Modeling and analysis of nest-site selection by honeybee swarms: the speed and accuracy trade-off

Nest-site selection in honeybees is a process of social decision making in which the scout bees in a swarm locate several potential nest sites, evaluate them, and select the best one by means of competitive signaling. We develop a model of this process and validate that the model possesses the key features of the bees' decision-making process, as revealed by prior empirical studies. Next, we use the model to study the “design” of the nest-site selection process, with a focus on how certain behavioral parameters have been tuned by natural selection to achieve a balance between speed and accuracy. First, we study the effects of the quorum threshold and the dance decay rate. We show that evolution seems to have settled on values for these two parameters that seek a balance between speed and accuracy of decision making by minimizing the time needed to achieve a consensus and maximizing the probability that the best site is chosen. Second, we study the adaptive tuning of the tendency of bees to explore for vs be recruited to a site. We show that this tendency appears to be tuned to regulate the positive feedback process of recruitment to ensure both a reasonably rapid choice and a low probability of a poor choice. Finally we show that the probability of choosing the best site is proportional to its quality, but that this proportionality depends on its quality relative to other discovered sites.

Resistance of the generalist moth Trichoplusia ni (Noctuidae) to a novel chemical defense in the invasive plant Conium maculatum

Summary. Conium maculatum is an apiaceous species native to Eurasia that is highly toxic to vertebrates due to the presence of piperidine alkaloids, including coniine and γ-coniceine. More than 200 years after invading the United States this species remains mostly free from generalist insect herbivores. The presence of novel chemical defenses in the introduced range could provide invasive species with a competitive advantage relative to native plants. The cabbage looper (Trichoplusia ni) is a generalist lepidopteran found throughout the US that occasionally feeds on C. maculatum. We evaluated the toxicity of piperidine alkaloids to T. ni and determined putative resistance mechanisms, both behavioral and physiological, that allows this insect to develop successfully on C. maculatum foliage. T. ni larvae raised on diets enriched with coniine and γ-coniceine showed a decrease in consumption and longer development time, but no effects on growth were found at any alkaloid concentration. In a diet choice experiment T. ni larvae showed no avoidance of alkaloid-enriched diets, suggesting that the deterrence produced by alkaloids was related to a post-ingestive metabolic response. The ability of T. ni to consume diets high in alkaloid content could be due to at least three different mechanisms: 1) a decreased consumption rate, 2) efficient excretion of at least 1/3 of ingested alkaloids unmetabolized in frass, and 3) partial detoxification of alkaloids by cytochrome P450 s, as shown by the decreased larval growth in the presence of piperonyl butoxide, a P450 inhibitor. Even though T. ni tolerates C. maculatum alkaloids, the use of this species as a host plant could be ecologically disadvantageous due to prolonged larval growth and thus increased exposure to predators. Novel plant secondary compounds do not guarantee increased resistance to generalist herbivores.     

Learning affects the preference for parasitized hosts of the facultative kleptoparasitoid <Emphasis Type="Italic">Eupelmus vuilleti</Emphasis>

For a facultative kleptoparasite, the decision to allocate time and energy to search for a prey or for a kleptoparasite opportunity could be influenced by its prior experience. In this study, we investigated the influence of experience on the proportion of the facultative kleptoparasitoid Eupelmus vuilleti females choosing an unparasitized host or a host parasitized by Dinarmus basalis. When exploiting hosts previously parasitized by D. basalis, a large proportion of E. vuilleti females reused the hole drilled by the D. basalis female, which led to a reduction in the time needed to reach the host. When submitted to successive choice tests between an unparasitized host and a host parasitized by D. basalis in a natural host-plant system, the proportion of E. vuilleti females which choose the parasitized host increased with the female experience. The same experiment carried out with an artificial host-plant system allowing an easy access to the host did not show a variation of the females’ choice with experience. A reward—here a faster access to the host through the hole drilled by the D. basalis female—seems necessary to mediate the associative learning expressed by the preferential choice of the parasitized host.     

Models of withdrawing renewable and non-renewable resources based on Odum's energy systems theory and Daly's quasi-sustainability principle

This paper presents a theoretical model of withdrawing resources based on Odum's energy systems diagrams. According to the theory of a general pulsing principle, withdrawing resources changes in time shifting from the initial phases of growth and maturity to the phases of descent and low energy restoration. A simulation was performed in order to hypothesize potential future trends of withdrawing renewable and non-renewable resources and to show some aspects of their sustainability–unsustainability, respectively. According to Odum's theory, after the rapid growth of the last century, our civilization is living in a climax transition phase and it is now approaching a descent phase. A “way down” will be necessary due to the exhaustion of non-renewable and to the limited use of renewable resources. An integrated “renew–non-renew” model was developed by Odum to show how a “business as usual” trend will bring us to a drastic transition to a world that uses scarce renewable resources. Nevertheless, a different choice is possible, based on Daly's concept of quasi-sustainability that can inspire a new model. The latter argued that the exploitation of a non-renewable resource must be paired with a compensating investment in a renewable substitute. Our model shows that we can use non-renewable resources better to considerably improve our capacity of capturing renewable resources in the future. We present this as a necessary condition to address a sustainable environmental policy.     

Ancillary benefits of reduced air pollution in the US from moderate greenhouse gas mitigation policies in the electricity sector

Actions to slow atmospheric accumulation of greenhouse gases also would reduce conventional air pollutants yielding “ancillary” benefits that tend to accrue locally and in the near-term. Using a detailed electricity model linked to an integrated assessment framework to value changes in human health, we find a tax of $25 per metric ton of carbon emissions would yield NO_x-related health benefits of about $8 per metric ton of carbon reduced in the year 2010 (1997 dollars). Additional savings of $4–$7 accrue from reduced investment in NO_x and SO₂ abatement in order to comply with emission caps. Total ancillary benefits of a $25 carbon tax are $12–$14, which appear to justify the costs of a $25 tax, although marginal benefits are less than marginal costs. At a tax of $75, greater total benefits are achieved but the value per ton of carbon reductions remains roughly constant at about $12.     

Extra-pair paternity in alpine marmots, <Emphasis Type="Italic">Marmota marmota</Emphasis>: genetic quality and genetic diversity effects

Assuming that a male’s genetic characteristics affect those of his offspring, extra-pair copulation has been hypothesized to increase heterozygosity of the progeny—the “genetic compatibility” hypothesis—and the genetic diversity within litters—the “genetic diversity” hypothesis. We tested these two hypotheses in the alpine marmot (Marmota marmota), a socially monogamous mammal showing a high rate of extra-pair paternity (EPP). In a first step, we tested the assumption that a male’s genetic characteristics (heterozygosity and genetic similarity to the female) affect those of his offspring. Genetic similarity between parents influenced offspring heterozygosity, offspring genetic similarity to their mother, and litter genetic diversity. The father’s heterozygosity also influenced litter genetic diversity but did not affect offspring heterozygosity. Hence, heterozygosity seems not to be heritable in the alpine marmot. In a second step, we compared genetic characteristics of extra-pair young (EPY) and within-pair young (WPY). EPY were less genetically similar to their mother but not more heterozygous than WPY. EPY siblings were also less genetically similar than their WPY half siblings. Finally, the presence of EPY promoted genetic diversity within the litter. Thus, our data support both the “genetic compatibility” and the “genetic diversity” hypotheses. We discuss further investigations needed to determine the primary causes of EPP in this species.     

Modeling estuarine-shelf exchanges in a deltaic estuary: Implications for coastal carbon budgets and hypoxia

The export of wetland-derived materials to the coastal ocean (i.e., the “Outwelling” hypothesis) has received considerable attention over the past several decades. While a number of studies have shown that estuaries export appreciable amounts of nutrients and carbon, few studies have attempted to estimate the importance of estuarine sources for the coastal carbon budgets in river-dominated coastal ecosystems. A novel tidal prism model was developed to examine estuarine-shelf exchanges in the Barataria estuary, a deltaic estuary located in the north-central Gulf of Mexico. This estuary has been the site of a massive wetland loss, and it has been hypothesized that carbon export from the eroding coastal wetlands supports the development of a large hypoxic zone in the coastal Gulf of Mexico. The model results show that the Barataria estuary receives nitrogen through the tidal passes and releases carbon to the coastal ocean. The mean calculated tidal water discharge of 6930 m³ s⁻¹ is equivalent to about 43% of the lower Mississippi River discharge. The annual total organic carbon (TOC) export is 109 million kg, or 57 gC m² yr⁻¹ when prorated to the total water area of the estuary. This carbon export is equivalent to a loss of 0.5 m of wetland soil horizon over an area of 8.4 km², and accounts for about 34% of the observed annual wetland loss in the estuary between 1978 and 2000. Compared to the lower Mississippi River, the Barataria estuary appears to be a very small source of TOC for the northern Gulf of Mexico (2.7% of riverine TOC), and is unlikely to have a significant influence on the development of the Gulf's hypoxia.     

The effect of pup vocalisations on food allocation in a cooperative mammal, the meerkat (Suricata suricatta)

In the meerkat (Suricata suricatta), a cooperative mongoose, pups follow potential feeders while the group is foraging and emit incessant calls when soliciting food from them. In contrast to a ’stationary’ brood of chicks, in which nestlings are fed at a fixed location, meerkat pups are ’mobile’ and become spread out. The question arises whether meerkat pups that experience different constraints to those facing chicks have evolved similar begging strategies. This paper describes the vocalisations that meerkat pups emit in the context of begging and investigates the influence of these calls on food allocation by older group members and on the behaviour of littermates. Meerkat pups use two types of calls when soliciting food from a potential feeder. The most common is a ’repeat’ call, which pups emit continuously when following an older forager over several hours a day. In addition, when a potential feeder finds a prey item, the pups next to it emit a bout of calls with increased calling rate, amplitude and fundamental frequency, termed ’high-pitched’ calls. Observations, together with playback experiments, showed that more prey was allocated to pups that called longer and more intensely. The pup closest to a feeder was almost always fed. The probability of emitting high-pitched calls did not depend on the time since a pup had received food, and the change from repeat to high-pitched calls occurred suddenly. The main function of the high-pitched call, therefore, does not appear to be to signal a pup’s hunger state. More likely, the two calls, in the context of begging, may be an adaptation to energetic constraints in a mobile feeding system. Pups, which are dispersed during foraging, may emit repeat calls over long periods to prevent potential feeders from eating all the prey themselves. At the moment a potential feeder finds prey, pups may give the more intense high-pitched calls to direct feeders to bring the food item to them and not to a littermate. Therefore, unlike the stationary feeding system where chicks emit one type of begging call when the feeder approaches the nest, meerkats, with a mobile feeding system, have evolved two discrete types of vocalisations in the context of begging. Received: 22 November 1999 / Revised: 1 July 2000 / Accepted: 17 July 2000     

New hydroepidemiological models of indicator organisms and zoonotic pathogens in agricultural watersheds

Simple analytical models are derived to assess how a series of cattle animal farms affect the transport and fate of an indicator organism (Escherichia coli) and a zoonotic pathogen (Campylobacter) in a stream. Separate steady-state mass-balance models are developed and solved for the ultimate minimum and maximum concentrations for the two organisms. The E. coli model assumes that the organism is ubiquitous and abundant in the animals’ digestive tracts. In contrast, a simple dose-response model is employed to relate the Campylobacter prevalence to drinking water drawn from the stream. Because faecal indicators are commonly employed to assess the efficacy of best management practice (BMP) interventions, we also employ the models to assess how BMPs impact pathogen levels. The model provides predictions of (a) the relative removal efficacy for Campylobacter and (b) the prevalence of Campylobacter infection among farm animals after implementation of BMPs. Dimensionless numbers and simple graphs are developed to assess how prevalence is influenced by a number of factors including animal density and farm spacing. A significant outcome of this model development is that the numerous dimensional input and parameter variables are reduced to a group of just four dimensionless Campylobacter-related quantities, characterizing: animal density; in-stream attenuation; animal-to-animal transmission; and infection recovery. Calculations reveal that for some constellations of these four quantities there can be a greater-than-expected benefit in that the proportional reduction of stream Campylobacter concentrations post-BMP can substantially exceed the proportional reduction of concentrations of E. coli in that stream. In addition, a criterion for system sterility (i.e., the conditions required for the farm infection rate to decrease with downstream distance) is derived.     

Proposing an information criterion for individual-based models developed in a pattern-oriented modelling framework

Individual-based models (IBMs) have been improved in quality and reliability in recent years with an approach called pattern-oriented modelling (POM). POM proposes guidelines to develop models reproducing multiple patterns observed on the field and to test systematically how well the IBMs reproduce them. POM studies used generally traditional methods of goodness of fit such as the sum of squares evaluation or ad hoc comparisons of fitting errors and variations. Model selection, however, can be a rigorous statistical approach based on information theory and information criteria such as the Akaike's information criterion (AIC) or the deviance information criterion (DIC). So far, it has not been tried to link POM to these rigorous techniques. The main problems to achieve that are: (a) the difficulty to have likelihood functions for IBMs’ parameters and (b) the possibility to obtain posterior distributions of IBMs’ parameters given the patterns to reproduce. In a first part, this paper answers problem (a) by proposing and explaining how to calculate a deviance measure (POMDEV) for models developed in a context of POM. And while answering the second problem, a second part of the paper proposes an information criterion for model selection in a POM context (the pattern-oriented modelling information criterion: POMIC). This criterion does not yet have the same theoretical foundation as, e.g., AIC, but uses formal analogies to the DIC. In a third part POMIC is tested with a modelling exercise. This exercise shows the potential of POMIC to use multiple patterns for selecting among multiple potential submodels and eventually select the most parsimonious and well fitting model version. We conclude that POMIC, although being a heuristically derived approach, can greatly improve the POM framework.     

How humans shape wolf behavior in Banff and Kootenay National Parks, Canada

This paper describes the conceptualization and implementation of an agent-based model to investigate how varying levels of human presence could affect elements of wolf behavior, including highway crossings; use of areas in proximity to roads and trails; size of home ranges; activities, such as hunting, patrolling, resting, and feeding pups; and survival of individuals in Banff and Kootenay National Parks, Canada. The model consists of a wolf module as the primary component with five packs represented as cognitive agents, and grizzly bear, elk, and human modules that represent dynamic components of the environment. A set of environmental data layers was used to develop a friction model that serves as a base map representing the landscape over which wolves moved. A decision model was built to simulate the sequence of wolf activities. The model was implemented in a Java Programming Language using RePast, an agent-based modeling library. Six months of wolf activities were simulated from April 16 to October 15 (i.e., a season coherent with regard to known wolf behaviors), and calibrated with GPS data from wolf radiocollars (n = 15) deployed from 2002 to 2004. Results showed that the simulated trajectories of wolf movements were correlated with the observed trajectories (Spearman's rho 0.566, P < 0.001); other critical behaviors, such as time spent at the den and not traveling were also correlated. The simulations revealed that wolf movements and behaviors were noticeably affected by the intensity of human presence. The packs’ home ranges shrank and wolves crossed highways less frequently with increased human presence. In an extreme example, a wolf pack whose home range is traversed by a high-traffic-volume highway was extirpated due to inability to hunt successfully under a scenario wherein human presence levels were increased 10-fold. The modeling prototype developed in this study may serve as a tool to test hypotheses about human effects on wolves and on other mammals, and guide decision-makers in designing management strategies that minimize impacts on wolves and on other species functionally related to wolves in the ecosystem.     

Unusual echolocation behavior in a small molossid bat, <Emphasis Type="Italic">Molossops temminckii,</Emphasis> that forages near background clutter

When searching for flying insects, Molossops temminckii uses unusual echolocation calls characterized by upward modulation of frequency vs time (UFM). Call frequency increases asymptotically in the relatively long (∼8 ms) pulses from a starting frequency of ∼40 kHz to a long narrowband tail at ∼50 kHz. When approaching a prey, the bat progressively increases the duration of calls and intersperses in the sequence broadband downwardly frequency-modulated signals with a terminal frequency of about 53 kHz, which totally replaces the UFM signals at the end of the approach phase. The sequence progresses to a capture buzz resembling those from other molossid and vespertilionid bats. The M. temminckii wing morphology is characterized by an average aspect ratio and a high wing loading, suggesting that it is more maneuverable than the typical Molossidae but less than typical Vespertilionidae. M. temminckii regularly forages near clutter, where it needs to pay attention to the background and might face forward and backward masking of signals. We hypothesize that the UFM echolocation signals of M. temminckii represent an adaptation to foraging near background clutter in a not very maneuverable bat needing a broad attention window. The broadband component of the signal might serve for the perception of the background and the narrowband tail for detection and perhaps classification of prey. Bats may solve the signal masking problems by separating emission and echoes in the frequency domain. The echolocation behavior of M. temminckii may shed light on the evolution of the narrowband frequency analysis echolocation systems adopted by some bats foraging within clutter.     

Recycling flows in emergy evaluation: A mathematical paradox?

This paper is a contribution to the emergy evaluation of systems involving recycling or reuse of waste. If waste exergy (its residual usefulness) is not negligible, wastes could serve as input to another process or be recycled. In cases of continuous waste recycle or reuse, what then is the role of emergy? Emergy is carried by matter and its value is shown to be the product of specific energy with mass flow rate and its transformity. This transformity (τ) given as the ratio of the total emergy input and the useful available energy in the product (exergy) is commonly calculated over a specific period of time (usually yearly) which makes transformity a time dependent factor. Assuming a process in which a part of the non-renewable input is an output (waste) from a previous system, for the waste to be reused, an emergy investment is needed. The transformity of the reused or recycled material should be calculated based on the pathway of the reused material at a certain time (T) which results in a specific transformity value (τ). In case of a second recycle of the same material that had undergone the previous recycle, the material pathway has a new time (T + T₁) which results in a transformity value (τ₁). Recycling flows as in the case of feedback is a dynamic process and as such the process introduces its own time period depending on its pathway which has to be considered in emergy evaluations. Through the inspiration of previous emergy studies, authors have tried to develop formulae which could be used in such cases of continuous recycling of material in this paper. The developed approach is then applied to a case study to give the reader a better understanding of the concept. As a result, a ‘factor’ is introduced which could be included on emergy evaluation tables to account for subsequent transformity changes in multiple recycling. This factor can be used to solve the difficulties in evaluating aggregated systems, serve as a correction factor to up-level such models keeping the correct evaluation and also solve problems of memory loss in emergy evaluation. The discussion deals with the questions; is it a pure mathematical paradox in the rules of emergy? Is it consistent with previous work? What were the previous solutions to avoid the cumulative problem in a reuse? What are the consequences?     

Evaluating multi-stakeholder perceptions of project impacts: a participatory value-based multi-criteria approach

This paper describes a participatory multi-stakeholder impact assessment approach based on the concept of a value tree (VT), generated using the value focused thinking method (VFT) developed by Keeney (1992 Keeney, R. 1992. Value-focused thinking, Cambridge, MA: Harvard University Press.  [Google Scholar]). This approach allows stakeholders to specify impact parameters to be evaluated, which in the context of the VT, are organized as goals, objectives and alternatives. The approach is implemented in two phases: Phase I is designed to formulate stakeholders' collective VT following VFT concepts and the cognitive mapping method, and Phase II is a participatory valuation approach based on the VT formed in Phase I. The VT and the VFT concepts are adopted as tools to capture different stakeholders' values, goals and perceptions. The resulting VT is structured as a hierarchy between goals and objectives, and a network consisting of relationships, linkages and cross-impacts of the different alternatives and objectives. The hierarchy and network structure enables stakeholders to decompose complex assessment problem into ‘smaller’ units, which makes for easier and clearer assessment context, without ignoring linkages of the units or assessment elements. The second phase allows stakeholders to express their preferences with respect to each assessment element, through a voting system that ultimately leads to measures of importance or relative weights associated with each element. A modified Analytic Hierarchy Process (called Analytic Network Process) was used to distill relative weights from the voting results. Results obtained from a case study in a Zimbabwean community forest show that the proposed approach is easy to implement and can address questions about whether a project can lead to a positive change in attitudes, and whether the changes actually lead to a propensity to adopt alternatives that the project supports (e.g. conservation-oriented alternatives).     

敌草快对藻类和溞类的急性毒性及其水中含量测定

敌草快是一种非选择性、广谱的联吡啶类触杀性除草剂,主要通过干扰植物细胞膜、破坏光合作用而快速发挥效果。为探究敌草快对水生生物的毒性,测定了该化合物对羊角月芽藻和大型溞的急性毒性,并建立了高效液相色谱法测定水中敌草快含量的方法。结果表明:检测方法在1.00×10-2~3.00×10-2mg a.i.·L-1范围内的线性相关系数为0.99995,添加回收率在90.3%~109%之间,相对标准偏差(RSD)为1.10%~10.3%,保留时间在7.2 min左右。按实测浓度和理论浓度分别计算敌草快对羊角月芽藻的72 h的半数效应浓度EyC50(72 h-EyC50),分别为3.16×10-2mg a.i.·L-1和3.32×10-2mg a.i.·L-1,均为高毒;对大型溞48 h的半数效应浓度EC50(48 h-EC50)分别为1.18×10-2mg a.i.·L-1和1.33×10-2mg a.i.·L-1,均为剧毒。     

Modelling the effects of eutrophication, mitigation measures and an extreme flood event on estuarine benthic food webs

Human-mediated and natural disturbances such as nutrient enrichment, habitat modification, and flood events often result in significant shifts in species composition and abundance that translate into changes in the food web structure. Six mass-balanced models were developed using the “Ecopath with Ecosim” software package to assess changes in benthic food web properties in the Mondego estuarine ecosystem (Portugal). Field, laboratory and literature information were used to construct the models. The main study objective was to assess at 2 sites (a Zostera meadow and a bare sediment area) the effects of: (1) a period of anthropogenic enrichment, which led to excessive production of organic matter in the form of algal blooms (1993/1994); (2) the implementation of mitigation measures, following a long period of eutrophication (1999/2000); and (3) a centenary flood (winter 2000/2001). Different numbers of compartments were identified at each site and in each time period. In general, the Zostera site, due to its complex community, showed a higher number of compartments and a higher level of system activity (i.e. sum of consumptions, respiration, flow to detritus, production, total system throughput, net primary production and system omnivory index). The differences at the two sites in the three time periods in the breakdown of throughput were mainly due to differences in the biomass of the primary producers (higher primary production at the Zostera site). Consumption, respiration and flow to detritus were dominated by the grazers Hydrobia ulvae and Scrobicularia plana at the Zostera and bare sediment sites respectively. At both sites, after recovery measures were implemented there was an increase in S. plana and Hediste diversicolor biomass, consumption, respiration and flows to detritus, and a decrease in H. ulvae biomass and associated flows, which increased again after the flood event. The mass-balanced models showed that the trophic structure of the benthic communities in Mondego estuary was affected differently by each disturbance event. Interestingly, in our study a high system throughput seems to be associated with higher stress levels, which contradicts the idea that higher system activity is always a sign of healthier conditions.     

Risk taking during parental care: a test of three hypotheses applied to the pied flycatcher

According to life-history theory, there will often be a conflict between investment in current versus future reproduction. If a predator appears during breeding, parents must make a compromise between ensuring the growth and survival of offspring (nest defence, feeding and brooding of young), and reducing the risk of predation to ensure their own survival. We model three hypotheses for the outcome of this conflict which are particularly relevant for altricial birds. They are not mutually exclusive, but focus on different costs and benefits. (1) Parental investment is determined by the parents’ own risk of predation. This hypothesis predicts that a lone parent should take smaller risks than a parent that has a mate. (2) Parental investment is related to the reproductive value of the offspring: Parents are predicted to take greater risks for larger broods, larger-sized or older offspring. (3) Finally, we present the new hypothesis that parental investment is related to the harm that offspring would suffer during a period of no parental care (incubation, brooding, feeding). This hypothesis predicts that parents should take greater risks for younger offspring, or for offspring in poorer condition, because the marginal benefit of parental care is largest in such cases. Hence, one may also expect that lone parents should take greater risks than two parents because their offspring are more in need of care. We tested these hypotheses on the pied flycatcher (Ficedula hypoleuca) by presenting a stuffed predator of the parents (a sparrowhawk, Accipiter nisus) close to the nest when parents were feeding the young. Risk taking was measured as the time that elapsed until the first visit to the nest. Most support was found for the ‘‘harm to offspring’’ hypothesis. Previous studies have usually measured the intensity of nest defence against typical nest predators, and have found evidence for the ‘‘reproductive value of offspring’’ hypothesis. However, our model predicts that the importance of the reproductive value of the offspring should decrease relative to the harm that offspring would suffer if they were not cared for when the predator type changes from a nest predator to a predator of adults, and when conditions for breeding turn from good to bad. Received: 13 April 1995/Accepted after revision: 11 March 1996     

A revised calculation of the econometric factors α- and β for the Extended Exergy Accounting method

Extended Exergy Accounting (“EEA”) is a method to compute the space- and time integral of the primary exergy required to produce a good or a service: the extended exergy of a commodity measures its “embodied exergy”, including externalities (Labour, Capital and Environmental Costa). A crucial point of the method is the calculation of two econometric coefficients, commonly referred to as “α” and “β”,used to calculate the extended exergy equivalents of Labour and Capital respectively. In previous applications of the EEA method, these coefficients have been assigned approximate values estimated on the basis of global system considerations. In this paper, a novel procedure is described that leads to the calculation of “exact” values of both econometric coefficients, based on detailed exergy- and monetary balances of the Society to which the EEA is applied. It is shown that both α and β depend in a non-trivial way from the consumption patterns, the technological level and the life- and socio-economic standards of each Country. It is also shown that the values are substantially different for developed (OECD) and underdeveloped Countries, and representative samples of values are calculated and critically analysed. On the basis of these new model coefficients, the specific exergy equivalents of Labour (ee_L) and of Capital (ee_K) are calculated, and shown to differ from the values used in previous EEA calculations.     

Modelling forest management within a global vegetation model—Part 1: Model structure and general behaviour

This article describes a new forest management module (FMM) that explicitly simulates forest stand growth and management within a process-based global vegetation model (GVM) called ORCHIDEE. The net primary productivity simulated by ORCHIDEE is used as an input to the FMM. The FMM then calculates stand and management characteristics such as stand density, tree size distribution, tree growth, the timing and intensity of thinnings and clear-cuts, wood extraction and litter generated after thinning. Some of these variables are then fed back to ORCHIDEE. These computations are made possible with a distribution-based modelling of individual tree size. The model derives natural mortality from the relative density index (rdi), a competition index based on tree size and stand density. Based on the common forestry management principle of avoiding natural mortality, a set of rules is defined to calculate the recurrent intensity and frequency of forestry operations during the stand lifetime. The new-coupled model is called ORCHIDEE-FM (forest management).The general behaviour of ORCHIDEE-FM is analysed for a broadleaf forest in north-eastern France. Flux simulation throughout a forest rotation compare well with the literature values, both in absolute values and dynamics.Results from ORCHIDEE-FM highlight the impact of forest management on ecosystem C-cycling, both in terms of carbon fluxes and stocks. In particular, the average net ecosystem productivity (NEP) of 225 gC m⁻² year⁻¹ is close to the biome average of 311 gC m⁻² year⁻¹. The NEP of the “unmanaged” case is 40% lower, leading us to conclude that management explains 40% of the cumulated carbon sink over 150 years. A sensitivity analysis reveals 4 major avenues for improvement: a better determination of initial conditions, an improved allocation scheme to explain age-related decline in productivity, and an increased specificity of both the self-thinning curve and the biomass-diameter allometry.     

Different proxies for the reactivity of aquatic sediments towards oxygen: A model assessment

Information on benthic carbon mineralization rates is often derived from the analysis of oxygen microprofiles in sediments. To enable a direct comparison of different sediment environments, it is often desirable to characterize sediments by a single proxy that expresses their “reactivity” towards oxygen. For this, there are three commonly used proxies: the oxygen penetration depth (OPD), the oxygen flux at the sediment-water interface (DOU), and the maximum volumetric oxygen consumption rate (R_max). The OPD can be directly determined from the oxygen depth profile, while the DOU is usually obtained by a linear fit to the oxygen gradient either in diffusive boundary layer. The oxygen consumption rate R_max requires the fitting of a reactive-transport model to the data profile. This article shows that the OPD alone is a suboptimal proxy, because it shows a strong dependence on the half-saturation constant K_s, and secondly, because it is sensitive to the particular re-oxidation conditions right above the oxic-anoxic interface. Similarly, the volumetric oxygen consumption rate R_max is rather strongly dependent on the kinetic model formulation employed. To show this we fitted three different (Bouldin, Blackman and Monod) kinetics to the same oxygen data profiles. When fitting these models, the R_max values obtained differed by 20% for exactly the same oxygen profile. Accordingly, if one reports R_max values, it is crucial to specify the kinetic model alongside. Overall, DOU emerges as sediment reactivity proxy which is the least model dependent.