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Ken A Paller

Publications and source records attributed to Ken A Paller.

At least 19 recordsLinked to original sources

Attention induces synchronization-based response gain in steady-state visual evoked potentials.

When attention is voluntarily directed to a spatial location, visual sensitivity increases at that location. What causes this improved sensitivity? Studies of single neuron spike rates in monkeys have provided mixed results in regard to whether attending to a stimulus increases its effective contrast (contrast gain) or multiplicatively boosts stimulus-driven neural responses (response or activity gain). We monitored frequency-tagged steady-state visual evoked potentials (SSVEPs) in humans and found that voluntary sustained attention multiplicatively increased stimulus-driven population electrophysiological activity. Analyses of intertrial phase coherence showed that this attentional response gain was at least partially due to the increased synchronization of SSVEPs to stimulus flicker. These results suggest that attention operates in a complementary manner at different levels; attention seems to increase single-neuron spike rates in a variety of ways, including contrast, response and activity gains, while also inducing a multiplicative boost on neural population activity via enhanced response synchronization.

Adult↗

Distinguishing source memory and item memory: brain potentials at encoding and retrieval.

Vivid memory for an episode generally includes memory for a central object or event plus memory for background context or source information. To assess neural differences between source and item memory, we used event-related potentials (ERPs) to monitor relevant memory processes at both encoding and retrieval. Participants fluent in Chinese studied Chinese words superimposed on a square or circular background during the study phase, followed by a 1-min delay. Then, memory was tested for both the words (items) and the corresponding background (source), or, in other blocks, tested for the words alone. ERPs to study-phase words differed as a function of whether the word was later remembered. These Dm effects in the interval from 400 to 600 ms, however, did not differ according to whether or not source was remembered. In contrast, ERPs to test-phase words showed clear old/new effects that did differ across conditions. When both item and source were remembered accurately, old/new effects emerged earlier and were larger in amplitude than when source memory was either incorrect or not queried. These results demonstrate that encoding processes indexed by ERPs may have primarily reflected encoding of the visual and semantic properties of these words, stressing item memory over source memory. Retrieval processes indexed by ERPs, in contrast, likely reflected a combination of item retrieval, source retrieval, and related processing engaged when people were remembering words seen earlier.

Adolescent↗

Fluent conceptual processing and explicit memory for faces are electrophysiologically distinct.

Implicit memory and explicit memory are fundamentally different manifestations of memory storage in the brain. Yet, conceptual fluency driven by previous experience could theoretically be responsible for both conceptual implicit memory and aspects of explicit memory. For example, contemplating the meaning of a word might serve to speed subsequent processing of that word and also make it seem familiar. We examined electrophysiological correlates of conceptual priming with 180 celebrity faces to determine whether or not they resemble electrophysiological correlates of explicit memory. Celebrity faces are ideal for this purpose because they carry with them preexisting conceptual information (i.e., biographical facts) that can selectively be brought to mind such that conceptual processing can be manipulated systematically. In our experiment, exposure to biographical information associated with only one-half of the celebrities yielded conceptual priming for those faces, whereas all faces were perceptually primed. Conceptual priming was indexed by positive brain potentials over frontal regions from approximately 250 to 500 ms. Explicit memory retrieval was associated with later brain potentials over posterior regions that were strikingly similar to potentials previously associated with pure familiarity for faces (when a face seems familiar in the absence of retrieval of any specific information about previous occurrence). Furthermore, the magnitude of conceptual priming was correlated across subjects with the amplitude of frontal but not posterior potentials, whereas the opposite was true for explicit memory. Distinct brain processes were thus associated with conceptual priming and conscious recognition of faces, thus providing a sharper focus on the border between implicit and explicit memory.

Adolescent↗

When memory does not fail: familiarity-based recognition in mild cognitive impairment and Alzheimer's disease.

Recognition can be guided by familiarity, a restricted form of retrieval devoid of contextual recall, or by recollection, which occurs when retrieval is sufficient to support the full experience of remembering an episode. Recollection and familiarity were disentangled by testing recognition memory using silhouette object drawings, high target-foil resemblance, and both yes-no and forced-choice procedures. Theoretically, forced-choice recognition could be mediated by familiarity alone. Alzheimer's disease and its preclinical stage, mild cognitive impairment (MCI), were associated with memory impairments that were greater on the yes-no test. Remarkably, forced-choice recognition was unequivocally normal in patients with MCI compared with age-matched controls. Neuropathology in hippocampus and entorhinal cortex, known to be present in MCI, presumably disrupted recollection while leaving familiarity-based recognition intact.

Aged↗

Neural correlates of perceptual contributions to nondeclarative memory for faces.

Face priming is a nondeclarative memory phenomenon that can be observed when recognition is facilitated for a recently encountered face. This data-driven form of priming is distinct from conceptually driven priming. Moreover, it includes two dissociable components, the facilitated access to pre-existing representations and facilitation in perceptual processing of faces. In the present study, we measured neural correlates of perceptual contributions to face priming with event-related brain potentials. Faces appeared two times (separated by 7-17 s), while participants discriminated familiar from unfamiliar faces. Half of the initial face stimuli were inverted, thereby disrupting perceptual face processing and making possible an assessment of perceptual contributions to face priming. Whereas none of the brain waves previously linked to perceptual processing of faces showed indications of priming, such effects were observed between 200 and 600 ms at left occipito-parieto-temporal recording sites. This electrical activity was present for both unfamiliar and familiar faces. The scalp topography of this effect was consistent with sources within the temporal and occipital cortices of the left hemisphere (based on a LORETA source localization). These findings suggest that priming of perceptual face processing is subserved by prolonged neural activity from 200 to 600 ms primarily in the left hemisphere. We propose that this priming reflects facilitated selection based on second-order relations among facial features.

Adolescent↗

Dissociating perceptual and representation-based contributions to priming of face recognition.

Repetition priming of object identification refers to the phenomenon whereby experience with an object induces systematic changes in subsequent processing of that same object. This data-driven form of priming is distinct from conceptually-driven priming. To date, considerable controversy exists about whether data-driven priming reflects facilitation in perceptual processing or mediation by preexisting object representations. The present study concerned priming of recognizing familiar and unfamiliar faces and how this priming is influenced by face inversion, which interferes with perceptual face processing. Perceptual and representation-based loci conjointly contributed to priming; the perceptual locus was operative similarly for familiar and unfamiliar faces, whereas the representation-based locus was only invoked for familiar faces and resulted in a response-time reduction triple the magnitude of that from the perceptual locus. The results constrain theoretical accounts of data-driven priming by indicating that improved identification can result from the combination of perceptual and representation-based facilitation.

Adult↗

Electrophysiological correlates of forming memories for faces, names, and face-name associations.

The ability to put a name to a face is a vital aspect of human interaction, but many people find this extremely difficult, especially after being introduced to someone for the first time. Creating enduring associations between arbitrary stimuli in this manner is also a prime example of what patients with amnesia find most difficult. To help develop a better understanding of this type of memory, we sought to obtain measures of the neural events responsible for successfully forming a new face-name association. We used event-related potentials (ERPs) extracted from high-density scalp EEG recordings in order to compare (1) memory for faces, (2) memory for names, and (3) memory for face-name associations. Each visual face appeared simultaneously with a unique spoken name. Signals observed 200-800 ms after the onset of face-name pairs predicted subsequent memory for faces, names, or face-name associations. Difference potentials observed as a function of subsequent memory performance were not identical for these three memory tests, nor were potentials predicting associative memory equivalent to the sum of potentials predicting item memory, suggesting that different neural events at the time of encoding are relevant for these distinct aspects of remembering people.

Adolescent↗

An electrophysiological investigation of memory encoding, depth of processing, and word frequency in humans.

Memory encoding can be studied by monitoring brain activity correlated with subsequent remembering. To understand brain potentials associated with encoding, we compared multiple factors known to affect encoding. Depth of processing was manipulated by requiring subjects to detect animal names (deep encoding) or boldface (shallow encoding) in a series of Chinese words. Recognition was more accurate with deep than shallow encoding, and for low- compared to high-frequency words. Potentials were generally more positive for subsequently recognized versus forgotten words; for deep compared to shallow processing; and, for remembered words only, for low- than for high-frequency words. Latency and topographic differences between these potentials suggested that several factors influence the effectiveness of encoding and can be distinguished using these methods, even with Chinese logographic symbols.

Adult↗

The neural basis of the butcher-on-the-bus phenomenon: when a face seems familiar but is not remembered.

A common distinction in contemporary research on episodic memory is between familiarity, an unsubstantiated impression that an event was experienced previously, and recollection, remembering some information plus the spatiotemporal context of the episode in which it was acquired. The epitome of pure familiarity--the butcher-on-the-bus phenomenon--occurs when one believes that a person is familiar (often upon seeing their face in an atypical context) while failing to recall any information about that person whatsoever. Prior research on familiarity and recollection has relied on verbal material. Whereas word meanings and pronunciations are well learned in advance, here we produced pure familiarity and recollection using photographs of faces never seen before the experiment. When participants recognized a face, recollection was inferred if they also remembered either the occupation associated with that face earlier in the experiment or any other episodic detail. Pure familiarity was inferred when recognition occurred in the absence of any such contextual retrieval. Analyses of brain potentials recorded during initial encoding showed that right-sided neural activity predicted subsequent face familiarity, whereas bilateral potentials predicted subsequent face recollection. Results during memory testing were inconsistent with the popular idea that familiarity is generically indexed by reduced frontal N400-like potentials. Instead, both memory experiences were associated with bilateral, parietal-maximum brain potentials, although with smaller amplitudes and for a shorter duration for familiarity. These similarities between electrophysiological correlates of pure familiarity and recollection suggest that familiarity with faces may arise by virtue of a subset of the neural processing responsible for recollection.

Adolescent↗

Neural evidence that vivid imagining can lead to false remembering.

The imperfect nature of memory is highlighted by the regularity with which people fail to remember, or worse, remember something that never happened. We investigated the formation of a particular type of erroneous memory by monitoring brain activity using functional magnetic resonance imaging during the presentation of words and photos. Participants generated a visual image of a common object in response to each word. Subsequently, they sometimes claimed to have seen photos of specific objects they had imagined but not actually seen. In precuneus and inferior parietal regions of the cerebral cortex, activations in response to words were greater when participants subsequently claimed to have seen the corresponding object than when a false memory for that object was not subsequently produced. These findings indicate that brain activity reflecting the engagement of visual imagery can lead to falsely remembering something that was only imagined.

Adolescent↗

Neural manifestations of memory with and without awareness.

Neurophysiological events responsible for different types of human memory tend to occur concurrently and are therefore difficult to measure independently. To surmount this problem, we produced perceptual priming (indicated by speeded responses) in the absence of conscious remembering. At encoding, faces appeared briefly while subjects' attention was diverted to other stimuli. Faces appeared again in either an implicit or explicit memory test. Neural correlates of priming were identified as brain potentials beginning 270 ms after face onset with more negative amplitudes for repeated than for new faces. Remembered faces, in contrast, activated a different configuration of intracranial sources producing positive potentials maximal at 600-700 ms. We thus disentangled and characterized distinct neural events associated with memory with and without awareness.

Adolescent↗

Neural correlates of the left-visual-field superiority in face perception appear at multiple stages of face processing.

Studies in healthy individuals and split-brain patients have shown that the representation of facial information from the left visual field (LVF) is better than the representation of facial information from the right visual field (RVF). To investigate the neurophysiological basis of this LVF superiority in face perception, we recorded event-related potentials (ERPs) to centrally presented face stimuli in which relevant facial information is present bilaterally (B faces) or only in the left (L faces) or the right (R faces) visual field. Behavioral findings showed best performance for B faces and, in line with the LVF superiority, better performance for L than R faces. Evoked potentials to B, L, and R faces at 100- to 150-msec poststimulus showed no evidence of asymmetric transfer of information between the hemispheres at early stages of visual processing, suggesting that this factor is not responsible for the LVF superiority. Neural correlates of the LVF superiority, however, were manifested in a shorter latency of the face-specific N170 component to L than R faces and in a larger amplitude to L than R faces at 220-280 and 400-600 msec over both hemispheres. These ERP amplitude differences between L and R faces covaried across subjects with the extent to which the face-specific N170 component was larger over the right than the left hemisphere. We conclude that the two hemispheres exchange information symmetrically at early stages of face processing and together generate a shared facial representation, which is better when facial information is directly presented to the right hemisphere (RH; L faces) than to the left hemisphere (LH; R faces) and best when both hemispheres receive facial information (B faces).

Adult↗

Brain networks for analyzing eye gaze.

The eyes convey a wealth of information in social interactions. This information is analyzed by multiple brain networks, which we identified using functional magnetic resonance imaging (MRI). Subjects attempted to detect a particular directional cue provided either by gaze changes on an image of a face or by an arrow presented alone or by an arrow superimposed on the face. Another control condition was included in which the eyes moved without providing meaningful directional information. Activation of the superior temporal sulcus accompanied extracting directional information from gaze relative to directional information from an arrow and relative to eye motion without relevant directional information. Such selectivity for gaze processing was not observed in face-responsive fusiform regions. Brain activations were also investigated while subjects viewed the same face but attempted to detect when the eyes gazed directly at them. Most notably, amygdala activation was greater during periods when direct gaze never occurred than during periods when direct gaze occurred on 40% of the trials. In summary, our results suggest that increases in neural processing in the amygdala facilitate the analysis of gaze cues when a person is actively monitoring for emotional gaze events, whereas increases in neural processing in the superior temporal sulcus support the analysis of gaze cues that provide socially meaningful spatial information.

Adult↗

Neural correlates of successful encoding identified using functional magnetic resonance imaging.

Neural activity that occurs during the creation of a new memory trace can be observed using functional magnetic resonance imaging (fMRI). Event-related designs have been used to demonstrate that activity in prefrontal and medial temporal lobe areas is associated with successful memory storage. Here we contrasted activity associated with encoding success and encoding effort. Participants viewed a series of 150 words but attempted to remember only half of them. Encoding effort was manipulated using a cue in the form of a letter (R or F) presented after each word to instruct participants either to remember or to forget that word. Increased activity in left inferior prefrontal cortex was observed when words were followed by the cue to remember. In contrast, increased left medial temporal lobe activity was observed for words that were successfully recalled later. These results show that fMRI correlates of the intention to encode a word are different from fMRI correlates of whether that encoding is successful. Prefrontal activation was strongly associated with intentional verbal encoding, whereas left medial temporal activation was crucial for the encoding that actually led to successful memory on the subsequent test.

Adolescent↗

Observing the transformation of experience into memory.

The ability to remember one's past depends on neural processing set in motion at the moment each event is experienced. Memory formation can be observed by segregating neural responses according to whether or not each event is recalled or recognized on a subsequent memory test. Subsequent memory analyses have been performed with various neural measures, including brain potentials extracted from intracranial and extracranial electroencephalographic recordings, and hemodynamic responses from functional magnetic resonance imaging. Neural responses can predict which events, and which aspects of those events, will be subsequently remembered or forgotten, thereby elucidating the neurocognitive processes that establish durable episodic memories.

Journal Article↗

Field potentials in the human hippocampus during the encoding and recognition of visual stimuli.

Intracranial field potentials were recorded from electrodes implanted in the hippocampus in 12 epileptic patients. Potentials were elicited by stimuli presented during a delayed matching-to-sample test. Each trial began with a sample stimulus composed of a 3 x 3 grid of rectangular color patches. The sample was followed by a sequence of similar but task-irrelevant stimuli and the sequential presentation of two test stimuli, one of which was identical to the sample. Patients indicated their recognition of the test stimulus that matched the sample with a button press. High-amplitude negative potentials were consistently elicited by sample and test stimuli. Peak amplitudes occurred 300-500 ms after stimulus onset and were larger for the sample in all cases. The patterns of potential gradients observed between adjacent hippocampal contacts and the locations of maximal amplitudes, as verified by magnetic resonance imaging in seven patients, suggest that these potentials were produced by neuronal activity in posterior hippocampus. These field potentials appear to index a memory storage function engaged in response to events that will later be remembered. The hippocampal contribution to storing declarative memories can thus begin, in some circumstances, within the first half-second after the presentation of a to-be-remembered stimulus.

Action Potentials↗

Mistaken memories: remembering events that never happened.

Our memories can be accurate, but they are not always accurate. Eyewitness testimony, for example, is notoriously unreliable. Insights into both veridical and false remembering have come from recent investigations of memory distortion. Behavioral measures have been used to demonstrate false memory phenomena in the laboratory, and neuroimaging measures have been used to provide clues about the relevant events in the brain that support remembering versus misremembering. A central category of misremembering results from confusion between memories for perceived and imagined events, which may result from overlap between particular features of the stored information comprising memories for perceived and imagined events.

Association↗

Neural correlates of person recognition.

Rapidly identifying known individuals is an essential skill in human society. To elucidate the neural basis of this skill, we monitored brain activity while experimental participants demonstrated their ability to recognize people on the basis of viewing their faces. Each participant first memorized the faces of 20 individuals who were not known to the participants in advance. Each face was presented along with a voice simulating the individual speaking their name and a biographical fact. Following this learning procedure, the associated verbal information could be recalled accurately in response to each face. These learned faces were subsequently viewed together with new faces in a memory task. Subjects made a yes-no recognition decision in response to each face while also covertly retrieving the person-specific information associated with each learned face. Brain activity that accompanied this retrieval of person-specific information was contrasted to that when new faces were processed. Functional magnetic resonance imaging in 10 participants showed that several brain regions were activated during blocks of learned faces, including left hippocampus, left middle temporal gyrus, left insula, and bilateral cerebellum. Recordings of event-related brain potentials in 10 other participants tracked the time course of face processing and showed that learned faces engaged neural activity responsible for person recognition 300-600 msec after face onset. Collectively, these results suggest that the visual input of a recently learned face can rapidly trigger retrieval of associated person-specific information through reactivation of distributed cortical networks linked via hippocampal connections.

Adult↗