"Paradigms and brain imaging technologies art becoming progressively more exacting, allowing cognitive scientists to investigate the ... neural circuitry responsible for abstract and uniquely human traits." R E M I N I S C I N G A N D D A Y D R E A M I N G Investigating the Similarities Between Episodic Memory Recall and Imagination K Y L E F I S C H E R T H E P O W E R O F I M A G I N A T I O N H A S L O N G B E E N T O U T E D A S A D I S T I N C T I V E P R O P E R T Y O F H U M A N C O G N I T I O N A N D R E M A I N S A H E A T E D T O P I C A M O N G S T P H I L O S O P H E R S , S C I E N T I S T S , A N D D A Y D R E A M E R S A L I K E . D E S P I T E P L A Y I N G A C R I T I C A L R O L E I N T H E M A J O R I T Y O F H U M A N I T Y ' S F I N E S T A C H I E V E M E N T S , L I T T L E IS K N O W N A B O U T H O W T H I S F A C U L T Y A C T U A L L Y W O R K S . R E C E N T F I N D I N G S F R O M L E S I O N P A T I E N T S P O I N T T O T H E I N V O L V E M E N T O F T H E H I P P O C A M P U S IN I M A G I N I N G N E W S C E N A R I O S . T H E H I P P O C A M P U S IS H I S T O R I C A L L Y A S S O C I A T E D W I T H L O N G - T E R M M E M O R Y C O N S O L I D A T I O N , A N D M E M O R Y M O D E L S A R I S I N G F R O M N E W F I N D I N G S IN T H E M O L E C U L A R P A T H W A Y S O F M E M O R Y A L L O W F O R A M O R E F L U I D T R A N S I T I O N B E T W E E N I M A G I N A T I O N A N D M E M O R Y . E L I Z A B E T H L O F T U S ' F A M O U S S T U D I E S O F F A L S E E P I S O D I C M E M O R Y R E C O V E R Y F U R T H E R D E M O N S T R A T E O V E R L A P B E T W E E N I M A G I N E D A N D R E C A L L E D E V E N T S . F I N A L L Y , A N F M R I S T U D Y I S P R O P O S E D T O C O M P A R E B R A I N A C T I V I T Y D U R I N G B O T H E P I S O D I C M E M O R Y R E C A L L A N D T A S K S T H A T R E Q U I R E I M A G I N A T I O N IN O R D E R T O P I N P O I N T R E G I O N S F O R F U R T H E R I N V E S T I G A T I O N . As the field of neuroscience expands, it is increasingly en- tering uncharted territory; the pieces of human experience that were once major arguments for Cartesian dualism, in- cluding morality, foresight, and theory of mind, are quickly finding neural homes. Paradigms and brain imaging tech- nologies are becoming progressively more exacting, allow- ing cognitive scientists to investigate the brain regions and neural circuitry responsible for abstract and uniquely human traits. One of these traits is imagination, the ability to produce vividly detailed episodic events. Recent studies of hippocampally-lesioned patients have shown that these injuries result in the inability to formulate new imaginative experiences,1 suggesting that these regions, classically linked with memory, are also necessary for imagination. New findings i n memory research have offered a modified memory model 1 1 suggesting a fluid continuum between real episodic memories and episodic imaginative experi- ences. The error-prone nature of episodic memories as evidenced by false memory experiments 1 1 1 supports the theory of a synonymous relationship between what is currently conceived as episodic memory and imagination. I n light of this evidence, we wi l l discuss a possible experi- mental paradigm that wi l l investigate similarities i n brain activity induced by episodic memory recall and the produc- tion of an imaginative narrative. IMAGIN ING A DRAGON IS A PROCESS WH ICH LIKELY OCCURS IN T H E H IPPOCAMPUS. Episodic memory is a uniquely human trait that expresses information i n a detailed narrative fashion, coding for who, what, where, when, and why . l v v When conventional speech refers to 'memories', it is most likely referring to episodic memories. For example, recall a childhood memory, per- haps your first baseball game. You may remember step- ping up to home plate, hearing your mom and dad cheer, and striking out in humiliation. Perhaps you may remem- ber the smell of the grass, the warm summer light, or the taste of the snacks back in the dug out. Now imagine a com- pletely fanciful event, perhaps a dragon attacking a ship full of Vikings. Again, you can hear the sea, taste the salt water, and feel the heat of the dragon's breath. As a final exercise, try integrating the two; while stepping up to the plate in your first baseball game, have a dragon stroll out into left field. Ridiculous, perhaps, but this brief exercise illustrates that as far as the nature of the mental experience is concerned, imagination and episodic memory seem, i f only anecdo- tally, identical. Although some nonhuman animals have displayed episodic-like memories,V 1 Homo sapiens are the only known species to possess this ability.V 1 1 As one might surmise from the previous exercise, classical false memory studies have been able to modify, falsify, and even produce new fictitious memories of fantastic proportions. One of the more classical studies, by Loftus, v m asked participants to recall a series of childhood memories that were given to the researchers by the participants' parents. During these sessions, the investigators would also ask the participants about having been lost i n the mall as a child, which the par- ents of the participants confirmed had never happened to their children. Nonetheless the participants recalled confi- dently the fictitious event, even creating precise details. 1 X The suggested nature of episodic memory supports the fluid model, which integrates imagination and episodic memory, but an understanding of the underlying mecha- nism of memory production is necessary to further investi- gate this theory. Memory is comprised of past events, but the function of memory is strictly future-based. A n E L E M E N T S F A L L 08 ever-changing environment behooves an organism to record responses in the chance that similar events occur in the future. In humans, the seat of memory is the hip- pocampus, a bilateral pair of horns that curl in the subcor- tical region of the brain, ending with the amygdala.x As lesion patient studies have shown, loss of the hippocampus results in profound short-term memory loss, as well as a deficit in the ability to create new long-term memories.X 1 The findings from these and other memory studies have led to the proposal of a memory model in which short-term memories, lasting from a few seconds to a few hours, are recorded i n the hippocampus and are later consolidated to the cortex as long-term memories, where they can remain for a l i fet ime. X 1 1 ' X 1 U synapses. Subsequent co-activation of these two synapses further increases A M PA numbers unti l these neurons fire swiftly with the smallest synaptic activity.™ Behaviorally, the Pavlovian response is observed; if, for ex- ample, a rat hears the tone that is followed by an electric shock, it wi l l quickly behave in response to the tone as i f it were being shocked. The activation provided by the neurons responding to "tone", after a repeated exposure to the tone-shock pairing that induced long-term potentia- tion, is enough to elicit a shock response without co-stimu- lation. The manner in which the hippocampus codes for the information that comprises such memories remains unknown. However, injection of a protein synthesis in- hibitor such as anisomycin during training sessions blocks "Memory is comprised of past events, but the function of memory is strictly future-based. An ever-changing environment behooves an organism to record responses in the chance that similar events occur in the future." Recent neurobiological research has challenged, or at least modified, this long-standing memory dogma. At the neu- ral level, memory encoding begins with a process called long-term potentiation, in which the co-stimulation of a neuron by two separate neurons at the same time molecu- larly increases the efficiency by which that neuron can later be activated under a similar circumstance. x i v This coinci- dental strengthening is often referred to as Hebb's Law, ex- pressed as "neurons that fire together, wire together." x v Co- stimulation leads to molecular changes facilitated by magnesium ions and voltage-gated NMDA channels, which respond to the excitatory neurotransmitter gluta- mate, causing an influx of calcium ions and, after a short metabolic pathway, protein synthesis that produces A M PA receptors that are inserted specifically in the activated the production of A M PA receptors, long-term potentiation, and subsequent memory of the association between tone and shock. x v n Surprisingly, recent findings from Nader x v m show that i f a protein synthesis inhibitor is injected when the learned behavior is recalled at a later date, the presenta- tion of a tone long after the inhibitor has worn off wi l l elicit very little behavioral response. In effect, the memory is erased when it is recalled in the presence of a protein syn- thesis inhibitor. Nader has proposed that instead of a short- term and long-term memory system, memories are rather in a labile, active state or an inactive, consolidated state, and the process of consolidation and reconsolidation are practically identical. x l x R E M I N I S C I N G A N D D A Y D R E A M I N G The study has caused much excitement and controversy in the psychology field due to its potential as a therapy to erase or dull traumatic memories, but it also raises questions as to the evolutionary selection of such a memory system. According to the active/inactive state memory model, every time a memory is recalled it is liable to be disrupted. The nature of this model seems very unstable, especially when one considers that those memories that are recalled the most are probably the most important. However the sys- tem seems highly evolutionarily conserved, with the same effects of protein synthesis inhibitors dulling or erasing be- havioral memory responses i n rats, chickens, and crabs. x x The positive attribute of this system is that the labile state achieved i n recalling a memory allows it to be brought back to the mental drawing board, where it is open to editing. Thus an individual can continually add to several memory scripts; a memory for "tone = shock = freeze" recalled to a labile state can easily have another stimulus added to it , in- creasing the efficiency of behavioral learning, a trait that could dramatically increase the survival of an individual. For example, a rat that remembers "tone = freeze" wi l l very quickly learn "light flash = freeze" when a light flash is placed directly before the tone/shock training. In humans, lesion patients without the facility of a hippocampus are unable to arrange remembered events in a correct order, supporting the idea of the hippocampus as the region responsible for the assembly of a memory narrative.™ It could also be possible that the only memories prone to error in an active/inactive memory model are episodic memories. Memories are split into two categories: declar- ative and nondeclarative. Declarative memories are further split into episodic and semantic memories, the latter of which deals with facts and certain language skills. Nondeclarative, or implicit memories, record procedural memory, perceptual representation systems, classical con- ditioning, and nonassociative learning. x x i i Few non- human animals display semantic memory capabilities, but the majority of cognoscente species have nondeclarative faculties.X X U 1 I n an active/inactive model of memory, the implicit information coded by these types of behavior, like a tone beep or the motions used for riding a bike, are un- likely to be falsely modified as the only equivalent data that can replace the coded information is derived from environ- mental stimuli. As we have seen from false memory studies, episodic memory seems to have the unique charac- teristic of being highly susceptible to thought-provoked re- consolidation error. Interestingly, it wasn't unti l 2007 that hippocampus lesion patients were tested for their ability to create mental images in response to a verbal prompt . x x i v Episodic memory pro- duction is practically nonexistent i n these individuals, and i f imagination is synonymous with episodic memory recall, one would expect to see retarded imaginative processes. Compared to control individuals of the same age and IQ, the responses of the amnesic patients were severely frag- mented, labored, and non-descriptive, leading the research team to conclude that imagining new events was impossi- ble for hippocampally-lesioned patients. To decisively test whether episodic memory and episodic imagery are the same wil l require a comparative study of brain activation during both tasks. Before the experiment begins, participants wi l l be asked to send their parents a short questionnaire about events from their child's past. While lying i n a function Magnetic Resonance Imaging (fMRI) machine, the participants wi l l be prompted with ei- ther one of their childhood memories or an event that never occurred. They wil l then be asked to recall the event and mentally relive the experience, regardless of its validity. I f the prompt is of a familiar event, the participant wi l l be asked to recall the specific memory. I f the prompt doesn't ring a bell i n the mind of the participant, they wi l l be asked to imagine the event as i f it had truly occurred. For the com- fort of the participant, the parents wi l l be asked for only pos- itive events. The participants wi l l be told to be as vivid as possible in recalling the memory, and upon finishing the session, lasting only a few minutes, they wil l need to de- scribe their mental responses. The participants wi l l also be asked to acknowledge which scenarios they recalled and E L E M E N T S : : F A L L 08 E N C O D I N G O F E P I S O D I C M E M O R I E S A C T I V A T E S N U M E R O U S PARTS O F T H E BRA I N,AS S E E N H E R E IN T H I S S E R I E S O F MR I S . which they imagined occurring, and brain activities be- tween these two groups wi l l be compared. Past positron emission tomography (PET) studies have found different activations during the encoding and re- trieval of episodic memories, x x v often termed 'hemispheric encoding-retrieval asymmetry > X X V 1. Encoding activity has been shown to activate left prefrontal cortex and right hip- pocampal regions in the majority of individuals. Retrieval has been shown to activate only the left prefrontal cortex, and i f imagining and episodic memory retrieval were of the same nature, one would expect similar activations. Despite these illuminating results, the majority of the paradigms that produced these patterns of activation used single still images as cues, which are poor real world examples of episodic memory. The paradigm proposed here uses real autobiographical episodes rather than single images. Also, retrieval required ac- knowledging familiarity when new and old images were presented to partici- pants, while this proposed experiment requires a mental recount of each episodic event. Although the cues used in these past experiments would qualify in content as episodic memories, they are poor examples of the powerfully expressive nature of human episodic memory. That being said, regions of acti- vation could vary widely from these previous results in lat- eralization, but wi l l most likely still occur within the left prefrontal cortex, hippocampus, and anterior cingulate gyrus xxvii>xxviii Other interesting considerations could be applied to fur- ther experiments; i f imagination and episodic memory uti- T H E BRAIN U N D E R G O E S A COMPLEX PROCESS WHEN CH ILDREN USE THE IR I M AG I N AT IONS. lize the same brain regions to function, does forcing one function hinder the production of the other? For instance, would accuracy in passive memory tasks decrease while a participant is daydreaming, or vice versa, while hippocam- pus activity remains the same? What roles do attention areas play in switching classically episodic regions from recording real life to reliving fictions? x x l x Do these areas play a role in the dream production of REM sleep? The frontal lobes are known to play a large role in global brain activities, including working memory, novel object recogni- tion, and planning for the future.30™*** 1 I f the frontal lobes are the region associated with Baddeley's central execu- tive, 5 0™ 1 is episodic memory one of the processes of a visuospatial sketchpad, the other being creative imagery? What role would the switch between the imagined and the real play i n a schizophrenic's notorious misinterpreta- tions of imagined events? Imagination is one of the cognitive abilities that, unt i l recently, most cognitive neuroscientists would not touch with a long pole; it re- minds one too much of the ambiguously defined subsets of the dualist mind. With in- creased understanding of our species cognitive abilities, re- search scientists are quickly closing i n on subjects once left only to the discussions of philosophers. Some of this re- search raises questions about the way we perceive imagina- tive processes. Findings describing the neural basis for memory reconsolidation allow for a more fluid memory model that explains the errors notorious to episodic mem- ory as seen in false memory studies. Hippocampus lesion amnesiacs not only display a loss i n short-term memory and long-term memory production, but also the inability to create episodic narratives. As abstract as it may sound at E L E M E N T S F A L L 08 first, imagination is a concise cognitive ability that may be synonymous with episodic memory recall. Understanding the mechanism underlying the similarities and differences between these two cognitive abilities could have implica- tions in attention theories, REM dream theory, and mem- ory models, as well as our perception of reality. E N D N O T E S i . Hassabis (2007) i i . Nader (2003) i i i . Loftus (1997) iv. Sudderdorf (2007) v. Gazzaniga (2002) vi . Gazzaniga (2002) v i i . Sudderdorf (2007) v i i i . Loftus (1997) ix. Loftus (1997) x. Gazzaniga (2002) xi . 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