episodic memory

What transient amnesia tells us about autobiographical memory and brain plasticity

November, 2011
  • Brain scans of those suffering from transient global amnesia indicate a permanent role of the hippocampus in autobiographical memory, and demonstrate the brain’s ability to self-repair.

When a middle-aged woman loses her memory after sex, it naturally makes the headlines. Many might equate this sort of headline to “Man marries alien”, but this is an example of a rare condition — temporary, you will be relieved to hear — known as transient global amnesia. Such abrupt, localized loss of autobiographical memory is usually preceded by strenuous physical activity or stressful events. It generally occurs in middle-aged or older adults, but has been known to occur in younger people. In those cases, there may be a history of migraine or head trauma.

Following an earlier study in which 29 of 41 TGA patients were found to have small lesions in the CA1 region of the hippocampus, scanning of another 16 TGA patients has revealed 14 had these same lesions. It seems likely that all the patients had such lesions, but because they are very small and don’t last long, they’re easy to miss. The lesion is best seen after 24-72 hours, but is gone after 5-6 days.

At the start of one of these attacks, memory for the first 30 years of life was significantly impaired, but still much better than memory for the years after that. There was a clear temporal gradient, with memory increasingly worse for events closer in time. There was no difference between events in the previous year and events in the previous five years, but a clear jump at that five-year point.

The exact location of the lesions was significant: when the lesion was in the anterior part of the region, memory for recent events was more impaired.

The hippocampus is known to be crucially involved in episodic memory (memory for events), and an integral part of the network for autobiographical memory. In recent years, it has come to be thought that such memories are only hosted temporarily by the hippocampus, and over a few years come to be permanently lodged in the neocortex (the standard consolidation model). Evidence from a number of studies of this change at the five-year mark has been taken as support for this theory. According to this, then, older memories should be safe from hippocampal damage.

An opposing theory, however, is that the hippocampus continues to be involved in such memories, with both the neocortex and the hippocampus involved in putting together reconsolidated memories (the multiple trace model). According to this model, each retrieval of an episodic memory creates a new version in the hippocampus. The more versions, the better protected a memory will be from any damage to the hippocampus.

The findings from this study show that while there is indeed a significant difference between older and more recent memories, the CA1 region of the hippocampus continues to be crucial for retrieving older memories, and for our sense of self-continuity.

Interestingly, some studies have also found a difference between the left and right hemispheres, with the right hippocampus showing a temporal gradient and the left hippocampus showing constant activation across all time periods. Such a hemisphere difference was not found in the present study. The researchers suggest that the reason may lie in the age of the participants (average age was 68), reflecting a reduction in hemispheric asymmetry with age.

There’s another message in this study. In these cases of TGA, memory function is restored within 24 hours (and generally sooner, within 6-10 hours). This shows how fast the brain can repair damage. Similarly, the fact that such tiny lesions have temporary effects so much more dramatic than the more lasting effects of larger lesions, is also a tribute to the plasticity of the brain.

The findings are consistent with findings of a preferential degeneration of CA1 neurons in the early stages of Alzheimer's disease, and suggest a target for treatment.

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Errorless learning not always best for older brains

October, 2011

New evidence challenges the view that older adults learn best through errorless learning. Trial-and-error learning can be better if done the right way.

Following a 1994 study that found that errorless learning was better than trial-and-error learning for amnesic patients and older adults, errorless learning has been widely adopted in the rehabilitation industry. Errorless learning involves being told the answer without repeatedly trying to answer the question and perhaps making mistakes. For example, in the 1994 study, participants in the trial-and-error condition could produce up to three errors in answer to the question “I am thinking of a word that begins with QU”, before being told the answer was QUOTE; in contrast, participants in the errorless condition were simply told “I am thinking of a word that begins with QU and it is ‘QUOTE’.”

In a way, it is surprising that errorless learning should be better, given that trial-and-error produces much deeper and richer encoding, and a number of studies with young adults have indeed found an advantage for making errors. Moreover, it’s well established that retrieving an item leads to better learning than passively studying it, even when you retrieve the wrong item. This testing effect has also been found in older adults.

In another way, the finding is not surprising at all, because clearly the trial-and-error condition offers many opportunities for confusion. You remember that QUEEN was mentioned, for example, but you don’t remember whether it was a right or wrong answer. Source memory, as I’ve often mentioned, is particularly affected by age.

So there are good theoretical reasons for both positions regarding the value of mistakes, and there’s experimental evidence for both. Clearly it’s a matter of circumstance. One possible factor influencing the benefit or otherwise of error concerns the type of processing. Those studies that have found a benefit have generally involved conceptual associations (e.g. What’s Canada’s capital? Toronto? No, Ottawa). It may be that errors are helpful to the extent that they act as retrieval cues, and evoke a network of related concepts. Those studies that have found errors harm learning have generally involved perceptual associations, such as word stems and word fragments (e.g., QU? QUeen? No, QUote). These errors are arbitrary, produce interference, and don’t provide useful retrieval cues.

So this new study tested the idea that producing errors conceptually associated with targets would boost memory for the encoding context in which information was studied, especially for older adults who do not spontaneously elaborate on targets at encoding.

In the first experiment, 33 young (average age 21) and 31 older adults (average age 72) were shown 90 nouns presented in three different, intermixed conditions. In the read condition (designed to provide a baseline), participants read aloud the noun fragment presented without a semantic category (e.g., p­_g). In the errorless condition, the semantic category was presented with the target word fragment (e.g. a farm animal  p­_g), and the participants read aloud the category and their answer. The category and target were then displayed. In the trial-and-error condition, the category was presented and participants were encouraged to make two guesses before being shown the target fragment together with the category. The researchers changed the target if it was guessed. Participants were then tested using a list of 70 words, of which 10 came from each of the study conditions, 10 were new unrelated words, and 30 were nontarget exemplars from the TEL categories. Those that the subject had guessed were labeled as learning errors; those that hadn’t come up were labeled as related lures. In addition to an overall recognition test (press “yes” to any word you’ve studied and “no” to any new word), there were two tests that required participants to endorse items that were studied in the TEL condition and reject those studied in the EL condition, and vice versa.

The young adults did better than the older on every test. TEL produced better learning than EL, and both produced better learning than the read condition (as expected). The benefit of TEL was greater for older adults. This is in keeping with the idea that generating exemplars of a semantic category, as occurs in trial-and-error learning, helps produce a richer, more elaborated code, and that this is of greater to older adults, who are less inclined to do this without encouragement.

There was a downside, however. Older adults were also more prone to falsely endorsing prior learning errors or semantically-related lures. It’s worth noting that both groups were more likely to falsely endorse learning errors than related lures.

But the main goal of this first experiment was to disentangle the contributions of recollection and familiarity to the two types of learning. It turns out that there was no difference between young and older adults in terms of familiarity; the difference in performance between the two groups stemmed from recollection. Recollection was a problem for older adults in the errorless condition, but not in the trial-and-error condition (where the recollective component of their performance matched that of young adults). This deficit is clearly closely related to age-related deficits in source memory.

It was also found that familiarity was marginally more important in the errorless condition than the trial-and-error condition. This is consistent with the idea that targets learned without errors acquire greater fluency than those learned with errors (with the downside that they don’t pick up those contextual details that making errors can provide).

In the second experiment, 15 young and 15 older adults carried out much the same procedure, except that during the recognition test they were also required to mention the context in which the words were learned was tested (that is, were the words learned through trial-and-error or not).

Once again, trial-and-error learning was associated with better source memory relative to errorless learning, particularly for the older adults.

These results support the hypothesis that trial-and-error learning is more beneficial than errorless learning for older adults when the trials encourage semantic elaboration. But another factor may also be involved. Unlike other errorless studies, participants were required to attend to errors as well as targets. Explicit attention to errors may help protect against interference.

In a similar way, a recent study involving young adults found that feedback given in increments (thus producing errors) is more effective than feedback given all at once in full. Clearly what we want is to find that balance point, where elaborative benefits are maximized and interference is minimized.

Reference: 

[2496] Cyr, A-A., & Anderson N. D.
(2011).  Trial-and-error learning improves source memory among young and older adults.
Psychology and Aging. No Pagination Specified - No Pagination Specified.

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Negative emotion can enhance memory for tested information

September, 2011

Images designed to arouse strong negative emotion can improve your memory for information you’re learning, if presented immediately after you’ve been tested on it.

In a recent study, 40 undergraduate students learned ten lists of ten pairs of Swahili-English words, with tests after each set of ten. On these tests, each correct answer was followed by an image, either a neutral one or one designed to arouse negative emotions, or by a blank screen. They then did a one-minute multiplication test before moving on to the next section.

On the final test of all 100 Swahili-English pairs, participants did best on items that had been followed by the negative pictures.

In a follow-up experiment, students were shown the images two seconds after successful retrieval. The results were the same.

In the final experiment, the section tests were replaced by a restudying period, where each presentation of a pair was followed by an image or blank screen. The effect did not occur, demonstrating that the effect depends on retrieval.

The study focused on negative emotion because earlier research has found no such memory benefit for positive images (including images designed to be sexually arousing).

The findings emphasize the importance of the immediate period after retrieval, suggesting that this is a fruitful time for manipulations that enhance or impair memory. This is consistent with the idea of reconsolidation — that when information is retrieved from memory, it is in a labile state, able to be changed. Thus, by presenting a negative image when the retrieved memory is still in that state, the memory absorbs some of that new context.

Reference: 

[2340] Finn, B., & Roediger H. L.
(2011).  Enhancing Retention Through Reconsolidation.
Psychological Science. 22(6), 781 - 786.

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Sleep deprivation eliminates fear generalization

January, 2011

In a small study, a sleepless night after trauma prevents the development of PTSD symptoms.

Given all the research showing the importance of sleep for consolidating memories, it should come as no great surprise that the reverse is also true: depriving yourself of sleep could help you forget experiences you would prefer not to remember.

In the study, 28 student volunteers were shown 14 short video clips, half of which showed safe driving down a city street, and half showed the car being involved in a nasty crash. Half of the volunteers were then deprived of sleep while the other half received a normal night's sleep. The next day, they were shown pictures and asked to indicate whether they had appeared in the clips they had seen. They were also asked to rate the fear evoked by the image, and their physiological responses measured. They were tested again 3 and 10 days later.

While there was no difference between the two groups in picture recognition, the control group rated the images from the crash videos as fearful, and these responses generalized over time to the other images. However, those who were sleep deprived showed such reactions only on the first day.

The finding suggests a possible therapy for PTSD or other anxiety disorders.

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Drug prevents post-traumatic stress syndrome

January, 2011

A new drug is successful in preventing PTSD in mice if delivered within 5 hours of the trauma.

A mouse study has revealed the brain becomes overly stimulated after a traumatic event causes an ongoing, frenzied interaction between two brain proteins long after they should have disengaged. However, the injection of newly developed drugs into the hippocampus within a five hour window calmed this process, and prevented the development of a post-traumatic fear response.

The new research shows the potential for PTSD occurs when a stressful event causes a flood of glutamate, which then interacts with a second protein (Homer1a). This protein continues to stimulate metabotropic glutamate receptor 5 [mGluR5] after the glutamate has dissipated. The new drugs bind mGluR5 and reverse its activity.

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Sleep reorganizes your memories

December, 2010

New studies show how sleep sculpts your memories, emphasizing what’s important and connecting it to other memories in your brain.

The role of sleep in consolidating memory is now well-established, but recent research suggests that sleep also reorganizes memories, picking out the emotional details and reconfiguring the memories to help you produce new and creative ideas. In an experiment in which participants were shown scenes of negative or neutral objects at either 9am or 9pm and tested 12 hours later, those tested on the same day tended to forget the negative scenes entirely, while those who had a night’s sleep tended to remember the negative objects but not their neutral backgrounds.

Follow-up experiments showed the same selective consolidation of emotional elements to a lesser degree after a 90-minute daytime nap, and to a greater degree after a 24-hour or even several-month delay (as long as sleep directly followed encoding).

These findings suggest that processes that occur during sleep increase the likelihood that our emotional responses to experiences will become central to our memories of them. Moreover, additional nights of sleep may continue to modify the memory.

In a different approach, another recent study has found that when volunteers were taught new words in the evening, then tested immediately, before spending the night in the sleep lab and being retested in the morning, they could remember more words in the morning than they did immediately after learning them, and they could recognize them faster. In comparison, a control group who were trained in the morning and re-tested in the evening showed no such improvement on the second test.

Deep sleep (slow-wave sleep) rather than rapid eye movement (REM) sleep or light sleep appeared to be the important phase for strengthening the new memories. Moreover, those who experienced more sleep spindles overnight were more successful in connecting the new words to the rest of the words in their mental lexicon, suggesting that the new words were communicated from the hippocampus to the neocortex during sleep. Sleep spindles are brief but intense bursts of brain activity that reflect information transfer between the hippocampus and the neocortex.

The findings confirm the role of sleep in reorganizing new memories, and demonstrate the importance of spindle activity in the process.

Taken together, these studies point to sleep being more important to memory than has been thought. The past decade has seen a wealth of studies establishing the role of sleep in consolidating procedural (skill) memory, but these findings demonstrate a deeper, wider, and more ongoing process. The findings also emphasize the malleability of memory, and the extent to which they are constructed (not copied) and reconstructed.

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Friends, family detect early Alzheimer's signs better than traditional tests

October, 2010

Cognitive tests only test you at a particular moment in time; early signs of Alzheimer's are more evident in declines in everyday behavior that are most visible to other people.

Confirming earlier research, a study involving 257 older adults (average age 75) has found that a two-minute questionnaire filled out by a close friend or family member is more accurate that standard cognitive tests in detecting early signs of Alzheimer’s.

The AD8 asks questions about changes in everyday activities:

  • Problems with judgment, such as bad financial decisions;
  • Reduced interest in hobbies and other activities;
  • Repeating of questions, stories or statements;
  • Trouble learning how to use a tool or appliance, such as a television remote control or a microwave;
  • Forgetting the month or year;
  • Difficulty handling complicated financial affairs, such as balancing a checkbook;
  • Difficulty remembering appointments; and
  • Consistent problems with thinking and memory.

Problems with two or more of these are grounds for further evaluation. The study found those with AD8 scores of 2 or more were very significantly more likely to have early biomarkers of Alzheimer’s (abnormal Pittsburgh compound B binding and cerebrospinal fluid biomarkers), and was better at detecting early stages of dementia than the MMSE. The AD8 has now been validated in several languages and is used in clinics around the world.

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New advice on how much cognitive abilities decline with age

October, 2010

A new study suggests that inconsistencies in rate of age-related cognitive decline may be partly due to practice effects, but though decline does occur it is slower than some have estimated.

Reports on cognitive decline with age have, over the years, come out with two general findings: older adults do significantly worse than younger adults; older adults are just as good as younger adults. Part of the problem is that there are two different approaches to studying this, each with their own specific bias. You can keep testing the same group of people as they get older — the problem with this is that they get more and more practiced, which mitigates the effects of age. Or you can test different groups of people, comparing older with younger — but cohort differences (e.g., educational background) may disadvantage the older generations. There is also argument about when it starts. Some studies suggest we start declining in our 20s, others in our 60s.

One of my favorite cognitive aging researchers has now tried to find the true story using data from the Virginia Cognitive Aging Project involving nearly 3800 adults aged 18 to 97 tested on reasoning, spatial visualization, episodic memory, perceptual speed and vocabulary, with 1616 tested at least twice. This gave a nice pool for both cross-sectional and longitudinal comparison (retesting ranged from 1 to 8 years and averaged 2.5 years).

From this data, Salthouse has estimated the size of practice effects and found them to be as large as or larger than the annual cross-sectional differences, although they varied depending on the task and the participant’s age. In general the practice effect was greater for younger adults, possibly because younger people learn better.

Once the practice-related "bonus points" were removed, age trends were flattened, with much less positive changes occurring at younger ages, and slightly less negative changes occurring at older ages. This suggests that change in cognitive ability over an adult lifetime (ignoring the effects of experience) is smaller than we thought.

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Change in our understanding of memory development

September, 2010

Children’s slowly developing memory for past events may not be due to the slow development of the prefrontal cortex, as was thought, but to changes in the hippocampus.

Children’s ability to remember past events improves as they get older. This has been thought by many to be due to the slow development of the prefrontal cortex. But now brain scans from 60 children (8-year-olds, 10- to 11-year-olds, and 14-year-olds) and 20 young adults have revealed marked developmental differences in the activity of the mediotemporal lobe.

The study involved the participants looking at a series of pictures (while in the scanner), and answering a different question about the image, depending on whether it was drawn in red or green ink. Later they were shown the pictures again, in black ink and mixed with new ones. They were asked whether they had seen them before and whether they had been red or green.

While the adolescents and adults selectively engaged regions of the hippocampus and posterior parahippocampal gyrus to recall event details, the younger children did not, with the 8-year-olds indiscriminately using these regions for both detail recollection and item recognition, and the 10- to 11-year-olds showing inconsistent activation. It seems that the hippocampus and posterior parahippocampal gyrus become increasingly specialized for remembering events, and these changes may partly account for long-term memory improvements during childhood.

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The problem of 'destination amnesia'

September, 2010

Two studies demonstrate why knowing whether you’ve told someone something is difficult for all of us, and that this is particularly so as you get older.

A number of studies have found that source memory (knowing where you heard/read/experienced something) is a particular problem for older adults. Destination memory (knowing who you’ve told) is an area that has been much less studied. Last year I reported on why destination memory is difficult for all of us (my report is repeated below). A follow-up study has found not only that destination memory is a particular problem for older adults, but that it is in fact a worse problem than source memory. Moreover, destination amnesia (falsely believing you've told someone something) is not only more common among older adults, but is associated with greater confidence in the false belief.

The study compared the performance of 40 students (aged 18-30) and 40 healthy older adults (aged 60-83). In the first task, the participant read out loud 50 interesting facts to 50 celebrities (whose faces appeared on a computer screen), and were then tested on their memory of which fact they told to which famous person. In the second task, they had to remember which famous person told them which particular fact. Older adults' performance was 21% worse than their younger counterparts on the destination memory test, but only 10% worse (50% vs 60%) on the source memory test. This latter difference was not statistically significant.

The 2009 study, involving 60 students, found good reason for destination memory to be so poor — apparently outgoing information is less integrated with context than incoming information is. In the study, 50 random facts were linked with the faces of 50 famous people; half the students then “told” each fact to one of the faces, reading it aloud to the celebrity’s picture. The other half read each fact silently and saw a different celebrity moments afterward. In the subsequent memory test, students who simulated telling the facts did 16% worse. In another experiment using personal facts, it was significantly worse.

However, the final experiment found that you could improve your destination memory by saying the name of the person you’re speaking to, as you tell them. The findings also suggest that self-focus is an important factor: increasing self-focus (e.g. by telling a personal story) worsened destination memory; reducing self-focus (e.g. by naming the listener) improved it.

Reference: 

[1809] Gopie, N., Craik F. I. M., & Hasher L.
(2010).  Destination memory impairment in older people..
Psychology and Aging.

[396] Gopie, N., & MacLeod C. M.
(2009).  Destination Memory: Stop Me if I've Told You This Before.
Psychological Science. 20(12), 1492 - 1499.

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