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Manns, J. R., Clark, R. E., & Squire, L. R. (2002). Standard delay eyeblink classical conditioning is independent of awareness. J Exp Psychol Anim Behav Process, 28(1), 32–37.
Abstract: P. F. Lovibond and D. R. Shanks (2002) suggested that all forms of classical conditioning depend on awareness of the stimulus contingencies. This article considers the available data for eyeblink classical conditioning, including data from 2 studies (R. E. Clark, J. R. Manns, & L. R. Squire, 2001; J. R. Manns, R. E. Clark, & L. R. Squire, 2001) that were completed too recently to have been considered in their review. In addition, in response to questions raised by P. F. Lovibond and D. R. Shanks, 2 new analyses of data are presented from studies published previously. The available data from humans and experimental animals provide strong evidence that delay eyeblink classical conditioning (but not trace eyeblink classical conditioning) can be acquired and retained independently of the forebrain and independently of awareness. This conclusion applies to standard conditioning paradigms; for example, to single-cue delay conditioning when a tone is used as the conditioned stimulus (CS) and to differential delay conditioning when the positive and negative conditioned stimuli (CS+ and CS-) are a tone and white noise.
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Nielsen, M., Collier-Baker, E., Davis, J. M., & Suddendorf, T. (2005). Imitation recognition in a captive chimpanzee (Pan troglodytes). Anim. Cogn., 8(1), 31–36.
Abstract: This study investigated the ability of a captive chimpanzee (Pan troglodytes) to recognise when he is being imitated. In the experimental condition of test 1a, an experimenter imitated the postures and behaviours of the chimpanzee as they were being displayed. In three control conditions the same experimenter exhibited (1) actions that were contingent on, but different from, the actions of the chimpanzee, (2) actions that were not contingent on, and different from, the actions of the chimpanzee, or (3) no action at all. The chimpanzee showed more “testing” sequences (i.e., systematically varying his actions while oriented to the imitating experimenter) and more repetitive behaviour when he was being imitated, than when he was not. This finding was replicated 4 months later in test 1b. When the experimenter repeated the same actions she displayed in the experimental condition of test 1a back to the chimpanzee in test 2, these actions now did not elicit those same testing sequences or repetitive behaviours. However, a live imitation condition did. Together these results provide the first evidence of imitation recognition in a nonhuman animal.
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Domjan, M. (1976). Determinants of the enhancement of flavored-water intake by prior exposure. J Exp Psychol Anim Behav Process, 2(1), 17–27.
Abstract: The intake of a 2.0% sodium saccharin solution in rats was observed to increase as a function of both the number (Experiment 1) and the duration (Experiment 3) of prior periods of access to the saccharin flavor, but did not increase when subjects were maintained on a fluid deprivation procedure in the absence of saccharin exposure (Experiment 2). The enhancement of intake was further influenced by the schedule of saccharin preexposures in the absence of variations in the amount of solution tasted (Experiment 4). The effect was not a function of the opportunity for subjects to determine their own pattern of contact with the saccharin flavor, the opportunity for association of the flavor with hunger and thirst reduction, or the amount of saccharin swallowed during preexposure (Experiment 5). These results suggest that mere exposure to a flavored solution is sufficient to increase subsequent intakes. The phenomenon is discussed in terms of the attenuation of neophobia elicited by the novelty of flavored solutions.
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Rudy, J. W., Iwens, J., & Best, P. J. (1977). Pairing novel exteroceptive cues and illness reduces illness-induced taste aversions. J Exp Psychol Anim Behav Process, 3(1), 14–25.
Abstract: Four experiments are reported that lead to the conclusion that pairing novel exteroceptive stimulation (placement into a black compartment) with a poison (lithium chloride) attenuates the development of an aversion to a taste (saccharin) subsequently paired with the poison. Such an attenuation effect occurs whether the exteroceptive cues are present or absent when the taste-poison pairing is administered. Interpretation and implications of this finding are discussed.
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Griffin, D. R., & Speck, G. B. (2004). New evidence of animal consciousness. Anim. Cogn., 7(1), 5–18.
Abstract: This paper reviews evidence that increases the probability that many animals experience at least simple levels of consciousness. First, the search for neural correlates of consciousness has not found any consciousness-producing structure or process that is limited to human brains. Second, appropriate responses to novel challenges for which the animal has not been prepared by genetic programming or previous experience provide suggestive evidence of animal consciousness because such versatility is most effectively organized by conscious thinking. For example, certain types of classical conditioning require awareness of the learned contingency in human subjects, suggesting comparable awareness in similarly conditioned animals. Other significant examples of versatile behavior suggestive of conscious thinking are scrub jays that exhibit all the objective attributes of episodic memory, evidence that monkeys sometimes know what they know, creative tool-making by crows, and recent interpretation of goal-directed behavior of rats as requiring simple nonreflexive consciousness. Third, animal communication often reports subjective experiences. Apes have demonstrated increased ability to use gestures or keyboard symbols to make requests and answer questions; and parrots have refined their ability to use the imitation of human words to ask for things they want and answer moderately complex questions. New data have demonstrated increased flexibility in the gestural communication of swarming honey bees that leads to vitally important group decisions as to which cavity a swarm should select as its new home. Although no single piece of evidence provides absolute proof of consciousness, this accumulation of strongly suggestive evidence increases significantly the likelihood that some animals experience at least simple conscious thoughts and feelings. The next challenge for cognitive ethologists is to investigate for particular animals the content of their awareness and what life is actually like, for them.
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Crystal, J. D. (1999). Systematic nonlinearities in the perception of temporal intervals. J Exp Psychol Anim Behav Process, 25(1), 3–17.
Abstract: Rats judged time intervals in a choice procedure in which accuracy was maintained at approximately 75% correct. Sensitivity to time (d') was approximately constant for short durations 2.0-32.0 s with 1.0- or 2.0-s spacing between intervals (n = 5 in each group, Experiment 1), 2.0-50.0 s with 2.0-s spacing (n = 2, Experiment 1), and 0.1-2.0 s with 0.1- or 0.2-s spacing (n = 6 in each group, Experiment 2). However, systematic departures from average sensitivity were observed, with local maxima in sensitivity at approximately 0.3, 1.2, 10.0, 24.0, and 36.0 s. Such systematic departures from an approximately constant d' are predicted by a connectionist theory of time with multiple oscillators and may require a modification of the linear timing hypothesis of scalar timing theory.
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Broom, D. M. (2010). Cognitive ability and awareness in domestic animals and decisions about obligations to animals. Appl. Anim. Behav. Sci., 126(1-2), 1–11.
Abstract: Observation of behaviour, especially social behaviour, and experimental studies of learning and brain function give us information about the complexity of concepts that animals have. In order to learn to obtain a resource or carry out an action, domestic animals may: relate stimuli such as human words to the reward, perform sequences of actions including navigation or detours, discriminate amongst other individuals, copy the actions of other individuals, distinguish between individuals who do or do not have information, or communicate so as to cause humans or other animals to carry out actions. Some parrots, that are accustomed to humans but not domesticated, can use words to have specific meanings. In some cases, stimuli, individuals or actions are remembered for days, weeks or years. Events likely to occur in the future may be predicted and changes over time taken into account. Scientific evidence for the needs of animals depends, in part, on studies assessing motivational strength whose methodology depends on the cognitive ability of the animals. Recognition and learning may be associated with changes in physiology, behaviour and positive or negative feelings. Learning and other complex behaviour can result in affect and affect can alter cognition. The demonstration of cognitive bias gives indications about affect and welfare but should be interpreted in the light of other information. All of the information mentioned so far helps to provide evidence about sentience and the level of awareness. The term sentience implies a range of abilities, not just the capacity to have some feelings. The reluctance of scientists to attribute complex abilities and feelings to non-humans has slowed the development of this area of science. Most people consider that they have obligations to some animals. However, they might protect animals because they consider that an animal has an intrinsic value, or because of their concern for its welfare. In social species, there has been selection promoting moral systems that might result in behaviours such as attempts to avoid harm to others, collaboration and other altruistic behaviour. An evaluation of such behaviour may provide one of the criteria for decisions about whether or not to protect animals of a particular species. Other criteria may be: whether or not the animal is known as an individual, similarity to humans, level of awareness, extent of feelings, being large, being rare, being useful or having aesthetic quality for humans. Cognitive ability should also be considered when designing methods of enriching the environments of captive animals.
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Baragli, P., Demuru, E., & Palagi, E. (2015). Mirror on the wall, who is the horsest of our all? Self-recognition in Equus caballus. In Proceedings of the 3. International Equine Science Meeting.
Abstract: Mirror Self-Recognition (MSR) is an extremely rare capacity in the animal kingdom that reveals the emergence of complex cognitive capacities (de Waal 2008). So far, MSR has been reported only in humans, chimpanzees (Gallup, 1970), bottlenose dolphins (Reiss and Marino, 2001) and Asian elephants (Plotnik et al, 2006), all species characterized by a highly developed cognition. There is growing evidence that domestic horses posses high cognitive abilities, such as cross-modal individual recognition (Proops et al, 2009), triadic post-conflict reunion to maintain social homeostasis (Cozzi et al, 2010), complex communicative systems (Whatan and McComb, 2014), flexibility in problem-solving (Lovrovich et al, 2015), and long-term memory (Hanggi and Ingersoll, 2009). All these capacities make horses a good candidate to test the ability of MSR in a domestic species. Through a classical MSR experimental paradigm (de Waal 2008) we tested eight horses living in social groups under semi-natural conditions (from the Italian Horse Protection rescue centre). Animals showing MSR typically go through four stages (Plotnik et al, 2006): (i) social response, (ii) physical mirror inspection (e.g., looking behind the mirror), (iii) repetitive mirror-testing behaviour (i.e., the beginning of mirror understanding), and (iv) self-directed behaviour (i.e., recognition of the mirror image as self). The final stage, known as the “mark-test”, is verified when a subject spontaneously uses the mirror to check for a coloured artificial mark on its own body which it cannot perceive otherwise. The horses underwent a three-phase “mark-test”: 1) with sham mark (transparent ultrasound water gel) positioned on both side at jaw level, 2) mark (yellow eye shadow mixed with ultrasound water gel) positioned on left side of jaw (with sham mark on the right), 3) mark (yellow eye shadow mixed with ultrasound water gel) positioned on right side of jaw (with sham mark on the left)
The mirror was one 0.5-cm-thick piece of 140x220-cm plexiglass glue on wood. Each test lasted one hour, horses were tested once a day, in consecutive days and at the same time. Our preliminary result on 1 horse shows some changes in self-directed behaviours which can be attributed to presence of the coloured mark. Firstly, the presence of the coloured mark significantly increased the frequency of scratching on both sides of the muzzle (p < 0.0001). The most intriguing result (p < 0.0001) comes from the comparison of the scratching rates directed towards the coloured mark side (N = 41) and the sham mark side (N = 23). Under the control condition (i.e. sham mark on both sides) no statistical difference was found for the scratching rates directed to the muzzle sides (dx N = 8; sx N = 5). Although further analyses are needed to confirm these preliminary results, our finding opens new scenarios about the evolution of Mirror Self-Recognition. The capacity of horses to recognize themselves in a mirror may be the outcome of an evolutionary convergence process driven by the cognitive pressures imposed by a complex social system and maintained despite thousands years of domestication.
Keywords:
Domestic horse · Mark test · Socio-cognitive skills · Self-awareness
References
De Waal FBM (2008) The thief in the mirror. PloS Biol 6(8):e201
Gallup GG Jr (1970) Chimpanzees: Self-recognition. Science 167: 86-87.
Reiss D, Marino L (2001). Mirror self-recognition in the bottlenose dolphin: A case of cognitive convergence. Proc Natl Acad Sci U S A 98:5937-5942.
Plotnik J, de Waal FBM, Reiss D (2006) Self-recognition in an Asian elephant. Proc Natl Acad Sci U S A 103: 17053-17057.
Proops L, McComb K, Reby D. (2009) Cross-modal individual recognition in domestic horses (Equus caballus). Proc Nat Acad Sci USA;106:947-951.
Cozzi A, Sighieri C, Gazzano A, Nicol CJ, Baragli P. Post-conflict friendly reunion in a permanent group of horses (Equus caballus). Behav Process 2010;85:185-190.
Wathan J, McComb K. The eyes and ears are visual indicators of attention in domestic horses. Curr Biol 2014;24(15): R677-R679.
Lovrovich P, Sighieri C, Baragli P (2015) Following human-given cues or not? Horses (Equus caballus) get smarter and change strategy in a delayed three choice task. Appl Anim Behav Sci, in press.
Hanggi EB, Ingersoll JF. (2009) Long-term memory for categories and concepts in horses (Equus caballus). Anim Cogn; 12:451-462.
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