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Dalla Costa, E., Rabolini, A., Scelsa, A., Canali, E., & Minero, M. (2012). A study on inter-observer reliability of castration pain assessment in horses. In K. Krueger, & (Ed.), Proceedings of the 2. International Equine Science Meeting (Vol. in press). Wald: Xenophon Publishing.
Abstract: Pain evaluation is a key issue for equine welfare and it is often cause of concern because it is difficult to determine its intensity and duration. This is essential when different people are looking after the animals and they need to decide when or not giving analgesics to guarantee the welfare of the subject. The most widely used technique to determine pain in horses is identifying pain related behaviors. The aim of this study was to determine inter-observer reliability of two different assessors evaluating pain related behaviors in horses undergoing castration. 8 stallions of different breed, aged between 2 and 4 years, were included in the study. All the subjects underwent routine castration (closed technique in general anesthesia). The subjects were placed in an observation box for 5 days and their behavior was recorded for 15 minutes before the surgery and 4, 8, 16, 24 and 40 hours after intervention. Two blind observers, using a given ethogram of horse pain related behaviors modified from literature (for a review Ashley, 2005), analyzed horses behavior at each interval. Descriptive statistics and K Kendall test were performed. Observers agreed significantly assessing agitation, reluctance to move, kicking the abdomen, lethargy, rolling, attention and curiosity (P<0.05), however agreement was low for head movements, stretching, flank watching, lowered head carriage, weight shifting, abnormal movement, fixed stare. Our results show that assessing pain in horses should be a cause of concern, because different pain related behaviors are difficult to identify and to have agreement between two observers. Training of care takers of horses on identification of specific behaviors is needed to standardize pain assessment. Acknowledgements: The authors wish to thank the EU VII Framework programme (FP7-KBBE-2010-4) for financing the Animal Welfare Indicators (AWIN) project and for providing funds for Emanuela Dalla Costa and Michela Minero to present this paper.
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Watanabe, N. M., Stahlman, W. D., Blaisdell, A. P., Garlick, D., Fast, C. D., & Blumstein, D. T. (2012). Quantifying personality in the terrestrial hermit crab: Different measures, different inferences. Behav. Process., 91(2), 133–140.
Abstract: There is much interest in studying animal personalities but considerable debate as to how to define and evaluate them. We assessed the utility of one proposed framework while studying personality in terrestrial hermit crabs (Coenobita clypeatus). We recorded the latency of individuals to emerge from their shells over multiple trials in four unique manipulations. We used the specific testing situations within these manipulations to define two temperament categories (shyness-boldness and exploration-avoidance). Our results identified individual behavioral consistency (i.e., personality) across repeated trials of the same situations, within both categories. Additionally, we found correlations between behaviors across contexts (traits) that suggested that the crabs had behavioral syndromes. While we found some correlations between behaviors that are supposed to measure the same temperament trait, these correlations were not inevitable. Furthermore, a principal component analysis (PCA) of our data revealed new relationships between behaviors and provided the foundation for an alternate interpretation: measured behaviors may be situation-specific, and may not reflect general personality traits at all. These results suggest that more attention must be placed on how we infer personalities from standardized methods, and that we must be careful to not force our data to fit our frameworks.
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Price, E. O. (1999). Behavioral development in animals undergoing domestication. App Anim Behav Sci, 65(3), 245–271.
Abstract: The process of domestication involves adaptation, usually to a captive environment. Domestication is attained by some combination of genetic changes occurring over generations and developmental mechanisms (e.g., physical maturation, learning) triggered by recurring environmental events or management practices in captivity that influence specific biological traits. The transition from free-living to captive status is often accompanied by changes in availability and/or accessibility of shelter, space, food and water, and by changes in predation and the social environment. These changes set the stage for the development of the domestic phenotype. Behavioral development in animals undergoing domestication is characterized by changes in the quantitative rather than qualitative nature of responses. The hypothesized loss of certain behavior patterns under domestication can usually be explained by the heightening of response thresholds. Increases in response frequency accompanying domestication can often be explained by atypical rates of exposure to certain forms of perceptual and locomotor stimulation. Genetic changes influencing the development of the domestic phenotype result from inbreeding, genetic drift, artificial selection, natural selection in captivity, and relaxed selection. Experiential contributions to the domestic phenotype include the presence or absence of key stimuli, changes in intraspecific aggressive interactions and interactions with humans. Man's role as a buffer between the animal and its environment is also believed to have an important effect on the development of the domestic phenotype. The domestication process has frequently reduced the sensitivity of animals to changes in their environment, perhaps the single-most important change accompanying domestication. It has also resulted in modified rates of behavioral and physical development. Interest in breeding animals in captivity for release in nature has flourished in recent decades. The capacity of domestic animals to survive and reproduce in nature may depend on the extent to which the gene pool of the population has been altered during the domestication process and flexibility in behavioral development. “Natural” gene pools should be protected when breeding wild animals in captivity for the purpose of reestablishing free-living natural populations. In some cases, captive-reared animals must be conditioned to live in nature prior to their release.
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Gil, M., Bhatt, R., Picotte, K. B., & Hull, E. M. (2013). Sexual experience increases oxytocin receptor gene expression and protein in the medial preoptic area of the male rat. In Psychoneuroendocrinology (Vol. 38, pp. 1688–1697). Pergamon Press.
Abstract: Oxytocin (OT) promotes social and reproductive behaviors in mammals, and OT deficits may be linked to disordered social behaviors like autism and severe anxiety. Male rat sexual behavior is an excellent model for OT regulation of behavior, as its pattern and neural substrates are well characterized. We previously reported that OT microinjected into the medial preoptic area (MPOA), a major integrative site for male sexual behavior, facilitates copulation in sexually experienced male rats, whereas intra-MPOA injection of an OT antagonist (OTA) inhibits copulation. In the present studies, copulation on the day of sacrifice stimulated OTR mRNA expression in the MPOA, irrespective of previous sexual experience, with the highest levels observed in first-time copulators. In addition, sexually experienced males had higher levels of OTR protein in the MPOA than sexually naïve males and first-time copulators. Finally, intra-MPOA injection of OT facilitated mating in sexually naive males. Others have reported a positive correlation between OT mRNA levels and male sexual behavior. Our studies show that OT in the MPOA facilitates mating in both sexually naive and experienced males, some of the behavioral effects of OT are mediated by the OTR, and sexual experience is associated with increased OTR expression in the MPOA. Taken together, these data suggest a reciprocal interaction between central OT and behavior, in which OT facilitates copulation and copulation stimulates the OT/OTR system in the brain.
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Hoffmann, G., Bentke, A., Rose-Meierhöfer, S., Berg, W., Mazetti, P., & Hardarson, G. H. (2012). Influence of an active stable system on the behavior and body condition of Icelandic horses. animal, 6(10), 1684–1693.
Abstract: Horses are often stabled in individual boxes, a method that does not meet their natural needs and may cause psychical and
musculoskeletal diseases. This problem is particularly evident in Iceland, where horses often spend the long winter periods in cramped
boxes. The aim of this study was to analyze the suitability of a group housing system in Iceland, but the results are also applicable to
horses of other regions. Eight Icelandic horses were observed in an active stable system, and their behavior and time budget were
recorded. Movement and lying behavior were studied with ALT (Activity, Lying, Temperature detection) pedometers. The effect of an
automatic concentrate feeding station (CFS) on the horses’ behavior was examined. In the first period of investigation, the horses
were fed concentrates manually, and in the second period, they were fed with the CFS. Additional behavioral observations and a
determination of social hierarchy occurred directly or by video surveillance. The physical condition of the horses was recorded by body
weight (BW) measurement and body condition scoring (BCS). The results showed a significant increase between the first and second
trial periods in both the activity (P,0.001) and the lying time (P50.003) of the horses with use of the CFS. However, there was no
significant change in BW during the first period without the CFS (P50.884) or during the second period with the CFS (P50.540).
The BCS of the horses was constant at a very good level during both trial periods, and the horses showed a low level of aggression, a
firm social hierarchy and behavioral synchronization. This study concludes that group housing according to the active stable principle is
a welfare-friendly option for keeping horses and is a suitable alternative to conventional individual boxes.
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Obergfell, J. (2012). Einf lus s v on St ruktur e l ement en auf da s Li eg ev e rha l t en v on Pf e rden in Gruppenha l tung unt e r Be rücks i cht igung de s Ag g r e s s i ons v e rha l t ens. Ph.D. thesis, , Karlsruhe.
Abstract: Durch die vorliegende Studie wurde der Einfluss von Strukturelementen auf das Liege- und Aggressionsverhalten von Pferden in Gruppenhaltung untersucht. Die Strukturelemente sollten Rückzugsmöglichkeiten bieten, sowie in ihrem Bereich zur Aufhebung der Individualdistanz führen und dadurch die Fläche relativ vergrößern. Für die Versuche stellte das Haupt- und Landesgestüt Marbach drei unabhängige Pferdegruppen mit verschiedener Herdengröße zur Verfügung, die jeweils in Einraum- Innenlaufställen gehalten wurden. Die Datenerfassung fand zwischen 23 und sieben Uhr statt. Insgesamt wurden 366 Stunden Videomaterial ausgewertet. Beim Ruheverhalten wurden mit Hilfe des event-sampling- Verfahrens die Parameter Gesamtliegedauer, Dauer in Seitenlage, Dauer der Einzelphasen in
Seitenlage und Abliegehäufigkeit pro Nacht erfasst. Es wurden Versuchsphasen ohne
Strukturelemente und mit hängenden Planen als Strukturierung durchgeführt. Im ersten Stall
kamen außerdem über einander gestapelte Strohballen zum Einsatz. Diese Art der
Strukturierung stellte sich jedoch als nicht praktikabel heraus und führte im Vergleich zu den Planen zu einer signifikanten Verschlechterung der Gesamtliegedauer. In Stall 1 konnte man eine tendenzielle Verbesserung der Parameter Gesamtliegedauer und Gesamtdauer in Seitenlage durch das Anbringen von Planen sehen. In Stall 3 dagegen verschlechterte sich das Ruheverhalten in den Versuchsphasen mit Planen gegenüber den Versuchsphasen ohne Struktur. Die Werte der Gesamtdauer in Seitenlage nahmen signifikant ab. In Stall 2 zeigten sich keine Unterschiede in den verschiedenen Versuchsphasen. Möglicherweise ist die Wirkung der Strukturelemente auf das Liegeverhalten abhängig von der Flächengröße. Stall 1 hatte bezogen auf die Leitlinien des BMELV die größte und Stall 3 die kleinste Fläche. Bei den anderen Parametern des Liegeverhaltens gab es keine signifikanten Unterschiede in den verschiedenen Versuchsphasen. Mit Hilfe des time-sampling-Verfahrens wurde die Anzahl gleichzeitig liegender Pferde und gleichzeitig liegender Pferde in Seitenlage bestimmt. Auch hier zeigten sich keine Unterschiede in den verschiedenen Versuchsphasen. Bei der Gegenüberstellung der Werte der Gesamtliegedauer und der Gesamtdauer in Seitenlage mit dem Alter der Pferde (Stall 1 und Stall 3) und mit dem Integrationszeitpunkt (Stall 1) konnte kein Zusammenhang festgestellt werden. Beim Aggressionsverhalten wurden mit Hilfe des
event-sampling-Verfahrens in den Ställen 2 und 3 verschiedene Arten von Aggressionen
erfasst, die dann in die drei Intensitätsgrade Low-Level-, Mid-Level- und High-Level-
Aggressionen unterteilt wurden. Neben der Anzahl wurde die Dauer der verschiedenen
Aggressionen bewertet. Insgesamt konnte eine positive Wirkung der Planen auf das
Aggressionsverhalten beobachtet werden. Die Gesamtanzahl an Aggressionen nahm in beiden
Ställen tendenziell in den Versuchsphasen mit Planen ab. In Stall 3 konnte, wenn man die
Aggressionen stundenweise betrachtet, ein signifikanter Unterschied festgestellt werden.
Auch der Hinterhandschlag und die Aggressionen, welche das Ruheverhalten stören,
verringerten sich tendenziell nach dem Anbringen von Strukturelementen. In beiden Ställen
nahm die relative Häufigkeit von Mid-Level-Aggressionen nach dem Anbringen von Planen
zu. Dagegen konnte bei den High-Level-Aggressionen und in Stall 3 bei den Low-Level-
Aggressionen eine relative Abnahme beobachtet werden. Sowohl die Anzahl als auch die
Dauer der Mid-Level-Aggressionen verringerten sich in Stall 3 stundenweise betrachtet
signifikant in den Versuchsphasen mit Planen. In Stall 2 war bei den High-Level-
Aggressionen sowohl bei der Dauer als auch bei der Anzahl eine signifikante Abnahme zu
sehen. Wenn man das Aggressionsverhalten in Bezug zu der Fläche in den zwei Ställen
betrachtet, schien diese vor allem einen Einfluss auf die High-Level-Aggressionen zu
nehmen. Mit Hilfe des Rangindex der Pferde im Stall 3 wurde eine Rangordnung aufgestellt.
Zwischen dem Platz der Pferde in der Rangordnung und den Parametern des Ruheverhaltens
(Gesamtliegedauer, Gesamtdauer in Seitenlage) sowie dem Alter der Pferde konnte kein
Zusammenhang festgestellt werden. Zwischen dem Rangindex und der Gesamtanzahl an
Aggressionen bestand dagegen ein hoch signifikanter Zusammenhang. Im Rahmen dieser
Studie ist das Anbringen von Strukturelementen in Bezug auf das Aggressionsverhalten in
Einraum-Innenlaufställen von Pferden zu empfehlen. Die Wirkung auf das Liegeverhalten der
Pferde sollte in weiteren Studien untersucht werden.
[In this study the influence of structural elements on lying and aggression behavior in horses
kept in groups was exanimated. Structural elements should offer the possibility of retreat.
Furthermore, within the respective area, the individual distance should be limited, thus
increasing the overall space relatively. For the studies the “Haupt- und Landesgestüt
Marbach” provided three non-related groups of different sized horses, each kept in separate
“Einraum-Innenlaufställen”. Data collection took place between 11 pm and 7 am. Altogether,
366 hours of video material were analyzed. Using the event-sampling-method, the following
parameters within the horses´ resting behavior were determined: The total lying period, the
total lying period in lateral position, the periods of the single lying in lateral position and the
frequency of lying down. Test phases were carried out without structural elements and with
hanging canvases as structural elements. In the first stable there were also three bales of straw
piled up on top of each other. This kind of structuring emerged as impractical and in
comparison to the canvas, led to a significant deterioration to the total period of lying. After
the canvases were fixed in the first stable, a slight improvement with the parameters of total
lying period and total lying period in the lateral position could be seen. In contrast, the resting
behavior in stable 3 worsened in the test phases with the canvases in comparison to the test phases without structural elements. The values for the total lying period in the lateral position
decreased significantly. In stable 2, there were no differences in the different test phases.
Possibly the effect of the structural elements is dependent of the space. Stable 1 had the
largest space and stable 3 the least space referred to the guideline of the BMELV. The other
parameters of the lying behavior showed no significant differences in the different test phases.
The number of horses lying at the same time, and the number of horses lying at the same time
in the lateral position were detected with the aid of the time-sampling-method. There were
also no differences between the different test phases. If the values of the total lying period and
the total lying period in lateral position were compared to the age of the horses (stable 1 and
stable 3) and to the point of integration (stable 1), there was no correlation.
Different kinds of aggression were detected by the means of the event-sampling-method in
stable 2 and 3 and were subdivided into the three levels of intensity: Low-level-, mid-level
and high-level-aggressions. Both the number and the duration of the different aggressions
were evaluated. All in all a positive influence of the canvas on the aggression behavior could
be observed. In both stables, the total number of aggressions decreased in the test phases with
canvases. In stable 3 a significant difference could be detected, if the aggressions were
observed per hour. Also hind limb kicks and aggressions disturbing the resting behavior,
decreased by trend, after fixing the canvases in the stable. In both stables the relative
frequency of mid-level-aggressions increased after the canvases were mounted. In contrast
there was a relative decrease at the high-level-aggressions and in stable 3 at the low-levelaggressions.
The number and the duration of the mid-level-aggressions in stable 3 decreased,
when considered by the hour in the test phases with canvases. In stable 2 a significant
decrease in duration and number of high-level-aggressions was observed. When examining
aggression behavior in reference to space in the two stables, high-level-aggressions seemed to
be mostly influenced. In stable 3 a rank order was established with the aid of the rank index of
the horses. There was no correlation between the horses´ rank and the parameters of the
resting behavior (the total lying period, the total lying period in lateral position) and between
horses´ rank and age. In contrast, there was a high significant correlation between the rank
index and the total number of aggressions. In context to this study, structural elements in
“Einraum-Innenlaufställen” (BMELV 2009) seem to have a positive influence on the
aggression behavior and thus seem to be recommendable. The influence on the lying behavior
should be investigated in further studies.]
Keywords: Pferde, Ruheverhalten, Liegeverhalten, Aggressionsverhalten, Strukturelemente, Rangordnung [horses, rest behavior, lying behavior, aggression behavior, structural elements, rank order]
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Shen, Y. - Q., Hebert, G., Lin, L. - Y., Luo, Y. - L., Moze, E., Li, K. - S., et al. (2005). Interleukine-1β and interleukine-6 levels in striatum and other brain structures after MPTP treatment: influence of behavioral lateralization. Journal of Neuroimmunology, 158(1–2), 14–25.
Abstract: MPTP (N-methyl-4-phenyl-1,2,3,6-tetrahydropyridine) induces diminution of the dopamine in nigrostriatal pathway and cognitive deficits in mice. MPTP treatment also increases pro-inflammatory cytokine production in substantia nigra and striatum. Since, pro-inflammatory cytokines influence striatal dopamine content and provoke cognitive impairments, the cognitive defects induced by MPTP may be partly due to brain cytokine induction in other structures than nigrostriatal pathway. Furthermore, behavioral lateralization, as assessed by paw preference, influences cytokine production at the periphery and in the central nervous system. Behavioral lateralization may thus influence brain cytokine levels after MPTP. In order to address these issues, mice selected for paw preference were injected with 25 mg/kg MPTP i.p. for five consecutive days after which striatal dopamine and DOPAC contents were measured by HPLC and IL-1β and IL-6 quantified by ELISA in the striatum, cerebral cortex, hippocampus and hypothalamus. The results showed that MPTP treatment induced dramatic loss of DA in striatum, simultaneously, IL-6 levels decreased in the striatum and increased in hippocampus and hypothalamus, while IL-1β levels decreased in the striatum, cerebral cortex and hippocampus. Interestingly, striatal dopamine turnover under basal conditions as well as striatal IL-1β and IL-6 levels under basal conditions and after MPTP depended on behavioral lateralization. Left pawed mice showed a higher decrease in dopamine turnover and lower cytokine levels as compared to right pawed animals. Behavioral lateralization also influenced IL-6 hippocampal levels under basal conditions and IL-1β cortical levels after MPTP. From these results, it can be concluded that MPTP-induced cognitive defects are accompanied by an alteration of pro-inflammatory cytokine levels in brain structures other than those involved in the nigrostriatal pathway. In addition, MPTP-induced dopamine decrease is influenced by behavioral lateralization, possibly through an effect on brain cytokine levels.
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Siniscalchi, M., Sasso, R., Pepe, A. M., Dimatteo, S., Vallortigara, G., & Quaranta, A. (2010). Catecholamine plasma levels following immune stimulation with rabies vaccine in dogs selected for their paw preferences. Neuroscience Letters, 476(3), 142–145.
Abstract: Epinephrine and norepinephrine plasma levels were assessed in dogs in relation to paw preference following an immune challenge with rabies vaccine. The results showed that both catecholamines increased after the vaccine administration, confirming the main role of the sympathetic nervous system in the modulation activity between the brain and the immune system. Moreover, ambidextrous dogs showed a significantly higher increase of epinephrine levels 8 days after immunization with respect to right- and left-pawed dogs, suggesting that the biological activity of this molecule could be key for a different immune response with regard to laterality.
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Kienapfel, K. (2011). Und was meinen die Pferde dazu? – Über das Ausdrucksverhalten von Pferden bei verschiedenen Halsstellungen [ And what are the opinions of the horses? – On the expressive behaviour of horses in different neck positions]. Pferdeheilkunde, 27(4), 357–372-380.
Abstract: The purpose of this study is to find convincing arguments, how horses should be ridden correctly, especially concerning the best head and neck position. Currently in the riding world, the opinions differ: The traditional way is to ride a horse with its nose in front of the vertical, while a new method claims that pulling the nose behind the vertical with an extreme flexion of the poll implies “gymnastic movements”. This article is focussed on the welfare and comfort of the horse in different head-neck positions, expressed by behaviours which document discomfort. As a first step, the behaviour of standing horses was observed while their heads and necks were put into seven postulated or frequently assumed positions. In the hyperflexed position the horses displayed most discomfort (with 49,7% of all reactions, like open mouths, head tossing, tail movements and other signs of resentment). The second strongest reactions of discomfort were shown in the highly raised position of the head (34,9%). The moderately rounded neck position as in traditional dressage lessons was not without signs of discomfort either, but less than the other two positions (11,2%). The high position and the extended position did not cause discomfort at all (0,17% and 0,23% respectively). In a second step, horses moving in all gaits under a rider were observed at competitions without the riders´ knowledge. The total number of observations was divided into two categories (each containing 30 horses) on the basis of the common rules of the FN: ridden with the horse’s nose in front of the vertical and ridden with the horse’s nose behind the vertical. The proportion of horses showing discomfort with the nose behind the vertical was clearly much higher (89,3%) compared to those ridden with the nose in front of the vertical (10,7%). Thus, the horses in the first category displayed signs of discomfort 8 times more frequent than those in the second group. In addition, scans carried out at tournaments showed that 92.8% of the horses were ridden with the nose behind the vertical. A clear result of this study is that the reality of riding differs widely from the rules. These results also show that riding in the hyperflexed position causes a great deal of discomfort for the horses and therefore should be abolished.
Die aktuellen Diskussionen in der Reiterwelt, welche Art und Weise des Reitens, besonders welche Kopf-Hals-Stellung zu erstreben ist, werfen die Frage auf, ob und wie die Pferde selbst ihr Befinden zum Ausdruck bringen. Über die Empfindungen der Pferde in verschiedenen Kopfhaltungen ist bisher sehr wenig bekannt. Deswegen wurde zunächst an stehenden Pferden das Ausdrucksverhalten beobachtet. Missfallensäußerungen häuften sich (mit 49,7% aller gezeigten Verhaltensauffälligkeiten wie Sperren, Rückwärtsgehen und Kopfschlagen) in der aufgerollten, hyperflektierten Stellung des Halses. An zweiter Stelle folgten Unmutsäußerungen in der absolut aufgerichteten Haltung (34,9%). Auch die beigezäumte Haltung wurde nicht ohne Unmutsbekundungen hingenommen, hier war deren Anzahl aber wesentlich geringer (11,2%). Die hohe Kopfstellung (0,17%) und die Dehnungshaltung (0,23%) bereiteten den Tieren kaum Unbehagen. Auch das Ausdrucksverhalten der Pferde unter dem Reiter wurde untersucht. Hierfür wurden, unter Berücksichtigung der schriftlich fixierten Regeln für das Turnierwesen der FN, je 30 Pferde in zwei Kategorien beobachtet: mit der Stirnlinie vor der Senkrechten und Pferde mit der Stirnlinie hinter der Senkrechten. Die Beobachtungen wurden unbemerkt von den Reitern auf den Abreiteplätzen von Turnieren durchgeführt. Die Anzahl an Verhaltensauffälligkeiten der Pferde mit der Stirnlinie hinter der Senkrechten war deutlich (89,3 %) erhöht im Gegensatz zu der anderen Gruppe (10,7 %). Die Pferde mit der Stirnlinie hinter der Senkrechten zeigten 8 Mal mehr Unmutsäußerungen als die mit der Stirnlinie vor der Senkrechten. Entgegen den Regeln der FN zeigten die durchgeführten Scans, dass unmittelbar vor den Prüfungen auf Turnieren 92,8% der Pferde mit der Stirnnasen-Linie hinter der Senkrechten geritten wurden. Ein Befund dieser Studie ist die Feststellung, dass die Praxis deutlich von den Regeln abweicht. Das Reiten mit der Stirnlinie hinter der Senkrechten ist nach diesen Befunden abzulehnen, da die Pferde deutliches Unwohlsein in dieser Haltung signalisieren.
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Schino, G., di Sorrentino, E. P., & Tiddi, B. (2007). Grooming and coalitions in Japanese macaques (<em>Macaca fuscata</em>): Partner choice and the time frame reciprocation. Journal of Comparative Psychology, 121(2), 181–188.
Abstract: Evidence of a reciprocal exchange of grooming and agonistic support in primates is mixed. In this study, the authors analyzed a large database of grooming and coalitions in captive female Japanese macaques (Macaca fuscata) to investigate their within-group distribution and temporal relations. Macaques groomed preferentially those individuals that groomed them most and supported preferentially those individuals that supported them most. They also supported preferentially those individuals that groomed them most and groomed preferentially those individuals that supported them most. These results were not explained by covariation of grooming and support with third variables such as kinship, rank, or time spent in proximity. However, receiving grooming did not increase the short-term probability of supporting a partner, and being supported did not increase the short-term probability of grooming a partner. The proximate mechanisms underlying the exchange of services were discussed in relation to the time frame of the behavioral choices made by the monkeys. (PsycINFO Database Record (c) 2012 APA, all rights reserved)
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