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Dissociating intentional learning from relative novelty responses in the medial temporal lobe.

The establishment of a role for medial temporal lobe (MTL) structures in episodic memory has led to an investigative focus on the specific contributions and interactions between constituent MTL regions, including the hippocampus and surrounding medial temporal cortices. By dissociating an intentional stimulus-category learning condition from a passive viewing condition, we demonstrate, using fMRI, that novelty- and familiarity-driven responses in human anterior and posterior hippocampus, respectively, only occur during intentional learning. With increasing familiarity of stimulus-category associations, there is a shift in neuronal responses from anterior to posterior hippocampal regions. This anterior/posterior response gradient may reflect a weighting of functional hippocampal architecture related to encoding of novel and retrieval of familiar information. By contrast, perirhinal cortex is engaged by novel stimuli irrespective of task, highlighting this region as a component of a generic familiarity discrimination system. By introducing distinct stimulus types, we further demonstrate that these MTL responses are independent of stimulus complexity. Different patterns of activity for intentional learning vs. passive viewing indicate that intentional encoding/retrieval of stimulus-category associations and automatic novelty/familiarity assessment of stimuli are processed in anatomically dissociable neuronal ensembles within the MTL memory system.

Adult↗

The nature of declarative and nondeclarative knowledge for implicit and explicit learning.

Using traditional implicit and explicit artificial-grammar learning tasks, the author investigated the similarities and differences between the acquisition of declarative knowledge under implicit and explicit learning conditions and the functions of the declarative knowledge during testing. Results suggested that declarative knowledge was not predictive of or required for implicit learning but was related to consistency in implicit learning performance. In contrast, declarative knowledge was predictive of and required for explicit learning and was related to consistency in performance. For explicit learning, the declarative knowledge functioned as a guide for other behavior. In contrast, for implicit learning, the declarative knowledge did not serve as a guide for behavior but was instead a post hoc description of the most commonly seen stimuli.

Adult↗

Stimulus class membership established via stimulus-reinforcer relations.

In an arbitrary matching-to-sample procedure, two mentally retarded subjects learned conditional discriminations with two sets of stimuli. Each set included a spoken name (N1 or N2), an object (O1 or O2), and a printed symbol (S1 or S2). One subject selected conditionally (a) O1 upon N1, and O2 upon N2, and (b) S1 upon O1, and S2 upon O2. The other subject selected conditionally (a) S1 upon N1, and S2 upon N2, and (b) O1 upon S1, and O2 upon S2. For both subjects, selections of O1 and S1 produced one type of food, F1; selections of O2 and S2 produced a different type of food, F2. Both subjects also learned identity-matching performances, selecting O1, O2, S1, S2, F1, and F2 conditionally upon those stimuli as samples; F1 followed selections of O1, S1, and F1; F2 followed selections of O2, S2, and F2. Matching performances consistent with stimulus class formation involving the names, objects, symbols, and foods were demonstrated on probe trials, even though these performances had not been taught explicitly. Next, new objects, X1 and X2, were presented on identity-matching trials, producing F1 and F2, respectively. Without further training, X1 was selected conditionally upon N1, S1, and O1, and X2 was selected upon N2, S2, and O2. When the contingencies were changed so that selections of X1 and X2 were now followed by F2 and F1, respectively, X2 was selected conditionally upon N1, S1, and O1, and X1 was selected upon N2, S2, and O2. Class membership of X1 and X2 had apparently changed. This study provides evidence that reinforcers may become members of stimulus classes, and that new stimuli may become class members through relations with reinforcers.

Adult↗

Cerebellar lesions and the nictitating membrane reflex: performance deficits of the conditioned and unconditioned response.

Unilateral cerebellar lesions abolished the occurrence of ipsilateral conditioned nictitating membrane responses during the 285 msec interval between onset of the conditioned and unconditioned stimuli on paired trials. This effect was obtained in 15 animals sustaining damage to the dorsolateral aspects of the interpositus nucleus and the adjoining white matter. However, conditioned responses did occur during the 800 msec observation interval employed on tone-alone test trials, and these responses exhibited the classic performance deficits normally associated with cerebellar damage: a low frequency of occurrence (14%, as compared with 96% before the lesion); a 3.1 mm decrease in amplitude; a 236 msec increase in onset latency; a 563 msec increase in latency of peak amplitude; and a 327 msec increase in rise time. Four of the 15 animals failed to demonstrate greater than 5% responding during the test trials. These performance deficits were not specific to the learned, conditioned response. Unconditioned responses were also reduced in frequency and increased in latency of peak amplitude and rise time, especially when elicited at lower air-puff intensities. These deficits in the unconditioned response were observed in animals that failed to exhibit conditioned responses on either paired or test trials, as well as in animals demonstrating conditioned responses only during test trials. We conclude that the cerebellum has a general role in regulating the nictitating membrane reflex so that deficits in learned responses observed after cerebellar lesions are secondary to a broader deficit in performance. The performance deficits appear to consist of a sensory component, as reflected by an increase in stimulus threshold for elicitation of the nictitating membrane reflex, and a motor component, as reflected by the altered topography of the evoked response. The results of this study thus reaffirm the role of the cerebellum in regulating the sensorimotor processes necessary for the optimal performance of both conditioned and unconditioned responses and extends this role to the expression of a simple cranial nerve reflex.

Animals↗

Age-related effects of the nicotinic antagonist mecamylamine on cognition and behavior.

Studies of the neurochemical pathology of Alzheimer's disease and Parkinson's disease reveal a severe and specific loss of central nicotinic cholinergic receptors. We have investigated the functional significance of this finding for cognitive functioning by studying the effects of the centrally active nicotinic antagonist mecamylamine. Single oral doses of mecamylamine were administered to 12 healthy young males and 15 healthy elderly subjects in doses of 5, 10, and 20 mg in a placebo-controlled, double-blind study. In both groups, the 20-mg dose caused a significant increase in errors in the learning condition of the Repeated Acquisition Task, producing a slower acquisition curve. There was no effect of drug on the performance component (retrieval of previously learned information). However, elderly subjects showed enhanced sensitivity to mecamylamine, with 10-mg dose producing significant impairment of learning not seen in the young normals. On a recognition memory task, there was an age-associated shift in response bias, with the elderly subjects becoming more liberal with increasing dose. Reaction-time measures suggested a dose-related slowing of reaction time on several tasks. Behavioral effects were minimal and physiologic effects were consistent with dose-related ganglionic blockade. These results indicate that acute blockade of nicotinic receptor function can produce measurable and significant cognitive impairment similar to some deficits seen in dementing illnesses, and that there is an age-related increase in sensitivity to nicotinic blockade.

Adult↗

Influence of sleep position experience on ability of prone-sleeping infants to escape from asphyxiating microenvironments by changing head position.

OBJECTIVE: Several studies have found that back- or side-sleeping infants who are inexperienced in prone sleeping are at much higher risk for sudden infant death syndrome (SIDS) when they turn to prone or are placed prone for sleep compared with infants who normally sleep prone. Moreover, such inexperienced infants are more likely to be found in the face-down position at death after being placed prone compared with SIDS infants who are experienced in prone sleeping. We hypothesized that lack of experience in prone sleeping is associated with increased difficulty in changing head position to avoid an asphyxiating sleep environment. METHODS: We studied 38 healthy infants while they slept prone. Half of these were experienced and half were inexperienced in prone sleeping. To create a mildly asphyxiating microenvironment, we placed infants to sleep prone with their faces covered by soft bedding. We recorded inspired CO2 (CO2I), electrocardiogram, and respiration, and we videotaped head movements. Also, we assessed gross motor development (Denver Development Scale). RESULTS: When sleeping prone, with their faces covered by bedding, all infants experienced mild asphyxia as a result of rebreathing. All aroused and attempted escape from this environment. Infants used 3 stereotyped head-repositioning strategies. The least effective was nuzzling into the bedding with occasional brief head lifts. More effective were head lifts combined with a head turn. Some infants, however, could turn only to 1 side, right or left. Infants who were inexperienced in prone sleeping had less effective protective behaviors than experienced infants. Infant age did not correlate with efficacy of protective behaviors. Infants who were experienced in prone sleep had advanced gross motor development compared with inexperienced infants. CONCLUSION: Infants who are inexperienced in prone sleeping have decreased ability to escape from asphyxiating sleep environments when placed prone. These observations potentially explain the increased risk associated with prone sleep in infants who are inexperienced. The increased occurrence of the face-down position in such infants is also potentially explained. These findings suggest that airway protective behaviors may be acquired through the mechanism of operant conditioning (learning).

Aging↗

Locomotion, incidental learning, and the selection of spatial reference systems.

In three experiments, we examined the effects of locomotion and incidental learning on the formation of spatial memories. Participants learned the locations of objects in a room and then made judgments of relative direction, using their memories (e.g., "Imagine you are standing at the clock, facing the jar. Point to the book"). The experiments manipulated the number of headings experienced, the amount of interaction with the objects, and whether the participants were informed that their memories of the layout would be tested. When participants were required to maintain a constant body orientation during learning (Experiment 1), they represented the layout in terms of a single reference direction parallel to that orientation. When they were allowed to move freely in the room (Experiment 2), they seemed to use two orthogonal reference axes aligned with the walls of the enclosing room. Extensive movement under incidental learning conditions (Experiment 3) yielded a mixture of these two encoding strategies across participants. There was no evidence that locomotion, interaction with objects, or incidental learning led to the formation of spatial memories that differed from those formed from static viewing.

Adult↗

The power law repealed: the case for an exponential law of practice.

The power function is treated as the law relating response time to practice trials. However, the evidence for a power law is flawed, because it is based on averaged data. We report a survey that assessed the form of the practice function for individual learners and learning conditions in paradigms that have shaped theories of skill acquisition. We fit power and exponential functions to 40 sets of data representing 7,910 learning series from 475 subjects in 24 experiments. The exponential function fit better than the power function in all the unaveraged data sets. Averaging produced a bias in favor of the power function. A new practice function based on the exponential, the APEX function, fit better than a power function with an extra, preexperimental practice parameter. Clearly, the best candidate for the law of practice is the exponential or APEX function, not the generally accepted power function. The theoretical implications are discussed.

Data Interpretation, Statistical↗

The mnemonic mechanisms of errorless learning.

Errorless learning enhances memory relative to errorful, trial-and-error learning, but the extent to which this advantage relies on implicit or explicit memory processes is not clear. Previous attempts to determine the mnemonic mechanisms of errorless learning have relied on contrasts between patient groups or between tasks, but both approaches are problematic. In this study, healthy younger and older adults were engaged in errorless or errorful learning using a process dissociation procedure that provides separate estimates of explicit recollection and implicit familiarity within-subjects and within-task (Hay & Jacoby, 1996). Consistent with much prior research, we found an age-related decrement in recollection, but age-invariance in familiarity. In the young adults, errorless learning reduced the misleading familiarity of prior errors, but this benefit was offset by the non-elaborative nature of the errorless learning condition that also reduced recollection. In the older adults, who are less able to oppose familiarity-based errors because of their lower recollection, errorless learning only reduced the misleading impact of previous errors. Our results support Baddeley and Wilson's (1994) position that the errorless learning effect is mediated by implicit memory processes: individuals with reduced explicit memory benefit from errorless learning because errorless learning bypasses the need to engage explicit error elimination processes. We do not recommend standard errorless learning for individuals with intact explicit memory, such as students trying to learn information in preparation for an examination.

Adolescent↗

Learning to write letters: transfer in automated movements indicates modularity of motor programs in human subjects.

Many automatic movements are open-loop, feed-forward motor programs (MP) that are kinematically well characterized by smooth speed and acceleration curves. However, it is unclear whether their internal representation consists of monolithic blocks or subroutines. This question was investigated using a learning paradigm of a writing task. Fifty-nine normal subjects were presented with two similar, but different new letters. Every subject practiced each letter in a series of 60 trials, with the order of letter series randomized. Every session was continuously recorded by a digitizing tablet. Using kinematic analysis, we measured the number of vertical acceleration peaks as an indication of the number of corrective movements (COM). Since COM declined as automatization was approached, we could quantitatively infer progress in motor learning under natural learning conditions. In the case of modular storage of MP, transfer in-between letters was expected due to the re-use of pre-learned motor subroutines. Statistical analysis showed that the exponential model described the data much better than the linear model (residual error: P<0.88 and P<0.00001, respectively), as expected for a learning paradigm. There was no difference between letters per se (P<0.77). Motor improvement differed significantly (P<0.02) between the first and the second series; there was a much greater reduction of COM in the second series (50.1 vs. 41.1%). This difference can be logically ascribed to transfer, indicating that automated movements are stored in motor subroutines.

Biomechanical Phenomena↗

The effect of periarcuate lesions in the monkey on the performance of symmetrically and asymmetrically reinforced visual and auditory go, no-go tasks.

Monkeys with lesions restricted to the periarcuate region of the frontal cortex were impaired on go, no-go tasks in which, depending on the stimulus present on any given trial, they were rewarded either for pushing a manipulandum or for withholding this response for a given period of time. By contrast, these animals were able to learn at a normal rate go, no-go tasks in which they were rewarded if they responded in the presence of a "positive" stimulus, but not in the presence of a "negative" stimulus. In the latter situation, responding in the presence of the "negative" stimulus was extinguished through non-reward, whereas the tasks on which impairments were demonstrated required solution on the basis of a conditional rule: If stimulus A, respond, and if stimulus B, withhold responding, to receive reward. Because the 2 experimental situations were identical in every respect except for the one critical difference in the testing procedure, the present set of experiments provides a powerful demonstration of the involvement of the periarcuate cortex in conditional learning with exteroceptive stimuli.

Animals↗

Lesions of the midline thalamic nuclei impair classical conditioning under partial but not continuous reinforcement conditions.

Rabbits with lesions of the midline thalamic nuclei were compared to rabbits with sham lesions on classical eyeblink (EB) and heart rate (HR) conditioning. Separate groups of sham and lesioned animals received either 25, 50 or 100% reinforcement with a periorbital shock unconditioned stimulus. Animals with lesions showed slightly impaired EB conditioning, compared to sham animals, under the partial but not continuous reinforcement schedules. Midline lesions also reduced the magnitude of the decelerative HR conditioned response under partial but not continuous reinforcement conditions. These findings suggest that the midline thalamic nuclei, like the mediodorsal nucleus of the thalamus, process information required for response selection under non-optimal learning conditions.

Animals↗

Learning style preferences of occupational therapy students.

Fifty-five undergraduate and 22 graduate students in occupational therapy were surveyed by using the Canfield-Lafferty Learning Styles Inventory. The results of the study indicate that their learning style preferences favor the learning conditions that permit knowing and liking the instructor personally, setting one's own objectives, and working alone and independently. The highest content and mode preferences for learning were working with people and engaging in direct experience, respectively. The lowest content and mode preferences were for working with numbers and for reading. Achievement expectations for grades were A and B. The findings are discussed.

Florida↗

stathmin, a gene enriched in the amygdala, controls both learned and innate fear.

Little is known about the molecular mechanisms of learned and innate fear. We have identified stathmin, an inhibitor of microtubule formation, as highly expressed in the lateral nucleus (LA) of the amygdala as well as in the thalamic and cortical structures that send information to the LA about the conditioned (learned fear) and unconditioned stimuli (innate fear). Whole-cell recordings from amygdala slices that are isolated from stathmin knockout mice show deficits in spike-timing-dependent long-term potentiation (LTP). The knockout mice also exhibit decreased memory in amygdala-dependent fear conditioning and fail to recognize danger in innately aversive environments. By contrast, these mice do not show deficits in the water maze, a spatial task dependent on the hippocampus, where stathmin is not normally expressed. We therefore conclude that stathmin is required for the induction of LTP in afferent inputs to the amygdala and is essential in regulating both innate and learned fear.

Amygdala↗

The origin of exemplar effects in rule-driven categorization.

S. W. Allen and L. R. Brooks (1991) have shown that exemplar memory can affect categorization even when participants are provided with a classification rule. G. Regehr and L. R. Brooks (1993) argued that stimuli must be individuated for such effects to occur. In this study, the authors further analyze the conditions that yield exemplar effects in this rule application paradigm. The results of Experiments 1-3 show that interchangeable attributes, which are not part of the rule, influence categorization only when attention is explicitly drawn on them. Experiment 4 shows that exemplar effects can occur in an incidental learning condition, whether stimulus individuation is preserved or not. The authors conclude that the influence of exemplar learning in rule-driven categorization stems from the attributes specified in the rule or in the instructions, not from the stimulus gestalts.

Cognition↗

Intra-amygdala administration of polyamines modulates fear conditioning in rats.

Amygdalar NMDA receptor activation has been implicated in the acquisition of fear memories in rats. However, little is known about the role of endogenous modulators of the NMDA receptor, such as polyamines, in pavlovian fear-conditioning learning. Therefore, in the present study we investigated whether the immediate pretraining or post-training bilateral infusion of arcaine, an antagonist of the NMDA receptor polyamine-binding site, or spermidine, an agonist of the NMDA receptor polyamine-binding site, into the amygdala affected classical fear conditioning in rats. Bilateral microinjections of arcaine (0.0002-0.2 nmol) decreased, whereas spermidine (0.002-20 nmol) increased, contextual and auditory fear conditioning. Arcaine coadministration, at a dose that had no effect per se, reversed the facilitatory effect of spermidine. These results provide evidence that endogenous and exogenous polyamines modulate the acquisition or early consolidation (or both) of the fear-conditioning task in the amygdala.

Amygdala↗

State-dependent modulation of feeding behavior by proopiomelanocortin-derived beta-endorphin.

Feeding behavior can be divided into appetitive and consummatory phases, differing in neural substrates and effects of deprivation. Opioids play an important role in the appetitive aspects of feeding, but they also have acute stimulatory effects on food consumption. Because the opioid peptide beta-endorphin is co-synthesized and released with melanocortins from proopiomelanocortin (POMC) neuronal terminals, we examined the physiological role of beta-endorphin in feeding and energy homeostasis using a strain of mutant mice with a selective deficiency of beta-endorphin. Male beta-endorphin-deficient mice unexpectedly became obese with ad libitum access to rodent chow. Total body weight increased by 15% with a 50-100% increase in the mass of white fat. The mice were hyperphagic with a normal metabolic rate. Despite the absence of endogenous beta-endorphin, the mutant mice did not differ from wild-type mice in their acute feeding responses to beta-endorphin or neuropeptide Y administered intracerebroventricularly or naloxone administered intraperitoneally. Additional mice were studied using an operant behavioral paradigm to examine their acquisition of food reinforcers under increasing work demands. Food-deprived, beta-endorphin-deficient male mice emitted the same number of lever presses under a progressive ratio schedule compared to wild-type mice. However, the mutant mice worked significantly less than did the wild-type mice for food reinforcers under nondeprived conditions. Controls for nonspecific effects on acquisition of conditioned learning, activity, satiety, and resistance to extinction revealed no genotype differences, supporting our interpretation that beta-endorphin selectively affects a motivational component of reward behavior under nondeprived conditions. Therefore, we propose that beta-endorphin may function in at least two primary modes to modulate feeding. In the appetitive phase, beta-endorphin release increases the incentive value of food as a primary reinforcer. In contrast, it appears that endogenous beta-endorphin may inhibit food consumption in parallel with melanocortins and that the orexigenic properties previously ascribed to it may actually be due to other classes of endogenous opioid peptides.

Adrenocorticotropic Hormone↗