Energy Reallocation Research Articles

Genotypic variation in host response to infection affects parasite reproductive rate

Parasite fitness is largely influenced by a variation in host response due to the host’s genetic background. Here we investigated the impact of host genotype on pathogen success in the snail vector of its castrating parasite, Schistosoma mansoni. We infected five inbred lines of Biomphalaria glabrata with two infection doses and followed their growth, reproductive output and parasite production throughout the course of infection. There was no difference in resistance to infection among inbred lines, but lines varied in their responses to infection and the numbers of parasites produced. Snails did not compensate for castration by increasing their fecundity during the early phase of infection (fecundity compensation). However, some lines were able to delay parasite shedding for up to 30weeks, thus prolonging reproduction before the onset of castration. Here we propose this strategy as a novel defense against castrating pathogens in snails. Gigantism, a predicted outcome of castration due to energy reallocation, occurred early in infection (<15weeks) and was not universal among the snail lines. Lines that did not show gigantism were also characterised by a high parasite production rate and low survivorship, perhaps indicating energy reallocation into parasite production and costly immune defense. We observed no differences in total parasite production among lines throughout the entire course of infection, although lines differed in their parasite reproductive rate. The average rate of parasite production varied among lines from 1300 to 2450 cercariae within a single 2h shedding period, resulting in a total production of 6981–29,509 cercariae over the lifetime of a single snail. Regardless of genetic background, snail size was a strong predictor of parasite reproduction: each millimetre increase in snail size at the time of the first shed resulted in up to 3500 more cercariae over the lifetime of the snail. The results of this study provide a detailed picture of variation in hosts’ responses to infection and the resulting impacts on parasite fitness, further defining the intricacies of snail-schistosome compatibility.

Depression as sickness behavior? A test of the host defense hypothesis in a high pathogen population.

Sadness is an emotion universally recognized across cultures, suggesting it plays an important functional role in regulating human behavior. Numerous adaptive explanations of persistent sadness interfering with daily functioning (hereafter "depression") have been proposed, but most do not explain frequent bidirectional associations between depression and greater immune activation. Here we test several predictions of the host defense hypothesis, which posits that depression is part of a broader coordinated evolved response to infection or tissue injury (i.e. "sickness behavior") that promotes energy conservation and reallocation to facilitate immune activation. In a high pathogen population of lean and relatively egalitarian Bolivian forager-horticulturalists, we test whether depression and its symptoms are associated with greater baseline concentration of immune biomarkers reliably associated with depression in Western populations (i.e. tumor necrosis factor alpha [TNF-α], interleukin-1 beta [IL-1β], interleukin-6 [IL-6], and C-reactive protein [CRP]). We also test whether greater pro-inflammatory cytokine responses to ex vivo antigen stimulation are associated with depression and its symptoms, which is expected if depression facilitates immune activation. These predictions are largely supported in a sample of older adult Tsimane (mean±SD age=53.2±11.0, range=34-85, n=649) after adjusting for potential confounders. Emotional, cognitive and somatic symptoms of depression are each associated with greater immune activation, both at baseline and in response to ex vivo stimulation. The association between depression and greater immune activation is therefore not unique to Western populations. While our findings are not predicted by other adaptive hypotheses of depression, they are not incompatible with those hypotheses and future research is necessary to isolate and test competing predictions.

Energy Reallocation in a Multi-User Network With a Shared Harvesting Module and Storage Battery

This work considers a novel multi-user system where transmitter nodes have shared access to a common energy harvesting (EH) module and storage battery. Users do not possess individual EH modules and can only withdraw energy from the shared battery which has a randomly varying energy level. An auction-based centralized energy allocation mechanism is proposed with the objective of maximizing the system social welfare. It is shown analytically that the mechanism converges to a unique maximum, is Pareto optimal, and truthful. Simulations show that overall, it outperforms uniform and incremental energy allocation for a range of scenarios.

Age-dependent survival and selected gene expression in Daphnia magna after short-term exposure to low dissolved oxygen

Accumulation and breakdown of large algal blooms often causes serious hypoxia in lakes, representing a major potential threat to zooplankton grazers such as Daphnia. Generally, populations comprise different age groups, whose molecular responses to environmental stressors are different. The aim of our study was to investigate the age-dependent effects of oxygen depletion on D. magna survival and selected gene expressions of energy-relevance and stress-defense. Four age groups of D. magna (1, 4, 7 and 14 days) were exposed to 0.0625, 0.125, 0.25 mM dissolved oxygen (DO) for 48 h. Results showed that survival rates of 14-day-old individuals in 0.0625 mM DO and 0.125 mM DO were much lower than those in 0.25 mM DO, while no significant difference was found in remaining age groups. Low DO up-regulated hemoglobin expression compared with the controls in all age groups; the similar pattern in a-esterase was observed when 1-day-old individuals were exposed to 0.0625 mM, 4and 7-day-old individuals at 0.125 mM, respectively. HSP70 and CAT expression were significantly upregulated in 1and 7-day-old individuals exposed to 0.125 mM. The findings indicate that age structure should be taken into account when interpreting dynamics of Daphnia populations affected by oxygen depletion. Age may modify energy reallocation and alter susceptibility of the organisms.

Phytoplankton strategies for photosynthetic energy allocation.

Phytoplankton physiology is dynamic and highly responsive to the environment. Phytoplankton acclimate to changing environmental conditions by a complex reallocation of carbon and energy through metabolic pathways to optimize growth. Considering the tremendous diversity of phytoplankton, it is not surprising that different phytoplankton taxa use different strategies to partition carbon and energy resources. It has therefore been satisfying to discover that general principles of energetic stoichiometry appear to govern these complex processes and can be broadly applied to interpret phytoplankton distributions, productivity, and food web dynamics. The expectation of future changes in aquatic environments brought on by climate change warrants gathering knowledge about underlying patterns of photosynthetic energy allocation and their impacts on community structure and ecosystem productivity.

From receptor balance to rational glucocorticoid therapy.

Corticosteroids secreted as end product of the hypothalamic-pituitary-adrenal axis act like a double-edged sword in the brain. The hormones coordinate appraisal processes and decision making during the initial phase of a stressful experience and promote subsequently cognitive performance underlying the management of stress adaptation. This action exerted by the steroids on the initiation and termination of the stress response is mediated by 2 related receptor systems: mineralocorticoid receptors (MRs) and glucocorticoid receptors (GRs). The receptor types are unevenly distributed but colocalized in abundance in neurons of the limbic brain to enable these complementary hormone actions. This contribution starts from a historical perspective with the observation that phasic occupancy of GR during ultradian rhythmicity is needed to maintain responsiveness to corticosteroids. Then, during stress, initially MR activation enhances excitability of limbic networks that are engaged in appraisal and emotion regulation. Next, the rising hormone concentration occupies GR, resulting in reallocation of energy to limbic-cortical circuits with a role in behavioral adaptation and memory storage. Upon MR:GR imbalance, dysregulation of the hypothalamic-pituitary-adrenal axis occurs, which can enhance an individual's vulnerability. Imbalance is characteristic for chronic stress experience and depression but also occurs during exposure to synthetic glucocorticoids. Hence, glucocorticoid psychopathology may develop in susceptible individuals because of suppression of ultradian/circadian rhythmicity and depletion of endogenous corticosterone from brain MR. This knowledge generated from testing the balance hypothesis can be translated to a rational glucocorticoid therapy.

Combined effects of different CO2 levels and N sources on the diazotrophic cyanobacterium Trichodesmium.

To predict effects of climate change and possible feedbacks, it is crucial to understand the mechanisms behind CO2 responses of biogeochemically relevant phytoplankton species. Previous experiments on the abundant N2 fixers Trichodesmium demonstrated strong CO2 responses, which were attributed to an energy reallocation between its carbon (C) and nitrogen (N) acquisition. Pursuing this hypothesis, we manipulated the cellular energy budget by growing Trichodesmium erythraeum IMS101 under different CO2 partial pressure (pCO2) levels (180, 380, 980 and 1400 µatm) and N sources (N2 and NO3−). Subsequently, biomass production and the main energy-generating processes (photosynthesis and respiration) and energy-consuming processes (N2 fixation and C acquisition) were measured. While oxygen fluxes and chlorophyll fluorescence indicated that energy generation and its diurnal cycle was neither affected by pCO2 nor N source, cells differed in production rates and composition. Elevated pCO2 increased N2 fixation and organic C and N contents. The degree of stimulation was higher for nitrogenase activity than for cell contents, indicating a pCO2 effect on the transfer efficiency from N2 to biomass. pCO2-dependent changes in the diurnal cycle of N2 fixation correlated well with C affinities, confirming the interactions between N and C acquisition. Regarding effects of the N source, production rates were enhanced in NO3− grown cells, which we attribute to the higher N retention and lower ATP demand compared with N2 fixation. pCO2 effects on C affinity were less pronounced in NO3− users than N2 fixers. Our study illustrates the necessity to understand energy budgets and fluxes under different environmental conditions for explaining indirect effects of rising pCO2.

Trichodesmium’s strategies to alleviate phosphorus limitation in the future acidified oceans.

Global warming may exacerbate inorganic nutrient limitation, including phosphorus (P), in the surface waters of tropical oceans that are home to extensive blooms of the marine diazotrophic cyanobacterium, Trichodesmium. We examined the combined effects of P limitation and pCO(2), forecast under ocean acidification scenarios, on Trichodesmium erythraeum IMS101 cultures. We measured nitrogen acquisition,glutamine synthetase activity, C uptake rates, intracellular Adenosine Triphosphate (ATP) concentration and the pool sizes of related key proteins. Here, we present data supporting the idea that cellular energy re-allocation enables the higher growth and N(2) fixation rates detected in Trichodesmium cultured under high pCO(2). This is reflected in altered protein abundance and metabolic pools. Also modified are particulate organic carbon and nitrogen production rates,enzymatic activities, and cellular ATP concentrations. We suggest that adjusting these cellular pathways to changing environmental conditions enables Trichodesmium to compensate for low P availability and to thrive in acidified oceans. Moreover, elevated pCO(2) could provide Trichodesmium with a competitive dominance that would extend its niche, particularly in P-limited regions of the tropical and subtropical oceans.

Energy Reallocation to Breeding Performance through Improved Nest Building in Laboratory Mice

Mice are housed at temperatures (20-26°C) that increase their basal metabolic rates and impose high energy demands to maintain core temperatures. Therefore, energy must be reallocated from other biological processes to increase heat production to offset heat loss. Supplying laboratory mice with nesting material may provide sufficient insulation to reduce heat loss and improve both feed conversion and breeding performance. Naïve C57BL/6, BALB/c, and CD-1breeding pairs were provided with bedding alone, or bedding supplemented with either 8g of Enviro-Dri, 8g of Nestlets, for 6 months. Mice provided with either nesting material built more dome-like nests than controls. Nesting material improved feed efficiency per pup weaned as well as pup weaning weight. The breeding index (pups weaned/dam/week) was higher when either nesting material was provided. Thus, the sparing of energy for thermoregulation of mice given additional nesting material may have been responsible for the improved breeding and growth of offspring.

Single and combined effects of cadmium and arsenate in Gammarus pulex (Crustacea, Amphipoda): Understanding the links between physiological and behavioural responses

This study aimed at investigating the individual and interactive effects of cadmium (Cd) and arsenate (AsV) in Gammarus pulex (Crustacea, Amphipoda) through the use of several biomarkers. Individuals were exposed for 240h to two concentrations of AsV or Cd alone, and all the possible binary mixtures of these concentrations of AsV and Cd in a complete factorial design. The pattern of the biomarkers’ responses to Cd and AsV alone or in mixture was similar in Gammarus pulex, even if the response intensity varied depending on the tested conditions. G. pulex responded to contamination with increased mobilization of the detoxification systems [i.e. γ-glutamyl-cystein ligase activity (GCL), reduced glutathione content (GSH) and metallothionein concentrations (MT)]. This response seems to imply changes in energy reserve utilization (total lipids and proteins are used prior to glycogen reserves), but also a possible energy reallocation from locomotion to detoxification processes. The observed increase in lipid peroxidation could be relied to the increasing gammarid mortality, despite the higher mobilization of detoxification systems. Even if the outcome of the complex interactions between AsV and Cd remains difficult to unravel, such studies are critically important for better assessing the effects of stressors on organisms, populations and communities in a multi-contamination context of ecosystems.

Time Course and Metabolic Costs of a Humoral Immune Response in the Little Ringed Plover Charadrius dubius

Despite host defense against parasites and pathogens being considered a costly life-history trait, relatively few studies have assessed the energetic cost of immune responsiveness. Knowledge of such energetic costs may help to understand the mechanisms by which trade-offs with other demanding activities occur. The time course and associated metabolic costs of mounting a primary and secondary humoral immune response was examined in little ringed plovers Charadrius dubius challenged with sheep red blood cells. As was expected, the injection with this antigen increased the production of specific antibodies significantly, with peaks 6 d postinjection in both primary and secondary responses. At the peak of secondary antibody response, the antibody production was 29% higher than that observed during the primary response, but the difference was nonsignificant. Mounting the primary response did not significantly increase the resting metabolic rate (RMR) of birds, whereas the secondary response did by 21%, suggesting that the latter was more costly in terms of RMR. In spite of the fact that the primary response did not involve an increase in RMR, birds significantly decreased their body mass. This could imply an internal energy reallocation strategy to cope with the induced immune challenge. Last, we found that RMR and antibody production peaks were not coupled, which could help to conciliate the variable results of previous studies. Collectively, the results of this study support the hypothesis that humoral immunity, especially the secondary response, entails energetic costs that may trade-off with other physiological activities.

An adaptive post-filtering method producing an artificial Lombard-like effect for intelligibility enhancement of narrowband telephone speech

Post-filtering can be used in mobile communications to improve the quality and intelligibility of speech. Energy reallocation with a high-pass type filter has been shown to work effectively in improving the intelligibility of speech in difficult noise conditions. This paper introduces a post-filtering algorithm that adapts to the background noise level as well as to the fundamental frequency of the speaker and models the spectral effects observed in natural Lombard speech. The introduced method and another post-filtering technique were compared to unprocessed telephone speech in subjective listening tests in terms of intelligibility and quality. The results indicate that the proposed method outperforms the reference method in difficult noise conditions.

Genes related to ion-transport and energy production are upregulated in response to CO2-driven pH decrease in corals: new insights from transcriptome analysis.

Since the preindustrial era, the average surface ocean pH has declined by 0.1 pH units and is predicted to decline by an additional 0.3 units by the year 2100. Although subtle, this decreasing pH has profound effects on the seawater saturation state of carbonate minerals and is thus predicted to impact on calcifying organisms. Among these are the scleractinian corals, which are the main builders of tropical coral reefs. Several recent studies have evaluated the physiological impact of low pH, particularly in relation to coral growth and calcification. However, very few studies have focused on the impact of low pH at the global molecular level. In this context we investigated global transcriptomic modifications in a scleractinian coral (Pocillopora damicornis) exposed to pH 7.4 compared to pH 8.1during a 3-week period. The RNAseq approach shows that 16% of our transcriptome was affected by the treatment with 6% of upregulations and 10% of downregulations. A more detailed analysis suggests that the downregulations are less coordinated than the upregulations and allowed the identification of several biological functions of interest. In order to better understand the links between these functions and the pH, transcript abundance of 48 candidate genes was quantified by q-RT-PCR (corals exposed at pH 7.2 and 7.8 for 3 weeks). The combined results of these two approaches suggest that pH≥7.4 induces an upregulation of genes coding for proteins involved in calcium and carbonate transport, conversion of CO2 into HCO3 − and organic matrix that may sustain calcification. Concomitantly, genes coding for heterotrophic and autotrophic related proteins are upregulated. This can reflect that low pH may increase the coral energy requirements, leading to an increase of energetic metabolism with the mobilization of energy reserves. In addition, the uncoordinated downregulations measured can reflect a general trade-off mechanism that may enable energy reallocation.

Oxygen and Heterotrophy Affect Calcification of the Scleractinian Coral Galaxea fascicularis

Heterotrophy is known to stimulate calcification of scleractinian corals, possibly through enhanced organic matrix synthesis and photosynthesis, and increased supply of metabolic DIC. In contrast to the positive long-term effects of heterotrophy, inhibition of calcification has been observed during feeding, which may be explained by a temporal oxygen limitation in coral tissue. To test this hypothesis, we measured the short-term effects of zooplankton feeding on light and dark calcification rates of the scleractinian coral Galaxea fascicularis (n = 4) at oxygen saturation levels ranging from 13 to 280%. Significant main and interactive effects of oxygen, heterotrophy and light on calcification rates were found (three-way factorial repeated measures ANOVA, p<0.05). Light and dark calcification rates of unfed corals were severely affected by hypoxia and hyperoxia, with optimal rates at 110% saturation. Light calcification rates of fed corals exhibited a similar trend, with highest rates at 150% saturation. In contrast, dark calcification rates of fed corals were close to zero under all oxygen saturations. We conclude that oxygen exerts a strong control over light and dark calcification rates of corals, and propose that in situ calcification rates are highly dynamic. Nevertheless, the inhibitory effect of heterotrophy on dark calcification appears to be oxygen-independent. We hypothesize that dark calcification is impaired during zooplankton feeding by a temporal decrease of the pH and aragonite saturation state of the calcifying medium, caused by increased respiration rates. This may invoke a transient reallocation of metabolic energy to soft tissue growth and organic matrix synthesis. These insights enhance our understanding of how oxygen and heterotrophy affect coral calcification, both in situ as well as in aquaculture.

Signal-to-noise ratio adaptive post-filtering method for intelligibility enhancement of telephone speech

Post-filtering can be utilized to improve the quality and intelligibility of telephone speech. Previous studies have shown that energy reallocation with a high-pass type filter works effectively in improving the intelligibility of speech in difficult noise conditions. The present study introduces a signal-to-noise ratio adaptive post-filtering method that utilizes energy reallocation to transfer energy from the first formant to higher frequencies. The proposed method adapts to the level of the background noise so that, in favorable noise conditions, the post-filter has a flat frequency response and the effect of the post-filtering is increased as the level of the ambient noise increases. The performance of the proposed method is compared with a similar post-filtering algorithm and unprocessed speech in subjective listening tests which evaluate both intelligibility and listener preference. The results indicate that both of the post-filtering methods maintain the quality of speech in negligible noise conditions and are able to provide intelligibility improvement over unprocessed speech in adverse noise conditions. Furthermore, the proposed post-filtering algorithm performs better than the other post-filtering method under evaluation in moderate to difficult noise conditions, where intelligibility improvement is mostly required.

Growth and Reproductive Performance of Triploid Yesso Scallops (Patinopecten yessoensis) Induced by Hypotonic Shock

The successful induction of triploid embryos or larvae has been performed in Patinopecten yessoensis during the past 2 decades. However, no research has been reported about the performance of triploid P. yessoensis cultured in the field. This study induced triploidy in P. yessoensis by hypotonic shock and compared the growth and reproductive performance of triploids and diploids reared under commercial conditions for up to 24 mo. The main results of this study are as follows: Triploid scallops were smaller in size and weight compared with diploids and had a retarded absolute growth rate (AGR). After 24 mo of cultivation, the mean shell height, shell length, shell width, and body weight of triploids were 9%, 10%, 9%, and 25% less than diploids, respectively (P < 0.01). Although normal in sex ratio, the reproductive potential of triploids was significantly reduced. Only 87% of the triploids exhibited sex-discernible gonads during the breeding season. None of the male triploids spawned, and the percentage of female spawners among the triploid population was only 27% of that for the diploid population. The relative fecundity of triploid females was only 4% of diploid females. Triploid eggs produced mostly aneuploid larvae and had an extremely small chance of generating viable offspring when fertilized by sperm from diploid males. Most aneuploid larvae died before the D-shaped stage, and no survival exceeded 7 days. The potential to yield viable offspring from the triploid population was estimated to be only 4% × 10-6 of that of the diploid population. Despite the growth disadvantage of triploids, this study may support, in part, the energy reallocation hypothesis, because triploid AGR was similar to diploid AGR (2% variance) during the sexual maturation season. Our results also indicate that there would be no growth advantage, but instead a disadvantage, for triploid P. yessoensis growing at the experiment site. In addition, this research provides the first evidence that viable triploid molluscs can be induced by hypotonic shock, of which the practical and evolutionary implications are also discussed.

Role of peripheral nerve fibres in acute and chronic inflammation in arthritis

The peripheral nervous system takes an active part in inflammatory processes by regulating effector cell function and reallocation of energy to the immune system. During acute inflammation, rapid neuronal reorganization and change of activity takes place. The hallmarks of this process are an increase in systemic sympathetic activity, a decrease in systemic parasympathetic activity and loss of sympathetic nerve fibres from sites of inflammation concomitant with increased innervation with sensory nerve fibres and increased sensory nerve fibre activity. On a systemic level, the increase in sympathetic activity (and decrease in parasympathetic activity) is necessary to provide enough energy to nourish the activated immune system. In locally inflamed tissue, the decrease in sympathetic nerve fibre density results in reduced anti-inflammatory signalling and, together with neuropeptides released from sensory nerve fibres, promotes local inflammation. In acute inflammation, this 'inflammatory configuration' of the peripheral nervous system favours the rapid clearance of antigenic threats. However, in chronic autoimmune inflammation, these changes of the peripheral nervous system lead to an unfavourable situation with ongoing energy reallocation and continuous local destruction. As an example of a chronic inflammatory condition, we discuss evidence for neuroimmune regulation in autoimmune arthritis with a focus on the sympathetic nervous system.

Retrofit of steam power plants in a steel mill

This article presents a systematic methodology for the analysis and design of steam power plants in a typical steel mill. Therein most of the steam is produced by applying synthesis gas, a side-product from the coke processing plants. This offers the opportunity of energy integration with vicinal companies using fuel oil or natural gas for generating steam. Components of this work include the energy reallocation and the retrofitted design of steam systems. In this study, the current strategy of energy utilization for existing sites in a steel mill is first assessed. In this regard, the energy policy is adjusted in order to enhance the performance and reduce the costs of the steam power plants. Thereafter, the retrofit is taken into account to evaluate the potential energy for the integration. The problems are formulated as a mixed integer nonlinear program based on a superstructure approach. The results of an industrial case study show that the energy integration among plants is benefit, which provides incentive to promote the cooperation of neighboring companies in an eco-industrial park.

Temperature-dependent consumer-resource dynamics: A coupled structured model for Gammarus pulex (L.) and leaf litter

The annual variation in population density of the stream macroinvertebrate Gammarus pulex is frequently linked to annual courses of both, resource availability and temperature, which is calling for a mechanistic analysis of the system. To this effect, we coupled a size structured population model for the consumer and a quality structured resource model for its main resource, leaf litter standing stock. We introduced temperature dependence of processes and a lumped internal energy storage for reproduction and energy reallocation. One-year's field data on body length frequencies of G. pulex, in-stream leaf litter standing stock, leaf fall and temperature were used to calibrate the model for parameters not yet determined in independent experiments. The model reproduced the seasonal pattern of both dynamics acceptably. Independently measured feeding rates of G. pulex were used to confirm the simulated relationship between resource and consumer. Such, we revealed the contribution of temperature and food response to changes in population density. Temperature effects of mortality and reproduction roughly concur with each other, while the energy response of reproduction is pivotally distinct. In the face of model application for projection of climate change effects, further attention should particularly be paid to the specific temperature response of G. pulex and to the contribution of non-shredding decomposition (physical abrasion and microbial colonization) to loss of leaf litter standing stock.

Assessing Coral Reefs on a Pacific-Wide Scale Using the Microbialization Score

The majority of the world's coral reefs are in various stages of decline. While a suite of disturbances (overfishing, eutrophication, and global climate change) have been identified, the mechanism(s) of reef system decline remain elusive. Increased microbial and viral loading with higher percentages of opportunistic and specific microbial pathogens have been identified as potentially unifying features of coral reefs in decline. Due to their relative size and high per cell activity, a small change in microbial biomass may signal a large reallocation of available energy in an ecosystem; that is the microbialization of the coral reef. Our hypothesis was that human activities alter the energy budget of the reef system, specifically by altering the allocation of metabolic energy between microbes and macrobes. To determine if this is occurring on a regional scale, we calculated the basal metabolic rates for the fish and microbial communities at 99 sites on twenty-nine coral islands throughout the Pacific Ocean using previously established scaling relationships. From these metabolic rate predictions, we derived a new metric for assessing and comparing reef health called the microbialization score. The microbialization score represents the percentage of the combined fish and microbial predicted metabolic rate that is microbial. Our results demonstrate a strong positive correlation between reef microbialization scores and human impact. In contrast, microbialization scores did not significantly correlate with ocean net primary production, local chla concentrations, or the combined metabolic rate of the fish and microbial communities. These findings support the hypothesis that human activities are shifting energy to the microbes, at the expense of the macrobes. Regardless of oceanographic context, the microbialization score is a powerful metric for assessing the level of human impact a reef system is experiencing.