Quantitative criteria for estimation of natural and artificial ecosystems functioning.

Using biotic turnover of substances in trophic chains, natural and artificial ecosystems are similar in functioning, but different in structure. It is necessary to have quantitative criteria to evaluate the efficiency of artificial ecosystems (AES). These criteria are dependent on the specific objectives for which the AES are designed. For example, if AES is considered for use in space, important criteria are efficiency in use of mass, power, volume (size) and human labor and reliability. Another task involves the determination of quantitative criteria for the functioning of natural ecosystems. To solve the problem, it is fruitful to use a hierarchical approach suitable for both individual links and the ecosystem as a whole. Energy flux criteria (principles) were developed to estimate the functional activities of biosystems at the population, community and ecosystem levels. A major feature of ecosystems as a whole is their biotic turnover of matter the rate of which is restricted by the lack of limiting substances. Obviously, the most generalized criterion is to take into account the energy flux used by the biosystem and the quantity of limiting substance included in its turnover. The use of energy flux by ecosystem, E(USED)--is determined from the photoassimilation of CO2 by plants (per time unit). It can be approximately estimated as the net primary production of photosynthesis (NPP). So, the ratio of CO2 photoassimilation rate (sometimes, measured as NPP) to the total mass of limiting substrate can serve as a main universal criterion (MUC). This MUC characterizes the specific cycling rate of limiting chemical elements in the system and effectiveness of every ecosystem including the global Biosphere. Comparative analysis and elaboration of quantitative criteria for estimation of natural and artificial ecosystems activities is of high importance both for theoretical considerations and for real applications.     

The effect of cannibalism intensity on net primary production and dynamics of trophic links in aquatic ecosystems.

A mathematical model was used to investigate the effect of cannibalism intensity on the net primary production and the dynamics of trophic links in an aquatic ecosystem characterized by cannibalism at the upper trophic level. A mathematical model of an aquatic ecosystem has been constructed, with the following principal trophic links: limiting nutrient concentration, producers (phytoplankton), nonpredatory and predatory zooplankton. The model takes into account the age structure of the predator and includes two age groups (the young and adults). The adult predators are cannibals feeding on both nonpredatory zooplankton and their own young, which consume phytoplankton. It has been found that when cannibalism intensity increases above a certain level, the concentrations of both adults and the young of the predators decrease. At the same time, the concentrations of the nonpredatory zooplankton and of nutrients increase, while the biomass of producers decreases. When the cannibalism intensity is low, the net primary production of the system increases to a certain level correlated with the increase in cannibalism intensity and drops sharply when the level of consumption of young is high. There is an optimal intensity of cannibalism, at which the productivity in the photosynthesis link is maximal.     

System analysis of links interactions and development of ecosystems of different types.

The anthropogenic impact on the Earth's ecosystems are leading to dramatic changes in ecosystem functioning and even to destruction of them. System analysis and the use of heuristic modeling can be an effective means to determine the main biological interactions and key factors that are of high importance for understanding the development of ecosystems. Cycling of limiting substances, induced by the external free energy flux, and trophic links interaction is the basis of the mathematical modeling studies presented in this paper. Mathematical models describe the dynamics of simplified ecosystems having different characteristics: 1) different degrees of biotic turnover closure (from open to completely closed); 2) different numbers of trophic links (including both "top-down", "bottom-up" regulation types); 3) different intensities of input-output flows of the limiting nutrient and its total amount in the system. Adaptive values of the changes of lower hierarchical levels (populational, trophic chain level) are to be estimated by integrity indices for total system functioning (e.g. NPP, total photosynthesis). The approach developed can be used for evaluating the contributions of lower hierarchical levels to the functioning of the higher hierarchical levels of the system. This approach may have value for determining biomanipulation management and their assessment.     

Mathematical modeling of response of ecosystems with different structure to external impact.

A mathematical model was used to study the response of ecosystems of different structures to external impact. The response was measured as a sensitivity coefficient: the magnitude of the system's response vs. the change of the factor in the inflow. The formula has been obtained to calculate the sensitivity coefficient for ecosystems containing different numbers of trophic links. The derived sensitivity coefficients demonstrate that the degree of compensation for the external impact can differ depending on the type of system regulation and the length of the trophic chain. E. g. the sensitivity coefficient decreases with complexity of trophic links in an ecosystem for top-down controlled systems and impact of degree of openness on sensitivity e.g. closed ecosystems show higher sensitivity then fully open ecosystem to impacts also bottom-up control system show less sensitivity then top-down. Grant numbers: N99-04-96017, N25.     

Key factors in development of man-made and natural ecosystems.

Key factors of ecosystem functioning are of the same nature for artificial and natural types. An hierarchical approach gives the opportunity for estimation of the quantitative behavior of both individual links and the system as a whole. At the organismic level we can use interactions of studied macroorganisms (man, animal, higher plant) with selected microorganisms as key indicating factors of the organisms immune status. The most informative factor for the population/community level is an age structure of populations and relationships of domination/elimination. The integrated key factors of the ecosystems level are productivity and rates of cycling of the limiting substances. The key factors approach is of great value for growth regulations and monitoring the state of any ecosystem, including the life support system (LSS)-type.     

Increase of atmospheric CO2: response patterns of a simple terrestrial man-made ecosystem.

Simple models of terrestrial ecosystems with a limited number of components are an efficient tool to study the main laws of functioning of populations, including microbial ones, and their communities, as components of natural ecosystems, under variable environmental conditions. Among other factors are the increase of carbon dioxide in the atmosphere and limitation of plants' growth by biogenic elements. The main types of ecosystems' responses to changes in environmental conditions (a change in CO2 concentration) have been demonstrated in a "plants-rhizospheric microorganisms-artificial soil" simple experimental system. The mathematical model of interactions between plants and microorganisms under normal and elevated atmospheric CO2 and limitation by nutrients (nitrogen and phosphorus) yielded a qualitative agreement between calculated and experimental values of limiting substances concentrations and release rates of exudates.     

“Biospherics” approach for studies of natural and artificial ecosystems

The main unifying feature of natural and artificial ecosystems is their biotic turnover (cycling) of substances which is induced with energy fluxes. A new integrating scientific discipline – Biospherics – studies biotic cycles (both in experiments and in mathematical models) of different degree of closure and complexity. By its origin, Biospherics is to be connected with extensive studies of Biosphere by Russian academician Vladimir Vernadsky. He developed and used “empirical generalizations” based on innumerous observations, comparisons and reflections. His “bio-geo-chemical principles” of Biosphere and ecosystems development have more qualitative than quantitative nature. Quantitative criteria to evaluate the efficiency of natural and artificial ecosystems are to take into account energy fluxes and their use in ecosystems of different types. At least, three of them are of value for estimation of natural and artificial ecosystems’ functional activities. Energy principle of extensive development (EPED), energy principle of intensive development (EPID) and main universal (generalized) criterion (MUC). The last criterion (Principle) characterizes the specific cycling rate of limiting chemical elements in multi-organism systems, developing under external energy fluxes. Its value can be a quantitative measure of effectiveness for every ecosystem functioning, including our global Biosphere. Different examples of these (above-mentioned) integrated criteria actions are presented and analyzed in the paper.     

Understanding plant-soil relationships using controlled environment facilities.

Although soil is a component of terrestrial ecosystems, it is comprised of a complex web of interacting organisms, and therefore can be considered itself as an ecosystem. Soil microflora and fauna derive energy from plants and plant residues and serve important functions in maintaining soil physical and chemical properties, thereby affecting net primary productivity (NPP), and in the case of contained environments, the quality of the life support system. We have been using 3 controlled-environment facilities (CEF's) that incorporate different levels of soil biological complexity and environmental control, and differ in their resemblance to natural ecosystems, to study relationships among plant physiology, soil ecology, fluxes of minerals and nutrients, and overall ecosystem function. The simplest system utilizes growth chambers and specialized root chambers with organic-less media to study the physiology of plant-mycorrhizal associations. A second system incorporates natural soil in open-top chambers to study soil bacterial and fungal population response to stress. The most complex CEF incorporates reconstructed soil profiles in a "constructed" ecosystem, enabling close examination of the soil foodweb. Our results show that closed ecosystem research is important for understanding mechanisms of response to ecosystem stresses. In addition, responses observed at one level of biological complexity may not allow prediction of response at a different level of biological complexity. In closed life support systems, incorporating soil foodwebs will require less artificial manipulation to maintain system stability and sustainability.     

Principles of biological adaptation of organisms in artificial ecosystems to changes of environmental factors.

Studying material transformations and biotic cycling in artificial ecosystems (AES), we need to know the principles of biological adaptation of active organisms to change in the environment. Microorganisms in AES for water purification are the most active transforming organisms and consumers of the organic substances contained in wastes. Utilization of organic substances is directly connected with the energy fluxes used by AES. According to energy criteria, the energy fluxes used by a biological system tend to reach maximum values under stable conditions. Unutilized substrate concentration decreases as a result of biological adaptations. After a dramatic change in environmental factors, for example, after a sharp increase in the flow rate of organic substances, the biological system is not able to react quickly. The concentration of unutilized substrate increases and the energy flux used by the biological system decreases. The structure of the microbial community also changes, with a decrease in biological diversity. The efficiency of energy use by simple terrestrial ecosystems depends on the energetic intensity and interactions between plants and rhizospheric microorganisms.     

Accumulation and effect of volatile organic compounds in closed life support systems.

Bioregenerative life support systems (BLSS) being considered for long duration space missions will operate with limited resupply and utilize biological systems to revitalize the atmosphere, purify water, and produce food. The presence of man-made materials, plant and microbial communities, and human activities will result in the production of volatile organic compounds (VOCs). A database of VOC production from potential BLSS crops is being developed by the Breadboard Project at Kennedy Space Center. Most research to date has focused on the development of air revitalization systems that minimize the concentration of atmospheric contaminants in a closed environment. Similar approaches are being pursued in the design of atmospheric revitalization systems in bioregenerative life support systems. in a BLSS one must consider the effect of VOC concentration on the performance of plants being used for water and atmospheric purification processes. In addition to phytotoxic responses, the impact of removing biogenic compounds from the atmosphere on BLSS function needs to be assessed. This paper provides a synopsis of criteria for setting exposure limits, gives an overview of existing information, and discusses production of biogenic compounds from plants grown in the Biomass Production Chamber at Kennedy Space Center.     

Limitations on relating ocean surface chlorophyll to productivity

An important potential use of ocean color chlorophyll data is to determine other important properties of the marine biosphere, such as primary productivity, new production, and particulate fluxes at spatial scales larger and temporal scales longer than those possible with ground-based observations. Such determinations will likely progress from relatively simple empirical correlations to algorithms that are actually predictive models of ecosystem dynamics. As an exmaple, this paper demonstrates how an empirical correlation between nitrate concentration and new production can be understood by a simple productivity model. Several models are then constructed to examine the functional relationship between total production and surface chlorophyll. The empirical correlation is substantially different than the analogous relation in the model. Understanding the relationship between surface chlorophyll and productivity on a global scale will probably require families of models for various marine ecosystems.     

ECOSIMP2 model: prediction of CO2 concentration changes and carbon status in closed ecosystems.

The ECOSIMP2 model, simulating the Plant-Soil-Atmosphere interactions, was developed as a tool for the management of an experimental artificial ecosystem. It consists in three main carbon compartments for production, consumption and decomposition of the biomass. The main biological parameters concern photosynthesis (apparent Km, CO2 compensation point), the harvest index, the rate of consumption, and the kinetics of litter decomposition. From realistic assumptions of kinetics of soil compartments, a steady-state case was obtained, simulating a terrestrial ecosystem. The stability of the atmospheric CO2 concentration was studied after a virtual enclosure of the system in a 20-m high greenhouse. In natural lighting the conditions of stability are severe because of the small size of the atmospheric compartment which amplifies any imbalance between carbon fluxes. The positive consequence of that amplification for research on artificial ecosystems was emphasized.     

Effects of CO2 concentration and light intensity on photosynthesis of a rootless submerged plant, Ceratophyllum demersum L., used for aquatic food production in bioregenerative life support systems.

In addition to green microalgae, aquatic higher plants are likely to play an important role in aquatic food production modules in bioregenerative systems for producing feed for fish, converting CO2 to O2 and remedying water quality. In the present study, the effects of culture conditions on the net photosynthetic rate of a rootless submerged plant, Ceratophyllum demersum L., was investigated to determine the optimum culture conditions for maximal function of plants in food production modules including both aquatic plant culture and fish culture systems. The net photosynthetic rate in plants was determined by the increase in dissolved O2 concentrations in a closed vessel containing a plantlet and water. The water in the vessel was aerated sufficiently with a gas containing a known concentration of CO2 gas mixed with N2 gas before closing the vessel. The CO2 concentrations in the aerating gas ranged from 0.3 to 10 mmol mol-1. Photosynthetic photon flux density (PPFD) in the vessel ranged from 0 (dark) to 1.0 mmol m-2 s-1, which was controlled with a metal halide lamp. Temperature was kept at 28 degrees C. The net photosynthetic rate increased with increasing PPFD levels and was saturated at 0.2 and 0.5 mmol m-2 s-1 PPFD under CO2 levels of 1.0 and 3.0 mmol mol-1, respectively. The net photosynthetic rate increased with increasing CO2 levels from 0.3 to 3.0 mmol mol-1 showing the maximum value, 75 nmol O2 gDW-1 s-1, at 2-3 mmol mol-1 CO2 and gradually decreased with increasing CO2 levels from 3.0 to 10 mmol mol-1. The results demonstrate that C. demersum could be an efficient CO2 to O2 converter under a 2.0 mmol mol-1 CO2 level and relatively low PPFD levels in aquatic food production modules.     

Closed Artificial ecosystems as a means of ecosystem studies for Earth and space needs.

Closed Artificial ecosystems (CAES) have good prospects for wide use as new means for quantitative studies of different types of both natural ecosystems and man-made ones. The paper deals with the discussion of three points of CAES applications. The first one is of importance for theoretical ecology development and is connected with bringing together "holistic" and "merological" approaches in ecosystems studies. Using CAES, we can combine both approaches, taking into account the biotic turnover of limiting substrates which few in number even for complicated natural ecosystems. The second CAES use concerns the development of "ecosystems health" concept and application of a key-factor-approach for the indication and measurement of healthy unhealthy state and functioning of ecosystems or their links. The third use is more of an applied nature, oriented to the intensification of bioremediation or biodepollution processes in different types of ecosystems, including the global biosphere. Grant numbers: N 99-04-96017, N25.     

Examination of a smallest CELSS (microcosm) through an individual-based model simulation.

Research of the effect of space environment on an ecosystem consisting of plants and animals is essential when they are to be positively used in space. Although there have been experiments on various organisms under space environment in the past, they mainly studied the effect of space environment on an individual organism or a single species. Microcosm is drawing attention as an experimental material of an ecosystem consisting of multiple species. The object in this research is to understand the nature of this network system called ecosystem. Thus, a mixed microorganism culturing system consisting of three types of microorganisms which form a minimum food chain system as a closed ecosystem (chlorella as the producer, bacteria as the decomposer, and rotifer as the consumer) was taken for the subject, on which to research the universal characteristics of ecosystems. From the results of experiments under the terrestrial environment, formation of colonies, which is an ecological structure, has been observed at its mature stage. The organisms form an optimal substance circulation system. Therefore, formation of colonies in simulation models is important. Many attempts have been made to create ecosystem models. For example, the Lotka-Volterra model forms a simultaneous equation with the differential equation expressing predator and prey relationship and many numerical calculations have been conducted on various ecosystems based on expanded L-V models. Conventionally, these top-down methods have been used. However, since this method only describes the average concentration of organisms that are distributed uniformly throughout the system and cannot express the spatial structure of the system, it was difficult to express ecosystem structures like colonies and density distributions. In actual ecosystems, there is heterogeneity in the number of individuals and in substance density, and this is thought to have great significance in ecosystems. Consequently, an individual-based model was used that applies rules to predator-prey relationship, suppression, production, self suppression, etc., of each species. It enabled the emergence of the overall system only by its local rules, and it was possible to reproduce colony generation. In addition, the transition and the ratio of populations for each species match well with experimental results.     

A simple closed aquatic ecosystem (CAES) for space

A closed aquatic ecosystem (CAES) was developed to study the effects of microgravity on the function of closed ecosystems aboard the Chinese retrieved satellite and on the spacecraft SHENZHOU-II. These systems housed a small freshwater snail (Bulinus australianus) and an autotrophic green algae (Chlorella pyrenoidosa). The results of the test on the satellite were that the concentration of algae changed little, but that the snails died during the experiments. We then sought to optimize the function of the control system, the cultural conditions and the data acquisition system and carried out an experiment on the spacecraft SHENZHOU-II. Using various sensors to monitor the CAES, real-time data regarding the operation of the CAES in microgravity was acquired. In addition, an on-board 1g centrifuge was included to identify gravity-related factors. It was found that microgravity is the major factor affecting the operation of the CAES in space. The change in biomass of the primary producer during each day in microgravity was larger than that of the control groups. The mean biomass concentration per day in the microgravity group decreased, but that of the control groups increased for several days and then leveled off. Space effects on the biomass of a primary producer may be a result of microgravity effects leading to increasing metabolic rates of the consumer combined with decreases in photosynthesis.     

The health of biological life support systems

In developing different types of biological life support systems for use in space or extreme environments on Eart, researchers should pay attention to the long term health or functional state of such systems. The difficulty of the task is compounded by the complexity of the links and structure to be found in biological systems. To solve the problem, a hierarchical approach may be used to estimate and monitor the health of the system as a whole and its individual links. Three levels in a typical hierarchy have been considered:

1. a. the organismic.

2. b. populations and communities.

3. c. the ecosystem.

Special attention has been given to the interactions between macro- and microorganisms. Microorganisms are considered the most suitable indicators of a system's health and its component links.     

Possible nutrient limiting factor in long term operation of closed aquatic ecosystem

To investigate nutrient limitation effect on the community metabolism of closed aquatic ecosystem and possible nutrient limiting factors in the experimental food chains, depletion of inorganic chemicals including carbon, nitrogen and phosphorous was tested. A closed aquatic ecosystem lab module consisting of Chlorella pyrenoidosa and Chlamydomonas reinhardtii, Daphnia magna and associated unidentified microbes was established. Closed ecological systems receive no carbon dioxide; therefore, we presumed carbon as a first limiting factor. The results showed that the algae population in the nutrient saturated group was statistically higher than that in the nutrient limited groups, and that the chlorophyll a content of algae in the phosphorus limited group was the highest among the limited groups. However, the nitrogen limited group supported the most Daphnia, followed by the carbon limited group, the nutrient saturated group and the phosphorus limited group. Redundancy analysis showed that the total phosphorus contents were correlated significantly with the population of algae, and that the amount of soluble carbohydrate as feedback of nutrient depletion was correlated with the number of Daphnia. Thus, these findings suggest that phosphorus is the limiting factor in the operation of closed aquatic ecosystem. The results presented herein have important indications for the future construction of long term closed ecological system.     

On monitoring the bacterial component as an indicator of the state of small man-made ecosystems.

High reproduction rates make the bacterial component of ecosystems a good indicator of the state of the system on the whole. This determines the necessity to develop rapid monitoring of the functional state of the bacterial component of small ecosystems. Information about substrate concentration in the population is indicative of the state of the bacterial culture. Conventional methods of monitoring the concentration of integral substrate in the system take time much longer than the changes in the ecosystem. The paper presents theoretical foundations for the logical sequence "catalase activity--intracellular substrate concentration--estimate of substrate consumed by bacteria" for experimental verification and as a consequence of development of the integral method of monitoring the bacterial population on the basis of determining bacterial catalase activity. Grant numbers: 04-96017, N25.     

Functional, regulatory and indicator features of microorganisms in man-made ecosystems.

Functional, regulatory and indicator features of microorganisms in development and functioning of the systems and sustaining stability of three man-made ecosystem types has been studied. 1) The functional (metabolic) feature was studied in aquatic ecosystems of biological treatment of sewage waters for the reducer component. 2) The regulatory feature of bacteria for plants (producer component) was studied in simple terrestrial systems "wheat plants-rhizospheric microorganisms-artificial soil" where the behavior of the system varied with activity of the microbial component. For example with atmospheric carbon dioxide content elevated microbes promote intensification of photosynthesis processes, without binding the carbon in the plant biomass. 3) The indicator feature for the humans (consumer component) was studied in Life Support Systems (LSS). High sensitivity of human microflora to system conditions allowed its use as an indicator of the state of both system components and the entire ecosystem. Grant numbers: N99-04-96017, N15.