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Hampton, R. R. (2001). Rhesus monkeys know when they remember. Proc. Natl. Acad. Sci. U.S.A., 98(9), 5359–5362.
Abstract: Humans are consciously aware of some memories and can make verbal reports about these memories. Other memories cannot be brought to consciousness, even though they influence behavior. This conspicuous difference in access to memories is central in taxonomies of human memory systems but has been difficult to document in animal studies, suggesting that some forms of memory may be unique to humans. Here I show that rhesus macaque monkeys can report the presence or absence of memory. Although it is probably impossible to document subjective, conscious properties of memory in nonverbal animals, this result objectively demonstrates an important functional parallel with human conscious memory. Animals able to discern the presence and absence of memory should improve accuracy if allowed to decline memory tests when they have forgotten, and should decline tests most frequently when memory is attenuated experimentally. One of two monkeys examined unequivocally met these criteria under all test conditions, whereas the second monkey met them in all but one case. Probe tests were used to rule out “cueing” by a wide variety of environmental and behavioral stimuli, leaving detection of the absence of memory per se as the most likely mechanism underlying the monkeys' abilities to selectively decline memory tests when they had forgotten.
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Allen, C. (1998). Assessing animal cognition: ethological and philosophical perspectives. J. Anim Sci., 76(1), 42–47.
Abstract: Developments in the scientific and philosophical study of animal cognition and mentality are of great importance to animal scientists who face continued public scrutiny of the treatment of animals in research and agriculture. Because beliefs about animal minds, animal cognition, and animal consciousness underlie many people's views about the ethical treatment of nonhuman animals, it has become increasingly difficult for animal scientists to avoid these issues. Animal scientists may learn from ethologists who study animal cognition and mentality from an evolutionary and comparative perspective and who are at the forefront of the development of naturalistic and laboratory techniques of observation and experimentation that are capable of revealing the cognitive and mental properties of nonhuman animals. Despite growing acceptance of the ethological study of animal cognition, there are critics who dispute the scientific validity of the field, especially when the topic is animal consciousness. Here, a proper understanding of developments in the philosophy of mind and the philosophy of science can help to place cognitive studies on a firm methodological and philosophical foundation. Ultimately, this is an interdisciplinary task, involving scientists and philosophers. Animal scientists are well-positioned to contribute to the study of animal cognition because they typically have access to a large pool of potential research subjects whose habitats are more controlled than in most field studies while being more natural than most laboratory psychology experiments. Despite some formidable questions remaining for analysis, the prospects for progress in assessing animal cognition are bright.
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Shettleworth, S. J. (2003). Memory and hippocampal specialization in food-storing birds: challenges for research on comparative cognition. Brain Behav Evol, 62(2), 108–116.
Abstract: The three-way association among food-storing behavior, spatial memory, and hippocampal enlargement in some species of birds is widely cited as an example of a new 'cognitive ecology' or 'neuroecology.' Whether this relationship is as strong as it first appears and whether it might be evidence for an adaptive specialization of memory and hippocampus in food-storers have recently been the subject of some controversy [Bolhuis and Macphail, 2001; Macphail and Bolhuis, 2001]. These critiques are based on misconceptions about the nature of adaptive specializations in cognition, misconceptions about the uniformity of results to be expected from applying the comparative method to data from a wide range of species, and a narrow view of what kinds of cognitive adaptations are theoretically interesting. New analyses of why food-storers (black-capped chickadees, Poecile Atricapilla) respond preferentially to spatial over color cues when both are relevant in a memory task show that this reflects a relative superiority of spatial memory as compared to memory for color rather than exceptional spatial attention or spatial discrimination ability. New studies of chickadees from more or less harsh winter climates also support the adaptive specialization hypothesis and suggest that within-species comparisons may be especially valuable for unraveling details of the relationships among ecology, memory, and brain in food-storing species.
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Pepperberg, I. M. (2002). In search of king Solomon's ring: cognitive and communicative studies of Grey parrots (Psittacus erithacus). Brain Behav Evol, 59(1-2), 54–67.
Abstract: During the past 24 years, I have used a modeling technique (M/R procedure) to train Grey parrots to use an allospecific code (English speech) referentially; I then use the code to test their cognitive abilities. The oldest bird, Alex, labels more than 50 different objects, 7 colors, 5 shapes, quantities to 6, 3 categories (color, shape, material) and uses 'no', 'come here', wanna go X' and 'want Y' (X and Y are appropriate location or item labels). He combines labels to identify, request, comment upon or refuse more than 100 items and to alter his environment. He processes queries to judge category, relative size, quantity, presence or absence of similarity/difference in attributes, and show label comprehension. He semantically separates labeling from requesting. He thus exhibits capacities once presumed limited to humans or nonhuman primates. Studies on this and other Greys show that parrots given training that lacks some aspect of input present in M/R protocols (reference, functionality, social interaction) fail to acquire referential English speech. Examining how input affects the extent to which parrots acquire an allospecific code may elucidate mechanisms of other forms of exceptional learning: learning unlikely in the normal course of development but that can occur under certain conditions.
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Marino, L. (2002). Convergence of complex cognitive abilities in cetaceans and primates. Brain Behav Evol, 59(1-2), 21–32.
Abstract: What examples of convergence in higher-level complex cognitive characteristics exist in the animal kingdom? In this paper I will provide evidence that convergent intelligence has occurred in two distantly related mammalian taxa. One of these is the order Cetacea (dolphins, whales and porpoises) and the other is our own order Primates, and in particular the suborder anthropoid primates (monkeys, apes, and humans). Despite a deep evolutionary divergence, adaptation to physically dissimilar environments, and very different neuroanatomical organization, some primates and cetaceans show striking convergence in social behavior, artificial 'language' comprehension, and self-recognition ability. Taken together, these findings have important implications for understanding the generality and specificity of those processes that underlie cognition in different species and the nature of the evolution of intelligence.
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Heschl, A., & Burkart, J. (2006). A new mark test for mirror self-recognition in non-human primates. Primates, 47(3), 187–198.
Abstract: For 30 years Gallup's (Science 167:86-87, 1970) mark test, which consists of confronting a mirror-experienced test animal with its own previously altered mirror image, usually a color mark on forehead, eyebrow or ear, has delivered valuable results about the distribution of visual self-recognition in non-human primates. Chimpanzees, bonobos, orangutans and, less frequently, gorillas can learn to correctly understand the reflection of their body in a mirror. However, the standard version of the mark test is good only for positively proving the existence of self-recognition. Conclusive statements about the lack of self-recognition are more difficult because of the methodological constraints of the test. This situation has led to a persistent controversy about the power of Gallup's original technique. We devised a new variant of the test which permits more unequivocal decisions about both the presence and absence of self-recognition. This new procedure was tested with marmoset monkeys (Callithrix jacchus), following extensive training with mirror-related tasks to facilitate performance in the standard mark test. The results show that a slightly altered mark test with a new marking substance (chocolate cream) can help to reliably discriminate between true negative results, indicating a real lack of ability to recognize oneself in a mirror, from false negative results that are due to methodological particularities of the standard test. Finally, an evolutionary hypothesis is put forward as to why many primates can use a mirror instrumentally – i.e. know how to use it for grasping at hidden objects – while failing in the decisive mark test.
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Overli, O., Sorensen, C., Pulman, K. G. T., Pottinger, T. G., Korzan, W., Summers, C. H., et al. (2007). Evolutionary background for stress-coping styles: relationships between physiological, behavioral, and cognitive traits in non-mammalian vertebrates. Neurosci Biobehav Rev, 31(3), 396–412.
Abstract: Reactions to stress vary between individuals, and physiological and behavioral responses tend to be associated in distinct suites of correlated traits, often termed stress-coping styles. In mammals, individuals exhibiting divergent stress-coping styles also appear to exhibit intrinsic differences in cognitive processing. A connection between physiology, behavior, and cognition was also recently demonstrated in strains of rainbow trout (Oncorhynchus mykiss) selected for consistently high or low cortisol responses to stress. The low-responsive (LR) strain display longer retention of a conditioned response, and tend to show proactive behaviors such as enhanced aggression, social dominance, and rapid resumption of feed intake after stress. Differences in brain monoamine neurochemistry have also been reported in these lines. In comparative studies, experiments with the lizard Anolis carolinensis reveal connections between monoaminergic activity in limbic structures, proactive behavior in novel environments, and the establishment of social status via agonistic behavior. Together these observations suggest that within-species diversity of physiological, behavioral and cognitive correlates of stress responsiveness is maintained by natural selection throughout the vertebrate sub-phylum.
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Cerutti, D. T., & Staddon, J. E. R. (2004). Immediacy versus anticipated delay in the time-left experiment: a test of the cognitive hypothesis. J Exp Psychol Anim Behav Process, 30(1), 45–57.
Abstract: In the time-left experiment (J. Gibbon & R. M. Church, 1981), animals are said to compare an expectation of a fixed delay to food, for one choice, with a decreasing delay expectation for the other, mentally representing both upcoming time to food and the difference between current time and upcoming time (the cognitive hypothesis). The results of 2 experiments support a simpler view: that animals choose according to the immediacies of reinforcement for each response at a time signaled by available time markers (the temporal control hypothesis). It is not necessary to assume that animals can either represent or subtract representations of times to food to explain the results of the time-left experiment.
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Pickens, C. L., & Holland, P. C. (2004). Conditioning and cognition. Neurosci Biobehav Rev, 28(7), 651–661.
Abstract: Animals' abilities to use internal representations of absent objects to guide adaptive behavior and acquire new information, and to represent multiple spatial, temporal, and object properties of complex events and event sequences, may underlie many aspects of human perception, memory, and symbolic thought. In this review, two classes of simple associative learning tasks that address these core cognitive capacities are discussed. The first set, including reinforcer revaluation and mediated learning procedures, address the power of Pavlovian conditioned stimuli to gain access, through learning, to representations of upcoming events. The second set of investigations concern the construction of complex stimulus representations, as illustrated in studies of contextual learning, the conjunction of explicit stimulus elements in configural learning procedures, and recent studies of episodic-like memory. The importance of identifying both cognitive process and brain system bases of performance in animal models is emphasized.
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Sarter, M. (2004). Animal cognition: defining the issues. Neurosci Biobehav Rev, 28(7), 645–650.
Abstract: The assessment of cognitive functions in rodents represents a critical experimental variable in many research fields, ranging from the basic cognitive neurosciences to psychopharmacology and neurotoxicology. The increasing use of animal behavioral tests as 'assays' for the assessment of effects on learning and memory has resulted in a considerable heterogeneity of data, particularly in the field of behavioral and psycho pharmacology. The limited predictive validity of changes in behavioral performance observed in standard animal tests of learning and memory indicates that a renewed effort to scrutinize the validity of these tests is warranted. In humans, levels of processing (effortful vs. automatic) and categories of information (procedural vs. episodic/declarative) are important variables of cognitive operations. The design of tasks that assess the recall of 'episodic' or 'declarative' information appears to represent a particular challenge for research using laboratory rodents. For example, the hypothesis that changes in inspection time for a previously encountered place or object are based on the recall of declarative/episodic information requires substantiation. In order to generalize findings on the effects of neuronal or pharmacological manipulations on learning and memory, obtained from one species and one task, to other species and other tasks, the mediating role of important sets of variables which influence learning and memory (e.g. attentional, affective) needs to be determined. Similar to the view that a neuronal manipulation (e.g. a lesion) represents a theory of the condition modeled (e.g. a degenerative disorder), an animal behavioral task represents a theory of the behavioral/cognitive process of interest. Therefore, the test of hypotheses regarding the validity of procedures used to assess cognitive functions in animals is an inherent part of the research process.
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