|
|
Viscido, S. V., Miller, M., & Wethey, D. S. (2002). The dilemma of the selfish herd: the search for a realistic movement rule. J. Theor. Biol., 217(2), 183–194.
Abstract: The selfish herd hypothesis predicts that aggregations form because individuals move toward one another to minimize their own predation risk. The “dilemma of the selfish herd” is that movement rules that are easy for individuals to follow, fail to produce true aggregations, while rules that produce aggregations require individual behavior so complex that one may doubt most animals can follow them. If natural selection at the individual level is responsible for herding behavior, a solution to the dilemma must exist. Using computer simulations, we examined four different movement rules. Relative predation risk was different for all four movement rules (p<0.05). We defined three criteria for measuring the quality of a movement rule. A good movement rule should (a) be statistically likely to benefit an individual that follows it, (b) be something we can imagine most animals are capable of following, and (c) result in a centrally compact flock. The local crowded horizon rule, which allowed individuals to take the positions of many flock-mates into account, but decreased the influence of flock-mates with distance, best satisfied these criteria. The local crowded horizon rule was very sensitive to the animal's perceptive ability. Therefore, the animal's ability to detect its neighbors is an important factor in the dynamics of group formation.
|
|
|
|
Viscido, S. V., Miller, M., & Wethey, D. S. (2001). The response of a selfish herd to an attack from outside the group perimeter. J. Theor. Biol., 208(3), 315–328.
Abstract: According to the selfish herd hypothesis, animals can decrease predation risk by moving toward one another if the predator can appear anywhere and will attack the nearest target. Previous studies have shown that aggregations can form using simple movement rules designed to decrease each animal's Domain of Danger. However, if the predator attacks from outside the group's perimeter, these simple movement rules might not lead to aggregation. To test whether simple selfish movement rules would decrease predation risk for those situations when the predator attacks from outside the flock perimeter, we constructed a computer model that allowed flocks of 75 simulated fiddler crabs to react to one another, and to a predator attacking from 7 m away. We attacked simulated crab flocks with predators of different sizes and attack speeds, and computed relative predation risk after 120 time steps. Final trajectories showed flight toward the center of the flock, but curving away from the predator. Path curvature depended on the predator's size and approach speed. The average crab experienced a greater decrease in predation risk when the predator was small or slow moving. Regardless of the predator's size and speed, however, predation risk always decreased as long as crabs took their flock-mates into account. We conclude that, even when flight away from an external predator occurs, the selfish avoidance of danger can lead to aggregation.
|
|
|
|
Suzuki, Y., & Toquenaga, Y. (2005). Effects of information and group structure on evolution of altruism: analysis of two-score model by covariance and contextual analyses. J. Theor. Biol., 232(2), 191–201.
Abstract: An altruistic individual has to gamble on cooperation to a stranger because it does not know whether the stranger is trustworthy before direct interaction. Nowak and Sigmund (Nature 393 (1998a) 573; J. Theor. Biol. 194 (1998b) 561) presented a new theoretical framework of indirect reciprocal altruism by image scoring game where all individuals are informed about a partner's behavior from its image score without direct interaction. Interestingly, in a simplified version of the image scoring game, the evolutionarily stability condition for altruism became a similar form of Hamilton's rule, i.e. inequality that the probability of getting correct information is more than the ratio of cost to benefit. Since the Hamilton's rule was derived by evolutionarily stable analysis, the evolutionary meaning of the probability of getting correct information has not been clearly examined in terms of kin and group selection. In this study, we applied covariance analysis to the two-score model for deriving the Hamilton's rule. We confirmed that the probability of getting correct information was proportional to the bias of altruistic interactions caused by using information about a partner's image score. The Hamilton's rule was dependent on the number of game bouts even though the information reduced the risk of cooperation to selfish one at the first encounter. In addition, we incorporated group structure to the two-score model to examine whether the probability of getting correct information affect selection for altruism by group selection. We calculated a Hamilton's rule of group selection by contextual analysis. Group selection is very effective when either the probability of getting correct information or that of future interaction, or both are low. The two Hamilton's rules derived by covariance and contextual analyses demonstrated the effects of information and group structure on the evolution of altruism. We inferred that information about a partner's behavior and group structure can produce flexible pathways for the evolution of altruism.
|
|
|
|
Hamilton, W. D. (1971). Geometry for the selfish herd. J. Theor. Biol., 31(2), 295–311.
Abstract: This paper presents an antithesis to the view that gregarious behaviour is evolved through benefits to the population or species. Following Galton (1871) and Williams (1964) gregarious behaviour is considered as a form of cover-seeking in which each animal tries to reduce its chance of being caught by a predator.
It is easy to see how pruning of marginal individuals can maintain centripetal instincts in already gregarious species; some evidence that marginal pruning actually occurs is summarized. Besides this, simply defined models are used to show that even in non-gregarious species selection is likely to favour individuals who stay close to others.
Although not universal or unipotent, cover-seeking is a widespread and important element in animal aggregation, as the literature shows. Neglect of the idea has probably followed from a general disbelief that evolution can be dysgenic for a species. Nevertheless, selection theory provides no support for such disbelief in the case of species with outbreeding or unsubdivided populations.
The model for two dimensions involves a complex problem in geometrical probability which has relevance also in metallurgy and communication science. Some empirical data on this, gathered from random number plots, is presented as of possible heuristic value.
|
|
|
|
Seyfarth, R. M. (1977). A model of social grooming among adult female monkeys. J. Theor. Biol., 65(4), 671–698.
Abstract: Grooming networks among adult female monkeys exhibit two similar features across a number of different species. High-ranking animals receive more grooming than others, and the majority of grooming occurs between females of adjacent rank. A theoretical model which duplicates these features is presented, and the properties of the model are used to explain the possible causation and function of female grooming behaviour. The model illustrates how relatively simple principles governing the behaviour of individuals may be used to explain more complex aspects of the social structure of non-human primate groups.
|
|
|
|
McGreevy, P. D., & McLean, A. N. (2007). Roles of learning theory and ethology in equitation. Journal of Veterinary Behavior: Clinical Applications and Research, 2(4), 108–118.
Abstract: By definition, ethology is primarily the scientific study of animal behavior, especially as it occurs in a natural environment; applied ethology being the study of animal behavior in the human domain. The terms equine ethology and ethological training are becoming commonplace in the equestrian domain, yet they seem to be used with a conspicuous lack of clarity and with no mention of learning theory. Most of what we do to train horses runs counter to their innate preferences. This article summarizes the ethological challenges encountered by working horses and considers the merits and limitations of ethological solutions. It also questions the use of terms such as “alpha” and “leader” and examines aspects of learning theory, equine cognition, and ethology as applied to horse training and clinical behavior modification. We propose 7 training principles that optimally account for the horse's ethological and learning abilities and maintain maximal responsivity in the trained horse. These principles can be summarized as: (1) use learning theory appropriately; (2) train easy-to-discriminate signals; (3) train and subsequently elicit responses singularly; (4) train only one response per signal; (5) train all responses to be initiated and subsequently completed within a consistent structure; (6) train persistence of current operantly conditioned responses; and (7) avoid and disassociate flight responses. Adherence to these principles and incorporating them into all horse training methodologies should accelerate training success, reduce behavioral wastage of horses, and improve safety for both humans and horses.
|
|
|
|
Tobin, T., & Combie, J. D. (1982). Performance testing in horses: a review of the role of simple behavioral models in the design of performance experiments. J Vet Pharmacol Ther, 5(2), 105–118.
|
|
|
|
Carroll, J., Murphy, C. J., Neitz, M., Hoeve, J. N., & Neitz, J. (2001). Photopigment basis for dichromatic color vision in the horse. J Vis, 1(2), 80–87.
Abstract: Horses, like other ungulates, are active in the day, at dusk, dawn, and night; and, they have eyes designed to have both high sensitivity for vision in dim light and good visual acuity under higher light levels (Walls, 1942). Typically, daytime activity is associated with the presence of multiple cone classes and color-vision capacity (Jacobs, 1993). Previous studies in other ungulates, such as pigs, goats, cows, sheep and deer, have shown that they have two spectrally different cone types, and hence, at least the photopigment basis for dichromatic color vision (Neitz & Jacobs, 1989; Jacobs, Deegan II, Neitz, Murphy, Miller, & Marchinton, 1994; Jacobs, Deegan II, & Neitz, 1998). Here, electroretinogram flicker photometry was used to measure the spectral sensitivities of the cones in the domestic horse (Equus caballus). Two distinct spectral mechanisms were identified and are consistent with the presence of a short-wavelength-sensitive (S) and a middle-to-long-wavelength-sensitive (M/L) cone. The spectral sensitivity of the S cone was estimated to have a peak of 428 nm, while the M/L cone had a peak of 539 nm. These two cone types would provide the basis for dichromatic color vision consistent with recent results from behavioral testing of horses (Macuda & Timney, 1999; Macuda & Timney, 2000; Timney & Macuda, 2001). The spectral peak of the M/L cone photopigment measured here, in vivo, is similar to that obtained when the gene was sequenced, cloned, and expressed in vitro (Yokoyama & Radlwimmer, 1999). Of the ungulates that have been studied to date, all have the photopigment basis for dichromatic color vision; however, they differ considerably from one another in the spectral tuning of their cone pigments. These differences may represent adaptations to the different visual requirements of different species.
|
|
|
|
Lyda, R. O., Hall, J. R., & Kirkpatrick, J. F. (2005). A comparison of Freund's Complete and Freund's Modified Adjuvants used with a contraceptive vaccine in wild horses (Equus caballus). J Zoo Wildl Med, 36(4), 610–616.
Abstract: Fifteen captive wild mares (Equus caballus) were treated with porcine zona pellucida contraceptive vaccine and either Freund's Complete Adjuvant (n = 7) or Freund's Modified Adjuvant (n = 8). All mares received a booster inoculation of porcine zona pellucida plus Freund's Incomplete Adjuvant a month later. Anti-porcine zona pellucida antibodies were measured over 10 mo following the initial inoculation. There were no significant differences in antibody titers at any point during the 10 mo, and seven of the eight mares in the Freund's Modified Adjuvant group were above the 60% level at the end of the study, which is considered to be the contraceptive threshold for horses. There were no significant differences in titers between pregnant and nonpregnant horses, nor was there a significant correlation between age and titers. One local injection site reaction occurred after booster treatment with Freund's Incomplete Adjuvant, and 11 healthy foals were born during the course of the study. These data suggest that Freund's Modified Adjuvant is an acceptable substitute for Freund's Complete Adjuvant in certain free-ranging and captive wildlife species.
|
|
|
|
Keay, J. M., Singh, J., Gaunt, M. C., & Kaur, T. (2006). Fecal glucocorticoids and their metabolites as indicators of stress in various mammalian species: a literature review. J Zoo Wildl Med, 37(3), 234–244.
Abstract: Conservation medicine is a discipline in which researchers and conservationists study and respond to the dynamic interplay between animals, humans, and the environment. From a wildlife perspective, animal species are encountering stressors from numerous sources. With the rapidly increasing human population, a corresponding increased demand for food, fuel, and shelter; habitat destruction; and increased competition for natural resources, the health and well-being of wild animal populations is increasingly at risk of disease and endangerment. Scientific data are needed to measure the impact that human encroachment is having on wildlife. Nonbiased biometric data provide a means to measure the amount of stress being imposed on animals from humans, the environment, and other animals. The stress response in animals functions via glucocorticoid metabolism and is regulated by the hypothalamic-pituitary-adrenal axis. Fecal glucocorticoids, in particular, may be an extremely useful biometric test, since sample collection is noninvasive to subjects and, therefore, does not introduce other variables that may alter assay results. For this reason, many researchers and conservationists have begun to use fecal glucocorticoids as a means to measure stress in various animal species. This review article summarizes the literature on many studies in which fecal glucocorticoids and their metabolites have been used to assess stress levels in various mammalian species. Variations between studies are the main focus of this review. Collection methods, storage conditions, shipping procedures, and laboratory techniques utilized by different researchers are discussed.
|
|
