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Ecological Stoichiometry Another Currency

Ecological stoichiometry is well established in aquatic ecology but not yet in terrestrial ecology. Two recent papers on stoichiometry (Anderson et al., 2004, Moe et al., 2005) argue persuasively that ecologists interested in animal population response to resource availability need to consider the currency with which they examine plant-animal interactions. They argue that ecological stoichiome-try provides a multiple currency approach to understand the effects of resource quality. By multiple currency, they mean that rather than abstracting populations as aggregations of individuals or biomass, organisms are represented by carbon (C), phosphorous (P), and other trace elements that allow key feedbacks, such as consumer-driven nutrient recycling processes (Anderson et al., 2004 p. 884). The argument is that both food quantity and quality can be incorporated into a single framework. The concept of currency here has two related parts one meaning refers to the difference between the...

Resource Acquisition and Animal Response in Dynamic Landscapes

Quantification in ecology has been the sine qua non that has differentiated rigorous science from something less. It is how we have kept the books. Quantifying the quantifiable to account for population response to resource availability usually has meant that the quantity of some resource (e.g., for herbivores, plant biomass, or areal coverage of the plant community or habitat) has been assumed to have some causal effect on some quantitative measure of animal response (e.g., the number or organisms surviving and recruited into the population). As scale effects have been recognized as important, landscape ecologists have followed a similar methodology and have assumed that the habitat area coverage (quantity) bears some relation to population and species performance and health. The explanatory power of the spatial amount of habitat elements seems inconsistent, and available metrics to assess the effects of spatial arrangement are problematic. Further, organisms respond to the...

Temporal Discontinuities

Obvious discontinuities in resource availability in time is the fruiting of plants. The availability of acorns is a good example of a resource that exhibits strong temporal discontinuities (Abrahamson and Layne, 2003). Wolff (1996) found that rodent densities were positively correlated with oak mast production over a period of 14 years. However, much temporal heterogeneity of resources is gradual i.e., it concerns variation in resource quality. In this ecological sense, all resources may be considered pulsed or discontinuous to some degree (see Ostfeld and Keesing, Chapter 2, this volume). Most resources appear seasonally in temperate and tropical environments. Additionally, resource quality changes over time as well as over spatial gradients. There is a voluminous literature on the response of species to environment resource gradients. These were largely a result of two papers by Tilman (1980, 1982) who proposed what has come to be known as the resource-ratio theory. The theory...

Two Important Distinctions

This has been referred to in a general sense as a single currency approach (Moe et al., 2005), where the clear implication is that currency refers to either quantitative or qualitative effects, but not both. Regardless, if the objective is to learn how species respond to changing landscapes and hence changing resource availability (quantity) and quality, attention to spatial details gets us only part way there. Attention to temporal effects as well as consideration of the qualitative differences in resources is necessary.

Resource Quality Keeping the Books

So then, how might we improve our keeping of the books in ecology Keeping the books, i.e., accounting in ecology, implies that we are capturing the essence of the interactions so that understanding is enhanced. Specifically, it implies that our observations are buttressed by a conceptual understanding that makes sense. Put another way, the assumption is that the variables we measure, i.e. the currencies, are appropriate and up to the task. Studies of habitat fragmentation that have addressed animal responses to resources availability have used almost exclusively the currency of quantity of resource as the explanatory variable. Indeed, many habitat use preference studies appear to be based on the hypothesis that the amount of habitat is more or less directly causally related to response variables such as animal density, growth, reproduction, survivorship, and birth and death rates. Additionally, even though habitat types themselves are often assumed to represent areas of different...

Resource Quality and Population Response

Stoichiometric theory has formalized these constraints (Anderson et al., 2004) by what is known as the threshold elemental ratio (TER). This is the carbon element threshold where the resource limitation shifts from carbon (C) to nutrient (P, N), that is, where the quality of the plant resource makes a difference. With plant C element ratios < 1, plant quality is always adequate for the herbivore and a single currency approach based on quantity of food will not deviate significantly from a stoichiometrical approach (Urabe and Watanabe, 1992 Urabe and Sterner, 1996). In these cases, ecologists have correctly used quantity to reflect herbivore response. It is when TER ratios > 1 that a stoichiometric model approach can be illuminating.

Different Predictions

Perhaps the fundamental key for population ecologists is that because stoichiometric models incorporate both food quantity (which ecologists usually measure) and quality, which is inferred but much less frequently incorporated into the measurements, there may appear empirical phenomena that cannot be predicted by single currency models. Examples from laboratory experiments include the observations of a (1) positive density dependence and a shift in the nature of the interaction from competition to facilitation (Sommer, 1992), similar to the Allee effect (2) coexistence of more than one predator on a single prey item in contrast to predictions based on the single (quantity)-currency theory (Grover, 2003 Hall, 2004 Hall et al., 2005) and (3) the diversity enhancing effects on herbivores of poor food quality (Anderson et al., 2004). Although these results come primarily from aquatic system experiments, terrestrial ecologists may find that Food quality may provide a better explanation for...

Method And Empirical Results

In the second experiment I offered a one-shot choice between a quarter-liter cup of cooking oil available now versus three cups after a delay of several days. This experiment offered real rewards, meaning that I actually gave the participant the cups of oil they chose after the promised delay. Cooking oil served well as a reward currency because it represents food value (cash is often spent on tobacco, which has its own unique utility curve) cooking oil is also easily storable and transportable, and highly desired. To limit potential jealousy related to unequal distribution of rewards, I offered the same choice to all participants in a given village and changed the delay to receive the three-cup option at each village visited. The three-cup option was delayed by two weeks at the first village, one week at the next site, and three days at the final location. A median indifference value was estimated for the population as a whole at 0.3-0.7 day (N 49).

Conclusions The Truth Is Always Beyond the Perception of Truth

Throughout this chapter, the theme has been to try to find a way to get closer to understanding the true state of nature as it applies to resource availability and animal population response. However, in science generally, and in ecology specifically, the idea of truth is an elusive concept. What we know or what we think we know is always based on (often unstated) assumptions is filtered through our methodological approaches, and is always constrained by the observation set we employ. Put in different terms, truth as a science concept is nuanced, and it is so because science is the one enterprise where we continually attempt to falsify our hypotheses and predictions, and examine our premises in order to test what we know. When one thinks about individual animal or population response to the spatial and temporal distribution of required and necessary resources, it seems reasonable that future advances in our understanding of animals that live in dynamical landscapes may be facilitated...

Historical materialism and the remaking of environments

Both metabolism and circulation have long conceptual and material histories. Circulation gained wide currency after William Harvey's postulation of the double circulation of blood in the body. Movement, flux and conduits rapidly thereafter became formative metaphors that would shape radically new visions of and practices for acting in the world. The concept of metabolism arose in the early nineteenth century, particularly in relationship to the material exchanges in the body with respect to respiration. It became extended later to include material exchanges between organisms and the environment as well as the bio-physical processes within living (and non-living or decaying) entities. For example, in the writings of Jacob Moleschott (1857) and Justus von Liebig (1840 1842), metabolism denoted not only the exchange of energy and substances between organisms and the environment, but the totality of biochemical reactions in a living thing. In fact, von Liebig's analysis turned organisms...

The invention of circulation

Alongside the emergence of the notion of metabolism in the natural and social sciences (an emergence not wholly disassociated with the rising metabolic rift caused by industrialization and urbanization), the notion of circulation began to gain greater and wider currency. For example, the idea of water circulation, that water piped into the city must leave the city by its sewers is not older than the nineteenth century (in the west). Circulating water, following a given path and finally returning to its source, remained foreign to western urban imaginations, spatial representations and engineering systems until then. Modern urbanization, highly dependent on the mastery of circulating flows, was linked with the representation of cities as consisting of and functioning through complex networks of circulatory systems (Kaika and Swyngedouw 1999). metabolism through considering the metabolic rift, circulation gained greater socio-ecological currency exactly when it became seen as an...

Traits invariants and theories of everything

In general we might expect certain values of a trait to be favoured rather universally. For example, it is clearly best, in a Darwinian sense, to produce many offspring, all other things being equal. Happily for those who like diversity, all other things are not equal high fecundity must come at a cost to some other trait of importance to the organism's fitness, such as offspring size. This phenomenon is known as a trade-off. The organism is thus faced with the more complex choice of finding the value of the trait that maximizes fitness in the face of trade-offs with other traits of importance. In the above example, evolution would essentially select between organisms that have large offspring but few of them, and organisms that have many but small offspring. Following this, our evolutionary model will generally make some assumptions about the nature of any trade-offs involved (a constraint), and about the consequences of each value of the trait for fitness (a currency). In general,...

Discussion and conclusion

Despite their paramount importance, fitness gain curves are exceedingly difficult to measure empirically (Brunet 1992 Klinkhamer et al. 1997 Campbell 2000 Klinkhamer and de Jong 2002), hindering the development and maturation of theory. Measurement of male or female fertility often requires parentage or paternity analysis with genetic markers, which is time consuming and feasible only in small populations with the precision necessary to describe a gain function. Another difficulty is that if direct effects of plant size are important, different-sized plants cannot be used to determine the gain curve, as in the very few attempts to measure gain curves in natural populations (Emms 1993 Klinkhamer et al. 1997 Klin-khamer and de Jong 2002). Furthermore, it is often unclear what resources should be measured to assess allocation. Most studies consider dry mass as a currency, but use of other currencies, such as nitrogen or phosphorus, can give different perspectives (e.g., McKone et al....

Terrestrial habitat along sea coasts

Their environment is important, especially for conservation management. A large amount of research has been published on this, mostly on the Eurasian otter, the most common currency being the occurrence of otter spraints in different types of vegetation along banks and coasts.

The Patch Departure Decision

Flowers visited before returning to the hive supports the assumption that honeybees maximize efficiency (net energy gain energy expenditure) rather than the more conventional currency of net energy gain (Schmid-Hempel et al. 1985 see also section 8.3). In order to respond to the travel time between flowers, foraging honeybees must monitor this variable in some way and then base their decision to cease foraging on their current estimate of travel time, stored in working memory. Memory for travel times between flowers is an important part of honeybee foraging.

Thermodynamic And Other Constraints

Once different environmental impacts are calculated, they must be weighted and balanced against each other, as well as other concerns, such as cost and long-term sustain-ability. These multiple, often conflicting, goals pose significant challenges to process optimization and design. How can designs be identified that best satisfy multiple objectives Multi-objective optimization algorithms provide a particularly useful approach, aimed at determining the set of non-dominant non-dominated ('Pareto') designs where a further improvement for one objective can only be made at the expense of another. This determines the set of potentially 'best' designs and explicitly identifies the trade-offs between them. This is in contrast to cost-benefit analysis, which deals with multiple objectives by identifying a single fundamental objective and then converting all the other objectives into this single currency. The multi-objective approach is particularly valuable in situations where there are a...

Musth as advertisement of male quality to females

Musth is an expensive proposition in the currency of energy. The reduced amount of time spent in feeding and greater time spent in searching for estrous females also means that a bull has to be in good condition in the first place to sustain the rigors of the musth period. High levels of androgens (e.g., testosterone) also increase metabolic rates and consume more energy. Thus, captive males in musth lose body condition even if they are chained and given normal quantities of forage. The constant dribbling of urine also exacts a physiological cost. A large male in musth may lose 350 liters of urine each day. When water is limiting, as in semiarid habitats or during the dry season, the physiological costs of urine dribbling for signaling can be substantial if the male is to leave uninterrupted trails.

Ecosystem Components and Properties

Typically, ecologists quantifying ecosystem dynamics use carbon as their currency to describe material flow and energy to quantify energy flux. Material flow and energy flow differ in one important property, namely their ability to be recycled. Chemical materials within an ecosystem are recycled through an ecosystem's component. In contrast, energy moves through an ecosystem only once and is not recycled (Figure 3). Most energy is transformed to heat and ultimately lost from the system. Consequently, the continual input of new solar energy is what keeps an ecosystem operational.

Fecundity and Fertility in Population Dynamical Models

Stage-structured models provide an estimate of the stable stage distribution, which is the theoretical proportional allocation of individuals within the population to the defined stages (e.g., age groups, size classes, developmental stages) resulting from constant demographic rates. The stable stage distribution is a useful metric because it provides the theoretical composition of a population exhibiting a fixed birth rate, so factors such as environmental variation or intrinsic regulation that alter this theoretical distribution can be ranked for their effects on the future composition of a population. This introduces the concept of'reproductive value' - a measure of the combined effects of fecundity, fertility, and survival that takes an individual's proportional contribution to the future status of its population into account. It is formally expressed as the sum of the current and future reproductive values, and is considered the currency used by natural selection to produce a...

Implications of the Bacteroid Urate System

The bacteroid-assisted ability of cockroaches to store, mobilize, and in some cases, transfer urates uniquely allows them to utilize nitrogen that is typically lost via excretion in the vast majority of insects (Cochran, 1985). These symbionts thus have a great deal of power in structuring the nutritional ecology and life history strategies of their hosts. Bacteroids damp out natural fluctuations in food availability, allowing cockroaches a degree of independence from the current food supply. An individual can engorge prodigiously at a single nitrogenous bonanza, like a bird dropping or a dead conspecific, then later, when these materials are required for reproduction, development, or maintenance, slowly mobilize the stored reserves from the fat body like a time-release vitamin. The legendary ability of cockroaches to withstand periods of starvation is at least in part based on this storage-mobilization physiology. The beauty of the system, however, is that stored urates are not only...

Measurement of lightdependent photosynthetic carbon fixation

On the basis of an equimolecular photosynthetic quotient (PQ 1), the light efficiency of photosynthesis could be calculated from the oxygen evolution data to be 5.0mgC (mg chla)-1 (mol photon)-1 m2. This is actually below the average of a large number of extrapolated values, which fall mainly in the range 6-18 mg C (mg chla)-1 (mol photon)-1 m2 (Harris, 1978). However, for over 50 years now, it has been possible to work directly in the currency of carbon, applying the so-called 14C method of Steemann Nielsen (1952)

Economic Impacts China

Aside from SARS-induced disruptions in the tourism and retail sectors, the fear-induced hoarding of essential goods and currency threatened fiscal liquidity. Moreover, calls by other countries for the quarantine of Chinese goods threatened China's export-driven manufacturing sector. If SARS had persisted and resulted in the disruption the production and supply chains, the increased risk profile associated with doing business in China would have led to a reduction in foreign investment and exports, damaging China's manufacturing sector. The end result would be a decline in the economic growth rate, on which the regime's legitimacy hinges.

Global Biogeochemical Cycling

The intensity of cycling is determined by two main factors. The first key factor is the amount of available biogenic elements, which will be considered below. The second and the main one is the power of the producers block. An amount of energy that can be put into the cycle per unit time by producers determines both the cycle intensity and length. The energetic concept of A. Lotka and H. Odum, based on the consideration of energy as a peculiar ecological 'currency' is very useful for understanding functioning of biogeo-chemical cycling.

Embracing the Complexity of Social Foraging

There have been numerous modifications of the original model. Relaxing the ideal and free assumptions of the original model can result in undermatching, or lower use of high-quality patches than expected based on resource distribution (Fretwell 1972 Abrahams 1986). Undermatching is a common finding in tests of the IFD (Kennedy and Gray 1993 but see Earn and Johnstone 1997). Other modifications include changing the form of competition and the currency assumed in the model. In this box I briefly review these ideas. Extensive reviews of IFD models and empirical tests can be found in Parker and Sutherland (1986), Milinski and Parker (1991), Kennedy and Gray (1993), Tregenza (1995), Tregenza et al. (1996), van der Meer and Ens (1997), and Giraldeau and Caraco (2000). The previous models all use net intake rate as the currency on which decisions are based. The IFD has also provided fertile ground for models exploring how animals balance energetic gain and safety (Moody et al. 1996 Grand and...

The paradox of deliberate simplicity

There obviously exists a whole series of other means that are, in turn, instruments and objectives and all reinforce each other reciprocally. We can think of reappropriating currency through the use of local currencies, melting currencies or non-convertible currencies (such as meal tickets or holiday vouchers).10

Benefits and Costs of Mutualism

Ecologists now recognize that one of the few generalizations that can be made about mutualisms is that nearly all of them involve both benefits and costs for each interacting species (Table 1). Mutualistic outcomes arise when the benefits of an interaction outweigh costs for both interacting species, such that the net effects of the interaction equal benefits minus costs. Currencies used as measures of benefits and costs often vary among mutualisms, but commonly include physiological or behavioral responses to various direct and indirect measures of growth, survival, and reproduction. Whatever currency is used to measure benefits and costs, they both are implicitly understood to ultimately affect reproduction and or survival, or possibly some energetic currency, as these are the fundamental units for ecological and evolutionary processes.

Manipulation as a Functional Adaptation

About the combinations likely to maximize the net reproductive output. This approach has been particularly fruitful in the area of foraging ecology (Stephens and Krebs, 1986). In the context of host manipulation by parasites, and assuming that the currency to be maximized is the transmission success of the parasite to its definitive host, optimality theory allows us to predict the optimal investment into manipulation of the intermediate host that a parasite should make (Poulin, 1994 Brown, 1999). From both the benefits and the costs of manipulation, the optimal level of manipulation can be derived as the one that achieves the greatest net benefits (benefits minus costs).

Expenditure per Progeny Further Reading

Natural selection recognizes only one currency successful offspring. Yet even though all living organisms have presumably been selected to maximize their own lifetime reproductive success, they vary greatly in exact modes of reproduction. Some, such as most annual plants, a multitude of insects, and certain fish like the Pacific salmon, reproduce only once during their entire lifetime. These 'big-bang' or semelparous reproducers typically exert a tremendous effort in this one and only opportunity to reproduce - in fact their exceedingly high investment in reproduction may contribute substantially to their own demise.

Biogeochemical Cycling

Organisms use the energy available to them as currency to acquire, concentrate, and organize chemical resources for growth and reproduction (Sterner and Elser 2002 see Chapter 4). Even sedentary organisms living in or on their material resources must expend energy to acquire resources against chemical gradients or to make these resources useable (e.g., through oxidation and reduction reactions necessary for digestion and assimilation). Energy gains must be greater than energy expenditures, or resource acquisition, growth, and reproduction cannot be maintained.

Problems of the rK Concept

The parameter K is not directly biologically interpretable. While r is the difference between per capita birth rate and death rate at very low population densities and can be directly related to the life history of individuals, K is quite a complex parameter it is meant to give the maximum number of individuals that a given environment can sustain under constant conditions. This phenomenological parameter cannot be determined in natural populations and is thus not directly biologically interpretable. In models, K is defined as the unstable or stable point of equilibrium where death rates equal birth rates (dN dt 0 in time-continuous models, Nt+1 Nt in time-discrete models). In real populations, such points of equilibrium are rarely constant over time. How K relates to life-history traits is indefinable, too. Stearns in 1977 wrote ''K is not a population parameter, but a composite of a population, its resources, and their interaction. Calling K a population trait is an artifact of...

Society and Landscape

Fig. 9.5 Needs and resources are connected by a semiotic interface the eco-field. All the eco-fields of an individual or a society create the cognitive landscape. The external component of this landscape represents the geographical landscape. Currency or other conventions can exclude the semiotic interface of the landscape from resource tracking causing landscape degradation. Currency or other conventions Currency or other conventions

Chemosynthesis Oxidation Reduction Reactions

To generate the vital cellular energy currency, ATP, nonphotosynthetic microorganisms use cellular processes to oxidize inorganic compounds (e.g., hydrogen gas, elemental sulfur, ammonia - to water, sulfate, and nitrate, respectively) or organic compounds (to CO2) and deliver the released electrons to the respiratory chain. The particular ecological context (habitat) of a microorganism determines which of a variety of possible final electron acceptors will be available for terminating its electron-transport chain. If the habitat is aerobic (rich in O2) then oxygen will be the dominant final electron acceptor, and

Marine Ecosystem Modeling Approaches

Marine Ecosystem

Marine pelagic ecosystem models are similar to ecological models that have been applied to lakes in many respects (see Lake Models) that is, they are typically constructed in terms of a set of ordinary differential equations that express the time rate of change of the mass in a set of state variables that represent different components of the pelagic ecosystem. However, there are some important differences between marine and freshwater ecosystems and models. In particular, phosphorus is much more likely to limit primary production in lakes as opposed to nitrogen in marine systems. Thus, nitrogen is often used as the primary model currency in marine modeling studies whereas phosphorus is a more obvious choice for lakes. But phosphorus is used in The state variables in these models vary in response to source and sink terms that for the living components represent growth, mortality, and sinking-out of the model domain. For example, a simple class of N-based pelagic ecosystem model...

Fixation of atmospheric nitrogen in mutualistic plants

Nodule Meristem Cells Plants

The costs and benefits of this mutualism need to be considered carefully. From the plant's point of view, we need to compare the energetic costs of alternative processes by which supplies of fixed nitrogen might be obtained. The route for most plants is direct from the soil as nitrate or ammonium ions. The metabol-ically cheapest route is the use of ammonium ions, but in most soils ammonium ions are rapidly converted to nitrates by micro-bial activity (nitrification). The energetic cost of reducing nitrate from the soil to ammonia is about 12 mol of adenosine triphosphate (ATP) per mol of ammonia formed. The mutualistic process (including the maintenance costs of the bacteroids) is energetically slightly more expensive to the plant about 13.5 mol of ATP. However, to the costs of nitrogen fixation itself we must also add the costs of forming and maintaining the nodules, which may be about 12 of the plant's total photosynthetic output. It is this that makes nitrogen fixation...

Single trait optimization reproductive lifespan

Some organisms display 'big bang' reproduction, after which they die (semelparous), while others reproduce more consistently over a long period of time (iteroparous) (Figure 4.1). The question of whether to be semelparous or iteroparous is a rather simple one because there are only two strategies to consider. We simply have to calculate, for any given organism, which should lead to the highest fitness. Charnov and Schaffer (1973) produced a simple calculation to help, which is derived by asking which strategy produces the highest rate of population growth (their fitness currency). They assumed a plant could display either an annual or a perennial strategy. The fitness of the annual is determined by its own survival to maturity and fecundity once mature. However, that of a perennial is determined by both its juvenile survival, annual fecundity, and adult survival from year-to-year. These are the constraints. If annuals are to be more fit than perennials, annuals must have higher annual...

Evolution of Biogeochemical Cycling

The abundance of carbon gas is quite important for development of vegetation. It is the main biomass-gener-ating substance, a peculiar ecological 'currency', for which, similarly to the real one, little inflation should take place. Volcanic activity permanently adds the gas to the cycling and creates such inflation. One of the pessimistic ecological forecasts is connected with gradual decrease of geological activity. When the carbonic gas entry to the biosphere finishes, 'stagnation' of the biogeo-chemical cycling is inevitable.

The Structure of This Book

This book is divided into three sections (a) Relevant Temporal Theory (Chapters 15), (b) Statistics of Time (Chapters 6 and 7), and (c) Temporally Focused Case Studies (Chapters 8-14). In the first chapter of section 1, Relevant Temporal Theory, Bissonette makes the argument that an enhanced understanding of animal response to resource availability may be possible if two elements are added to the standard, single currency quantity approach. The first element relates to measuring resource quality and requires adding an additional currency to our ecological ledger book. The second element incorporates the idea of temporal discontinuity in resource quantity and quality. Bissonette suggests that a broader incorporation of these two elements into wildlife ecology will enhance our understanding of animal response to resource availability at both small and larger spatial extents. In the second chapter, Ostfeld and Keesing argue that pulses of resources are major bottom-up drivers in...

Growth and Impact of Tourism

By the mid-1990s, tourism had become the largest industry in the world. Each year, more than half a billion people spend some time as international tourists, while uncounted millions more travel domestically. The advance of tourism has reached farther and deeper into more isolated areas of the planet than all the invasions and migrations of history. Many areas are becoming dependent on the tourism sector as their primary source of income. The infrastructure built to facilitate this transnational flow of people, goods, services, and currency includes air, rail, land, and sea carriers, roads and airports, hotels, restaurants, and resort complexes. Tourist ministries, international agencies, and entrepreneurs also restore heritage sites, promote commercial handicraft enterprises, and commodify nature and culture for mass consumption.

Mortality and morbidity patterns in modern conflicts

In addition to causing the displacement of millions of people, conflict also takes an important toll on both agrarian and industrial economies. Although there is some dispute as to the magnitude of the impact of conflict on national economies, some estimate that a 15-year civil war would reduce gross domestic product by as much as 30 percent (Collier, 1999 Imai and Weinstein, 2000). Local economies may be even more devastated, with more immediate consequences for the health of the population - particularly through food shortages resulting in a high prevalence of under-nutrition. In rural areas, where a relatively high proportion of food is derived from subsistence farming, farmers may be physically unable to plant as much as they might in the absence of conflict. Where even small-scale commercial food production is a way of life, farmers may be obstructed from bringing their produce to market. Resulting scarcities can be responsible for higher food prices, which, combined with the...

Eukaryotic Cells

Mitochondrion-independent pathway (glycolysis and fermentation) uses glucose molecules which are oxidized to two molecules of pyruvate, with a net yield of two adenosine triphosphate (ATP) molecules. Pyruvate can be oxidized further to smaller molecules (such as acetate, lactate or ethanol) to yield further energy. Glycolysis does not require oxygen and does not yield CO2. The glucose and other substrates for this set of reactions is obtained from digestion of food, from de novo synthesis from other molecules, or transport into the cell in species where that is possible. Normally, it is assumed that most free-living protozoa do not transport glucose through the membrane, but acetate and other soluble 2-3 carbon molecules often are. In species with mitochondria, a further sequence of reactions is possible which produce > 30 ATP molecules from each glucose equivalent. The mitochondrion provides the cell with enzymes for the Krebs cycle, the electron transport chain and oxidative...

Consumer Choice

Stephens and Krebs's chapter 5, entitled The economics of choice, considered situations in which foragers face trade-offs. In such situations, increasing the gain of one thing important for fitness (say, food intake) compromises the attainment of another (say, safety). The chapter provided a brief introduction to microeconomic consumer choice theory, which provides a framework for analyzing trade-off problems by assigning utility so that their value can be measured on a common scale. Animal psychologists had used this approach in operant conditioning experiments, with some success, to study the choices made by animals between, for example, different food types obtainable by varying amounts of bar pressing, or different delays to reinforcements of different sizes. Behavioral ecologists had far less success with this theoretical structure because it was difficult to express the fitness value of very different things (e.g., food and safety) in a common currency. When Foraging Theory was...

Ecosystem ecology

Imposed on matter circulation by the articulation of flows, and it is quantified by the AMI. As said before, ecological flow networks are ecosystem images. Given the high number of actors that play the game in an ecosystem, processes such as feeding, decay, excretion, and so forth give rise to a vast array of matter and energy passages between species or nutrient pools. This multiplicity of exchanges can be captured by a network representation, whose connections are in terms of some currency that is either material (nitrogen, carbon, phosphorus) or energetic (joule). In this scenario, it is straightforward to ask in what way ascendency can contribute to characterize ecosystem status. Each arrow in network represents a flow of a given currency and the magnitude associated to the link quantifies the intensity of that passage. The summation of all the flows in a network yields the total amount of material or energy that the ecosystem handles. This quantity estimates the level of activity...

Sustainability

Accordingly, cities also can be seen as ecological networks. By studying energy matter exchanges through network architectures, ecologists have been able to unveil how efficiently an ecosystem uses energy and the major constraints which curb the maximization of this efficiency. In addition, they have been able to determine the ecosystem potential for development and ability to maintain structure and functions over time in the face of external stress. Since achieving sustainability critically depends upon making better use of natural resources and creating new patterns of development in which protection and stability are maximized, the ecosystem perspective is more than a simple conceptual analogy. Criteria and tools which form the apparatus of ecosystem analysis can thus be applied to human systems to obtain clues for sustainable strategies. In natural ecosystems, there can be as many networks as currencies in use (carbon, nitrogen, phosphorus, and so forth) in urban ecosystems, this...

Beyond Kcals

Early applications of foraging models, especially the diet breadth model, adopted a straightforward energy currency to measure the costs and benefits of options in the alternative set. The value of a moose was the weight of its edible tissue represented as kcals. This is consistent with a prime methodological predilection of behavioral ecologists (Winterhalder 2002a) begin simply. Once you understand how the A more radical variation on currency is evident in models devised to help explain an anomaly in early field studies of foraging behavior although each sex often could do better by harvesting the same set of resources, men sometimes specialize on large game and women on plants and small animals (Hill et al. 1987), each at a cost to their potential foraging efficiency. The show-off (Hawkes and Bliege Bird 2002) and costly signaling (Smith et al. 2003) models assume that resource values and hence their patterns of acquisition and distribution will sometimes be predicated on the...

Human Capital

The concept of human capital, however, was not explicitly elaborated until the twentieth century. The term gained currency through the work of the neo-classical Columbia-Chicago school of labor economics in the 1960s, particularly through the work of Mincer (1958), Schultz (1961, 1962), and Becker (1964). The concept has since played a prominent role in the evolution and application of both labor economics and development economics.

Conclusions

Accordingly, if the environment does not affect the population in a systematic manner, all types of biological populations show exponential growth. Traditionally, ecologists have treated populations with discrete generations differently from those with overlapping generations. Difference equations such as 1.4 and 1.5 have been used in the first case. By contrast differential equations (1.3) and their solved forms (1.8) have been employed to describe populations with overlapping generations. In both cases we use the finite rate of increase, l, or the intrinsic rate of increase, r, as a common currency for comparing population growth potentials.

Territories

Defend territories after reproduction has ceased for the year. If bears, hummingbirds, and other animals use food as an index for the potential to produce offspring, then food can legitimately be considered to be at least a proximately limiting resource. Fitness is the ultimate currency in biology, and fitness may be affected by one or more limiting resources that need not be offspring or other direct components of reproduction. Evolution via natural selection requires heritable variation that affects reproductive output among individuals in a population. The effects can be via offspring, or they can be via food, nest sites, tunnel systems, or other potentially limiting resources.

Structural Domain

Odum created another set of symbols to model systems based on the energy flows through them. He called them energy diagrams, and used six main icons shown in Figure 6. All systems were described in terms of energy, assuming that for all variables and processes we can calculate the 'embodied' energy. In this case energy works as a general currency to measure all processes and 'things'.

Theoretical Basis

In addition to empirical studies, attempts have been made to estimate theoretical migration speeds of particular species from knowledge of their likely flight speeds, and rates of fuel deposition and use (Boxes 8.1 and 8.2, Figure 8.1 Hedenstrom & Alerstam 1998, Alerstam 2003). Flight speeds can be taken from radar measurements (ideally corrected to their still-air values, Bruderer & Boldt 2001) or from theoretical estimates (Pennycuick 1969, 1975). Rates of fuel deposition can be measured from the repeat weighing of individual birds before migration or at stopover sites. Such measurements are usually taken over periods of days, so include sleeping and other non-feeding times. Rates of fuel use during flight can be measured only with difficulty, and are more easily obtained as theoretical estimates, based on body mass and other features of the bird (see Appendix 5.1). Rates of fuel (energy) gain and loss are best expressed in some common currency, such as multiples of BMR (basal...

Cooperate or else

There is a sense of 'fair play' in human societies. For example, people regularly challenge others when they 'queue jump' (typically but not always when they have jumped ahead of the challenger, indicating that self-interest is part of the motivation). In the well-known 'ultimatum' game in economics, two players A and B have to agree on how a monetary reward has to be shared.48 Player A (the proposer) has one chance to suggest how the money is to be shared (e.g. 60 to player A, 40 to player B), but player B (the responder) can accept or reject the proposed division. If the bid is rejected, then both receive nothing, but if the bid is accepted then the proposal is implemented. What would you propose if you were given 100 dollars (or whatever your local currency) The logical optimum is to offer the responder an almost negligible amount (1 cent say, because 1 cent is better than nothing). Perhaps you would be rather more generous and keep 90 yourself, providing a 10 'sweetener' to keep...

Ecosystem Services

The idea of paying for ecosystem services has been gaining momentum. Yet, because ecosystem services are typically not sold in markets, they usually lack a market value. Given the value of natural capital, nonmarket valuation approaches are being developed by economists and ecologists to account for ecosystem services in decision-making processes. The notion being that economic valuation gives decision makers a common currency to assess the relative importance of ecosystem processes and other forms of capital.

Tremulation

Eremochrysa (including Chrysopiella) and Chrysoperla are two closely related members of the Mallada species group (Figure 10.2). Polite male-female duetting using identical signals is characteristic of both genera. Although Eremochrysa seems to have retained simple tremulation signals (e.g. E. minora Banks see Henry and Johnson, 1989), Chrysoperla has become the tremulation champion of the green lacewings. Tremulation signals have been found whenever looked for in this genus, in representatives of each of the four recognised species groups (Brooks, 1994). As in Chrysopa and Eremochrysa, their low-frequency signals are sexually monomorphic and organised into repeating volleys of abdominal vibration, which in turn are grouped into identical SRUs which serve as the currency of exchange during heterosexual duets (Figure 10.3). A SRU consists of just one, several or many volleys and may include more than one type of volley, e.g. of different durations or pitches (Figure 10.4, Figure...

Network Models

The complex of flows related to a compartment in a trophic network may be graphically indicated by a figure whose nodes represent the biomass of various biological groups and whose arrows represent flows of matter or energy, usually called currency, that belong to one of the processes listed above. Figure 5.1 shows how such flows may occur. Tir Th between compartments , from outside to a compartment (import) ,. out of the system in form of currency that is still usable (export) Rt out of the system in form of non-usable currency (dissipation of energy also called costs of maintenance, a synonym for respiration in an environmental system).

Diet Breadth Models

Resources are ranked by their post-encounter return rates, which are simply the measure of energy return (as measured by some currency) after search and processing costs have been considered (Kelly 1995, 78). The decision to take a resource upon encounter thus depends on the quality of the resource, its density (which affects search costs), and its processing costs (which affects return rates). The key assumption of the model is that the forager is attempting to maximize energy returns. As Kelly (1995, 86-87) notes, the model makes a number of important observations about the composition of human diets resource abundance alone cannot be used to predict the presence of a resource in a diet, as search costs for high-return-rate resources decrease, diets will expand regardless of the abundance of

National

Deforestation to finance development can produce short-term benefits for the developing country. The benefits can include easing of debt crisis obtaining currency for foreign exchange securing national boundaries attracting international investment easing of social problems in some overcrowded areas (due to transmigration programs) and increasing political control in frontier areas (benefit for governing party, not for opposition parties).

Currencies

IBMs contain much more information about the state of all the individuals and their environment than we can process. We thus need to aggregate this information into one or more aggregated system-level state variables. We need to define currencies (often also referred to as 'indices') that allow us to characterize a single run of the model over a certain time horizon by, ideally, one single number. This number is then used as a currency to compare different parametrizations or formulations of the model. For example, in models addressing extinction risk of small populations, the risk of extinction over a certain time horizon is such a currency or, if we want to compare model output to an observed time series, we can use root mean square deviation as a currency. In general, any quantitative measure of how good a certain observed pattern is reproduced is a good currency. Often, it is not clear from the outset which currency allows for deepest insights, so currencies are experimental and...

The Ethogram

Early in any study a researcher must classify the behavior patterns to be documented. This classification is an ethogram, and constitutes a dictionary of the researcher's language without an ethogram, meaning is not fixed. Ethograms have become unpopular because (like dictionaries) they take up large amounts of space and are dull to read. However, they are vital reference material for those wishing to assess a study's conclusions critically. As a corollary, it is not uncommon for the brunt of reviewers' comments now to fall on statistics rather than data acquisition, reflecting the same shift in emphasis away from data, and onto interpretation, as the hard currency of behavioral science. By analogy, consider the shift from real objects to paper representations in financial systems 50 years on, would you rather find a stash of gold or war bonds

Pollinator economics

Its relative contribution to the forager's fitness. However, animals probably cannot evaluate the fitness consequences of different foraging options rather, they must assess opportunities based on the proximate benefits and costs associated with current physiological and ecological conditions. Often, the behavior of experienced feeding animals maximizes a single variable, or foraging currency, that integrates foraging benefits and costs (Stephens & Krebs 1986). Such behavior bears diverse consequences for pollination, because it affects a pollinator's choice of plant species (e.g., Rasheed & Harder 1997a), choice of individual plants (e.g., Heinrich 1979 Waser & Price 1983 Thomson 1988), and behavior on those plants (e.g., Galen & Plowright 1985 Hodges 1985 Rasheed & Harder 1997b). In addition to handling costs, the relevant foraging currency must incorporate the time and energy expended on other activities. These additional costs always include travel within and...

Inflorescences

We conclude by emphasizing two essential features of the selection of floral design and display. The first feature arises from recognition that floral and inflorescence characteristics create the environment within which pollinators tend to maximize a specific foraging currency. Because of this role, plant evolution could improve foraging benefits or alleviate costs however, it will do so only to the extent that such changes promote plant mating (e.g., Harder & Cruzan 1990 Harder & Barclay 1994). Therefore, the evolutionary relevance of specific floral or inflorescence traits must extend beyond their impact on pollinator behavior to realized mating outcomes. The second feature deserving emphasis is that, despite the key role of individual flowers in controlling pollen exchange with pollinators, mating fundamentally involves entire plants. For example, contrary to the expectation that a plant's pollen export increases mono-tonically with the number of pollinator visits received...

Territoriality

Early theoretical work on territorial behavior was based on Brown's idea of economic defendability whether animals act to defend a territory should depend on whether the benefits gained from the territory outweigh the costs associated in maintaining it. This is difficult to test because the costs of defense and the benefits to be gained from a territory are not easy to measure in the same currency. Consequently, explicit tests of this approach have either been qualitative or based on relatively simple systems (e.g., nectar foragers like hummingbirds, where costs and benefits can both be measured in energy).

Reproductive value

Natural selection favors those individuals that make the greatest proportionate contribution to the future of the population to which they belong. All life history components affect this contribution, ultimately through their effects on fecundity and survival. It is necessary, though, to combine these effects into a single currency so that different life histories may be judged and compared. A number of measures of fitness have been used. All the better ones have made use of both fecundity and survival schedules, but they have done so in different ways, and there has often been marked disagreement as to which of them is the most appropriate. The intrinsic rate of natural increase, r, and the basic reproductive rate, R0 (see above) have had their advocates, as has 'reproductive value' (Fisher, 1930 Williams, 1966), especially reproductive value at birth (Kozlowski, 1993 de Jong, 1994). For an exploration of the basic patterns in life histories, however, the similarities between these...

Qenetic Diversity

Genetic diversity refers to any variation in the nucleotides, genes, chromosomes, or whole genomes of organisms. This is the fundamental currency of diversity (Williams and Humphries, 1996) and the basis for all other organismal diversity. Approximately 1 billion different genes are recognized from all the known species on earth (World Conservation Monitoring Center, 1992). But not all species have the same number of genes. The potential genetic diversity of a species can be measured by the total number and type of genes present within its entire DNA or genome. However, a greater total number of genes might not correspond with a greater observable complexity in the anatomy and physiology of the organism (that is, greater phenotypic complexity). For example, the genome of the cultivated subspecies of rice, Oryza sativa L. ssp. indica, is estimated at 46,022 to 55,615 genes (Yu et al., 2002), and the total size of the human genome is currently predicted to be not much larger, at...

Ecotourism

Big-game safaris and trophy hunting are major businesses in eastern and southern Africa. Game parks, wildlife reservations, and nature preserves attract substantial foreign currency. Africans are building more lodges and modern guest facilities, in a delicate balancing act between preservation and overuse of land and water. The endangered large mammals of Africa are at the mercy of governments, local needs, tourists, and the travel industry. Collaboration and cooperative planning among the various stakeholders need to support set-asides of land and water, conservation activi

Provisioning

What should central place foragers maximize Kacelnik (1984) compared the load sizes that his European starlings collected with the predictions offour objective functions, or currencies. The currency he called delivery is the total delivery of prey energy to the nest on each trip, divided by round-trip time. The currency called yield subtracts from the total delivery the amount ofener-gy spent by the parent on each trip, all divided by round-trip time while that called family gain further subtracts the energy spent by the young during each trip. These three closely related measures are all rates and are all expressed in units ofwatts ( joules per second). The fourth currency is somewhat different it takes the total delivery and divides by the energy expended by the parent. We call this currency efficiency (joules delivered to the nest per joule expended by the parent), and it has no units. Statistically, the family gain currency matched Kacelnik's observations best, but all four...

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