Isolating structures of gerbils’ digestive tract (Gerbillidae, <Emphasis Type="Italic">Rhombomys,Meriones</Emphasis>) and their functional significance

The morphology of the digestive tract of three species of gerbils characterized by different food specialization—Rhombomys opimus, Meriones tamariscinus and M. meridianus—is considered. The correlation between areas of glandular and cornified epithelium is variable and depends greatly on the type of stomach filling and distribution of food mass in it. The bordering fold and ileocecal junction is not only able to isolate the forestomach and caecum significantly but also may be a mechanism of regulation of food movement through the digestive tract. The close location of ileum’s opening to the caecum and its outlet to the colon, as well as isolation of caecum’s cavity by a circular fold, could provide direct food transit from the ileum to the colon.     

Nutritional Ecology of <Emphasis Type="Italic">Ateles chamek</Emphasis> in lowland Bolivia: How Macronutrient Balancing Influences Food Choices

All free-living animals must make choices regarding which foods to eat, with the choices influencing their health and fitness. An important goal in nutritional ecology is therefore to understand what governs animals’ diet selection. Despite large variation in the availability of different food items, Peruvian spider monkeys (Ateles chamek) maintain a relatively stable daily protein intake, but allow total energy intake to vary as a function of the composition of available food items. This is referred to as protein-dominated macronutrient balancing. Here we assess the influence of this nutritional strategy on daily and seasonal nutritional intakes, estimate the nutritional value of different foods, and interpret unusual food choices. We conducted continuous all-day observations of focal spider monkeys inhabiting a semideciduous forest in Bolivia. We recorded feeding events, collected foods, and analyzed their nutrient content. By using the Geometric Framework for nutrition, we show that individuals reached their daily end-point in nutrient space —balance between protein and nonprotein energy intake— by consuming nutritionally balanced foods or by alternating between nutritionally complementary foods. The macronutritionally balanced figs of Ficus boliviana were their primary staple food and therefore dominated their overall nutritional intake. Our results also demonstrate that spider monkeys consumed a diverse array of ripe fruits to overcome periods of fig scarcity rather than vice versa; they could obtain sufficient protein on a diet of pure fruit; and unripe figs constituted a nutritionally rewarding and reliable food resource. We hope that the approaches taken and the conclusions reached in this study will catalyze further inquiries into the nutritional ecology of frugivorous primates.     

Proteomic analysis of human skeletal muscle (<Emphasis Type="Italic">m. vastus lateralis</Emphasis>) proteins: Identification of 89 gene expression products

Proteins from bioptates and autoptates of human skeletal muscle m. vastus lateralis were separated by O’Farrell two-dimensional gel electrophoresis (2DE). MALDI-TOF MS and MS/MS enabled identification of 89 protein spots as expression products of 55 genes. A modification of the O’Farrell’s method including non-equilibrium electrophoresis in a pH gradient allowed detection — among major sarcomeric, mitochondrial, and cytosolic proteins — of several proteins, such as PDZ- and LIM domain-containing ones (pI > 8.70), fragments of known proteins, and a stable complex of heavy and light ferritin chains. The data underlie further studies of human skeletal muscle proteins in terms of molecular mechanisms of some physiological and pathological processes.     

Left-hand Preference for a Complex Manual Task in a Population of Wild Siamangs (<Emphasis Type="BoldItalic">Symphalangus syndactylus</Emphasis>)

Many hypotheses have been proposed linking the emergence of lateralized activities—such as handedness—in primates, with hemispheric specialization and the evolution of complex communication such as human language. Although data to test these ideas are rapidly accumulating for many primate taxa, some species are still largely unexplored, especially under natural conditions. I present the first data on a population of wild siamangs (Symphalangus syndactylus), highly arboreal small apes. Preference for the left hand is shown both at the individual and population levels for a complex manual task: collecting and drinking water from tree holes. There was no difference in hand preference between males and females, and immature individuals showed more variable patterns than adults. These results are consistent with the postural origins theory, allow a new interpretation of the findings of comparable studies, and indicate a useful behavior for future investigations of laterality in wild primates.     

Vocal exchanges of signature whistles in bottlenose dolphins (<Emphasis Type="Italic">Tursiops truncatus</Emphasis>)

Bottlenose dolphins (Tursiops truncatus) produce individually distinctive vocalizations—referred to as “signature whistles”—that are thought to function as an individual and conspecific recognition system for maintenance of consistent contact between individuals. Observations and playback experiments were conducted at aquariums to study these whistle–vocal exchanges in bottlenose dolphins. Temporal patterns of vocalization were examined by analyzing the intercall intervals between two consecutive whistles. When a second individual produced a call that was different from the first individual’s vocalization, most of these calls were shorter than 1 s. However, when two consecutive calls were produced by the same individual, the second call rarely occurred within 1 s of the first. These results suggest that a second whistle may be produced by a different caller in response to the first whistle; however, in the case of an absence of a response, the first caller is likely to give further whistles. The results of this acoustic analysis show that the dolphins used in this study mostly used signature whistles during the recorded vocal exchanges.     

Consciousness is Cheap, Even if Symbols are Expensive; Metabolism and the Brain’s Dark Energy

Use of symbols, the key to the biosemiotics field as to many others, required bigger brains which implied a promissory note for greater energy consumption; symbols are obviously expensive. A score years before the current estimate of 18–20% for the human brain’s metabolic demand on the organism, it was known that neural tissue is metabolically dear. This paper first discusses two evolutionary responses to this demand, on both of which there is some consensus. The first, assigning care of altricial infants with burgeoning brains (and in human infants the metabolic demand peaks at 65% of the total) to “allomothers” is not unique to humans. The second, using relatively small neurons as primates do, risks misfires past a certain minimal value. Moreover, in apparent paradox, there is an increasing consensus that large “Von Economo” neurons are critical for communication. This paper’s main contribution is the discussion of two further evolutionary tricks. The first is the use of self-similarity in the cortex, both in structure and process, to allow the cortex readily—and in energetic terms, parsimoniously—to shift between states in a high-dimensional space. This leads to discussion of the kind of formalism appropriate to model these shifts, a formalism which—it is tentatively suggested—may do double duty for the modeling of symbolic thought. The second trick is the superimposition on the background “white noise” of neural firing of EEG-detected waves like gamma. The paper describes a method, using the Hilbert transform, of calculating the dips in energy consumption as the brain is transitioned by gamma waves. It is hypothesized that consciousness may be a spandrel, the incidental result of a neurodynamic imperative that the brain enter a maximally sensitive (in sensory terms) “zero power” state a few times a second. If that is the case, then there are obvious benefits for health in meditation, which can be viewed as a state of consciousness extended over time by limiting afferent stimuli.     

Simple rules result in the adaptive turning of food items to reduce drag during cooperative food transport in the ant <Emphasis Type="Italic">Pheidole oxyops</Emphasis>

Insect workers cooperate to carry out a variety of tasks. One example is cooperative transport of food items by two or more ant workers, which is important in foraging in many species. We predicted that natural selection would result in strategies that improve the performance of this task and tested this in Pheidole oxyops, a Neotropical ant in which ca. 70% of the biomass of dead insects brought back to the nest is transported cooperatively. We specifically tested the hypothesis that groups would re-orient food items to reduce drag, given that legs, wings, and other projections should affect the ease of dragging prey in different orientations. By presenting ants with artificial food items and dead cockroaches, both of which required approximately twice as much force to drag backwards as forwards, and a control which was equally easy to move in both orientations, we showed that natural groups of 3–20 food-transporting ants usually turned items that were facing backwards (72 and 83% of trials for artificial food items or cockroaches, respectively), the orientation requiring greater force, but not items facing forwards (10 and 12% of trials, respectively). Turning usually involved a single ‘steering’ ant. The key role of the ‘steering’ ant was shown by removing either the current steering ant or a randomly chosen ‘non-steering’ ant during turning. In 100% of the trials in which the steering ant was removed, turning stopped until another ant took its place. Conversely, turning stopped in only 17% of trials in which a ‘non-steering’ ant was removed. Turning is an emergent property of the system and may not have been directly selected for. Rather, turning seems to occur through a combination of pre-existing retrieval behaviour and the underlying physics of large loads. Points where the food item catches the ground can act as a fulcrum or pivot around which the item can rotate. Ants furthest from the fulcrum have more leverage and so are more likely to play a key role in turning. A simple rule relevant to individual transport of food items such as “grasp the food item and move towards the nest”, when used in the context of cooperative transport, has allowed the ants to solve a seemingly complicated problem requiring coordination.     

What Role Do Mothers Play in the Gestural Acquisition of Bonobos (Pan paniscus) and Chimpanzees (Pan troglodytes)?

Contemporary research hypothesizes that biological inheritance and ontogenetic factors shape the development of gestural communication in nonhuman great apes. However, little is known about the specific role that mothers play in the acquisition of their infants’ gestures. We observed 6 bonobo (Pan paniscus) and 4 chimpanzee (Pan troglodytes) mother–infant dyads and recorded their gesture types and frequency. We analyzed all behavioral contexts in which gestures occurred as well as the play context alone. Infants of both species were unlikely to share gestures with their mother or unrelated adult females. However, gestural sharing was prevalent within age groups. Within and across species, infant–infant and mother–mother groups were homogeneous regarding the types of gestures they shared, although there was individual variation in the frequency of gesture use. Our findings provide limited evidence that infants learned their gestures by imitating their mothers. Phylogenetic influences seem to be vital in gestural acquisition but, we suggest, repertoire development cannot be disentangled from individual social encounters during life.     

The Concept of Animal Welfare at the Interface between Producers and Scientists: The Example of Organic Pig Farming

In organic farming animal welfare is one important aspect included in the internationally agreed organic principles of health, ecology, fairness and care (IFOAM 2006), reflecting expectation of consumers and farmers. The definition of organic animal welfare includes—besides traditional terms of animal welfare—‘regeneration’ and ‘naturalness’. Organic animal welfare assessment needs to reflect this and use complex parameters, include natural behaviour and a systemic view. Furthermore, various parties with seemingly conflicting interests are involved, causing ethical dilemmas, such as the use of nose rings for outdoor sows (impaired animal welfare vs. destruction of humus). Solutions can only be found when foundational concepts are translated and applied to practical situations. On-farm animal welfare assessment and implementation of improvement strategies are increasingly relevant scientific areas. They combine on-farm welfare assessment, identification of key problem areas and connected risk factors. Constant communication between all parties is crucial for success. Animal health and welfare planning is one application of this approach, which was carried out on Austrian organic pig farms as well as organic dairy farms in seven European countries. The projects included welfare assessment, feedback and benchmarking as a tool for communication between farmers, advisors and scientists. Finally goals were set by the farmer and improvement strategies applicable to organic farming were implemented. This included prevention of disease by management strategies instead of routine treatment with pharmaceutical products. It appeared that next to problem structuring, multidisciplinary problem solving demands good communications skills to relate animal welfare science to value reflections.     

Nationalization and Globalization Trends in the Wild Mushroom Commerce of Italy with Emphasis on Porcini (<Emphasis Type="Italic">Boletus edulis</Emphasis> and Allied Species)

Nationalization and Globalization Trends in the Wild Mushroom Commerce of Italy with Emphasis on Porcini ( Boletus edulis and Allied Species). This paper presents an historical overview of wild mushroom commerce in Italy, with a focus on recent trends in the production of porcini (Boletus edulis and closely allied species). Over the past century, two major trends—nationalization and globalization—have been apparent in the wild mushroom commerce of Italy. First, a simplified national mushroom menu has emerged through processes of governmental regulation and culinary fashion, but it has come at the expense of differing, localized mushroom traditions which may suffer under the European Union’s free trade principles. Second, Italy has emerged as a focal point of a global market for a small number of mushroom species—particular porcini. While the name porcini has become synonymous with Italian cuisine, and in spite of a vibrant tradition of recreational mushroom collecting in Italy, most of the porcini commercially available in Italy or exported by Italy are no longer of Italian origin. Porcini and other mushrooms now flow into Italy from all over the world—especially from China and eastern Europe—and are then often exported as “Italian porcini.” This globalization of the wild mushroom trade, while offering significant income to rural producers and processors around the globe, has other effects as well, for example, a kind of national branding as “Italian” of globally-produced products, of which porcini is one, that is in direct opposition to some of the European Union’s rules for regional denominations.

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Processi di Nazionalizzazione e Globalizzazione nel Commercio Italiano dei Funghi spontanei, con Particolare Riguardo ai Porcini ( Boletus edulis e Specie Affini). Questo articolo presenta una panoramica storica sul commercio dei funghi spontanei in Italia, con particolare riguardo alle recenti tendenze nella produzione dei porcini (Boletus edulis e specie affini). Nello scorso secolo si sono osservate due tendenze principali—di nazionalizzazione e di globalizzazione—nel commercio dei funghi spontanei in Italia. In primo luogo si è affermata nel territorio nazionale una tradizione limitata al consumo di un numero contenuto di specie, sia per effetto di alcune normative che di mode culinarie, ma ciò è avvenuto a discapito di tradizioni locali più ricche, che potrebbero ulteriormente risentire dei principi per il libero scambio all’interno dell’Unione Europea. In secondo luogo, l’Italia si è posta in evidenza come un punto nodale per il mercato globale di alcune specie fungine, in particolare dei porcini. Nonostante quest’ultimo termine sia tradizionalmente associato alla cucina italiana, e nonostante esista in Italia una vivace e radicata tradizione nella raccolta amatoriale dei funghi, la maggior parte dei porcini ivi commercializzati (allo stato fresco, essiccati o variamente conservati) o esportati verso altri paesi non sono più di origine locale. I porcini e altre specie fungine giungono attualmente in Italia da ogni parte del mondo—in modo particolare dalla Cina e dall’Europa orientale—e sono successivamente spesso esportati come “prodotti Italiani.” Questo processo di globalizzazione del mercato dei funghi spontanei, pur offrendo un significativo introito ai raccoglitori e commercianti rurali su tutto il globo, è responsabile di altri effetti, come per l’appunto una sorta di marchio nazionale “Italiano” su alcuni prodotti di provenienza globale, come per l’appunto i porcini, cosa che contrasta con alcune delle regole dell’Unione Europea in materia di denominazioni regionali.

     

Feeding Competition and Group Size in Alouatta pigra

Researchers consider group size in primates to be determined by complex relationships among numerous ecological forces. Antipredator benefits and better resource defense are the primary pressures for large groups. Conversely, intragroup limited food availability, can result in greater intragroup feeding competition and individual energy expenditure in larger groups, creating energetic advantages for individuals in small groups and placing an upper limit group size. However, the extent to which food availability constrains group size remains unclear for many species, including black howlers (Alouatta pigra), which ubiquitously live in small social groups (≤10 individuals). We studied the relationship between group size and 2 key indices of feeding competition—day journey length and activity budgets—in 3 groups of wild Alouatta pigra at a hurricane-damaged site in Belize, Central America. We controlled for differences in food availability between home ranges (food tree density) and compared both indicators of feeding competition directly with temporal variation in food availability for each group. Our results show no consistent association between resource availability, group size, and either index of competition, indicating that feeding competition does not limit group size at the site—i.e., that larger groups can form without increased costs of feeding competition. The results support the search for other explanations, possibly social ones, for small group size in the primates, and we conclude with suggestions and evidence for such alternative explanations.     

Variation in Spatial Cohesiveness in a Group of Japanese Macaques (<Emphasis Type="Italic">Macaca fuscata</Emphasis>)

The spatial cohesiveness of a group is an important element that characterizes the social structure of group-living species. Moreover, remaining cohesive is crucial if individuals are to coordinate their activities and reach collective decisions. We measured interindividual spacing in a group of wild Japanese macaques (Macaca fuscata) to assess the spatial cohesiveness of a social group quantitatively. We used simultaneous focal animal sampling, with 2 observers recording individuals’ locations with a global positioning system (GPS) during 3 seasons. Interindividual distances differed among seasons; they were short in autumn (mean ± SD: 25.6 ± 20.1 m), intermediate in winter (mean ± SD: 46.3 ± 35.7 m), and long in summer (mean ± SD: 62.3 ± 47.1 m). Measurements taken in summer revealed extremely wide spacing (maximum: 1225 m), suggesting subgrouping. Distances also varied with activity during each season; they were short during resting and grooming, intermediate during foraging, and long during moving. Group cohesion was also influenced by food distribution. More group members were ≤20 m of the focal individual during foraging on clumped food than foraging on scattered food in each season, and the group foraged on clumped food most frequently in autumn. Individuals were also likely to aggregate at resting/grooming sites and clumped food patches and to disperse when moving within a day. These results demonstrate that Japanese macaques show considerable variation in spatial cohesiveness both within short time periods, e.g., 1 d, and among seasons, and that they adjust group cohesiveness flexibly depending on the food conditions and foraging tactics.     

How much is Europe spending on invasive alien species?

Over the last 15 years, despite the lack of a specific strategy or a dedicated financial instrument to deal with invasive alien species (IAS), the European Commission (EC) has contributed to financing almost 300 projects addressing this issue, for a total budget exceeding 132 million EUR. Such figures are based on projects funded under two specific EU financial tools: LIFE and the RTD Framework Programmes. The contribution of the two programmes has been characterised by an overall positive trend over the years, in terms of both the number of projects and the budget spent. Such trend can be assumed to reflect an overall increase in both the awareness of the problem among wildlife managers and scientific institutions, and the willingness to pay by the EC institutions and the EU citizens in general. Such data might contribute to the development of a response indicator measuring ‘Trends in invasive alien species in Europe’, useful to assess progress toward the target of halting the loss of biodiversity by 2010—as a part of the SEBI 2010 process. The results may also contribute to assess the economic impact of IAS in Europe—in terms of costs for reduction and/or prevention of damages—and to support policy decisions and communication campaigns. Finally, the results are encouraging and support the need for the development and the implementation of a sound EU strategy on IAS, so as to regulate and optimise the administration of the available financial resources—whenever appropriate—on the basis of specific priorities.     

Saltatory Search in Free-Living <Emphasis Type="Italic">Callithrix jacchus</Emphasis>: Environmental and Age Influences

Researchers generally categorize motile foraging behavior into 3 strategies: ambush, cruise, and saltatory searching. During saltatory searches, predators move from one location to the next, frequently pausing to scan for prey that are hard to detect and widely distributed. We investigated whether 1) the foraging strategy of free-living common marmosets conforms to the strategy; 2) scanning occurs solely when the individual is stationary; 3) the environment (dense and sparse vegetation) influences foraging behavior; and 4) the age of the marmosets is related to their foraging behavior. Bezerra carried out the observations in a 32-ha fragment of Atlantic Forest in the Northeast of Brazil and in an adjacent condominium. Using the focal sampling method, we observed 31 common marmosets, including adults, juveniles, and infants, Bezerra recorded the following behaviors (in dense and sparse vegetation): locomotion (subdivided into minor locomotion—moving distances ≤1 m; major locomotion—moving distances >1 m), scan, pause, and capture. The frequency of scanning behavior was significantly greater when individuals were stationary. Adults and juveniles exhibited the clearest differentiation in terms of locomotion, both adjusting the behavior in accordance with the environment; periods of major locomotion were more frequent in sparse vegetation, whereas minor locomotion was more frequent in dense vegetation. In contrast, infants exhibited major locomotion more frequently in dense vegetation. We conclude that common marmosets use a saltatory strategy when foraging, and that their foraging behavior is plastic, changing both with the age of the individual and with the density of the vegetation.     

Foraging ecology and reproductive biology of the Stonechat Saxicola torquata: comparison between a revitalized, intensively cultivated and a historical, traditionally cultivated agro-ecosystem

An effective strategy to reintegrate biodiversity within otherwise intensively cultivated agroecosystems is to create set-aside and wildflower areas. It remains largely unknown, however, whether the ecological performance of an agroecosystem revitalized in this manner is comparable—from a species’ population biology perspective—to traditionally exploited farmland. To address this question we compared, during two successive years, the trophic ecology and breeding performance of an insectivorous, indicator passerine (the Stonechat Saxicola torquata) in a revitalized intensively cultivated farmland (RIC) and a traditional, extensively cultivated farmland (TEC) in southern Switzerland. The chicks' diet and prey abundance did not differ between the RIC and TEC, with orthopterans, caterpillars (Lepidoptera) and coleopterans predominating (approx. 80% of diet biomass). Although Stonechat pairs initiated more broods in TEC than in the RIC, reproductive success (number of fledglings/territory × year) did not differ significantly between the TEC and RIC. The chicks’ condition (body mass) was slightly better in TEC than in RIC, while no such effect could be shown for chick constitution (tarsus length) in either year. The inter-site (RIC vs. TEC) variation fell well within the inter-annual variation of breeding parameters, indicating that environmental stochasticity could be a greater determinant of reproductive output and young quality than agroecosystem type. Although in need of replication, these results suggest that incentives for setting aside farmland and creating wildflower areas within agroecosystems may not only enhance plant and invertebrate diversity, as has been demonstrated earlier, but can also support functioning populations of vertebrates situated at higher trophic levels along the food chain.     

How does pattern of feeding and rate of nutrient delivery influence conditioned food preferences?

Ruminant herbivores have been shown to learn about food properties by associating food flavours with the food’s post-ingestive consequences. Previous experimentation supporting the conditioned food aversion/preference hypothesis has generally employed very simple diet learning tasks which do not effectively represent the wide range of foods selected within single bouts typical of wild, free-ranging ruminant herbivores. We tested the ability of a ruminant herbivore to associate a food with artificially administered nutrient rewards in a designed experiment where we altered the temporal pattern of encounter with the food as well as the nature (fast or slow reward) of the post-ingestive outcome. Twenty-four goats were offered branches of Sitka spruce (SS) and Norway spruce (NS) for 4 h per day on two days per week for five weeks. The pattern of feeding varied with treatment such that the species on offer changed every hour (short) or every 2 h (long). The energy treatment altered the reward delivered during Sitka consumption so that animals were dosed either with predominantly sugar (rapidly fermented), predominantly starch (slower fermentation rate), or with water (placebo). Preference was measured on the day following each learning day. We expected that goats would find it easier to associate SS with post-ingestive rewards when the duration of encounter was longest, and that associations would be stronger with the most rapidly digested post-ingestive reward. In the event, goats did not alter their consumption of SS in response to the treatments. Our results suggest that at the scale of temporal resolution of encounters with different plant species (1–2 h), and at the different rates of experiencing post-ingestive consequences tested in this experiment, ruminants do not appear to discriminate the nutritive properties of foods predominantly through a post-ingestive feedback mechanism. They must, instead, use a range of cues—including post-ingestive consequences—to assess food properties. An erratum to this article can be found at     

Diet composition and foraging activity of <Emphasis Type="Italic">Pipistrellus pygmaeus</Emphasis> in a floodplain forest

In 2000–2002 bat droppings were collected under the emerging crevice of a nursery colony of Pipistrellus pygmaeus. The locality was situated in a floodplain forest at the confluence of the Dyje and Morava rivers (S Moravia, Czech Republic). In total, 27 samples (20 pellets in one sample) of droppings were used to analyze prey remains. In the diet, 40 taxonomic groups of invertebrates were found. As expected, small dipteran insects were the main food item in which Nematocera dominated. Besides Chironomidae and Ceratopogonidae also a high percentage of nematoceran eggs were recorded. Surprisingly, a relatively high percentage of Brachycera was recorded. Further frequent prey items belonged to the orders of Trichoptera, Hymenoptera, Coleoptera and Sternorrhyncha. A heterodyne bat detector was used to follow foraging activity of P. pygmaeus on line transects in forest and water habitats in the vicinity of the colony. A significant decrease in foraging activity over water habitats and in forest sites during the late pregnancy (mid-May — early June) and an increase during the lactation and post-lactation periods (mid-June — early August) were found. Changes in the frequency of occurrence of Chironomidae, Neuroptera, Trichoptera, Aphidinea and Simuliidae were correlated with the bats’ foraging activity.     

Grazing Increases Soil Warming in Headwater Wetlands: Importance to Land Managers and Water Users

Ecosystems - The earth’s mountains continue to lose water. Glaciers are melting and mountain snow/rain balance tilts increasingly liquescent. Water is running off sooner—sometimes...     

Does larder-hoarding affect the condition of chicks in urban kestrels?

Hoarding by urban kestrels and its subsequent exploitation significantly increases the frequency and regularity of feeding during the early stage of the nestling period. Twelve kestrels’ nests situated in various part of the city were chosen to test whether hoarding could affect the chicks’ condition. Four nests (with 22 nestlings) were provided with food — one dead mouse per chick per day (F group) and 8 (43 nestlings) were used as a control group (NF). Nestlings were weighed daily, and the length of their tarsus was also measured when the oldest chick in the nest was 11 days old. 50% of the NF nests experienced a decrease in the number of nestlings but no nestlings were lost in any of the F nests. However, no significant differences were found between the groups either in body mass or tarsus length. A possible reason was the behaviour of the adults which could be interpreted as the regulation of the frequency of prey delivery depending on its presence or absence in the nest.