Effects of an Ecosystem Engineer on Belowground Movement of Microarthropods

Ecosystem engineers affect other species by changing physical environments. Such changes may influence movement of organisms, particularly belowground where soil permeability can restrict dispersal. We investigated whether earthworms, iconic ecosystem engineers, influence microarthropod movement. Our experiment tested whether movement is affected by tunnels (i.e., burrows), earthworm excreta (mucus, castings), or earthworms themselves. Earthworm burrows form tunnel networks that may facilitate movement. This effect may be enhanced by excreta, which could provide resources for microarthropods moving along the network. Earthworms may also promote movement via phoresy. Conversely, negative effects could occur if earthworms alter predator-prey relationships or change competitive interactions between microarthropods. We used microcosms consisting of a box connecting a “source” container in which microarthropods were present and a “destination” container filled with autoclaved soil. Treatments were set up within the boxes, which also contained autoclaved soil, as follows: 1) control with no burrows; 2) artificial burrows with no excreta; 3) abandoned burrows with excreta but no earthworms; and 4) earthworms (Lumbricus rubellus) present in burrows. Half of the replicates were sampled once after eight days, while the other half were sampled repeatedly to examine movement over time. Rather than performing classical pairwise comparisons to test our hypotheses, we used AIC_c to assess support for three competing models (presence of tunnels, excreta, and earthworms). More individuals of Collembola, Mesostigmata, and all microarthropods together dispersed when tunnels were present. Models that included excreta and earthworms were less well supported. Total numbers of dispersing Oribatida and Prostigmata+Astigmata were not well explained by any models tested. Further research is needed to examine the impact of soil structure and ecosystem engineering on movement belowground, as the substantial increase in movement of some microarthropods when corridors were present suggests these factors can strongly affect colonization and community assembly.     

Shining Light on Benthic Macroalgae: Mechanisms of Complementarity in Layered Macroalgal Assemblages

Phototrophs underpin most ecosystem processes, but to do this they need sufficient light. This critical resource, however, is compromised along many marine shores by increased loads of sediments and nutrients from degraded inland habitats. Increased attenuation of total irradiance within coastal water columns due to turbidity is known to reduce species'' depth limits and affect the taxonomic structure and architecture of algal-dominated assemblages, but virtually no attention has been paid to the potential for changes in spectral quality of light energy to impact production dynamics. Pioneering studies over 70 years ago showed how different pigmentation of red, green and brown algae affected absorption spectra, action spectra, and photosynthetic efficiency across the PAR (photosynthetically active radiation) spectrum. Little of this, however, has found its way into ecological syntheses of the impacts of optically active contaminants on coastal macroalgal communities. Here we test the ability of macroalgal assemblages composed of multiple functional groups (including representatives from the chlorophyta, rhodophyta and phaeophyta) to use the total light resource, including different light wavelengths and examine the effects of suspended sediments on the penetration and spectral quality of light in coastal waters. We show that assemblages composed of multiple functional groups are better able to use light throughout the PAR spectrum. Macroalgal assemblages with four sub-canopy species were between 50–75% more productive than assemblages with only one or two sub-canopy species. Furthermore, attenuation of the PAR spectrum showed both a loss of quanta and a shift in spectral distribution with depth across coastal waters of different clarity, with consequences to productivity dynamics of diverse layered assemblages. The processes of light complementarity may help provide a mechanistic understanding of how altered turbidity affects macroalgal assemblages in coastal waters, which are increasingly threatened by diminishing light quantity and altered spectral distributions through sedimentation and eutrophication.     

An Inventory of Marine Benthic Macroalgae of Hainan Island,China

In this review, the results of the study of marine benthic algal flora of Hainan Island, China, during the period from 1933 to 2015 are considered. It has been found that the local floristic diversity and taxonomic composition are similar to those of macroalgae from other large islands of the Indo-Pacific, where coral reefs are the main ecosystem of the shallow-water zone. An analysis of historical (decadal) changes in the marine benthic flora of Hainan has shown the following pattern of variations in the species diversity of macroalgae since the middle of the 20th century: a decline in low-productive epilithic algae with voluminous forms of thalli and an increase in highly productive opportunistic species with filamentous, fine filamentous, and membranous forms. These historical changes are thought to be caused by human impacts: inappropriate exploitation of coral reefs and eutrophication of coastal waters due to household sewage from large cities and discharge of mariculture wastes.     

Effects of Consumer Interactions on Benthic Resources and Ecosystem Processes in a Neotropical Stream

The effect of consumers on their resources has been demonstrated in many systems but is often confounded by trophic interactions with other consumers. Consumers may also have behavioral and life history adaptations to each other and to co-occurring predators that may additionally modulate their particular roles in ecosystems. We experimentally excluded large consumers from tile periphyton, leaves and natural benthic substrata using submerged electrified frames in three stream reaches with overlapping consumer assemblages in Trinidad, West Indies. Concurrently, we assessed visits to (non-electrified) control frames by the three most common large consumers–primarily insectivorous killifish (Rivulus hartii), omnivorous guppies (Poecilia reticulata) and omnivorous crabs (Eudaniela garmani). Consumers caused the greatest decrease in final chlorophyll a biomass and accrual rates the most in the downstream reach containing all three focal consumers in the presence of fish predators. Consumers also caused the greatest increase in leaf decay rates in the upstream reach containing only killifish and crabs. In the downstream reach where guppies co-occur with predators, we found significantly lower benthic invertebrate biomass in control relative to exclosure treatments than the midstream reach where guppies occur in the absence of predators. These data suggest that differences in guppy foraging, potentially driven by differences in their life history phenotype, may affect ecosystem structure and processes as much as their presence or absence and that interactions among consumers may further mediate their effects in these stream ecosystems.     

Microbial Response of a Freshwater Benthic Community to a Simulated Diatom Sedimentation Event: Interactive Effects of Benthic Fauna

Abstract The response of a sediment microbial assemblage to a pulse of diatoms was studied over 36 days by measuring bacterial activity and biomass, ATP concentration, and overall community respiration in laboratory microcosms. Also, the contribution of macrofaunal chironomids to the decomposition of settling diatoms in benthic communities, and the relative importance of benthic meiofauna in community metabolism, were determined. The addition of diatoms resulted in an immediate response by sediment bacteria, with higher bacterial production recorded after only 2 h, and a more than tenfold increase within one day. The rapid response by sediment bacteria was accompanied by relatively high initial concentrations of dissolved organic carbon. In treatments receiving diatoms, higher bacterial production was sustained throughout the experiment. Surprisingly, neither these elevated production estimates, nor the starvation of controls affected bacterial abundance. Mean bacterial cell volume, however, was markedly affected by the addition of diatoms. Combining community respiration measurements and bacterial production estimates showed that growth efficiencies for sediment bacteria ranged from 14.6 to 34.5%. The contribution of ambient meiozoobenthos to carbon metabolism was less than 1%. Carbon budgets showed that 1.3 mg C was cooxidized along with 4.3 mg added diatom C. Sediment reworking by Chironomus larvae initially enhanced bacterial production, but the presence of Chironomus resulted in lower bacterial production estimates after 16 and 36 days. This was interpreted as a result of faster decomposition of diatoms in treatments with chironomids, which was validated by a faster decline of ATP and chlorophyll a in the sediment. Our results indicate that Chironomus larvae compete with sediment bacteria for available organic substrates. Received: 11 June 1996; Accepted: 13 August 1996     

Consequences of Increasing Hypoxic Disturbance on Benthic Communities and Ecosystem Functioning

Disturbance-mediated species loss has prompted research considering how ecosystem functions are changed when biota is impaired. However, there is still limited empirical evidence from natural environments evaluating the direct and indirect (i.e. via biota) effects of disturbance on ecosystem functioning. Oxygen deficiency is a widespread threat to coastal and estuarine communities. While the negative impacts of hypoxia on benthic communities are well known, few studies have assessed in situ how benthic communities subjected to different degrees of hypoxic stress alter their contribution to ecosystem functioning. We studied changes in sediment ecosystem function (i.e. oxygen and nutrient fluxes across the sediment water-interface) by artificially inducing hypoxia of different durations (0, 3, 7 and 48 days) in a subtidal sandy habitat. Benthic chamber incubations were used for measuring responses in sediment oxygen and nutrient fluxes. Changes in benthic species richness, structure and traits were quantified, while stress-induced behavioral changes were documented by observing bivalve reburial rates. The initial change in faunal behavior was followed by non-linear degradation in benthic parameters (abundance, biomass, bioturbation potential), gradually impairing the structural and functional composition of the benthic community. In terms of ecosystem function, the increasing duration of hypoxia altered sediment oxygen consumption and enhanced sediment effluxes of NH₄ ⁺ and dissolved Si. Although effluxes of PO₄ ³⁻ were not altered significantly, changes were observed in sediment PO₄ ³⁻ sorption capability. The duration of hypoxia (i.e. number of days of stress) explained a minor part of the changes in ecosystem function. Instead, the benthic community and disturbance-driven changes within the benthos explained a larger proportion of the variability in sediment oxygen- and nutrient fluxes. Our results emphasize that the level of stress to the benthic habitat matters, and that the link between biodiversity and ecosystem function is likely to be affected by a range of factors in complex, natural environments.     

Complexities in Understanding Ecosystem Response to Ozone

Ecological risk assessment of O₃ impact requires consideration of many factors that, perhaps, are not of concern in human health risk assessment. The episodic nature of O₃ exposure, functional complexity of species assemblages, and the broad spatial and temporal scales characteristic of natural ecosystems make ecological risk assessment extremely difficult. The majority of exposure studies using plants have examined the sensitivity of individual species, growing under controlled conditions. Research has shown that individuals growing in plant mixtures may not respond the same way to O₃ as when growing alone. In addition, other naturally occurring stresses can modify plant response to O₃. Understanding the effect of O₃ on natural systems and protecting vegetation resources represent significant scientific and regulatory challenges. Here we review several factors that need to be considered when evaluating ecosystem response to O₃. Then we briefly present two examples of controlled seedling studies that were conducted to better understand mechanisms of tree response to O₃. In the first example controlled exposure studies revealed responses in tree roots that led to hypothesis testing in the field in ponderosa pine ecosystems. Field experiments have confirmed a similar response in root biomass and carbohydrates across a natural O₃ gradient in S. California, suggesting at least a partial role for O₃ in the response. The second example illustrates the difficulty of understanding mechanistic interactions to O₃ stress even in controlled chamber studies. The second example also illustrates the difficulty of using chamber studies to understand responses in the field. While our knowledge of vegetation response to O₃ is extensive and compelling, important questions remain about how to quantify these effects in the field, assess their magnitude, and establish a suitable standard that is protective of ecosystems.     

Effects of Obesity Phenotype on Coronary Heart Disease Risk Factors in Response to Weight Loss

Objective: To determine whether there is a difference in risk‐factor improvement for coronary heart disease (CHD) between the intra‐abdominal fat (IF) and subcutaneous fat (SF) obesity phenotypes after weight loss. Research Methods and Procedures: Subjects included 55 mildly obese women (body mass index, 25 to 36 kg/m²; age range, 34 to 63 years) who had at least two of three CHD risk factors [systolic blood pressure (SBP), >140 mm Hg; total cholesterol (TC), >220 mg/dL; fasting plasma glucose, >110 mg/dL). Using computed tomography, IF obesity was classified as ≥110 cm² of the IF area measured; subjects with <110 cm² were classified as having SF obesity. The IF and SF obesity groups were divided into diet‐only and diet‐plus‐exercise groups. Assays and measurements were performed before and after a 14‐week (98‐day) intervention. Results: Weight was reduced by 7 to 10 kg in each group. The IF and SF areas, SBP, diastolic blood pressure, TC, and low‐density lipoprotein‐cholesterol were significantly reduced in all groups (p < 0.01). Reduction in IF area was greater in IF obesity than in SF obesity, whereas no differences were observed in the improvement of CHD risk factors. Sample sizes needed for observing a significant difference for SBP, TC, triglycerides, and fasting plasma glucose were greater than the number of subjects in this study. Discussion: Our results suggest that the influence of the obesity phenotype on improving CHD risk factors is not apparent. A larger study is needed to prove the validity of this finding.     

River Ecosystem Response to Prescribed Vegetation Burning on Blanket peatland

Catchment-scale land-use change is recognised as a major threat to aquatic biodiversity and ecosystem functioning globally. In the UK uplands rotational vegetation burning is practised widely to boost production of recreational game birds, and while some recent studies have suggested burning can alter river water quality there has been minimal attention paid to effects on aquatic biota. We studied ten rivers across the north of England between March 2010 and October 2011, five of which drained burned catchments and five from unburned catchments. There were significant effects of burning, season and their interaction on river macroinvertebrate communities, with rivers draining burned catchments having significantly lower taxonomic richness and Simpson’s diversity. ANOSIM revealed a significant effect of burning on macroinvertebrate community composition, with typically reduced Ephemeroptera abundance and diversity and greater abundance of Chironomidae and Nemouridae. Grazer and collector-gatherer feeding groups were also significantly less abundant in rivers draining burned catchments. These biotic changes were associated with lower pH and higher Si, Mn, Fe and Al in burned systems. Vegetation burning on peatland therefore has effects beyond the terrestrial part of the system where the management intervention is being practiced. Similar responses of river macroinvertebrate communities have been observed in peatlands disturbed by forestry activity across northern Europe. Finally we found river ecosystem changes similar to those observed in studies of wild and prescribed forest fires across North America and South Africa, illustrating some potentially generic effects of fire on aquatic ecosystems.     

Marine Ecosystem Response to the Atlantic Multidecadal Oscillation

Against the backdrop of warming of the Northern Hemisphere it has recently been acknowledged that North Atlantic temperature changes undergo considerable variability over multidecadal periods. The leading component of natural low-frequency temperature variability has been termed the Atlantic Multidecadal Oscillation (AMO). Presently, correlative studies on the biological impact of the AMO on marine ecosystems over the duration of a whole AMO cycle (∼60 years) is largely unknown due to the rarity of continuously sustained biological observations at the same time period. To test whether there is multidecadal cyclic behaviour in biological time-series in the North Atlantic we used one of the world''s longest continuously sustained marine biological time-series in oceanic waters, long-term fisheries data and historical records over the last century and beyond. Our findings suggest that the AMO is far from a trivial presence against the backdrop of continued temperature warming in the North Atlantic and accounts for the second most important macro-trend in North Atlantic plankton records; responsible for habitat switching (abrupt ecosystem/regime shifts) over multidecadal scales and influences the fortunes of various fisheries over many centuries.     

Holocene Carbon Accumulation of Fen Peatlands in Boreal Western Canada: A Complex Ecosystem Response to Climate Variation and Disturbance

Understanding the long-term ecological dynamics of northern peatlands is essential for assessment of the possible responses and feedbacks of these carbon-rich ecosystems to climate change and natural disturbance. I used high-resolution macrofossil and lithological analyses of a fen peatland in western Canada to infer the Holocene developmental history of the peatland, to document the temporal pattern of long-term peat accumulation, and to investigate ecosystems responses to climate changes in terms of species composition and carbon accumulation. The peatland has been dominated by sedges and brown mosses during its 10,000-year history, despite interruption by tephra deposition. Peat accumulation rates vary by more than an order of magnitude and decline from 5500 to 1300 cal BP, resulting in a convex depth–age curve, which contrasts with the carbon accumulation patterns documented for oceanic peatlands. The synthesis of regional data from continental western Canada indicates that fens tend to accumulate more carbon than bogs of the same ages. These data suggest that the carbon sink potential of northern peatlands has varied dramatically in the past, so estimates of the present and projected carbon sink strengths of these peatlands need to take this temporal variation into consideration. Widespread slowdown of peat accumulation over the last 4000 years may have resulted from climate cooling in northern latitudes after the Holocene insolation maximum. The findings indicate that long-term peatland dynamics are modified by many local and regional factors and that gradual environmental change may be capable of triggering abrupt shifts and jumps in ecosystem states.     

Non-Linear Interactions Determine the Impact of Sea-Level Rise on Estuarine Benthic Biodiversity and Ecosystem Processes

Sea-level rise induced by climate change may have significant impacts on the ecosystem functions and ecosystem services provided by intertidal sediment ecosystems. Accelerated sea-level rise is expected to lead to steeper beach slopes, coarser particle sizes and increased wave exposure, with consequent impacts on intertidal ecosystems. We examined the relationships between abundance, biomass, and community metabolism of benthic fauna with beach slope, particle size and exposure, using samples across a range of conditions from three different locations in the UK, to determine the significance of sediment particle size beach slope and wave exposure in affecting benthic fauna and ecosystem function in different ecological contexts. Our results show that abundance, biomass and oxygen consumption of intertidal macrofauna and meiofauna are affected significantly by interactions among sediment particle size, beach slope and wave exposure. For macrofauna on less sloping beaches, the effect of these physical constraints is mediated by the local context, although for meiofauna and for macrofauna on intermediate and steeper beaches, the effects of physical constraints dominate. Steeper beach slopes, coarser particle sizes and increased wave exposure generally result in decreases in abundance, biomass and oxygen consumption, but these relationships are complex and non-linear. Sea-level rise is likely to lead to changes in ecosystem structure with generally negative impacts on ecosystem functions and ecosystem services. However, the impacts of sea-level rise will also be affected by local ecological context, especially for less sloping beaches.     

Laboratory Measure of an Ecosystem Response to a Sustained Stress

A new method is described for measuring environmental stress through the use of the duckweed (Lemna minor) rhizosphere.     

Increased Benthic Algal Primary Production in Response to the Invasive Zebra Mussel (Dreissena polymorpha) in a Productive Ecosystem, Oneida Lake, New York

Increased water clarity associated with zebra mussel (Dreissena polymorpha) populations may favor benthic algal primary production in freshwater systems previously dominated by pelagic phytoplankton production. While zebra mussel-mediated water clarity effects on benthic primary production have been implicated in published reports, few production estimates are available. This study estimates benthic primary production in Oneida Lake, NY before and after zebra mussel invasion (1992), using measured photosynthetic parameters ( , α^B and β) from sampled benthic algal communities. In the summers of 2003 and 2004, primary production was measured as O₂ evolution from algal communities on hard (cobble) and soft (sediment) substrate from several depths. We also backcast estimates of benthic primary production from measurements of light penetration since 1975. Estimates of whole-lake epipelic and epilithic algal primary production showed a significant (4%) increase and exhibited significantly less interannual variability subsequent to the establishment of zebra mussels. We applied our model to two lakes of differing trophic status; the model significantly overestimated benthic primary production in a hypereutrophic lake, but there was no significant difference between the actual and predicted primary production values in the oligotrophic lake. The hypereutrophic lake had higher zebra mussel densities than Oneida (224 vs. 41 per sample respectively). Though total community respiration (measured in total darkness) was factored into our model predictions of production, our model may need modification when heterotrophic respiration is a large portion of total community metabolism.     

Response of Benthic Invertebrates to Disturbance From Stream Restoration: The Importance of Bryophytes

The response of benthic invertebrates to disturbance from stream restoration was studied in the Reinikankoski rapids, central Finland. I hypothesized that stream bryophytes could act as a refugium for invertebrates. The restoration procedure destroyed nearly half of the bryophytes present at the study reach, and invertebrate densities decreased sharply immediately after the restoration. Within 2 weeks, invertebrates had recolonized the disturbed reach, and within 1 month peak numbers were attained. Invertebrates showed a clear association with bryophytes, especially after the restoration. My study shows that invertebrate recovery can be relatively fast in winter, and underlines the importance of stream mosses as invertebrate habitat and refugia. They should be taken in consideration in stream restoration projects by leaving patches of stream bottom intact.     

Physiological Response of Adipocytes to Weight Loss and Maintenance

Background

Metabolic processes in adipose tissue are dysregulated in obese subjects and, in response to weight loss, either normalize or change in favor of weight regain.

Objective

To determine changes in adipocyte glucose and fatty acid metabolism in relation to changes in adipocyte size during weight loss and maintenance.

Methods

Twenty-eight healthy subjects (12 males), age 20–50 y, and BMI 28–35 kg/m², followed a very low energy diet for 2 months, followed by a 10-month period of weight maintenance. Body weight, body composition (deuterium dilution and BodPod), protein levels (Western blot) and adipocyte size were assessed prior to and after weight loss and after the 10-month follow-up.

Results

A 10% weight loss resulted in a 16% decrease in adipocyte size. A marker for glycolysis decreased (AldoC) during weight loss in association with adipocyte shrinking, and remained decreased during follow-up in association with weight maintenance. A marker for fatty acid transport increased (FABP4) during weight loss and remained increased during follow-up. Markers for mitochondrial beta-oxidation (HADHsc) and lipolysis (ATGL) were only increased after the 10-month follow-up. During weight loss HADHsc and ATGL were coordinately regulated, which became weaker during follow-up due to adipocyte size-related changes in HADHsc expression. AldoC was the major denominator of adipocyte size and body weight, whereas changes in ATGL during weight loss contributed to body weight during follow-up. Upregulation of ATGL and HADHsc occured in the absence of a negative energy balance and was triggered by adipocyte shrinkage or indicated preadipocyte differentiation.

Conclusion

Markers for adipocyte glucose and fatty acid metabolism are changed in response to weight loss in line with normalization from a dysregulated obese status to an improved metabolic status.

Trial Registration

ClinicalTrials.gov {"type":"clinical-trial","attrs":{"text":"NCT01015508","term_id":"NCT01015508"}}NCT01015508     

Interactive Effects of Vegetation Type and Topographic Position on Nitrogen Availability and Loss in a Temperate Montane Ecosystem

Determining the fate of deposited nitrogen (N) in natural ecosystems remains a challenge. Heterogeneity of vegetation types and resulting plant–soil feedbacks interact with topo-hydrologic gradients to mediate spatial patterns of N availability and loss, yet net effects of variation in these two factors together across complex terrain remain unclear. Here we measured a suite of N-cycle pools and fluxes in sites that differed factorially in vegetation type (mixed forest vs. herbaceous) and topographic position (upslope vs. downslope) in a protected montane watershed near Salt Lake City, UT. Vegetation type was associated with large variation in N availability—herbaceous sites had larger NO₃ ^? pools, higher NO₃ ^?:NH₄ ⁺ ratios, higher nitrification potentials, lower soil C:N values, enriched δ¹⁵N values, and lower microbial biomass compared to forests, especially those upslope. Downslope sites tended to exhibit higher N availability and indicators of N-cycle openness, but patterns were moderated by vegetation type. In downslope forest, soil NO₃ ^? depth profiles and higher foliar N content suggested trees were accessing deep soil N and transferring it to the surface via litterfall, while more deep soil NO₃ ^? but no change in surface or foliar N suggested herbaceous cover was not N limited or deeper N pools were not accessible. Soil NO₃ ^? leaching from below the rooting zone closely tracked N availability, revealing a link between N status and hydrologic loss as well as an important role for roots in N retention. NO₃ ^? isotopes did not reveal a similar link for gaseous losses (that is, denitrification), instead reflecting nitrification and/or transport dynamics. Together, these results suggest a coupled ecological, topo-hydrologic perspective can help assess the fate of N in complex landscapes.     

Quantifying Patterns of Change in Marine Ecosystem Response to Multiple Pressures

The ability to understand and ultimately predict ecosystem response to multiple pressures is paramount to successfully implement ecosystem-based management. Thresholds shifts and nonlinear patterns in ecosystem responses can be used to determine reference points that identify levels of a pressure that may drastically alter ecosystem status, which can inform management action. However, quantifying ecosystem reference points has proven elusive due in large part to the multi-dimensional nature of both ecosystem pressures and ecosystem responses. We used ecological indicators, synthetic measures of ecosystem status and functioning, to enumerate important ecosystem attributes and to reduce the complexity of the Northeast Shelf Large Marine Ecosystem (NES LME). Random forests were used to quantify the importance of four environmental and four anthropogenic pressure variables to the value of ecological indicators, and to quantify shifts in aggregate ecological indicator response along pressure gradients. Anthropogenic pressure variables were critical defining features and were able to predict an average of 8-13% (up to 25-66% for individual ecological indicators) of the variation in ecological indicator values, whereas environmental pressures were able to predict an average of 1-5 % (up to 9-26% for individual ecological indicators) of ecological indicator variation. Each pressure variable predicted a different suite of ecological indicator’s variation and the shapes of ecological indicator responses along pressure gradients were generally nonlinear. Threshold shifts in ecosystem response to exploitation, the most important pressure variable, occurred when commercial landings were 20 and 60% of total surveyed biomass. Although present, threshold shifts in ecosystem response to environmental pressures were much less important, which suggests that anthropogenic pressures have significantly altered the ecosystem structure and functioning of the NES LME. Gradient response curves provide ecologically informed transformations of pressure variables to explain patterns of ecosystem structure and functioning. By concurrently identifying thresholds for a suite of ecological indicator responses to multiple pressures, we demonstrate that ecosystem reference points can be evaluated and used to support ecosystem-based management.