Influence of budget and reinforcement location on risk-sensitive preference

Little is known about the effect that procedural variables have on risk-sensitive preference. This study assessed the effect of procedural variables on pigeons' choice between a fixed and variable amount of reinforcement (amount risk) and, in a separate condition, between a fixed and variable delay until reinforcement (delay risk). Experiment 1 investigated the impact of water reinforcement and risk dimension when pigeons were in a restrictive budget, where access to water was less than that necessary to maintain current body weight, and a condition where the pigeons had ample access to water. Pigeons exhibited a greater tendency to prefer the variable alternative for delay risk than for amount risk in both restrictive and ample budgets. Varying water budget had no effect on risk preference. Experiment 2 investigated the influence of water reinforcer location while in a restrictive budget, in which reinforcers were delivered to a single location, two distinct locations, or a randomly selected location. With amount risk, pigeons were risk averse when reinforcers were delivered in separate or random locations and were indifferent to risk when delivered to a single location. With delay risk, pigeons were generally risk prone with no effect from reinforcement location. The finding that pigeons were risk averse when reinforcers were delivered to separate locations and were indifferent to risk when delivered to a single location offers a methodological explanation to the inconsistent findings in the literature with amount risk.     

Amount-dependent temporal discounting?

Amount-dependent temporal discounting has been demonstrated for human choice between outcomes differing in amount and delay. In the only study to date with non-humans, Grace reported no evidence for amount-dependent temporal discounting with pigeons in a concurrent-chains procedure. The present experiments repeated Grace's procedure but with modifications to enhance the discrimination between small and large magnitude outcomes. In Experiment 1, sensitivity of pigeons' initial-link choice to the terminal link delay ratio was greater with large reinforcer durations in the terminal links than with small reinforcer durations. This result is consistent with a greater rate of temporal discounting for larger reinforcers (the reverse of the result for humans), but can also be explained as enhanced discrimination of delay ratios with larger reinforcer durations. The results of a second experiment supported Grace's conclusion that amount-dependent temporal discounting does not characterize pigeons' choice in concurrent chains. Because reinforcer amount was held constant between choice alternatives in the present experiments and that of Grace, but varied in the human studies, our results question whether prior demonstrations of amount-dependent discounting reflect the effects of reinforcer delay or of reinforcer amount. Differences in the procedures used to study discounting in humans (titration procedures) and non-humans (concurrent chains) may contribute to the divergent results across species.     

The use of specialisation indices to predict vulnerability of coral‐feeding butterflyfishes to environmental change

In the absence of detailed assessments of extinction risk, ecological specialisation is often used as a proxy of vulnerability to environmental disturbances and extinction risk. Numerous indices can be used to estimate specialisation; however, the utility of these different indices to predict vulnerability to future environmental change is unknown. Here we compare the performance of specialisation indices using coral‐feeding butterflyfishes as a model group. Our aims were to 1) quantify the dietary preferences of three butterflyfish species across habitats with differing levels of resource availability; 2) investigate how estimates of dietary specialisation vary with the use of different specialisation indices; 3) determine which specialisation indices best inform predictions of vulnerability to environmental change; and 4) assess the utility of resource selection functions to inform predictions of vulnerability to environmental change. The relative level of dietary specialisation estimated for all three species varied when different specialisation indices were used, indicating that the choice of index can have a considerable impact upon estimates of specialisation. Specialisation indices that do not consider resource abundance may fail to distinguish species that primarily use common resources from species that actively target resources disproportionately more than they are available. Resource selection functions provided the greatest insights into the potential response of species to changes in resource availability. Examination of resource selection functions, in addition to specialisation indices, indicated that Chaetodon trifascialis was the most specialised feeder, with highly conserved dietary preferences across all sites, suggesting that this species is highly vulnerable to the impacts of climate‐induced coral loss on reefs. Our results indicate that vulnerability assessments based on some specialisation indices may be misleading and the best estimates of dietary specialisation will be provided by indices which incorporate resource availability measures, as well as assessing responses of species to changes in resource availability.     

Risk-sensitive foraging in rats: the effects of response-effort and reward-amount manipulations on choice behavior

The literature on risk-sensitive foraging theory provides several accounts of species that fluctuate between risk-averse and risk-prone strategies. The daily energy budget rule suggests that shifts in foraging strategy are precipitated by changes in the forager's energy budget. Researchers have attempted to alter the organism's energy budget using a variety of techniques such as food deprivation, manipulation of ambient temperatures, and delays to food reward; however, response-effort manipulations have been relatively neglected. A choice preparation using a wheel-running response and rats examined risk-sensitive preferences when both response effort and reward amounts were manipulated. Concurrently available reinforcement schedules (FI/60 and VI/60) yielded equivalent food amounts per unit time in all treatments. Two levels of response effort (20 or 120 g tangential resistance) and two levels of reward amount (three or nine pellets) were combined to form four distinct response-effort/reward-amount pairings. Increasing reward amounts significantly shifted choice toward the FI schedule in both response-effort conditions. The incidence of choice preference and the magnitude of shifts in choice were greater for the high response-effort conditions than for the low response-effort conditions. Implications of the significant interaction between response effort and reward amount are discussed in terms of a general energy-budget model.     

The feeding ecology of a selective browser, the giraffe (Giraffa camelopardalis tippelskirchi)

A herbivore can manipulate certain factors of its feeding behaviour in order to achieve the metabolic requirements for reproduction. These factors include the choice of habitat in which to feed, the selection criteria for choosing food items, and the time allocated to feeding or devoted to other energy-consuming activities. The manipulation of these behavioural factors by giraffe in the Serengeti National Park, Tanzania is analysed in relation to the seasonal changes in the quantity and quality of the food resource.
The pattern of habitat choice demonstrates seasonal movements by giraffe across the drainage catena in response to differences in the rates of browse production between woodland types.
Giraffe show a positive inter-specific selection for the minor food items with a low available biomass, and utilize the dominant species in approximate proportion to their availability. Plant part selection is shown for the flushing shoots with very high protein contents.
Selection criteria include a significant selection for phosphorus, and in the dry season a selection for high energy material by breeding females. With the high nutritional qualities of the diets, negative criteria such as secondary chemical compounds are important. Patterns of browse selection are correlated with inter-specific seasonal changes in plant phenology.
Giraffe devote more time to foraging as the biomass and quality of the food declines in the dry season. Energy is conserved by minimizing energy-consuming activities at the most demanding times of day.     

Fluctuations in prey availability and food limitation for a terrestrial salamander

Summary A Virginia population of the forest-dwelling salamander Plethodon cinereus was repeatedly sampled over four years to determine volume and number of prey ingested under varying conditions of moisture and temperature. The proportion of the population on a daily negative energy budget was estimated by comparing actual foraging success with laboratory determined values of energetic requirements at various temperatures.Prey became limited in availability during rainless periods, apparently because salamanders were not able to forage in dry leaf litter. Foraging success increased with increasing rainfall. Food was a limiting resource for a majority of the population on most sampling days, as determined by energy budget analysis. While ambient moisture regulated food availability, ambient temperature set the metabolic requirements and assimilation efficiencies for the population. Consequently, food was more limiting on dry, warm days and less so on wet, cool days. These data support the hypothesis that intraspecific competition frequently occurs for a food resource that is periodically limited in availability.     

Starvation reveals the cause of infection-induced castration and gigantism

Parasites often induce life-history changes in their hosts. In many cases, these infection-induced life-history changes are driven by changes in the pattern of energy allocation and utilization within the host. Because these processes will affect both host and parasite fitness, it can be challenging to determine who benefits from them. Determining the causes and consequences of infection-induced life-history changes requires the ability to experimentally manipulate life history and a framework for connecting life history to host and parasite fitness. Here, we combine a novel starvation manipulation with energy budget models to provide new insights into castration and gigantism in the Daphnia magna–Pasteuria ramosa host–parasite system. Our results show that starvation primarily affects investment in reproduction, and increasing starvation stress reduces gigantism and parasite fitness without affecting castration. These results are consistent with an energetic structure where the parasite uses growth energy as a resource. This finding gives us new understanding of the role of castration and gigantism in this system, and how life-history variation will affect infection outcome and epidemiological dynamics. The approach of combining targeted life-history manipulations with energy budget models can be adapted to understand life-history changes in other disease systems.     

Umbilical cord clamping and preterm infants: a randomised trial.

OBJECTIVE--To investigate the clinical effects of regulating umbilical cord clamping in preterm infants. DESIGN--A prospective randomised study. SETTING--The Queen Mother''s Hospital, Glasgow. SUBJECTS--36 vaginally delivered infants over 27 and under 33 weeks'' gestation. INTERVENTION--Holding the infant 20 cm below the introitus for 30 seconds before clamping the umbilical cord ("regulated" group, 17 patients), or conventional management ("random" group, 19 patients). MAIN OUTCOME MEASURES--Initial packed cell volume, peak serum bilirubin concentrations, red cell transfusion requirements, and respiratory impairment (assessed by ventilatory requirements, arterial-alveolar oxygen tension ratio over the first day in ventilated infants, and duration of dependence on supplemental oxygen). RESULTS--There were statistically significant differences between the two groups in mean initial packed cell volume (regulated group 0.564, random group 0.509) and median red cell transfusion requirements (regulated group zero, random group 23 ml/kg). 13 infants from each group underwent mechanical ventilation and showed significant differences in mean minimum arterial-alveolar oxygen tension ratio on the first day (regulated group 0.42, random group 0.22) and in median duration of dependence on supplemental oxygen (regulated group three days, random group 10 days). Differences in final outcome measures such as duration of supplemental oxygen dependence and red cell transfusion requirements were mediated primarily through arterial-alveolar oxygen tension ratio and also packed cell volume. CONCLUSIONS--This intervention at preterm deliveries produces clinical and economic benefits.     

Skeletal muscle hypertrophy in response to isometric, lengthening, and shortening training bouts of equivalent duration.

Movements generated by muscle contraction generally include periods of muscle shortening and lengthening as well as force development in the absence of external length changes (isometric). However, in the specific case of resistance exercise training, exercises are often intentionally designed to emphasize one of these modes. The purpose of the present study was to objectively evaluate the relative effectiveness of each training mode for inducing compensatory hypertrophy. With the use of a rat model with electrically stimulated (sciatic nerve) contractions, groups of rats completed 10 training sessions in 20 days. Within each training session, the duration of the stimulation was equal across the three modes. Although this protocol provided equivalent durations of duty cycle, the torque integral for the individual contractions varied markedly with training mode such that lengthening > isometric > shortening. The results indicate that the hypertrophy response did not track the torque integral with mass increases of isometric by 14%, shortening by 12%, and lengthening by 11%. All three modes of training resulted in similar increases in total muscle DNA and RNA. Isometric and shortening but not lengthening mode training resulted in increased muscle insulin-like growth factor I mRNA levels. These results indicate that relatively pure movement mode exercises result in similar levels of compensatory hypertrophy that do not necessarily track with the total amount of force generated during each contraction.     

Investment in defense and cost of predator-induced defense along a resource gradient

An organism’s investment in different traits to reduce predation is determined by the fitness benefit of the defense relative to the fitness costs associated with the allocation of time and resources to the defense. Inherent tradeoffs in time and resource allocation should result in differential investment in defense along a resource gradient, but competing models predict different patterns of investment. There are currently insufficient empirical data on changes in investment in defensive traits or their costs along resource gradients to differentiate between the competing allocation models. In this study, I exposed tadpoles to caged predators along a resource gradient in order to estimate investment in defense and costs of defense by assessing predator-induced plasticity. Induced defenses included increased tail depth, reduced feeding, and reduced swimming activity; costs associated with these defenses were reduced developmental rate, reduced growth, and reduced survival. At low resource availability, these costs predominately resulted in reduced survival, while at high resource availability the costs yielded a reduced developmental rate. Defensive traits responded strongly to predation risk, but did not respond to resource availability (with the exception of feeding activity), whereas traits construed as costs of defenses showed the opposite pattern. Therefore, defensive traits were highly sensitive to predation risk, while traits construed as costs of defense were highly sensitive to resource allocation tradeoffs. This difference in sensitivity between the two groups of traits may explain why the correlation between the expression of defensive traits and the expression of the associated defense costs was weak. Furthermore, my results indicate that genetic linkages and mechanistic integration of multiple defensive traits and their associated costs may constrain time and resource allocation in ways that are not addressed in existing models. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Adverse changes in mood and cognitive performance of house officers after night duty.

The effects of long hours of work by junior doctors are the subject of growing concern. Experimental investigations of the effects of night duty on young doctors are few and have given inconclusive results. To measure the effects of long hours of work and reduced sleep on cognitive performance and mood 20 house officers (14 men and six women; mean age 25, range 24-35) were examined for 35 minutes in one session towards the end of a normal working day and in a second session at the same time after working for up to 31 continuous hours with reduced sleep. The order of the sessions was counterbalanced across the subjects. Each session comprised a series of cognitive tests: choice reaction time, vigilance reaction time, and haptic sorting tests and completion of a profile of mood states and a general questionnaire. After night duty there was a significant slowing in cognitive processing together with a decline in reaction times in the vigilance test. Loss of sleep contributed only to increased variability in the choice reaction time. Significant deleterious changes in mood after night duty occurred in all the mood scales after night duty. Continuous working may adversely affect the cognitive function and mood to the detriment of the welfare of doctors and their patients.     

Post-rest adaptation of electrical and mechanical activity in the isolated guinea-pig papillary muscle

In guinea-pig papillary muscle the time course of the changes in contractile force and action potential duration (APD) were studied after periods of rest of variable duration. After a long period of rest, the force of contraction adapted to pre-rest control values in a monophasic manner whereas the time-course of the APD was clearly biphasic. The post-rest adaptation of the APD could be described mathematically by a simple model, which considers the action potential duration during steady state as the sum of a resting value (APDR) plus a lengthening effect of activation (LEA) minus a shortening effect of activation (SEA). LEA and SEA are assumed to occur immediately, with each excitation and to decay continuously. During repetitive stimulation, both effects will accumulate. Using the constants found for the post-rest adaptation of the APD, the steady-state frequency-dependence of the APD could also be described with this model.     

Pointing movement in short and long-term exposure to hypoxia.

Kinematics variables of pointing movements where assessed in five adult subjects exposed acutely (30 min) and chronically (10 days) to a low O2 mixture (13.5% O2 in N2). Amplitude of displacement did not vary in both experimental conditions but movement duration markedly increased compared to pre and post exposure conditions. While in acute hypoxia the times of acceleration and deceleration are almost equal, in chronic hypoxia deceleration time exceeded of 100 ms the time of acceleration. The time from the peak acceleration to the peak of deceleration ("switch" time) increased in both experimental conditions and was about 50% of the movement duration. This time lengthening at hypoxia may be explained either by alteration of propioceptive loops or by a different strategy elaborated by the CNS to generally slow accurate movements.     

Metabolic traits predict the effects of warming on phytoplankton competition

Understanding how changes in temperature affect interspecific competition is critical for predicting changes in ecological communities with global warming. Here, we develop a theoretical model that links interspecific differences in the temperature dependence of resource acquisition and growth to the outcome of pairwise competition in phytoplankton. We parameterised our model with these metabolic traits derived from six species of freshwater phytoplankton and tested its ability to predict the outcome of competition in all pairwise combinations of the species in a factorial experiment, manipulating temperature and nutrient availability. The model correctly predicted the outcome of competition in 72% of the pairwise experiments, with competitive advantage determined by difference in thermal sensitivity of growth rates of the two species. These results demonstrate that metabolic traits play a key role in determining how changes in temperature influence interspecific competition and lay the foundation for mechanistically predicting the effects of warming in complex, multi‐species communities.     

Effects of temperature elevation on a field population of Acyrthosiphon svalbardicum (Hemiptera: Aphididae) on Spitsbergen

A manipulation experiment was carried out on a field population of the aphid Acyrthosiphon svalbardicum near Ny Ålesund, on the high arctic island of Spitsbergen, using cloches to raise temperature. An average rise in temperature of 2.8 deg. C over the summer season markedly advanced the phenology of both the host plant Dryas octopetala and the aphid. Advanced aphid phenology, with concomitant increases in reproductive output and survival, and successful completion of the life-cycle led to an eleven-fold increase in the number of overwintering eggs. Thermal budget requirements in day degrees above 0°C were calculated for key life-cycle stages of the aphid. Temperature data from Ny Ålesund over the past 23 years were used to calculate thermal budgets for the field site over the same period and these were compared with the requirements of the aphid. Each estimated thermal budget was then adjusted to simulate the effect of a +2, +4, and –2deg. C change in average temperature on aphid performance. This retrospective analysis (i) confirms that the life-cycle of A. svalbardicum is well suited to exploit higher summer temperatures, (ii) indicates that the annual success of local populations are sensitive to small changes in temperature and (iii) suggests that the aphid is living at the limits of its thermal range at Ny Ålesund based on its summer thermal budget requirements.     

Resource availability modulates the effect of body size on reproductive development

Within-species variation in animal body size predicts major differences in life history, for example, in reproductive development, fecundity, and even longevity. Purely from an energetic perspective, large size could entail larger energy reserves, fuelling different life functions, such as reproduction and survival (the “energy reserve” hypothesis). Conversely, larger body size could demand more energy for maintenance, and larger individuals might do worse in reproduction and survival under resource shortage (the “energy demand” hypothesis). Disentangling these alternative hypotheses is difficult because large size often correlates with better resource availability during growth, which could mask direct effects of body size on fitness traits. Here, we used experimental body size manipulation in the freshwater cnidarian Hydra oligactis, coupled with manipulation of resource (food) availability to separate direct effects of body size from resource availability on fitness traits (sexual development time, fecundity, and survival). We found significant interaction between body size and food availability in sexual development time in both males and females, such that large individuals responded less strongly to variation in resource availability. These results are consistent with an energy reserve effect of large size in Hydra. Surprisingly, the response was different in males and females: small and starved females delayed their reproduction, while small and starved males developed reproductive organs faster. In case of fecundity and survival, both size and food availability had significant effects, but we detected no interaction between them. Our observations suggest that in Hydra, small individuals are sensitive to fluctuations in resource availability, but these small individuals are able to adjust their reproductive development to maintain fitness.     

Motivational state and reward content determine choice behavior under risk in mice

Risk is a ubiquitous feature of the environment for most organisms, who must often choose between a small and certain reward and a larger but less certain reward. To study choice behavior under risk in a genetically well characterized species, we trained mice (C57BL/6) on a discrete trial, concurrent-choice task in which they must choose between two levers. Pressing one lever (safe choice) is always followed by a small reward. Pressing the other lever (risky choice) is followed by a larger reward, but only on some of the trials. The overall payoff is the same on both levers. When mice were not food deprived, they were indifferent to risk, choosing both levers with equal probability regardless of the level of risk. In contrast, following food or water deprivation, mice earning 10% sucrose solution were risk-averse, though the addition of alcohol to the sucrose solution dose-dependently reduced risk aversion, even before the mice became intoxicated. Our results falsify the budget rule in optimal foraging theory often used to explain behavior under risk. Instead, they suggest that the overall demand or desired amount for a particular reward determines risk preference. Changes in motivational state or reward identity affect risk preference by changing demand. Any manipulation that increases the demand for a reward also increases risk aversion, by selectively increasing the frequency of safe choices without affecting frequency of risky choices.     

Temporal discounting predicts individual competitive success in a human analogue of group foraging

Twenty groups of college undergraduates (N=9-12) participated in a discrete-trials analogue of group foraging in which points, exchangeable for course credit, were the resource to be acquired. Group matching of foragers to patch resource availability was well described by the generalized Ideal Free Distribution, with undermatching (imperfect sensitivity to resource differentials) the norm. Individuals differed on several measures of competitive success, and a measure of temporal discounting (TD) accounted for a significant proportion of the variance in these measures. Tendencies in switch patterns and visit durations differed for the most-impulsive and least-impulsive individuals in a fashion that was consistent with a TD interpretation. When competitive weights, defined in terms of TD scores, were substituted for counts of individuals in the generalized Ideal Free Distribution, group-matching slopes more closely approximated perfect sensitivity than in standard analyses. This was true for groups incorporating all impulsive individuals, all non-impulsive individuals, or a broad range of TD scores. The results suggest that well-understood individual psychological processes are associated with competitive ability in group choice.     

Responses of canopy duration to temperature changes in four temperate tree species: relative contributions of spring and autumn leaf phenology

While changes in spring phenological events due to global warming have been widely documented, changes in autumn phenology, and therefore in growing season length, are less studied and poorly understood. However, it may be helpful to assess the potential lengthening of the growing season under climate warming in order to determine its further impact on forest productivity and C balance. The present study aimed to: (1) characterise the sensitivity of leaf phenological events to temperature, and (2) quantify the relative contributions of leaf unfolding and senescence to the extension of canopy duration with increasing temperature, in four deciduous tree species (Acer pseudoplatanus, Fagus sylvatica, Fraxinus excelsior and Quercus petraea). For 3 consecutive years, we monitored the spring and autumn phenology of 41 populations at elevations ranging from 100 to 1,600 m. Overall, we found significant altitudinal trends in leaf phenology and species-specific differences in temperature sensitivity. With increasing temperature, we recorded an advance in flushing from 1.9 ± 0.3 to 6.6 ± 0.4 days °C⁻¹ (mean ± SD) and a 0 to 5.6 ± 0.6 days °C⁻¹ delay in leaf senescence. Together both changes resulted in a 6.9 ± 1.0 to 13.0 ± 0.7 days °C⁻¹ lengthening of canopy duration depending on species. For three of the four studied species, advances in flushing were the main factor responsible for lengthening canopy duration with increasing temperature, leading to a potentially larger gain in solar radiation than delays in leaf senescence. In contrast, for beech, we found a higher sensitivity to temperature in leaf senescence than in flushing, resulting in an equivalent contribution in solar radiation gain. These results suggest that climate warming will alter the C uptake period and forest productivity by lengthening canopy duration. Moreover, the between-species differences in phenological responses to temperature evidenced here could affect biotic interactions under climate warming.