Selene Cansino

Episodic memory decay along the adult lifespan: A review of behavioral and neurophysiological evidence

The ability to learn and remember new information declines along life. Empirical evidence reveals that this deficit occurs unevenly with different types of memory. Episodic memory, which is referred to as the ability to remember our own experiences in a determined temporal and spatial context, is especially vulnerable to aging. Episodic information can be retrieved with or without the context information that took place when the episodic event was encoded. There is agreement that, with advancing age, the source information related to an episodic event is more susceptible to be forgotten than the event; however, there is no consensus regarding the age at which this decline begins, the speed of source-memory decline along life or the possible changes, due to aging, in neurophysiological activity during encoding of source information that is subsequently correctly retrieved. In an attempt to answer the first two issues, a behavioral study with 552 subjects from 20 to 80 years of age was conducted, which provided evidence of the exact age at which source memory starts to decline and of the speed of this memory loss along life. To address the last question, event-related potentials were recorded while young and old adults encoded source information, to investigate whether older adults generate memory traces different from young adults during encoding.

Neuromagnetic fields reveal cortical plasticity when learning an auditory discrimination task.

Auditory evoked neuromagnetic fields of the primary and association auditory cortices were recorded while subjects learned to discriminate small differences in frequency and intensity between two consecutive tones. When discrimination was no better than chance, evoked field patterns across the scalp manifested no significant differences between correct and incorrect responses. However, when performance was correct on at least 75% of the trials, the spatial pattern of magnetic field differed significantly between correct and incorrect responses during the first 70 ms following the onset of the second tone. In this respect, the magnetic field pattern predicted when the subject would make an incorrect judgment more than 100 ms prior to indicating the judgment by a button press. One subject improved discrimination for much smaller differences between stimuli after 200 h of training. Evidence of cortical plasticity with improved discrimination is provided by an accompanying decrease of the relative magnetic field amplitude of the 100 ms response components in the primary and association auditory cortices.

Age-related changes in neural activity during source memory encoding in young, middle-aged and elderly adults

Source memory, the ability to remember contextual information present at the moment an event occurs, declines gradually during normal aging. The present study addressed whether source memory decline is related to changes in neural activity during encoding across age. Event-related potentials (ERPs) were recorded in three groups of 14 subjects each: young (21-26 years), middle-aged (50-55 years) and older adults (70-77 years). ERPs were recorded while the subjects performed a natural/artificial judgment on images of common objects that were presented randomly in one of the quadrants of the screen (encoding phase). At retrieval, old images mixed with new ones were presented at the center of the screen and the subjects judged whether each image was new or old and, if old, were asked to indicate at which position of the screen the image was presented in the encoding session. The neurophysiological activity recorded during encoding was segregated for the study items according to whether their context was correctly retrieved or not, so as to search for subsequent memory effects (SME). These effects, which consisted of larger amplitude for items subsequently attracting a correct source judgment than an incorrect one, were observed in the three groups, but their onset was delayed across the age groups. The amplitude of the SME was similar across age groups at the frontal and central electrode sites, but was manifested more at the posterior sites in middle-aged and older adults, suggesting that source memory decline may be related to less efficient encoding mechanisms.

Neurophysiology of Successful Encoding and Retrieval of Source Memory

Event-related potentials were recorded during encoding, to identify whether brain activity predicts subsequent retrieval of spatial source information, and during retrieval, to investigate the neural correlates of successful and unsuccessful spatial context recollection. The amplitude registered during encoding for study items that were later recognized and assigned a correct source judgment was more positive than the amplitude for recognized items given incorrect source judgments; this difference started 480 msec poststimulus, predominantly at central and anterior sites. During retrieval, the waveform was more positive from 250 to 1,600 msec poststimulus when the brain had retrieved episodic information successfully than when it had failed. These findings indicate that brain electrical activity recorded during the first presentation of an item within its context predicts the subsequent retrieval of the specific spatial context. During retrieval, brain activity differed quantitatively at anterior sites and qualitatively at posterior sites according to the accuracy of source memory.

Tonotopic cortical representation of periodic complex sounds

Most of the sounds that are biologically relevant are complex periodic sounds, i.e., they are made up of harmonics, whose frequencies are integer multiples of a fundamental frequency (Fo). The Fo of a complex sound can be varied by modifying its periodicity frequency; these variations are perceived as the pitch of the voice or as the note of a musical instrument. The center frequency (CF) of peaks occurring in the audio spectrum also carries information, which is essential, for instance, in vowel recognition. The aim of the present study was to establish whether the generators underlying the 100m are tonotopically organized based on the Fo or CF of complex sounds. Auditory evoked neuromagnetic fields were recorded with a whole‐head magnetoencephalography (MEG) system while 14 subjects listened to 9 different sounds (3 Fo × 3 CF) presented in random order. Equivalent current dipole (ECD) sources for the 100m component show an orderly progression along the y‐axis for both hemispheres, with higher CFs represented more medially. In the right hemisphere, sources for higher CFs were more posterior, while in the left hemisphere they were more inferior. ECD orientation also varied as a function of the sound CF. These results show that the spectral content CF of the complex sounds employed here predominates, at the latency of the 100m component, over a concurrent mapping of their periodic frequency Fo. The effect was observed both on dipole placement and dipole orientation. Hum. Brain Mapping 20:71–81, 2003.

Neural correlates of source memory retrieval in young, middle-aged and elderly adults

Event-related potentials (ERPs) were recorded in young (21-27 years old), middle-aged (50-57 years old) and older adults (70-77 years old) to determine whether the decline in source memory that occurs with advancing age coincides with contemporaneous neurophysiological changes. Source memory for the spatial location (quadrant on the screen) of images presented during encoding was examined. The images were shown in the center of the screen during the retrieval task. Retrieval success for source information was characterized by different scalp topographies at frontal electrode sites in young adults relative to middle-aged and older adults. The right frontal effect during unsuccessful retrieval attempts showed amplitude and latency differences across age groups and was related to the ability to discriminate between old and new images only in young adults. These results suggest that the neural correlates of the retrieval success and attempt were affected by age and these effects were present by middle-age.

fMRI subsequent source memory effects in young, middle-aged and old adults.

The ability to remember the spatial context in which our experiences occur declines linearly across the adult lifespan. However, little is known about whether this source memory decline is associated with neural activity changes. In the present study, functional magnetic resonance imaging (fMRI) scans were recorded in young, middle-aged and old adults to investigate brain activity variations across the adult lifespan during encoding of subsequent spatial source memory retrieval. Twelve healthy individuals of both sexes were enrolled in each age group. During encoding, participants performed natural/artificial judgment of images of common objects that were randomly presented in one of the quadrants of the screen. During retrieval, the images presented at encoding were randomly mixed with new ones and displayed at the center of the screen. Participants judged whether each image was new or old and, if an image was old, they were instructed to indicate in which quadrant the image was presented in the encoding session. The contrast between study items that were later recognized and assigned a correct source judgment with those whose sources were subsequently forgotten revealed that positive subsequent memory effects disappear by middle age in the left medial orbitofrontal gyrus and appear in the left superior occipital gyrus. This under-recruitment and over-recruitment brain activity was also present in old adults. The results allowed us to identify the specific brain regions that first fail to encode spatial information into an episodic representation during the adult lifespan.

What does the brain do while playing scrabble?: ERPs associated with a short–long-term memory task

Event-related brain potentials (ERPs) were recorded while subjects performed the scrabble paradigm, a cued recall task that demands retrieving semantic memory information from long-term memory since subjects are not exposed to a previous study phase. The task combines short- and long-term memory processes and consists of forming words from a set of letters presented in random order. Short-term memory was manipulated by varying the number of letters (three, four and five) presented to the subject, while semantic memory was examined by comparing correct trials with no response trials. Behavioral results reveal that the subjects performed the task serially, as denoted by a linear reaction time increment as the number of random letters in the set increased. Short-term memory procedures were reflected by an amplitude increase of the N200 and by an amplitude decrease of the P300 increasing the number of letters. Successfully retrieving semantic information from long-term memory was indexed by a negative slow wave recorded at left frontal and left central sites, and by a positive slow wave predominant over right hemisphere sites. These findings provide evidence that semantic retrieval memory involves activity from both left and right hemispheres.

Brain activity during source memory retrieval in young, middle-aged and old adults.

We investigated neurofunctional changes associated with source memory decline across the adult life span using functional magnetic resonance imaging (fMRI). Young, middle-aged and old adults carried out a natural/artificial judgment of images of common objects that were randomly presented in one of the quadrants of the screen. At retrieval, the images were displayed at the center of the screen and the participants judged whether each image was new or old and, if old, they indicated in which quadrant of the screen the image had originally been presented. Comparing the items associated with correct versus incorrect source judgments revealed that no regions showed greater activity in young adults than in middle-aged adults; however, in young and middle-aged adults the activity in the left hippocampus and left anterior temporal cortex was of greater magnitude than in the older adults. Several regions also exhibited greater activity in young adults than in old adults. These results suggest that in middle age the recollection neural network, assessable by fMRI, is still preserved.

The Rate of Source Memory Decline across the Adult Life Span.

Previous studies have suggested that the ability to remember contextual information related to specific episodic experiences declines with advancing age; however, the exact moment in the adult life span when this deficit begins is still controversial. Source memory for spatial information was tested in a life span sample of 1,500 adults between the ages of 21 and 80. Initially, images of common objects were randomly presented on one quadrant of a screen while the participants judged whether they were natural or artificial. During the retrieval phase, these same images were mixed with new ones, and all images were displayed in the center of the screen. The participants were asked to judge whether each image was new or old, and whether it was old, to indicate in which quadrant of the screen it had originally been presented. Source accuracy decreased linearly with advancing age at a rate of 0.6% per year across all decades even after controlling for educational level; this decline was unaffected by sex. These results reveal that either spatial information becomes less efficiently bound to episodic representations over time or that the ability to retrieve this information decreases gradually throughout the adult life span.