The influence of ethanol on feeding in the frugivorous yellow-vented bulbul (Pycnonotus xanthopygos)

The sugars in fleshy fruits provide a rich source of energy to frugivorous animals. However, these carbohydrates also serve as a substrate for alcoholic fermentation by yeasts, ethanol being the main by-product of this process. Ethanol ingestion via frugivory thus occurs in a diverse assemblage of invertebrate and vertebrate taxa, including numerous species of birds. We tested the roles of ethanol as an odor cue for resource location by adult yellow-vented bulbuls (Pycnonotus xanthopygos) and as a possible appetite stimulant in feeding trials with artificial food. We hypothesized (1) that the odor of ethanol does not serve as a food-locating cue in diurnal frugivorous passerine birds, and predicted that the choice of food source and the mass of food eaten by such birds will not be influenced by the odor of ethanol. We further hypothesized (2) that food intake in passerine birds is affected by ingestion of ethanol according to its concentration [EtOH], and predicted that food intake will follow a bell-shaped curve in relation to [EtOH]. In accord with hypothesis (1) and its prediction, we found that the odor of ethanol did not affect food preferences, in either ethanol-naïve or ethanol-experienced yellow-vented bulbuls, when presented at concentrations found in naturally ripe fruit (0.0–1.0%); this suggests that the odor of ethanol is not a food-locating cue for the bulbuls. Hypothesis (2) was partially supported, namely at low [EtOH] (0–3%), food intake was constant and at high [EtOH] (3%) food intake decreased, following only the right half of the predicted bell-shaped response. Ethanol-naïve birds showed no preference towards any [EtOH] presented in two-way choice trials. However, daily food intake in ethanol-experienced bulbuls in single option trials decreased by an average of 36% when the artificial food contained the highest tested concentration of ethanol (3.0%). We suggest that decreasing food intake when food ethanol concentration is relatively high may be a means of avoiding intoxication and is related to the ethanol-metabolizing ability of the bird.     

State-dependent valuation learning in fish: banded tetras prefer stimuli associated with greater past deprivation

Humans and some birds and insects sometimes prefer alternatives associated with greater past cost or need, sometimes affording losses. It has been proposed that this is widespread because learning may include knowledge about both the physical properties of alternatives and state-dependent fitness gains. We examine the phenomenon for the first time in a fish, the banded tetra (Astyanax fasciatus). During training we paired two different color cues to identical food rewards, one under greater deprivation than the other. We then tested preference between these cues under both deprivation states. Consistent with previous results in other taxa, the fish preferred the cue associated with previous greater deprivation regardless of the condition under which they were tested. These results provide further support to the view that organisms assign value using state-dependent increments in fitness during learning. Although generally adaptive, under experimental conditions state-dependent valuation learning can lead to paradoxical choices.     

Chlorpyrifos: Ecotoxicological Risk Assessment for Birds and Mammals in Corn Agroecosystems

A tiered risk assessment was conducted for the use of granular and liquid formulations of chlorpyrifos in corn agroecosystems in the U.S. The initial screening Tier I assessment suggested that under high-exposure scenarios the granular and some spray formulations present potential hazards to birds. Higher tiered probabilistic risk assessments were conducted separately for the granular and liquid formulations. The probabilistic assessment indicated that risk to birds from exposure to granular formulation is small and that this route of exposure would not be a significant source of mortality. Similarly, the assessment of potential exposure of birds to food items contaminated with chlorpyrifos showed that the risk from exposure via food was small, even if it was assumed that birds feed only on the treated fields. Although they have potentially greater sensitivity to chlorpyrifos, effects in nestling birds consuming food items from fields treated with granular chlorpyrifos were negligible. However, risks to young birds may be greater where the major source of food is from fields treated with liquid formulations of chlorpyrifos. A review of field studies showed that wildlife mortality incidents associated with use of either granular or liquid formulations of chlorpyrifos are not widely apparent in agroecosystems. Based on the multiple lines of evidence, we conclude that the presumption that chlorpyrifos use in corn agroecosystems will result in extensive mortality of terrestrial wildlife, particularly birds and mammals, is not supported by the scientific evidence.     

What You See Is What You Get? Exclusion Performances in Ravens and Keas

Background

Among birds, corvids and parrots are prime candidates for advanced cognitive abilities. Still, hardly anything is known about cognitive similarities and dissimilarities between them. Recently, exclusion has gained increasing interest in comparative cognition. To select the correct option in an exclusion task, one option has to be rejected (or excluded) and the correct option may be inferred, which raises the possibility that causal understanding is involved. However, little is yet known about its evolutionary history, as only few species, and mainly mammals, have been studied.

Methodology/Principal Findings

We tested ravens and keas in a choice task requiring the search for food in two differently shaped tubes. We provided the birds with partial information about the content of one of the two tubes and asked whether they could use this information to infer the location of the hidden food and adjust their searching behaviour accordingly. Additionally, this setup allowed us to investigate whether the birds would appreciate the impact of the shape of the tubes on the visibility of food. The keas chose the baited tube more often than the ravens. However, the ravens applied the more efficient strategy, choosing by exclusion more frequently than the keas. An additional experiment confirmed this, indicating that ravens and keas either differ in their cognitive skills or that they apply them differently.

Conclusion

To our knowledge, this is the first study to demonstrate that corvids and parrots may perform differently in cognitive tasks, highlighting the potential impact of different selection pressures on the cognitive evolution of these large-brained birds.     

Attribution and Expression of Incentive Salience Are Differentially Signaled by Ultrasonic Vocalizations in Rats

During Pavlovian incentive learning, the affective properties of rewards are thought to be transferred to their predicting cues. However, how rewards are represented emotionally in animals is widely unknown. This study sought to determine whether 50-kHz ultrasonic vocalizations (USVs) in rats may signal such a state of incentive motivation to natural, nutritional rewards. To this end, rats learned to anticipate food rewards and, across experiments, the current physiological state (deprived vs. sated), the type of learning mechanism recruited (Pavlovian vs. instrumental), the hedonic properties of UCS (low vs. high palatable food), and the availability of food reward (continued vs. discontinued) were manipulated. Overall, we found that reward-cues elicited 50-kHz calls as they were signaling a putative affective state indicative of incentive motivation in the rat. Attribution and expression of incentive salience, however, seemed not to be an unified process, and could be teased apart in two different ways: 1) under high motivational state (i.e., hunger), the attribution of incentive salience to cues occurred without being expressed at the USVs level, if reward expectations were higher than the outcome; 2) in all experiments when food rewards were devalued by satiation, reward cues were still able to elicit USVs and conditioned anticipatory activity although reward seeking and consumption were drastically weakened. Our results suggest that rats are capable of representing rewards emotionally beyond apparent, immediate physiological demands. These findings may have translational potential in uncovering mechanisms underlying aberrant and persistent motivation as observed in drug addiction, gambling, and eating disorders.     

Seasonal modulation of diurnal food consumption in Indian songbirds

Seasonal changes in daily food consumption have a direct bearing with energy requirement of bird that is in turn associated with life history stage of birds. We compared seasonal changes in daily food intake in adult male migratory redheaded bunting (Emberiza bruniceps) that over winters in Indian subcontinent with those in non-migratory blackheaded munia to reiterate the same. We also compared daily food eating pattern (DFEP) in wintering blackheaded and redheaded buntings, closely related Emberizidae finches to establish circadian nature of feeding behavior and how it varied at species level. The birds were held under short days (8L:16D; 8 h of light and 16 h of darkness) and two hourly food consumption was measured to profile their DFEP. Further, we extended the study to establish how the circadian pattern of food consumption varied depending on birds’ physiological state and effect of photoperiod in adult male redheaded buntings. Redheaded buntings DFEP and locomotor activity were compared in pre-migratory months of February (spring) and September (autumn). The results suggest that September (photorefractory) birds exhibit clear bimodality in their feeding behavior as compared to (photosensitive) birds in February. Another experiment compared bird’s DFEP held under short (8L:16D) and long (16L:8D) days for 5 weeks and suggested that under long days, prolonged hours of photophase render adaptive advantage to birds for positive energy budgeting. The present study clearly establishes the circadian nature of feeding behavior and that it modulates over seasons. The bimodal i.e. morning and evening peaks of food consumption suggest morning–evening food entrainable oscillators, however this needs to be investigated with mechanistic approach in future studies.     

Singing as a handicap: the effects of food availability and weather on song output in the Australian reed warbler Acrocephalus australis

Bird song is generally regarded as a sexually selected trait, and may represent a reliable handicap signal under at least certain conditions. Females may use the degree of male song production as a reliable cue to male condition or territory quality. We investigated the effect of supplementary feeding on song output in the migratory Australian reed warbler Acrocephalus australis . We experimentally increased the food availability on alternate days, and recorded several weather variables. We measured song rate and song length independently. Supplementary fed birds sang more on feeding days than on non-feeding days, while control birds did not show this effect. Song output was not significantly associated with any of the weather variables examined. Our results indicate that singing has the potential to serve as a reliable handicap signal to territorial food availability irrespective of the prevailing weather conditions. We discuss the role of energetic constraints and behavioural flexibility on the signaling function of song.     

Irrational choices in hummingbird foraging behaviour

It is conventionally assumed that, when animals evaluate alternative options, the value assigned to an option is absolute and independent of the other options available. It follows that animal choices should exhibit the rational property of regularity whereby the proportion of choices for an option cannot be increased by the addition of further options to the choice set. However, violations of regularity occur in human decision making, suggesting that humans may use comparative evaluation mechanisms whereby the value of an option is computed relative to the other options available. For example, in the asymmetrically dominated decoy effect the preference for a target option over a competitor is altered by the addition of a decoy option that is inferior to the target and competitor on one attribute, but lies between them on a second. We tested whether foraging wild rufous hummingbirds, Selasphorus rufus, would demonstrate violations of regularity in response to an asymmetrically dominated decoy. Sixteen birds chose between three artificial flower types (Target: 15 μl, 40% sucrose; Competitor: 45 μl, 30%; Decoy: 10 μl, 35%) in Binary (Target versus Competitor) and Trinary (Target versus Competitor versus Decoy) treatments. We predicted higher preference for the Target in the Trinary treatment. The birds ranked the three options in the same order in the Binary and Trinary treatments (Competitor>Target>Decoy). Seven birds showed violations of regularity, six increasing their absolute preference for the Competitor in the Trinary treatment. Overall, relative preference for the Competitor over the Target was higher in the Trinary than in the Binary treatment. These changes in preference are incompatible with an absolute evaluation mechanism.     

Phenotypic flexibility in digestive system structure and function in migratory birds and its ecological significance

Birds during migration must satisfy the high energy and nutrient demands associated with repeated, intensive flight while often experiencing unpredictable variation in food supply and food quality. Solutions to such different challenges may often be physiologically incompatible. For example, increased food intake and gut size are primarily responsible for satisfying the high energy and nutrient demands associated with migration in birds. However, short-term fasting or food restriction during flight may cause partial atrophy of the gut that may limit utilization of ingested food energy and nutrients. We review the evidence available on the effects of long- and short-term changes in food quality and quantity on digestive performance in migratory birds, and the importance of digestive constraints in limiting the tempo of migration in birds. Another important physiological consequence of feeding in birds is the effect of diet on body composition dynamics during migration. Recent evidence suggests that birds utilize and replenish both protein and fat reserves during migration, and diet quality influences the rate of replenishment of both these reserves. We conclude that diet and phenotypic flexibility in both body composition and the digestive system of migratory birds are important in allowing birds to successfully overcome the often-conflicting physiological challenges of migration.     

Energy budgets and risk-sensitive foraging in starlings

The effect of energy budget on risk-sensitive foraging was assessedin a laboratory experiment using starlings (Sturnus vulgaris).Subjects chose between two options offering the same mean amountof food per trial, but differing in variance: a "fixed" optiongave 5 units food in every trial, and a "variable" option gave2 or 11 units food with probabilities 2/3 and 1/3, respectively.We manipulated energy budgets by controlling the cumulativeamount of food received by each bird at the end of a day. Inone treatment (positive budget) individuals were allowed toeat at the level of their own ad-libitum daily consumption,while for the other (negative budget), food was rationed toprovoke a steady drop in body weight during the experimentalperiod. No subject was allowed to drop below 80% of its ad libitumbody weight. Contrary to predictions from the "energy budgetrule" and contrary to reported results of some other studies, starlingssignificantly preferred the "fixed" option irrespective of energybudget conditions. Our results support the view that persistentrisk aversion for food amounts and risk proneness for food delaysare the norm, and shifts in risk attitude according to energybudget are exceptions. Several algorithms, which may have evolvedto maximize energetic pay off between variable food sources,can produce this trend as a side effect. We discuss two of thesealgorithms: (1) maximization of local (per trial) rate as opposedto global rate of gains, with longer handling time for largerrewards, and (2) choosing larger rewards and smaller delayssubject to Weber's law in the memory for the parameters of eachfood supply.     

Prefrontal coding of temporally discounted values during intertemporal choice

Reward from a particular action is seldom immediate, and the influence of such delayed outcome on choice decreases with delay. It has been postulated that when faced with immediate and delayed rewards, decision makers choose the option with maximum temporally discounted value. We examined the preference of monkeys for delayed reward in an intertemporal choice task and the neural basis for real-time computation of temporally discounted values in the dorsolateral prefrontal cortex. During this task, the locations of the targets associated with small or large rewards and their corresponding delays were randomly varied. We found that prefrontal neurons often encoded the temporally discounted value of reward expected from a particular option. Furthermore, activity tended to increase with [corrected] discounted values for targets [corrected] presented in the neuron's preferred direction, suggesting that activity related to temporally discounted values in the prefrontal cortex might determine the animal's behavior during intertemporal choice.     

Colour association influences honey bee choice between sucrose concentrations

Certain colours associated with floral food resources are more quickly learned by honey bees (Apis mellifera) than are other colours. But the impact of colour, and other floral cues, on bee choice behaviour has not yet been determined. In these experiments, colour association and sugar concentration of reward were varied to assess how they interact to affect bee choice behaviour. Thirty-five bees were individually given binary choices between blue and yellow artificial flowers that contained either the same rewards or rewards of different sucrose concentrations. Honey bee choice between sucrose concentrations was affected by colour association and this effect was greatest when absolute difference between rewards was the lowest. The honey bee's ability to maximize energetic profitability during foraging is constrained by floral cue effectiveness.     

Effects of demanding foraging conditions on cache retrival accuracy in food-caching mountain chickadees (Poecile gambeli)

Birds rely, at least in part, on spatial memory for recovering previously hidden caches but accurate cache recovery may be more critical for birds that forage in harsh conditions where the food supply is limited and unpredictable. Failure to find caches in these conditions may potentially result in death from starvation. In order to test this hypothesis we compared the cache recovery behaviour of 24 wild-caught mountain chickadees (Poecile gambeli), half of which were maintained on a limited and unpredictable food supply while the rest were maintained on an ad libitum food supply for 60 days. We then tested their cache retrieval accuracy by allowing birds from both groups to cache seeds in the experimental room and recover them 5 hours later. Our results showed that birds maintained on a limited and unpredictable food supply made significantly fewer visits to non-cache sites when recovering their caches compared to birds maintained on ad libitum food. We found the same difference in performance in two versions of a one-trial associative learning task in which the birds had to rely on memory to find previously encountered hidden food. In a non-spatial memory version of the task, in which the baited feeder was clearly marked, there were no significant differences between the two groups. We therefore concluded that the two groups differed in their efficiency at cache retrieval. We suggest that this difference is more likely to be attributable to a difference in memory (encoding or recall) than to a difference in their motivation to search for hidden food, although the possibility of some motivational differences still exists. Overall, our results suggest that demanding foraging conditions favour more accurate cache retrieval in food-caching birds.     

Hazel grouse (Bonasa bonasia) pairs during the nonbreeding season: mutual benefits of a cooperative alliance

During the nonbreeding season (autumn and winter), hazel grouse(Bonasa bonasia) males and females associate as loosely boundpairs, not as strongly bound pairs as previously thought. Thesepairs could be considered cooperative alliances, with each membergaining both direct and indirect benefits. The most importantdirect benefit appeared to be mutual vigilance against predators.This benefit was found at two levels: at the level of arborealfeeding sites, two birds could forage faster and farther fromcover than single birds, and at the territory scale, the pairwas together more in dangerous habitats. By foraging fartherfrom cover, two birds could use about 23% more of the preferredfood trees and 9%–10% more of the food available withina territory than a single bird, partially mitigating the costof having two birds feeding on limited winter food in the samespace. Defending a common territory appeared to be a less importantdirect benefit of the alliance because the members of a pairdid not defend the same territory and often associated withneighboring birds of the opposite sex. A future benefit of thealliance was having breeding partners in the spring; this wasmost beneficial for males, as the sex ratio was male biased.Members of the pair were often apart. At arboreal feeding sites,this separation was perhaps because the most preferred trees,black alders (Alnns glutinosa) with the most staminate catkins,were located in areas with the most cover, where one bird maybe safer from predators than two. Members of pairs in more securehabitats were more often with extrapair birds of the oppositesex. This suggested a trade-off; birds in safe habitats mayhave visited potential breeding partners, but birds in dangeroushabitats may have had to remain together, foregoing this option,to increase their survival probability. This social organizationappeared to be an adaptation to surviving in a heterogeneoushabitat, with some of the winter food located in dense coverand some located in more open and dangerous situations.     

Faster spring migration in northern wheatears is not explained by an endogenous seasonal difference in refueling rates

A widespread phenomenon in migrant birds is that they travel faster in spring than in autumn. During migration birds spend most time at stopover sites and, correspondingly, the faster spring migration is mainly explained by shorter stopovers in spring than autumn. Because a main purpose of stopovers is to replenish the fuel used in flight, a higher rate of fuel deposition (FDR) in spring is thought to explain the shorter stopovers and hence shorter total duration of migration in spring. Critical migratory processes, including the onset and extent of pre‐migratory fueling, are endogenously regulated. It is therefore not unlikely that refueling at stopover sites is, at least partly, also under endogenous control. We here tested whether there is an endogenous seasonal difference in food intake and FDR, which could contribute to shorter stopovers and hence faster migration in spring. We measured daily food intake and daily FDR in two subspecies of the northern wheatear Oenanthe oenanthe, temporarily confined at stopover under identical constant indoor conditions in spring and autumn. The two wheatear subspecies differed markedly in absolute food intake and FDR. Within subspecies, however, food intake and FDR did not differ between spring and autumn, indicating that faster spring migration in northern wheatears is not explained by an endogenously controlled seasonal difference in birds’ motivation to refuel. To further substantiate this claim, similar measurements should be taken at other locations along northern wheatears’ migration routes. Comparable experiments in other species could test the generality of our results.     

Chimpanzees (Pan troglodytes) fail a what-where-when task but find rewards by using a location-based association strategy

Recollecting the what-where-when of an episode, or episodic-like memory, has been established in corvids and rodents. In humans, a linkage between remembering the past and imagining the future has been recognised. While chimpanzees can plan for the future, their episodic-like memory has hardly been investigated. We tested chimpanzees (Pan troglodytes) with an adapted food-catching paradigm. They observed the baiting of two locations amongst four and chose one after a given delay (15 min, 1 h or 5 h). We used two combinations of food types, a preferred and a less preferred food that disappeared at different rates. The subjects had to base their choices on the time elapsed since baiting, and on their memory of which food was where. They could recover either their preferred food or the one that remained present. All animals failed to obtain the preferred or present foods above chance levels. They were like-wise unsuccessful at choosing baited cups above chance levels. The subjects, thus, failed to use any feature of the baiting events to guide their choices. Nonetheless, their choices were not random, but the result of a developed location-based association strategy. Choices in the second half of the study correlated with the rewards obtained at each location in the first half of the study, independent from the choices made for each location in the first half of the study. This simple location-based strategy yielded a fair amount of food. The animals' failure to remember the what-where-when in the presented set-up may be due to the complexity of the task, rather than an inability to form episodic-like memories, as they even failed to remember what was where after 15 minutes.     

Development of an agri-environment scheme option: seed-bearing crops for farmland birds

Winter mortality, resulting from reduced food supply during a period of agricultural intensification, is thought to have driven population declines for some farmland bird species. Planting of game crops has increased over this period in order to provide food and cover for gamebirds. We investigate the potential of this managed habitat for farmland songbird conservation, using intensive single-site studies, and an extensive national survey. Game crops were used more than other farmland habitats by a wide range of bird species. Kale and quinoa were used by many species, whereas maize was used by very few. Cereals such as triticale and millet were used by many species, including several not associated with brassicas such as kale. Crop species differed in the rate of seed shedding, and therefore in the amount of seed food that they provided through the winter. Crop location influenced use by some bird species, with crops close to hedges or other cover generally being favoured. Use of nitrogen fertilizer influenced seed yield, and therefore crop value as a source of food for birds. Our results suggest that, if managed and sited correctly, a combination of two or three crop species can provide a valuable winter food resource for many nationally declining farmland bird species, but further attention needs to be given to their agronomy. This form of management is now incorporated as an option within agri-environment schemes in England, Scotland and Wales. It enables farmers to apply existing skills to conservation and is compatible with their cultural values.     

Context-dependent foraging choices in risk-sensitive starlings

Existing models of risk-sensitive foraging assume that animals assign value to different options according to an absolute currency. It follows from this assumption that choices are expected to be both transitive and regular, because the value assigned to an option is independent of its context. I tested these predictions by comparing preferences obtained in binary and trinary contexts. European starlings, Sturnus vulgaris, were trained using an operant paradigm to forage for three options: Constant (C) that provided a fixed number of food pellets; Low variance (L) with a coefficient of variation of 71% in the number of pellets; High variance (H) with a coefficient of variation of 106%. The preferences of the birds were tested in three binary choice treatments (CL, CH, LH) and one trinary choice treatment (CLH). Overall, there was no evidence for violations of either transitivity or regularity. However, overall, a bird's relative preference for its most preferred option over its second most preferred option was significantly greater in the trinary treatment than in the comparable binary treatment. This effect of context on choice is compatible with starlings' use of comparative instead of absolute currencies in decision making. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

The inhibitory effect of pretrial reinforcement on instrumental conditioning

It has been shown previously that pretrial feeding retards acquisition of an instrumental response. The present experiment tested three hypotheses for this effect: priming, overshadowing and conditioned inhibition. Rats were trained in a runway task with a partial schedule of food reinforcement. Rats that were prefed immediately prior to rewarded trials showed a decrement in acquisition when compared with subjects prefed 180 seconds pretrial. Other subjects prefed immediately prior to nonrewarded trials showed a similar decrement in acquisition, even though the pretrial and goal reinforcers occurred on separate trials. Thus, the adverse effects of prefeeding occurred in the absence of conditions necessary for priming or overshadowing to occur. The results suggest there may be an inhibitory aftereffect of food rewards which can become conditioned to subsequent stimuli.     

Avian foraging studies: an overlooked source of distribution data for macroecological and conservation studies

Macroecological and biogeographical studies and their applicability for biological conservation vitally depend on distribution data. These are usually taken from distribution atlases and databases or maps found in identification guides. A previous study pointed to another little explored source of data — local faunistic studies. Here, I would like to draw attention to another potential source. Papers that analysed the food composition of some taxa (e.g. insectivorous birds) are an overlooked source of rich information on taxa distributions. These studies frequently include an 'Appendix' with a list of food items determined to the species level. These studies also contain data on abundances, number of samples, sampling time, and geographical location as a rule. Foraging birds naturally provide data on invertebrate distributions with good spatio-temporal coverage and reasonably large samples. Importantly, birds frequently collect rare and by entomological methods hardly detectable species (i.e. those living high, in tree canopies, in very dense vegetation, or with secretive lifestyle). Data from bird dietary studies may help to ameliorate one of the most serious problems in distribution studies — zero inflation. I briefly discuss pros and cons of this so far neglected source of biogeographical data.