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Spadavecchia, C., Arendt-Nielsen, L., Spadavecchia, L., Mosing, M., Auer, U., & van den Hoven, R. (2007). Effects of butorphanol on the withdrawal reflex using threshold, suprathreshold and repeated subthreshold electrical stimuli in conscious horses. Vet Anaesth Analg, 34(1), 48–58.
Abstract: OBJECTIVE: To assess the effects of a single intravenous dose of butorphanol (0.1 mg kg(-1)) on the nociceptive withdrawal reflex (NWR) using threshold, suprathreshold and repeated subthreshold electrical stimuli in conscious horses. STUDY DESIGN: 'Unblinded', prospective experimental study. ANIMALS: Ten adult horses, five geldings and five mares, mean body mass 517 kg (range 487-569 kg). METHODS: The NWR was elicited using single transcutaneous electrical stimulation of the palmar digital nerve. Repeated stimulations were applied to evoke temporal summation. Surface electromyography was performed to record and quantify the responses of the common digital extensor muscle to stimulation and behavioural reactions were scored. Before butorphanol administration and at fixed time points up to 2 hours after injection, baseline threshold intensities for NWR and temporal summation were defined and single suprathreshold stimulations applied. Friedman repeated-measures analysis of variance on ranks and Wilcoxon signed-rank test were used with the Student-Newman-Keul's method applied post-hoc. The level of significance (alpha) was set at 0.05. RESULTS: Butorphanol did not modify either the thresholds for NWR and temporal summation or the reaction scores, but the difference between suprathreshold and threshold reflex amplitudes was reduced when single stimulation was applied. Upon repeated stimulation after butorphanol administration, a significant decrease in the relative amplitude was calculated for both the 30-80 and the 80-200 millisecond intervals after each stimulus, and for the whole post-stimulation interval in the right thoracic limb. In the left thoracic limb a decrease in the relative amplitude was found only in the 30-80 millisecond epoch. CONCLUSION: Butorphanol at 0.1 mg kg(-1) has no direct action on spinal Adelta nociceptive activity but may have some supraspinal effects that reduce the gain of the nociceptive system. CLINICAL RELEVANCE: Butorphanol has minimal effect on sharp immediate Adelta-mediated pain but may alter spinal processing and decrease the delayed sensations of pain.
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Flack, J. C., Krakauer, D. C., & de Waal, F. B. M. (2005). Robustness mechanisms in primate societies: a perturbation study. Proc Biol Sci, 272(1568), 1091–1099.
Abstract: Conflict management mechanisms have a direct, critical effect on system robustness because they mitigate conflict intensity and help repair damaged relationships. However, robustness mechanisms can also have indirect effects on system integrity by facilitating interactions among components. We explore the indirect role that conflict management mechanisms play in the maintenance of social system robustness, using a perturbation technique to 'knockout' components responsible for effective conflict management. We explore the effects of knockout on pigtailed macaque (Macaca nemestrina) social organization, using a captive group of 84 individuals. This system is ideal in addressing this question because there is heterogeneity in performance of conflict management. Consequently, conflict managers can be easily removed without disrupting other control structures. We find that powerful conflict managers are essential in maintaining social order for the benefit of all members of society. We show that knockout of components responsible for conflict management results in system destabilization by significantly increasing mean levels of conflict and aggression, decreasing socio-positive interaction and decreasing the operation of repair mechanisms.
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Flack, J. C., de Waal, F. B. M., & Krakauer, D. C. (2005). Social structure, robustness, and policing cost in a cognitively sophisticated species. Am Nat, 165(5), E126–139.
Abstract: Conflict management is one of the primary requirements for social complexity. Of the many forms of conflict management, one of the rarest and most interesting is third-party policing, or intervening impartially to control conflict. Third-party policing should be hard to evolve because policers personally pay a cost for intervening, while the benefits are diffused over the whole group. In this study we investigate the incidence and costs of policing in a primate society. We report quantitative evidence of non-kin policing in the nonhuman primate, the pigtailed macaque. We find that policing is effective at reducing the intensity of or terminating conflict when performed by the most powerful individuals. We define a measure, social power consensus, that predicts effective low-cost interventions by powerful individuals and ineffective, relatively costly interventions by low-power individuals. Finally, we develop a simple probabilistic model to explore whether the degree to which policing can effectively reduce the societal cost of conflict is dependent on variance in the distribution of power. Our data and simple model suggest that third-party policing effectiveness and cost are dependent on power structure and might emerge only in societies with high variance in power.
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Flack, J. C., Girvan, M., de Waal, F. B. M., & Krakauer, D. C. (2006). Policing stabilizes construction of social niches in primates. Nature, 439(7075), 426–429.
Abstract: All organisms interact with their environment, and in doing so shape it, modifying resource availability. Termed niche construction, this process has been studied primarily at the ecological level with an emphasis on the consequences of construction across generations. We focus on the behavioural process of construction within a single generation, identifying the role a robustness mechanism--conflict management--has in promoting interactions that build social resource networks or social niches. Using 'knockout' experiments on a large, captive group of pigtailed macaques (Macaca nemestrina), we show that a policing function, performed infrequently by a small subset of individuals, significantly contributes to maintaining stable resource networks in the face of chronic perturbations that arise through conflict. When policing is absent, social niches destabilize, with group members building smaller, less diverse, and less integrated grooming, play, proximity and contact-sitting networks. Instability is quantified in terms of reduced mean degree, increased clustering, reduced reach, and increased assortativity. Policing not only controls conflict, we find it significantly influences the structure of networks that constitute essential social resources in gregarious primate societies. The structure of such networks plays a critical role in infant survivorship, emergence and spread of cooperative behaviour, social learning and cultural traditions.
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Rietmann, T. R., Stuart, A. E. A., Bernasconi, P., Stauffacher, M., Auer, J. A., & Weishaupt, M. A. (2004). Assessment of mental stress in warmblood horses: heart rate variability in comparison to heart rate and selected behavioural parameters. Appl. Anim. Behav. Sci., 88(1-2), 121–136.
Abstract: The aim of the study was to investigate whether heart rate variability (HRV) could assess alterations of the autonomic nervous system (ANS) at different levels of excitement. The behavioural and physiological responses of 20 warmblood horses to a challenging ground exercise task were studied. Prior to the experiment, the horses were evaluated at rest and during forward walking (FW). The horses were then forced to move backwards continuously during 3 min according to a standardised protocol (BW1). Subsequently, the horses were exposed to two training sessions, after which the backward walking (BW2) was re-evaluated. Heart rate (HR) and HRV-parameters such as the standard deviation of the beat-to-beat intervals (SDRR), the low (LF; sympathetic tone) and high frequency (HF) component of HRV (HF; parasympathetic tone) and their ratio (LF/HF; index representing the sympatho-vagal balance) were sampled at rest, and during FW, BW1 and BW2. Stress-related behaviour during BW1 and BW2 was determined from video recordings. The results of the different evaluations were compared to each other. Compared to rest and FW, the first backward experiment induced a significant rise in HR, LF and LF/HF and a significant decrease of HF. SDRR decreased from both FW and rest with only the latter reaching significance. In BW2 after the training sessions, HR and the parameters of the sympathetic branch of the ANS (LF, LF/HF) were decreased and the vagal tone (HF) increased compared to BW1; all changes were significant. The duration of stress indicating behavioural patterns revealed also a significant decrease of excitement after the training, when backward walking did not differ from forward walking in any parameter. Correlations between HRV-parameters and stress indicating behaviour as well as HR were found. We conclude that the HRV-parameters LF and HF are valuable measures to quantify sympatho-vagal balance, which allows a more precise assessment of the responses of HR and SDRR to mental stress during low intensity exercise.
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Brauer, J., Kaminski, J., Riedel, J., Call, J., & Tomasello, M. (2006). Making inferences about the location of hidden food: social dog, causal ape. J Comp Psychol, 120(1), 38–47.
Abstract: Domestic dogs (Canis familiaris) and great apes from the genus Pan were tested on a series of object choice tasks. In each task, the location of hidden food was indicated for subjects by some kind of communicative, behavioral, or physical cue. On the basis of differences in the ecologies of these 2 genera, as well as on previous research, the authors hypothesized that dogs should be especially skillful in using human communicative cues such as the pointing gesture, whereas apes should be especially skillful in using physical, causal cues such as food in a cup making noise when it is shaken. The overall pattern of performance by the 2 genera strongly supported this social-dog, causal-ape hypothesis. This result is discussed in terms of apes' adaptations for complex, extractive foraging and dogs' adaptations, during the domestication process, for cooperative communication with humans.
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Call, J., Brauer, J., Kaminski, J., & Tomasello, M. (2003). Domestic dogs (Canis familiaris) are sensitive to the attentional state of humans. J Comp Psychol, 117(3), 257–263.
Abstract: Twelve domestic dogs (Canis familiaris) were given a series of trials in which they were forbidden to take a piece of visible food. In some trials, the human continued to look at the dog throughout the trial (control condition), whereas in others, the human (a) left the room, (b) turned her back, (c) engaged in a distracting activity, or (d) closed her eyes. Dogs behaved in clearly different ways in most of the conditions in which the human did not watch them compared with the control condition, in which she did. In particular, when the human looked at them, dogs retrieved less food, approached it in a more indirect way, and sat (as opposed to laid down) more often than in the other conditions. Results are discussed in terms of domestic dogs' social-cognitive skills and their unique evolutionary and ontogenetic histories.
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Rietmann, T. R., Stauffacher, M., Bernasconi, P., Auer, J. A., & Weishaupt, M. A. (2004). The association between heart rate, heart rate variability, endocrine and behavioural pain measures in horses suffering from laminitis. J Vet Med A Physiol Pathol Clin Med, 51(5), 218–225.
Abstract: The objective of this study was to compare the stress response of horses suffering from laminitis after short- and long-term treatment with the intent to evaluate power spectral analysis of heart rate variability (HRV) for pain monitoring. Data were collected from 19 horses with acute or chronic exacerbating laminitis without known primary disease before and after treatment with non-steroidal anti-inflammatory drugs (NSAID). Recordings were carried out the day after admission to the equine hospital. Measurements were repeated on day 7 of the treatment. The recorded parameters included a clinical orthopaedic index (OLPI: Obel-grade plus hoof tester score), frequency of weight-shifting between contralateral limbs, mean beat-to-beat interval (R-R) duration, standard deviation of continuous R-R intervals, low- (LF) and high-frequency (HF) components of HRV, sympatho-vagal balance (LF/HF), and plasma concentration of cortisol, adrenalin and noradrenalin. The LF represents mainly sympathetic influences on the heart whereas HF is mediated by the parasympathetic tone. Weight-shifting and OLPI decreased significantly with treatment. The LF normalized units (n.u.) decreased after NSAID from 60.41 +/- 21.42 to 51.12 +/- 19.81 and was 49.33 +/- 22.64 on day 7, whereas HF n.u. increased from 35.07 +/- 20.02 to 43.14 +/- 18.30 and was 45.98 +/- 23.00 on day 7. Hormone levels showed no tendency to change with treatment. The OLPI was only correlated with LF/HF, LF and HF (R = 0.57, 0.55 and -0.54 respectively). Significant negative correlations existed between HFn.u. and weight-shifting frequency (R = -0.37), HFn.u. and adrenalin (R = -0.47), and HFn.u. and noradrenalin (R = 0.33). The LFn.u. only correlated positively with adrenalin. Cortisol levels were poorly associated with the other parameters. Determination of the sympatho-vagal influences on cardiac function may offer complementary information for reliable assessment of pain and may represent a valuable alternative method to catecholamine measurements.
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Puppe, B., Langbein, J., Bauer, J., & Hoy, S. (2008). A comparative view on social hierarchy formation at different stages of pig production using sociometric measures. Livestock Science, 113(2-3), 155–162.
Abstract: A standardised and comprehensive approach to describe dominance relationships in gregarious farm animals quantitatively was recently developed, incorporating a combination of appropriate sociometric measures. The present study applied this approach to a comparative analysis of the social hierarchies within 57 groups of domestic pigs at different age/production stages with a total of 496 animals. Unacquainted pigs were grouped to three age categories which correspond to the typical production stages: weaned pigs (PIG28, 12 groups), growing pigs (PIG80, 16 groups), and reproductive sows (SOW, 29 groups). Based on observed agonistic interactions, sociometric values were calculated both at the dyadic and at the group level and may be considered as preliminary reference values for further studies. As indicated by the respective values of the Kendall index (PIG28: 0.66, tested as significant in 69.0% of the observed groups; PIG80: 0.71, 87.5%; SOW: 0.61, 69.0%), and the improved Landau index (PIG28: 0.70, 75.0%; PIG80: 0.72, 93.7%; SOW: 0.71, 72.4%), a social organisation towards a quasi-linear social hierarchy was predominantly developed throughout all age/production categories. However, compared to weaned and growing pigs, sows were characterised by significant differences concerning establishment (fewer agonistic interactions) and kind (more unknown dyads, fewer two-way and significant dyads, higher directional consistency index) of their social hierarchy. It seems that sows have effectively adapted their agonistic behaviour towards pen-mates to regulate social dominance relationships, whereas younger pigs frequently display agonistic interactions also to gain additional experience on social cues (e.g. the fighting ability of an opponent). Hence, it is concluded that the effective experience of socialisation during sensitive periods may increase the social skills of pigs which in turn can improve their welfare and health, e.g. by adjusted aggressive behaviour. The consideration of comparable and standardised sociometric measures in livestock breeding may help to improve husbandry conditions.
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Bauer, I. E., McMorrow, J. Y.,, & D. W. (1994). The Historic Ranges of Three Equid Species in North-East Africa: A Quantitative Comparison of Environmental Tolerances. J Biogeogr, 21(2), 169–182.
Abstract: The historic ranges of three equid species native to north-east Africa are analysed with respect to annual rainfall, several temperature parameters and a satellite-derived multispectral index of primary productivity. Equus africanus Fitzinger, Equus grevyi Oustalet and Equus burchelli Gray used to largely replace each other, geographically, with narrow zones of range overlap occurring between E. africanus and E. grevyi in the Awash valley, and between E. grevyi and E. burchelli in southern Ethiopia and northern Kenya. The three species are shown to succeed each other along an environmental gradient. The position of each species on this gradient and the resulting location and extent of its range are discussed. Competitive exclusion, specific adaptations and historic events are likely determinants of equid distribution. In the area of sympatry between E. grevyi and E. burchelli, mixed habitat characters as well as environmental fluctuations seem to prevent either species from excluding the other. Different social organizations of E. grevyi and E. burchelli and the resulting migratory patterns may be adaptations to the environment in their allopatric ranges; in their sympatric range they could alleviate competition.
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