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Ethanol-maintained behavior in a foraging context: effects of search and procurement cost.

In a simulated foraging procedure, the effects of manipulating search and procurement costs of access to ethanol, food, and water on ethanol self-administration were determined in rats. Completion of a response ratio on one lever (search) resulted in the opportunity to complete a high- or low-cost response ratio on one of three other levers (procurement). Each procurement lever was exclusively associated with access to ethanol, food, or water, and the six procurement opportunities occurred in random order. Any procurement opportunity could be rejected, and a new search subsequently initiated. Percent acceptance of high-cost opportunities varied directly with increases in search cost and inversely with increases in high procurement cost. When search cost was increased, daily water and food consumption decreased, whereas daily ethanol consumption was unaffected. Regardless of these manipulations, rats consumed 1.0-2.0 g/kg ethanol daily. Ethanol consumption also was unaffected by increases in low or high food procurement cost values. These results indicate that ethanol self-administration can be analyzed within a foraging environment in which the animal initiates and determines the length and size of all drinking bouts and meals. Within this environment, patterns of ethanol self-administration were orderly and distinguishable from patterns of food- and water-maintained behavior.

Animals↗

Intake and digestibility of berseem (Medicago sativa) and Sorghum Abu 70 (Sorghum vulgare) forages by Sudan zebu cattle and desert sheep.

Two experiments were carried out with Sudan zebu cattle and desert sheep to study the voluntary intake and digestibility of berseem (Medicago sativa) and Abu 70 (Sorghum vulgare) forages when offered ad lib. alone and with supplements of molasses to the cattle and dura grain to the sheep. In both experiments the intake of berseem was significantly higher than Abu 70. Supplementation increased Abu 70 intake and while molasses significantly increased berseem intake by cattle, dura decreased its intake by sheep. On the basis of metabolic body size sheep showed a higher intake of these forages than cattle. There was no significant difference in DM digestibility between berseem and Abu 70 by cattle. Sheep digested berseem significantly better than Abu 70. Molasses and dura significantly increased the digestibility of Abu 70 by cattle and sheep but had no significant effect on the DM digestibility of berseem.

Animals↗

Foraging behaviour in tadpoles of the bronze frog Rana temporalis: experimental evidence for the ideal free distribution.

The ability of bronze frog Rana temporalis tadpoles (pure or mixed parental lines) to assess the profitability of food habitats and distribute themselves accordingly was tested experimentally using a rectangular choice tank with a non- continuous input design. Food (boiled spinach) was placed at two opposite ends of the choice tank in a desired ratio (1 : 1, 1 : 2 or 1 : 4) to create habitat A and B. The tadpoles in Gosner stage 28-33, pre-starved for 24 h, were introduced in an open ended mesh cylinder placed in the center of the choice tank, held for 4 min (for acclimation) and then released to allow free movement and habitat selection. The number of tadpoles foraging at each habitat was recorded at 10, 15, 20, 25 and 30 min time intervals. The actual suitability, Si (the food available in a habitat after colonization of tadpoles) of each habitat was obtained from the equation Si = Bi - fi (di) where Bi is basic suitability (amount of food provided at each habitat before release of tadpoles), fi is the rate of depletion of food (lowering effect) with introduction of each tadpole, and di is the density of tadpoles in habitat i. The expected number of tadpoles at each habitat was derived from the actual suitability. With no food in the choice tank, movement of the tadpoles in the test arena was random indicating no bias towards any end of the choice tank or the procedure. In tests with a 1 : 1 food ratio, the observed ratio of tadpoles (11.71 : 12.28) was comparable with the expected 12 : 12 ratio. The observed number of tadpoles in the habitats with a 1 : 2 food ratio was 8.71 : 15.29 and 7.87 : 16.13 for pure and mixed parental lines respectively. In both cases, the observed ratios were close to the expected values (7 : 17). Likewise, in experiments with a 1 : 4 food ratio, the observed number of tadpoles in the two habitats (10.78 : 37.22) did not differ significantly from the expected ratio of 7 : 41. In all tests, the number of R. temporalis tadpoles matched ideally with habitat profitability (undermatching index K approximate, equals 1). The study shows that tadpoles of the bronze frog exhibit an ideal free distribution while foraging regardless of whether they are siblings or non-siblings in a group, which correlates well with their group living strategy in nature

Animals↗

Chemical compounds of the foraging recruitment pheromone in bumblebees.

When the frenzied and irregular food-recruitment dances of bumblebees were first discovered, it was thought that they might represent an evolutionary prototype to the honeybee waggle dance. It later emerged that the primary function of the bumblebee dance was the distribution of an alerting pheromone. Here, we identify the chemical compounds of the bumblebee recruitment pheromone and their behaviour effects. The presence of two monoterpenes and one sesquiterpene (eucalyptol, ocimene and farnesol) in the nest airspace and in the tergal glands increases strongly during foraging. Of these, eucalyptol has the strongest recruitment effect when a bee nest is experimentally exposed to it. Since honeybees use terpenes for marking food sources rather than recruiting foragers inside the nest, this suggests independent evolutionary roots of food recruitment in these two groups of bees.

Animal Feed↗

High levels of linkage disequilibrium and associations with forage quality at a phenylalanine ammonia-lyase locus in European maize (Zea mays L.) inbreds.

Forage quality of maize is influenced by both the content and structure of lignin in the cell wall. Phenylalanine Ammonia-Lyase (PAL) catalyzes the first step in lignin biosynthesis in plants; the deamination of L-phenylalanine to cinnamic acid. Successive enzymatic steps lead to the formation of three monolignols, constituting the complex structure of lignin. We have cloned and sequenced a PAL genomic sequence from 32 maize inbred lines currently employed in forage maize breeding programs in Europe. Low nucleotide diversity and excessive linkage disequilibrium (LD) was identified at this PAL locus, possibly reflecting selective constrains resulting from PAL being the first enzyme in the monolignol, and other, pathways. While the association analysis was affected by extended LD and population structure, several individual polymorphisms were associated with neutral detergent fiber (not considering population structure) and a single polymorphism was associated with in vitro digestibility of organic matter (considering population structure).

Genetic Markers↗

PKA and PKC content in the honey bee central brain differs in genotypic strains with distinct foraging behavior.

Selection of honey bees for pollen storage resulted in high and low pollen-hoarding strains differing in foraging behavior traits including resource choice and quality, load size, sucrose responsiveness, age of foraging initiation, and learning performance. To determine how these genotypic differences correlate with changes at the level of proteins involved in neuronal function, we measured the content of protein kinase A, protein kinase C, and synapsin in the brains of high- and low-strain bees. In the central brain protein kinase A and protein kinase C levels were greater in high-strain bees and increased from emergence to 5 days in both strains. By 15 days, high-strain bees retained significantly higher levels of protein kinase C than low-strain bees, but overall protein kinase C content decreased in both strains. Synapsin levels increased from emergence to 5 days but did not differ between the two strains. In contrast to the protein kinase A content in the central brain, the basal protein kinase A activity did not differ between the strains or between the two age groups. This provides first evidence that the two genetic strains of honey bees show characteristic differences in the regulation of protein expression that may contribute to the behavioral differences between them.

Animals↗

Changes in the metabolic rate of the foraging honeybee: effect of the carried weight or of the reward rate?

The metabolic rate of free flying honeybees (Apis mellifera ligustica) foraging on a multiple automatic feeder was measured in complete absence of perturbation. Each time the sucrose flow rate was doubled, the metabolic rate increased by 18.2 +/- 2.0% (14.7 microl CO2 min(-1) and final crop load by 25.1 +/- 2.4% (7.04 microl). The possibility that the heavier load carried by the bees caused the increase in the metabolic rate was analyzed in detail. It was found that, for the same weight in the crop, the metabolic rate increased with the increasing reward rate. Therefore, a factor other than the carried weight might account for this increase: a motivational drive, whose intensity may depend on the reward rate at the food source. Although at higher reward rates metabolic rate increased during the visit, at lower reward rates it remained constant, suggesting that the effect of the carried weight on the metabolic rate might be controlled by this motivational drive. The hypothesis that honeybees maximize individual efficiency by reducing their crop load loses support, as foraging costs are not determined by the carried weight. The functional meaning of the reduction in crop load would be to increase the informational exchange at the hive.

Animals↗

Energetics of foraging and locomotion in the platypus Ornithorhynchus anatinus.

We measured the energy requirements of platypuses foraging, diving and resting in a swim tank using flow-through respirometry. Also, walking metabolic rates were obtained from platypuses walking on a conventional treadmill. Energy requirements while foraging were found to depend on water temperature, body weight and dive duration and averaged 8.48 W kg(-1). Rates for subsurface swimming averaged 6.71 W kg(-1). Minimal cost of transport for subsurface swimming platypuses was 1.85 J N(-1)m(-1) at a speed of 0.4 m s(-1). Aerobic dive limit of the platypus amounted to 59 s. Metabolic rate of platypuses resting on the water surface was minimal with 3.91 W kg(-1) while minimal RMR on land was 2.08 W kg(-1). The metabolic rate for walking was 8.80 W kg(-1) and 10.56 W kg(-1) at speeds of 0.2 m s(-1) and 0.3 m s(-1), respectively. A formula was derived, which allows prediction of power requirements of platypuses in the wild from measurements of body weight, dive duration and water temperature. Platypuses were found to expend energy at only half the rate of semiaquatic eutherians of comparable body sizes during both walking and diving. However, costs of transport at optimal speed were in line with findings for eutherians. These patterns suggest that underwater locomotion of semiaquatic mammals have converged on very similar efficiencies despite differences in phylogeny and locomotor mode.

Animals↗

Effects of opportunistic predation on anti-predator behavioural responses in a guild of ground foragers.

We studied factors that affect prey selection by a generalist predator that opportunistically attacks prey species, and the associated inter- and intra-specific responses of prey to this type of predation. Our model system was a guild of ground-foraging birds that are preyed upon by magpies ( Pica pica) during the breeding season. We found that magpies attacked up to 12 species during three consecutive breeding seasons. The overall capture success was estimated to be 4.9%. Magpies tended to attack from the air, targeting solitary prey, either on the ground or flying. Inter-specific prey responses to the risk of magpie predation included a reduction in the mean number of species occupying a foraging patch when magpies were present and a decrease in the distance between heterospecific neighbours. Intra-specific responses to magpie predation varied between species that were subject to different attack rates. Preferentially attacked prey enhanced their risk responses (increase in scanning time and scanning rate in the presence of magpies) relative to those species attacked in proportion to their abundance (increase only in scanning rate with magpies). Species attacked infrequently, relative to their abundance, showed no antipredator response. The probability of being attacked, rather than mortality rate, appears to be the factor to which prey species respond.

Animals↗

Bush selection along foraging pathways by sympatric impala and greater kudu.

In order to identify the selection mechanism of two sympatric African browsers, we analysed encounter rates and selection of bushes along foraging pathways. We monitored the tracks, left overnight, by kudu and impala on an experimental plot of natural Acacia nilotica and Dichrostachys cinerea in the highveld of Zimbabwe, and recorded the number of bushes attacked in each category. Both ungulates were selective for the bush categories, but kudu were consistently more selective than impala, and showed a higher preference for the larger A. nilotica and D. cinerea bushes, which had a significantly greater number of bites which were not reachable by impala. For both kudu and impala, the probability of attacking larger bushes increased significantly with the proportion of large bushes encountered along the foraging pathways, whereas the consumption of smaller bushes was apparently unpredictable. For the most abundant food item (medium D. cinerea), the probability of attack by impala along a pathway decreased with increasing proportions of larger bushes in the experimental area, but was also dependent on impala group size and season. In addition, we found that encounter rates with larger bushes were significantly higher for kudu than for impala. Experimentally reducing the availability of the larger bushes had little effect on both impala and kudu during the following rainy season. However, during the following cool dry season, kudu showed an increased selectivity with a strong preference for the remaining large bushes (large A. nilotica), followed by a sharp decrease in selectivity in the hot dry season when they also fed from significant numbers of medium trees. Impala had little reaction to the experimental changes in the availability of bush categories in either season. We suggest that both kudu and impala selected bushes on the basis of the potential number of bites they can provide, and this resulted in different search strategies. Kudu focussed on the larger bushes which have a larger number of twigs which are out of reach of impala and kudu also probably directed their path preferentially towards the few larger bushes to maximize encounter rates with this favoured bush category. These differences in bush selection process lead to a low overlap in resource use between the two browsers in this type of savanna.

Animals↗

Resource partitioning through oceanic segregation of foraging juvenile southern elephant seals ( Mirounga leonina).

In highly dynamic and unpredictable environments such as the Southern Ocean, species that have evolved behaviors that reduce the effects of intra-specific competition may have a selective advantage. This is particularly true when juveniles face disadvantages when foraging due to morphological or physiological limitation, which is the case for many marine mammals. We tracked the at-sea movements of 48 juvenile southern elephant seals ( Mirounga leonina) between the ages of 1 and 4 years from the population at Macquarie Island using locations derived from recorded light levels. There were significant differences in the total amount of the Southern Ocean covered by the different age-groups. The younger seals used a smaller area than the older seals. On average, the younger individuals also made more trips to sea than the older seals and did not travel as far on each trip. Females spent more time at sea than males and there were no significant differences between the total areas used by male and females. In summary, younger seals remained closer to the island at all times, and they spent more time in more northerly regions that older seals. These differences in behavior created temporal and spatial segregation between juveniles of different ages. Therefore, we suggest that these temporal and spatial separations help to avoid intra-specific competition for resources on land, space on beaches, and at-sea foraging areas. Such modifications of haul-out timing and behavior enable them to exploit a patchy and unpredictable environment.

Age Factors↗

Animal-plant-microbe interactions: direct and indirect effects of swan foraging behaviour modulate methane cycling in temperate shallow wetlands.

Wetlands are among the most important ecosystems on Earth both in terms of productivity and biodiversity, but also as a source of the greenhouse gas CH(4). Microbial processes catalyzing nutrient recycling and CH(4) production are controlled by sediment physico-chemistry, which is in turn affected by plant activity and the foraging behaviour of herbivores. We performed field and laboratory experiments to evaluate the direct effect of herbivores on soil microbial activity and their indirect effects as the consequence of reduced macrophyte density, using migratory Bewick's swans (Cygnus columbianus bewickii Yarrell) feeding on fennel pondweed (Potamogeton pectinatus L.) tubers as a model system. A controlled foraging experiment using field enclosures indicated that swan bioturbation decreases CH(4) production, through a decrease in the activity of methanogenic Archaea and an increased rate of CH(4) oxidation in the bioturbated sediment. We also found a positive correlation between tuber density (a surrogate of plant density during the previous growth season) and CH(4) production activity. A laboratory experiment showed that sediment sterilization enhances pondweed growth, probably due to elimination of the negative effects of microbial activity on plant growth. In summary, the bioturbation caused by swan grazing modulates CH(4) cycling by means of both direct and indirect (i.e. plant-mediated) effects with potential consequences for CH(4) emission from wetland systems.

Animals↗

Herbivore-induced infochemicals influence foraging behaviour in two intertidal predators.

Herbivore-induced defences appear ubiquitous across most biomes and habitats. Yet the direct correlation between induced changes in host plant chemistry and the population dynamics of the herbivore remain untested in many systems. In plant-herbivore interactions in the terrestrial environment, indirect or tritrophic interactions appear a successful way in which changes in the host plant chemistry induced by prior herbivory can impact on herbivore populations via increased success of natural enemies. This set of interactions remains untested in the marine system. Here, we present work from experiments using orthogonal contrasts of plants with different prior treatments (control, mechanical damage or herbivory) and the presence or absence of herbivores on the foraging behaviour of a crab, Carcinus maenas, and a fish, Lipophrys pholis. These experiments were carried out using a novel flow-through flume, i.e. as a choice chamber supplied by turbulent water from independent cue sources. Our results show that in the Ascophyllum nodosum (plant)-Littorina obtusata (herbivore) system infochemicals from induced plants can directly influence predator foraging behaviour. L. pholis was attracted to the presence of a feeding L. obtusata, but was also more attracted to odours from herbivore-induced tissue than odours from mechanically damaged or naïve A. nodosum. C. maenas was more attracted to odours from herbivore-induced tissue compared to naïve tissue, regardless of the presence of L. obtusata. This is the first demonstration of such behavioural consequences of herbivore-induced changes in plants for marine systems.

Analysis of Variance↗

Chunking versus foraging search patterns by rats in the hierarchically baited radial maze.

Rats were exposed to a radial maze containing six black smooth arms and six wire-grid-covered arms and a striped 'exit arm' in experiment 1. The probability of a black or grid arm being baited (5/6 vs 1/6) with sunflower seeds was associated with its proximal cue for some rats (the Relevant Arm Cue group) but not for others (the Irrelevant Arm Cue group). All rats could terminate a trial and receive a highly preferred morsel of apple by entering the exit arm only after having sampled all six seed-baited arms. Relevant Arm Cue rats usually chose some arms from the more densely baited set before choosing an arm from the less densely baited set and made fewer reentries than Irrelevant Arm Cue rats. Although such clustered, higher choice accuracy in the Relevant Arm Cue group corresponds to human clustered, better recall of verbal items from lists hierarchically organized by categories, these rats did not similarly exhaustively retrieve items (arm locations). That is, when required to terminate a trial by entering the 'exit' arm for an apple morsel after having sampled all seed-baited arms, both groups were equally unable to withhold making nonrewarded premature exits. This nonexhaustive foraging search pattern was maintained in the next two experiments in which the radial maze was reduced to three black and three grid arms along with the striped 'exit' arm and in which black and grid arm cues were paired with number of seeds (eight or one) in an arm for Relevant Arm Cue rats. Although Relevant Arm Cue rats displayed perfect clustering by entering all eight-seeded arms before a one-seeded arm, they made more premature exits and reentries into eight-seeded arms in experiment 2 or when forced to enter all eight-seeded arms in experiment 3 than did Irrelevant Arm Cue rats. These foraging tendencies prevent accurate estimations of the amount of information (i.e., arm locations) rats can 'chunk'.

Animals↗

Effects of dietary nutrients and foraging costs on meal patterns of rats.

Rats were fed either a cereal-based or a purified casein-based diet in a foraging paradigm in which the costs of procurement and consumption were varied. The group offered the cereal-based diet consumed about 10% more calories than the group offered the casein-based diet, but both groups grew at the same rate. The intake of a control group offered a choice between the two diets was approximately 80% from the casein diet, and the growth of this group did not differ from that of the experimental groups. Variations in the cost of procurement and the cost of consumption affected the patterning of meals differentially for the two diets: changes in meal patterns tended to control the time and/or energy spent feeding. These results show that (1) meal patterns in the foraging paradigm are sensitive to subtle differences in diets, and (2) the amount of diet consumed (acceptance) and the choice between diets (preference) are determined by the economics of feeding and the nutritive quality of the foods, as well as by their palatability.

Animal Feed↗

Macronutrient selection by foraging rats.

Rats were studied in a laboratory simulation of foraging for separate sources of carbohydrate, fat, and protein. Fixed ratios of bar-presses were required to search for a cue signalling the availability of a meal and to procure each meal. In the first phase, the costs of procuring all three foods were raised equally for three rats. Meal frequency declined and meal size increased for all three foods. Protein opportunities were accepted more frequently than those for other foods, and the proportion of calories taken from protein remained low and constant. Two rats were studied in the remaining three phases. When protein meal cost was raised while the costs of energy foods were low, protein intake was gradually reduced and at the highest cost was dropped from the diet. There were individual differences in the preferred source of energy calories; when each animal's preferred energy source was made more costly, these preferences persisted over the range of costs studied in one rat, but shifted abruptly in the other rat. Finally, when search costs were raised, the animals became less selective in accepting meal opportunities and composing their diets. These results suggest that diet selection in the rat is flexible and responsive to changes in the costs of acquiring food. This ability is consistent with the foraging behaviors required by omnivorous generalized feeders.

Animals↗

Dietary amino acid complementation as a foraging strategy for wild birds.

Foraging birds finding foods containing protein of various quality could satisfy their amino acid needs by either 1) choosing only those foods that contain a suitable array of essential amino acids, or 2) choosing foods in amounts that permit complementation of constituent amino acids. We studied the abilities of sparrows (Zonotrichia leuchophrys gambelii) to use dietary amino acid complementation as a foraging strategy. We conducted two series of tests: 1) to determine if sparrows could select adequate portions from food pairs in order to compensate for reciprocal dietary deficiencies of either valine and lysine or lysine and threonine, and 2) to evaluate to what extent fortuitous (passive) complementation of dietary amino acids was constrained in sparrows by the magnitude of the dietary amino acid deficiency and the schedule of daily intakes of complementary foods. Small birds appear to have limited abilities to actively exploit dietary amino acid complementation. Birds offered a choice of two foods deficient in either lysine, valine, or threonine but of complementary compositions, reduced their daily intakes of food and body masses, but to a lesser extent (ca. one-half to one-third, and ca. one-quarter as much, respectively) than when either of the deficient foods were offered alone. Even passive complementation of dietary amino acids was severely constrained by both the magnitude of the dietary amino acid deficiency and the time course of feeding. The greater the amino acid deficiency, the less effective the complementation of dietary amino acids. The time course for effective and efficient complementation of dietary essential amino acids in these small, metabolically active animals was less than 2 h.

Amino Acids↗

Frontal forebrain lesions: effects on the foraging and apomorphine pecking of pigeons.

The role of the nucleus basalis prosencephali (Bas), a frontal forebrain structure peculiar to birds, in the control of forage pecking and apomorphine-induced pecking was investigated. In a quasi-natural grit-grain selection task bilateral coagulations of the Bas and the associated neostratum frontolaterale (Nfl) caused a marked fall in grain per peck uptake and a simultaneous increase in grit per peck uptake. Bas lesions also had a reducing effect on the compulsive pecking elicited by systemic injections of apomorphine. This confirms that the Bas is involved in the control of forage pecking and dopaminergic pecking of birds.

Animals↗