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Cramer, P. Defense mechanisms in psychology today: Further processes for adaptation. American Psychologist, 55 , — Descartes, R. In The philosophical writings of descartes, vol. II, pp. Cambridge: Cambridge University Press. Freud, S. The Interpretation of Dreams. Standard Edition , vols.

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London: Hogarth. Introductory lectures on psycho-analysis.

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The ego and the id. Standard Edition , vol. New introductory lectures on psycho-analysis. Freud, A. The ego and the mechanisms of defence. London: The Hogarth Press. An outline of psycho-analysis. Jones, B. Repression: The evolution of a psycho-analytic concept from the s to the s. Journal of the American Psycho-analytic Association, 41 , 63— Kaplan-Solms, K. Clinical studies in Neuro-psychoanalysis: Introduction to a depth psychology. London: Karnac. Kihlstrom, J.

The cognitive unconscious. Science, , — MacIntyre, A. The unconscious: A conceptual analysis. Petocz, A. Freud, psychoanalysis, and symbolism. Talvitie, V. Priming is presumably advantageous because animals evolved in a world where things that are encountered once are likely to be encountered again. Priming improves the speed and efficiency with which organisms interact with a familiar environment and may influence feature-based attentional processes Hutchinson and Turk-Browne ; Theeuwes Evoked-potential studies indicate that the electrophysiological signature of priming occurs early and well before the activity that signals conscious recognition of a past event Paller et al.

In neuroimaging studies, priming is often associated with reduced activity in regions of neocortex relevant to the task Squire et al. A similar finding repetition suppression Desimone has been described in nonhuman primates a stimulus-specific attenuation in firing rate with repeated presentation of a stimulus , and may underlie the phenomenon of priming Wiggs and Martin Models have been proposed to explain how a net reduction in cortical activity could allow for faster perceptual processing i.

Some studies have found a correlation between behavioral measures of priming and reduced activity in the prefrontal cortex Maccotta and Buckner This result has not been found in ventral temporal cortex for either humans or nonhuman primates Maccotta and Buckner ; McMahon and Olson Changes in cortex also underlie the related phenomenon of perceptual learning Gilbert et al.

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  • Conscious and Unconscious Memory Systems.

Perceptual learning refers to gradual improvement in the detection or discrimination of visual stimuli with repeated practice. Changes in cortical circuitry during perceptual learning are detectable as early as primary visual cortex V1 and may depend in part on structural changes in the long-range horizontal connections formed by V1 pyramidal cells Gilbert and Li This circuitry is under the control of bottom-up processes as well as top-down influences related to attention and behavioral context Gilbert and Li Another early example of nondeclarative memory was simple classical conditioning, best illustrated in the literature of delay eyeblink conditioning.

In delay conditioning, a neutral conditioned stimulus CS , such as a tone, is presented just before an unconditioned stimulus US , such as an airpuff to the eye. The two stimuli then overlap and coterminate. Critically, delay eyeblink conditioning is intact in amnesia and is acquired independently of awareness Gabrieli et al. Participants who did not become aware of the relationship between the CS and US i. Indeed, when CS—US association strength was varied by changing the number of consecutive CS alone or CS—US presentations , the probability of a conditioned response increased with association strength but was inversely related to how much the US was expected Clark et al.

The inevitable contrast: Conscious vs. unconscious processes in action control

Largely on the basis of work with rabbits, delay eyeblink conditioning proved to depend on the cerebellum and associated brain stem circuitry Thompson and Krupa ; Thompson and Steinmetz Forebrain structures are not necessary for acquisition or retention of classically conditioned eyeblink responses. Evaluative information, that is, whether a stimulus has positive or negative value, is acquired largely as nondeclarative memory. Biological study of this kind of memory has focused especially on the associative learning of fear Davis ; Adolphs ; LeDoux Its nondeclarative status is illustrated by the fact that, in humans, associative fear learning proceeded normally after hippocampal lesions, even though the CS—US pairings could not be reported Bechara et al.

The amygdala has a critical role in fear learning, and its function as well as its connectivity appears to be conserved widely across species. In human neuroimaging studies, the amygdala was activated not only by fear but by strongly positive emotions as well Hamann et al. Thus, the amygdala appears to be critical for associating sensory stimuli with stimulus valence.

Ordinarily, animals express fear learning by freezing behavior immobility. However, in a task where learned fear must instead be expressed by executing an avoidance response an escape , freezing is maladaptive. In this case, prefrontal cortex inhibits defense behaviors such as freezing that are mediated by the amygdala, thereby allowing the animal to escape Moscarello and LeDoux Inhibitory action of the prefrontal cortex on the amygdala from infralimbic prefrontal cortex in rat or from ventromedial prefrontal cortex in humans has also been found to occur during the reversal of fear learning i.

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This work has relevance for clinical disorders, such as phobias and posttraumatic stress disorder Davis In addition to these functions, it is important to note that the amygdala also exerts a modulatory influence on both declarative and nondeclarative memory. This role of the amygdala is the basis for the fact that emotionally arousing events are typically remembered better than emotionally neutral events.

The mechanism for this effect is understood and depends on the release of stress hormones from the adrenal gland, which affects the forebrain via the vagus nerve, the nucleus of the solitary tract, and the locus coeruleus. Ultimately, the effect is mediated by the amygdala through its basolateral nucleus McGaugh and Roozendaal The gradual trial-and-error learning that leads to the formation of habits was proposed in the s to be supported by the striatum Mishkin et al.

Habit memory is characterized by automatized, repetitive behavior and, unlike declarative memory, is insensitive to changes in reward value Dickinson An early demonstration of the distinction between declarative memory and habit memory came from rats with fornix lesions or caudate lesions tested on two ostensibly similar tasks.

Rats with fornix lesions, which disrupt hippocampal function, failed when they needed to acquire a flexible behavior but succeeded when they needed to respond repetitively. Rats with caudate lesions showed the opposite pattern Packard et al. In the task, probabilistic classification, participants gradually learned which of two outcomes sun or rain would occur on each trial, given the particular combination of four cues that appeared. One, two, or three cues could appear on any trial, and each cue was independently and probabilistically related to the outcome.

Patients with hippocampal lesions learned the task at a normal rate but could not report facts about the task. Parkinson patients remembered the facts but could not learn the task. Tasks that can be learned quickly by memorization can also be learned by a trial-and-error, habit-based strategy, albeit much more slowly. Two severely amnesic patients with large medial temporal lobe lesions also learned but only gradually, requiring more than 25 test sessions and trials Bayley et al.

Unlike declarative memory, which is flexible and can guide behavior in different contexts, the acquired knowledge in this case was rigidly organized. Patient performance collapsed when the task format was altered by asking participants to sort the 16 objects into correct and incorrect groups a trivial task for controls.

In addition, although by the end of training the patients were consistently performing at a high level, at the start of each test day they were never able to describe the task, the instructions, or the objects. Indeed, during testing they expressed surprise that they were performing so well.

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These findings provide particularly strong evidence for the distinction between declarative conscious and nondeclarative unconscious memory systems. Reward-based learning of this kind depends on dopamine neurons in the midbrain substantia nigra and ventral tegmental area , which project to the striatum and signal the information value of the reward Schultz The dorsolateral striatum is crucial for the development of habits in coordination with other brain regions.

Neurophysiological studies in mice during skill learning documented that the dorsolateral striatum was increasingly engaged as performance became more automatic and habit-like Yin et al. In contrast, the dorsomedial striatum was engaged only early in training. Similarly, in rats learning a conditioned T-maze task, activity gradually increased in the dorsolateral striatum as training progressed, and this activity correlated with performance Thorn et al.

In the dorsomedial striatum, activity first increased but then decreased as training progressed. Increased activity in the dorsolateral striatum during the later stages of habit formation occurred together with late-developing activity in infralimbic cortex Smith and Graybiel Moreover, disruption of infralimbic cortex during late training prevented habit formation. Thus, these two regions dorsolateral striatum and infralimbic cortex appear to work together to support a fully formed habit.

The memory systems of the mammalian brain operate independently and in parallel to support behavior, and how one system or another gains control is a topic of considerable interest Poldrack and Packard ; McDonald and Hong ; Packard and Goodman In some circumstances, memory systems are described as working cooperatively to optimize behavior and in other circumstances are described as working competitively. However, it is not easy to pin down what should count for or against cooperativity, competition, or independence in any particular case.

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For example, the fact that the manipulation of one memory system can affect the operation of another has been taken as evidence for competition between systems Schwabe Yet, even for systems that are strictly independent, the loss of one system would be expected to affect the operation of another system by affording it more opportunity to control behavior. Much of the experimental work on the relationship between memory systems has focused on hippocampus-dependent declarative memory and dorsolateral striatum-dependent habit memory.

In an illustrative study Packard and McGaugh , rats were trained in a four-arm, plus-shaped maze to go left, always beginning in the south arm and with the north arm blocked.