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Searching on patch networks using correlated random walks: space usage and optimal foraging predictions using Markov chain models.

We describe a novel representation of a discrete correlated random walk as the transition matrix of a Markov chain with the displacements as the states. Such a representation makes it possible to utilize results from the theory of absorbing Markov chains, to make biologically interesting predictions without having to resort to Monte Carlo simulations. Our motivation for constructing such a representation is to explore the relationship between the movement strategy of an animal searching for resources upon a network of patches, and its consequent utilization of space and foraging success. As an illustrative case study, we have determined the optimal movement strategy and the consequent usage of space for a central place forager utilizing a continuous movement space which is discretized as a hexagonal lattice. The optimal movement strategy determines the size of the optimal home range. In this example, the animal uses mnemokinesis, which is a sinuosity regulating mechanism, to return it to the central place. The movement strategy thus refers to the choice of the intrinsic path sinuosity and the strength of the mnemokinetic mechanism. Although the movement space has been discretized as a regular lattice in this example, the method can be readily applied to naturally compartmentalized movement spaces, such as forest canopy networks. This paper is thus an attempt at incorporating results from the theory of random walk-based animal movements into Foraging Theory.

Animals↗

Geographic profiling and animal foraging.

Geographic profiling was originally developed as a statistical tool for use in criminal cases, particularly those involving serial killers and rapists. It is designed to help police forces prioritize lists of suspects by using the location of crime scenes to identify the areas in which the criminal is most likely to live. Two important concepts are the buffer zone (criminals are less likely to commit crimes in the immediate vicinity of their home) and distance decay (criminals commit fewer crimes as the distance from their home increases). In this study, we show how the techniques of geographic profiling may be applied to animal data, using as an example foraging patterns in two sympatric colonies of pipistrelle bats, Pipistrellus pipistrellus and P. pygmaeus, in the northeast of Scotland. We show that if model variables are fitted to known roost locations, these variables may be used as numerical descriptors of foraging patterns. We go on to show that these variables can be used to differentiate patterns of foraging in these two species.

Animals↗

Modelling the many-wrongs principle: the navigational advantages of aggregation in nomadic foragers.

We develop a simple individual-based model to gain an understanding of the drivers of aggregation behaviour in nomadic foragers. The model incorporates two key elements influencing nomadic foragers in variable environments: uncertainty regarding the location of food sources and variability in the spatio-temporal distribution of ephemeral food sources. A genetic algorithm is used to evolve parameters describing an individual's movement and aggregation strategy. We apply the aggregation model to a case study of the Bornean bearded pig (Sus barbatus). Bearded pigs are ideal for considering the foraging advantages of aggregation, because they are highly mobile and exhibit a variety of aggregation strategies, ranging from solitary and sedentary to mass aggregation and wide ranging migration. Our model demonstrates the "many-wrongs principle", and shows that environmental variability, uncertainty in the location of food sources, and local population density drive aggregation behaviour.

Animals↗

Foraging in the ant Camponotus mus: nectar-intake rate and crop filling depend on colony starvation.

The effects of colony starvation on the dynamics of nectar collection were studied in individual workers of the ant Camponotus mus. A laboratory colony was first deprived of carbohydrates for 15days, and thereafter fed daily ad libitum with diluted honey until satiation. During these two successive experimental phases, the probability of feeding, crop filling and fluid-intake rates were recorded daily for individual foragers collecting a 10% (w/w) sucrose solution. The feeding responses of individuals varied with the nutritional state of the colony. When the colony was deprived of sugar, acceptance of the sucrose solution was higher than under satiation. Feeding time increased with increasing starvation. During deprivation workers fed nearly continuously on the solution, whereas a number of feeding interruptions occurred under satiation. Crop filling also increased with increasing starvation, and showed a marked decrease when the colony was satiated. Fluid-intake rate during the deprivation phase was roughly twice that during the satiation phase. This matched well with the difference in sucking frequency recorded during ingestion in satiated and starved workers, which was also higher during starvation. Results indicate that the responsiveness of foragers, determined by the nutritional state of the colony, influenced both foraging decisions and the dynamics of fluid intake.

Journal Article↗

The transfer of polychlorinated dibenzo-p-dioxins and dibenzofurans from soil into eggs of foraging chicken.

The levels of the 2,3,7,8-chlorosubstituted PCDD/Fs (polychlorinated dibenzo-p-dioxins and dibenzofurans) were determined in eggs of foraging chicken and in the corresponding soils. Compared to eggs from hens kept without soil contact the determined levels were clearly elevated. These increased levels were attributed to soil related PCDD/F uptake and were used to deduce congener specific factors describing the transfer from soil into the eggs by the foraging. Based on these transfer factors an calculation method allowing the prediction of the 2,3,7,8-TCDD toxic equivalent (I-TEq) levels in eggs based on the PCDD/F concentrations in soil was developed. The comparison of predicted and determined PCDD/F levels was used to discuss limitations and uncertainties of the model. Based on a given soil contamination, this approach allows a conservative prediction of the I-Teq levels in eggs of foraging chicken.

Animal Feed↗

Shallow groundwater quality on dairy farms with irrigated forage crops.

California's dairies are the largest confined animal industry in the state. A major portion of these dairies, which have an average herd size of nearly 1000 animal units, are located in low-relief valleys and basins. Large amounts of liquid manure are generated and stored in these dairies. In the semi-arid climate, liquid manure is frequently applied via flood or furrow irrigation to forage crops that are grown almost year-round. Little is known about the impact of manure management practices on water quality of the extensive alluvial aquifers underlying these basins. The objective of this work is to assess nitrate and salt leaching to shallow groundwater in a relatively vulnerable hydrogeologic region and to quantify the impact from individual sources on dairies. The complex array of potential point and nonpoint sources was divided into three major source areas representing farm management units: (1) manure water lagoons (ponds); (2) feedlot or exercise yard, dry manure, and feed storage areas (corrals); and (3) manure irrigated forage fields (fields). An extensive shallow groundwater-monitoring network (44 wells) was installed in five representative dairy operations in the northeastern San Joaquin Valley, CA. Water quality (electrical conductivity, nitrate-nitrogen, total Kjehldahl nitrogen) was observed over a 4-year period. Nitrate-N, reduced nitrogen and electrical conductivity (EC, salinity) were subject to large spatial and temporal variability. The range of observed nitrate-N and salinity levels was similar on all five dairies. Average shallow groundwater nitrate-N concentrations within the dairies were 64 mg/l compared to 24 mg/l in shallow wells immediately upgradient of these dairies. Average EC levels were 1.9 mS/cm within the dairies and 0.8 mS/cm immediately upgradient. Within the dairies, nitrate-N levels did not significantly vary across dairy management units. However, EC levels were significantly higher in corral and pond areas (2.3 mS/cm) than in field areas (1.6 mS/cm) indicating leaching from those management units. Pond leaching was further inferred from the presence of reduced nitrogen in three of four wells located immediately downgradient of pond berms. The estimated minimum average annual groundwater nitrate-N and salt loading from manure-treated forage fields were 280 and 4300 kg/ha, respectively. Leaching rates for ponds are estimated to be on the order of 0.8 m/year, at least locally. Since manure-treated fields represent by far the largest land area of the dairy, proper nutrient management will be a key to protecting groundwater quality in dairy regions overlying alluvial aquifers.

Animals↗

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.

Journal Article↗

Spectral-tuning mechanisms of marine mammal rhodopsins and correlations with foraging depth.

It has been observed that deep-foraging marine mammals have visual pigments that are blue shifted in terms of their wavelength of maximal absorbance (lambda(max)) when compared to analogous pigments from terrestrial mammals. The mechanisms underlying the spectral tuning of two of these blue-shifted pigments have recently been elucidated and depend on three amino acid substitutions (83Asn, 292Ser, and 299Ser) in dolphin rhodopsin, but only one amino acid substitution (308Ser ) in the dolphin long-wavelength-sensitive pigment. The objective of this study was to investigate the molecular basis for changes in the spectral sensitivity of rod visual pigments from seven distantly related marine mammals. The results show a relationship between blue-shifted rhodopsins (lambda(max) < or = 490 nm), deep-diving foraging behavior, and the substitutions 83Asn and 292Ser. Species that forage primarily near the surface in coastal habitats have a rhodopsin with a lambda(max) similar to that of terrestrial mammals (500 nm) and possess the substitutions 83Asp and 292Ala, identical to rhodopsins from terrestrial mammals.

Animals↗

Reveal protein molecular structural-chemical differences between two types of winterfat (forage) seeds with physiological differences in low temperature tolerance using synchrotron-based Fourier transform infrared microspectroscopy.

Winterfat (Krascheninnikovia lanata) (forage seed) is a long-lived native shrub with superior forage quality for livestock and wildlife. The objectives of this study were to use advanced synchrotron technology [S-Fourier transform infrared microspectroscopy (FTIR)] as a novel approach to reveal protein molecular structural-chemical differences in terms of protein secondary structures between the two types of winterfat (forage) seeds, which show physiological differences in low-temperature tolerances. This experiment was performed at beamline U10B at the National Synchrotron Light Source (NSLS) in Brookhaven National Laboratory (BNL), U.S. Department of Energy (NSLS-BNL, New York). The results showed that with the synchrotron analytical technique (S-FTIR), the molecular structural-chemical makeup and characteristics of the winterfat seed tissues could be imaged and revealed. The protein secondary structures differed between the large and the small seed tissues. By using the multicomponent peaks modeling method, the results show that the large seeds contained no significant differences (P > 0.05) in percentage of beta-sheet (average 37.0%) and alpha-helix (average 24.1%). However, the large seeds contained a lower (P < 0.05) percentage of beta-turns (18.1 vs 20.1%) and a lower (P < 0.05) ratio of beta-turns to alpha-helices (0.8 vs 0.9) and beta-turns to beta-sheets (0.5 vs 0.6). Our results demonstrate the potential of highly spatially resolved synchrotron-based FTIR microspectroscopy to reveal differences of structural molecular chemistry and protein secondary structures, which are associated with seed size variation and may affect germination behaviors.

Amaranthaceae↗

Quantification of fatty acids in forages by near-infrared reflectance spectroscopy.

Near-infrared reflectance spectroscopy (NIRS) was evaluated as a possible alternative to gas chromatography (GC) for the quantitative analysis of fatty acids in forages. Herbage samples from 11 greenhouse-grown forage species (grasses, legumes, and forbs) were collected at three stages of growth. Samples were freeze-dried, ground, and analyzed by GC and NIRS techniques. Half of the 195 samples were used to develop an NIRS calibration file for each of eight fatty acids, with the remaining half used as a validation data set. Spectral data, collected over a wavelength range of 1100-2498 nm, were regressed against GC data to develop calibration equations for lauric (C12:0), myristic (C14:0), palmitic (C16:0), stearic (C18:0), palmitoleic (C16:1), oleic (C18:1), linoleic (C18:2), and alpha-linolenic (C18:3) acids. Calibration equations had high coefficients of determination for calibration (0.93-0.99) and cross-validation (0.89-0.98), and standard errors of calibration and cross-validation were < 20% of the respective means. Simple linear regressions of NIRS results against GC data for the validation data set had r2 values ranging from 0.86 to 0.97. Regression slopes for C12:0, C14:0, C16:0, C18:0, C16:1, C18:2, and C18:3 were not significantly different (P = 0.05) from 1.0. The regression slope for C18:1 was 1.1. The ratio of standard error of prediction to standard deviation was > 3.0 for all fatty acids except C12:0 (2.6) and C14:0 (2.9). Validation statistics indicate that NIRS has high prediction ability for fatty acids in forages. Calibration equations developed using data for all plant materials accurately predicted concentrations of C16:0, C18:2, and C18:3 in individual plant species. Accuracy of prediction was less, but acceptable, for fatty acids (C12:0, C14:0, C18:0, C16:1, and C18:1) that were less prevalent.

Animal Feed↗

Mode of foraging competition is related to tutor preference in Zenaida aurita.

This study compared the direction of social learning in 2 populations of Barbados Zenaida doves (Zenaida aurita). One population (St. James) is territorial; it competes aggressively with conspecifics but scramble competes with heterospecifics. The other population (Deep Water Harbour) forages in large homospecific flocks. Field observations were conducted to quantify intraspecific and interspecific patterns of foraging association and aggression. Wild-caught doves from both areas were then tested on novel foraging tasks demonstrated by either a conspecific or a heterospecific tutor. In all experiments, St. James doves learned more readily from the heterospecific tutor (Carib grackle -Quiscalus lugubris-), whereas Deep Water Harbour doves learned more readily from the conspecific tutor. The type of competitive feeding interaction in the field (i.e., scramble vs. interference) appears to better predict the pattern of social learning in an experiment than does species identity.

Aggression↗

Developmental changes in manipulation in tufted capuchins (Cebus apella) from birth through 2 years and their relation to foraging and weaning.

This study examined the contributions of physical and sensorimotor development to manipulation in capuchins (Cebus apella) from birth to 2 years. Between months 1-6 and 7-12, manipulation increased significantly in frequency, in the proportion that was vigorous or required fine motor control, and in the proportion directed at portable objects. Fine motor control, moving objects in relation to the body, and stamina are largely in place by 12 months, after which little changed. All elements of the manipulative repertoire have appeared, and vigorous and dexterous activities have peaked before fully independent foraging. Emergence of permanent dentition and achievement of approximately half of adult body size accompany the attainment of fully independent foraging at 15 months. Thereafter, increasing strength and specific knowledge probably contribute more to changing foraging competence in young capuchins than do stamina and sensorimotor development.

Aging↗

Use of visual, acoustic, and olfactory information during embedded invertebrate foraging in brown capuchins (Cebus apella).

Experiments were conducted to investigate which sensory cues are used by brown capuchins (Cebus apella) in embedded invertebrate foraging. The importance of visual, olfactory, and acoustic cues in such foraging was determined by presenting subjects with a stimulus log modified to block out given sensory cues. Experiment 1 was designed to investigate whether subjects could locate an invertebrate embedded in wood when only visual, acoustic, or olfactory information was available. Experiments 2 and 3 were designed to investigate extractive foraging behavior when two sensory cues were provided. It was hypothesized that the combination of visual and acoustic information would be necessary for subjects to successfully locate embedded invertebrates. Results indicated that subjects' performance was most successful when both visual and acoustic information was available.

Animals↗

Asymptotic prey profitability drives star-nosed moles to the foraging speed limit.

Foraging theory provides models for predicting predator diet choices assuming natural selection has favoured predators that maximize their rate of energy intake during foraging. Prey profitability (energy gained divided by prey handling time) is an essential variable for estimating the optimal diet. Time constraints of capturing and consuming prey generally result in handling times ranging from minutes to seconds, yet profitability increases dramatically as handling time approaches zero, providing the potential for strong directional selection for increasing predator speed at high encounter rates (tiny increments in speed increase profitability markedly, allowing expanded diets of smaller prey). We provide evidence that the unusual anatomical and behavioural specializations characterizing star-nosed moles resulted from progressively stronger selection for speed, allowing the progressive addition of small prey to their diet. Here we report handling times as short as 120 ms (mean 227 ms) for moles identifying and eating prey. 'Double takes' during prey identification suggest that star-nosed moles have reached the speed limit for processing tactile information. The exceptional speed of star-nosed moles, coupled with unusual specializations for finding and eating tiny prey, provide new support for optimal foraging theory.

Animals↗

Improved forage digestibility of tall fescue (Festuca arundinacea) by transgenic down-regulation of cinnamyl alcohol dehydrogenase.

Lignification of cell walls during plant development has been identified as the major factor limiting forage digestibility and concomitantly animal productivity. cDNA sequences encoding a key lignin biosynthetic enzyme, cinnamyl alcohol dehydrogenase (CAD), were cloned from the widely grown monocotyledonous forage species tall fescue (Festuca arundinacea Schreb.). Recombinant tall fescue CAD expressed in E. coli exhibited the highest V(max)/K(m) values when coniferaldehyde and sinapaldehyde were used as substrates. Transgenic tall fescue plants carrying either sense or antisense CAD gene constructs were obtained by microprojectile bombardment of single genotype-derived embryogenic suspension cells. Severely reduced levels of mRNA transcripts and significantly reduced CAD enzymatic activities were found in two transgenic plants carrying sense and antisense CAD transgenes, respectively. These CAD down-regulated transgenic lines had significantly decreased lignin content and altered ratios of syringyl (S) to guaiacyl (G), G to p-hydroxyphenyl (H) and S to H units. No significant changes in cellulose, hemicellulose, neutral sugar composition, p-coumaric acid and ferulic acid levels were observed in the transgenic plants. Increases of in vitro dry matter digestibility of 7.2-9.5% were achieved in the CAD down-regulated lines, thus providing a novel germplasm to be used for the development of grass cultivars with improved forage quality.

Journal Article↗

Influence of the method of forage conservation on feeding behaviour, intake and characteristics of reticulo-rumen content, in sheep fed ad libitum.

The influence of silage conservation methods on eating behaviour and the characteristics of reticulo-rumen contents was studied in sheep by comparing 3 forages, a silage without additive (WAS), one with additive (FAS) and hay (H), prepared from the same cut green forage. The sheep were rumen fistulated. WAS was very badly and FAS poorly preserved while the hay was dried under favourable conditions. The forages were fed ad libitum and the dry matter (DM) intake was 1,054, 1,241 and 1,469 g/day for WAS, FAS and H respectively. There was a single feeding in the morning. At the main meal, DM intake was 270, 317 and 388 g/day and ingestion rate 4.76, 4.56 and 4.16 g of DM/min for WAS, FAS and H respectively. There was slight recovery in ingestion around 16-19 h with hay and FAS but not with WAS. With hay, rumination lasted much longer than with the silages and began sooner after the end of the main meal. In contrast, overall rumination efficiency was the same for hay and FAS. There were fewer contractions of the reticulo-rumen with both WAS and hay. The amounts of reticulo-rumen contents were comparable for the 2 silages and higher for hay. The amounts of NH3 in the contents were the same with WAS and hay but greater with FAS. With WAS, contents were richer in butyric, valeric and caproic acids. The DM turnover rate of the contents was the same for FAS and hay and lower, but not significantly, for WAS. Accordingly, with silage, satiety seems to be rapidly reached but there was no evidence that organoleptic factors were involved. There were no problems with the digestion rate of silages. However, the factors limiting silage intake persisted throughout the diurnal cycle, and reticulo-rumen fill was affected. The more poorly preserved the silage the more these factors reduced the duration and volume of the meals, or both.

Animals↗

Infections and intoxications associated with animal feed and forage which may present a hazard to human health.

Animal feed or forage may be the source of a limited number of infections for farm animals that could lead to human illness. Likely organisms include Salmonella enterica, Toxoplasma gondii, Trichinella spiralis and possibly the agent of bovine spongiform encephalopathy. The risk to human health from other infectious agents which may contaminate either feed or forage appear to be either negligible, e.g. Bacillus anthracis and Mycobacterium bovis, or non-existent, e.g. Clostridium botulinum toxin and Listeria monocytogenes. Mycotoxins present in animal feed can result in foods of animal origin also containing them. This risk is well recognized but has yet to be quantified accurately and in some instances the risk may be of theoretical rather than practical importance. Pesticides, agricultural and industrial chemicals, heavy metals and radionuclides may pollute animal feed and forages. The methods available for controlling pollution from these sources are well understood from a technical point of view although the effective implementation of controls can be difficult.

Animal Diseases↗

Foraging biology predicts food web complexity.

Food webs, the networks of feeding links between species, are central to our understanding of ecosystem structure, stability, and function. One of the key aspects of food web structure is complexity, or connectance, the number of links expressed as a proportion of the total possible number of links. Connectance (complexity) is linked to the stability of webs and is a key parameter in recent models of other aspects of web structure. However, there is still no fundamental biological explanation for connectance in food webs. Here, we propose that constraints on diet breadth, driven by optimal foraging, provide such an explanation. We show that a simple diet breadth model predicts highly constrained values of connectance as an emergent consequence of individual foraging behavior. When combined with features of real food web data, such as taxonomic and trophic aggregation and cumulative sampling of diets, the model predicts well the levels of connectance and scaling of connectance with species richness, seen in real food webs. This result is a previously undescribed synthesis of foraging theory and food web theory, in which network properties emerge from the behavior of individuals and, as such, provides a mechanistic explanation of connectance currently lacking in food web models.

Animals↗