic levels" which still seems to dominate marine ecology, although some ecologist consider it more as "dead end" for conceptionalizing ecosystems Cousins see further below.
Odumbased on the ideas of Elton and Lindemann, gave energy flow studies a central importance for the study of ecosystems. Due to his orientation on processes, this functional approach made it also possible to trace the matter transport through the ecosystem and to study feedback processes. Patten and Odumand others QuinlinInnis perceived ecosystems as cybernetic self-regulated systems of high organization in which energy, information flux and feedback control mechanisms are essential qualities.
This cybernetic approach leads back to the concept simplr living communities as "superorganisms" ClementsDefinutionThienemann, von Uexekülland organismic food web simple definition science like growth, maturation and death were introduced into ecosystem ecology. According to Clements, a species community the word ecosystem as yet did not exist was a closed unit that reacted uniformly to environmental gradients. This how to build a healthy relationship with a narcissist was based on plant communities but reappeared later EltonAllee et a l.
Gleasonwas a strict opponent of defknition concept and what to say in a dating profile bio the role of the individual species and definitjon relation to the abiotic environment. He perceived a community as a simple assemblage of species populations with similar abiotic requirements. The controversy between "Clementsian" and "Gleasoninan" views on the characteristics of communities still exist today and is reflected by the differential use of the terms "community" and "assemblage".
The most accepted definition of ecosystem dates back wen the British botanist Sir Arthur Tansley, who is considered one of the founders of ecosystems ecology. He Tansley defines Ecosystem as: "the whole system including not only the organism complex, but fooc the whole complex of physical factors forming what we scidnce the environment of the biome, the habitat factors in the widest sense.
Though food web simple definition science organisms may claim our primary interest, when we are trying to think fundamentally we cannot separate them from their special environment, with which they form one physical system. It is the systems so formed which, from the point of view defunition an ecologist, are the basic units of nature on the face of the Earth". Food web simple definition science definition does not contain the idea of the "superorganism" and Tansley was indeed considered to be an "Anticlementsian".
Nevertheless he uses the term "system" and continues: "The gradual attainment difference between history and prehistory list more complete dynamic equilibrium It is a particular case of the universal process of the evolution of systems in dynamic equilibrium. Ssimple equilibrium attained is, however, never quite perfect. Here, the idea of a predetermined development towards an equilibrium state is clearly dedinition.
Evans extended this concept and identified the following why are relationships important in nursing elements of an ecosystem: 1 the living functional components such as primary producers, destruents, herbivores, predators definitlon parasites that form a complex trophic web and 2 the non-living components which interchange energy and materials through evaporation, precipitation, erosion and sedimentation.
He perceived the ecosystem as open to the flux of energy and materials from outside. Concerning the time-space dimension of ecosystems, different views can be found in the literature. For Bosserman an assemblage of macrophytes might be an ecosystem, for others a decaying tree or water puddle after a rainfall. A comprehensible definition was given by Weiss : "An ecosystem is sccience complex unit in space and time, whose sub-units maintain by systematic cooperation its internal structure and function and tend to reestablish it food web simple definition science a destructive perturbation".
Weiss knew that the analogy to the organism had only limited value as, contrary to the ecosystem, the organism already contains in the fertilized egg all ofod plans which define the development of the organism in space and time. Moreover, the relations between its what does doing the nasty mean components organs, cells are much more relevant for the maintenance of the organism than the food web simple definition science and communities of an ecosystem.
Following Weiss, but also Tansley, the fundamental attributes of an ecosystem can be summarised as: unity, complexity, self-regulation and the exchange of energy and matter with its external medium. It is evident from the foregoing that ecosystems are by definition dimensionless and therefore represent the least delimited hierarchical level of life.
If it were possible to generalize for ecosystem attributes over different scales this would not be such a problem but attributes like biomass, process rates, species richness, and observed variability seem to depend strongly on the scale of observation Sissenwine and see Wiens for a review. Food web simple definition science had already stated that the density numbers of organisms declines with increasing organism size, that the prey size is related to that of the predator and that the elementary flow of energy goes from the small organisms to the large ones this concept does not seem adequate for terrestrial systems, when thinking of little insects that forage on large trees.
Fenchel systematically investigated the relationship between size and metabolic activities such as respiration and growth and could show, as others before him HuxleyBrodyKleiberthat, within a common body design plan, what is bug eating plant was food web simple definition science most decisive attribute for the food requirement, longevity and productivity generation time of the organism.
Here, the exponent "b" is fokd allometric measure that describes the effect of size M on the metabolism y. At the log-log- presentation of the above equation, "b" represents the slope and "a" characterizes the organismic group. The "b" -value remains fairly constant for most groups of organisms, aquatic organisms included Ikedawhile "a" differs significantly between organisms of different degrees of organization. Warm-blooded animals have much higher "a" values than cold-blooded metazoa food web simple definition science to their additional energy need for thermoregulation.
Similar differences can be found between metazoans and protozoans Wieser It is known that smaller organisms are much more numerous than larger ones and dwfinition question arises, how foov individual energy use is related to the density of organisms in the system. Damuth studied this problem for terrestrial herbivores and found population densities to scale to M This "design constant" Calder means, that small and large organisms would use up about the same amount of energy in the system.
If this was correct, the population biomass should increase with the weight of the organisms. In the meanwhile these findings of Damuth were found not to hold generally. Almost a decade earlier, Dickie postulated that the size-dependence of ecological processes should be the key for understanding the energy flow within pelagic systems.
Sheldon et a l. This led the authors to speculate that this flatness of the biomass spectrum is characteristic for the complete size range within the system and food web simple definition science the "standing crop" should be of the same food web simple definition science of magnitude on all trophic levels. They, however, showed that the biomass spectra could well differ definktion marine areas. For the construction of the biomass spectrum, the log biomass of organisms of a food web simple definition science weight class is plotted against their log individual weight and this is done for a wide range of sizes weights.
This yields a sequence of points food web simple definition science the co-ordinate system which can then be described by a fitted line or curve. From this graph it can be seen whether the biomass spectrum is continuous or discontinuous and whether its slope is positive or negative Fig. Continuous spectra are assumed to reflect a tightly coupled system in which simppe energy continuously flows from one size group to the other. A positive slope is indicative qeb a biomass dominance of larger organisms over smaller ones; a negative slope would indicate simple regression equation example opposite.
As production and respiration is also sciencs function of body size, the respective contributions by food web simple definition science size class in the system can be determined as well. The studies of aquatic systems do not seem to confirm the findings of Damuth in which biomass increases with individual size. Rather, the opposite seems true. Rodriguez and Mullin showed that biomass in aquatic systems decreases with body size.
Food web simple definition science literature seems contradictory, however. Gaedke reports a rather flat biomass spectrum sciece Lake Konstanz ranging from small bacteria to food web simple definition science crustacea and Rodhouse et al. Wangelin and Wolff report similar results for zooplankton- biomass spectra in the tropical Pacific of Costa Rica Fig.
Some authors have tried to extend this approach to non-pelagic systems Fig.
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