Emotion can have a powerful effect on humans and animals. Numerous studies have shown that the most vivid autobiographical memories tend to be of emotional events, which are likely to be recalled more often and with more clarity and detail than neutral events.

The activity of emotionally enhanced memory retention can be linked to human evolution; during early development, responsive behavior to environmental events would have progressed as a process of trial and error. Survival depended on behavioral patterns that were repeated or reinforced through life and death situations. Through evolution, this process of learning became genetically embedded in humans and all animal species in what is known as flight or fight instinct.

Artificially inducing this instinct through traumatic physical or emotional stimuli essentially creates the same physiological condition that heightens memory retention by exciting neuro-chemical activity affecting areas of the brain responsible for encoding and recalling memory.Schacter, D. L. (1996). Searching for memory. New York: Basic Books. This memory-enhancing effect of emotion has been demonstrated in many laboratory studies, using stimuli ranging from words to pictures to narrated slide shows, as well as autobiographical memory studies. However, as described below, emotion does not always enhance memory.

Arousal and valence in memory One of the most common frameworks in the emotions field proposes that affective experiences are best characterized by two main dimensions: arousal and valence. The dimension of valence ranges from highly positive to highly negative, whereas the dimension of arousal ranges from calming or soothing to exciting or agitating.

The majority of studies to date have focused on the arousal dimension of emotion as the critical factor contributing to the emotional enhancement effect on memory. Different explanations have been offered for this effect, according to the different stages of memory formation and reconstruction. Memory has been shown to be better with arousal linked with emotion than without emotion. The use of a PET scan has allowed scientists to see that pictures with an "emotional-stimulus" have significantly larger amount of activity in the amygdala. In a study using fluoro-2-deoxyglucose (FDG-PET) to examine the brain during recall of films that were both neutral and aversive, there was a positive correlation between the brain glucose and metabolic rate in the amygdala. The activity in the amygdala is part of the episodic memory that was being created due to the adverse stimuli. Most recently, an intracranial EEG study found that the amygdala triggered more pronounced hippocampal sharp-wave ripples after the encoding of more arousing experiences, which are believed to play a critical role in memory consolidation.

However, a growing body of research is dedicated to the emotional valence dimension and its effects on memory. It has been claimed that this is an essential step towards a more complete understanding of emotion effects on memory. The studies that did investigate this dimension have found that emotional valence alone can enhance memory; that is, nonarousing items with positive or negative valence can be better remembered than neutral items.

Emotion and encoding From an information processing perspective, encoding refers to the process of interpreting incoming stimuli and combining the processed information. At the encoding level the following mechanisms have been suggested as mediators of emotion effects on memory:

Selectivity of attention Easterbrook's (1959) cue utilization theory predicted that high levels of arousal will lead to attention narrowing, defined as a decrease in the range of cues from the stimulus and its environment to which the organism is sensitive. According to this hypothesis, attention will be focused primarily on the arousing details (cues) of the stimulus, so that information central to the source of the emotional arousal will be encoded while peripheral details will not.

Accordingly, several studies have demonstrated that the presentation of emotionally-arousing stimuli (compared to neutral stimuli) results in enhanced memory for central details (details central to the appearance or meaning of the emotional stimuli) and impaired memory for peripheral details. Also consistent with this hypothesis are findings of weapon focus effect, in which witnesses to a crime remember the gun or knife in great detail but not other details such as the perpetrator's clothing or vehicle. In laboratory replications it was found that participants spend a disproportionate amount of time looking at a weapon in a scene, and this looking time is inversely related to the likelihood that individuals will subsequently identify the perpetrator of the crime. Other researchers have suggested arousal may also increase the duration of attentional focusing on the arousing stimuli, thus delaying the disengagement of attention from it. Ochsner (2000) summarized the different findings and suggested that by influencing attention selectivity and dwell time, arousing stimuli are more distinctively encoded, resulting in more accurate memory of those stimuli.

While these previous studies focused on how emotion affects memory for emotionally arousing stimuli, in their arousal-biased competition theory, Mather and Sutherland (2011) argue that how arousal influences memory for non-emotional stimuli depends on the priority of those stimuli at the time of the arousal. Arousal enhances perception and memory of high priority stimuli but impairs perception and memory of low priority stimuli. Priority can be determined by bottom-up salience or by top-down goals.

Prioritized processing Emotional items also appear more likely to be processed when attention is limited, suggesting a facilitated or prioritized processing of emotional information. This effect was demonstrated using the attentional blink paradigm in which 2 target items are presented in close temporal proximity within a stream of rapidly presented stimuli.

The typical finding is that participants often miss the second target item, as if there were a "blink" of attention following the first target's presentation, reducing the likelihood that the second target stimulus is attended. However, when the second target stimulus elicits emotional arousal (a "taboo" word), participants are less likely to miss the target's presentation, which suggests that under conditions of limited attention, arousing items are more likely to be processed than neutral items.

Additional support for the prioritized processing hypothesis was provided by studies investigating the visual extinction deficit. People suffering from this deficit can perceive a single stimulus in either side visual field if it is presented alone but are unaware of the same stimulus in the visual field opposed to the lesional side, if another stimulus is presented simultaneously on the lesional side.

Emotion has been found to modulate the magnitude of the visual extinction deficit, so that items that signal emotional relevance (e.g., spiders) are more likely to be processed in the presence of competing distractors than nonemotional items (e.g., flowers).

Emotion and storage In addition to its effects during the encoding phase, emotional arousal appears to increase the likelihood of memory consolidation during the retention (storage) stage of memory (the process of creating a permanent record of the encoded information). A number of studies show that over time, memories for neutral stimuli decrease but memories for arousing stimuli remain the same or improve.

Others have discovered that memory enhancements for emotional information tend to be greater after longer delays than after relatively short ones. This delayed effect is consistent with the proposal that emotionally-arousing memories are more likely to be converted into a relatively permanent trace, whereas memories for nonarousing events are more vulnerable to disruption.

A few studies have even found that emotionally arousing stimuli enhance memory only after a delay. The most famous of these was a study by Kleinsmith and Kaplan (1963) that found an advantage for numbers paired with arousing words over those paired with neutral words only at delayed test, but not at immediate test. As outlined by Mather (2007), the Kleinsmith and Kaplan effects were most likely due to a methodological confound. However, Sharot and Phelps (2004) found better recognition of arousing words over neutral words at a delayed test but not at an immediate test, supporting the notion that there is enhanced memory consolidation for arousing stimuli.McGaugh, J. L. (1992). Affect, neuromodulatory systems and memory storage. In S. Christianson (Ed.), The handbook of emotion and memory: Research and theory (pp. 269-288). According to these theories, different physiological systems, including those involved in the discharge of hormones believed to affect memory consolidation, become active during, and closely following, the occurrence of arousing events.

Another possible explanation for the findings of the emotional arousal delayed effect is post-event processing regarding the cause of the arousal. According to the post stimulus elaboration (PSE) hypothesis, an arousing emotional experience may cause more effort to be invested in elaboration of the experience, which would subsequently be processed at a deeper level than a neutral experience. Elaboration refers to the process of establishing links between newly-encountered information and previously-stored information.

It has long been known that when individuals process items in an elaborative fashion, such that meaning is extracted from items and inter-item associations are formed, memory is enhanced. Thus, if a person gives more thought to central details in an arousing event, memory for such information is likely to be enhanced. However, these processes could also disrupt consolidation of memories for peripheral details. Christianson (1992) suggested that the combined action of perceptual, attentional, and elaborative processing, triggered by an emotionally arousing experience, produces memory enhancements of details related to the emotion laden stimulus, at the cost of less elaboration and consolidation of memory for the peripheral details.

Emotion and elaboration The processes involved in this enhancement may be distinct from those mediating the enhanced memory for arousing items. It has been suggested that in contrast to the relatively automatic attentional modulation of memory for arousing information, memory for non-arousing positive or negative stimuli may benefit instead from conscious encoding strategies, such as elaboration. This elaborative processing can be autobiographical or semantic.

Autobiographical elaboration is known to benefit memory by creating links between the processed stimuli, and the self, for example, deciding whether a word would describe the personal self. Memory formed through autobiographical elaboration is enhanced as compared to items processed for meaning, but not in relation to the self.

Since words such as "sorrow" or "comfort" may be more likely to be associated with autobiographical experiences or self-introspection than neutral words such as "shadow", autobiographical elaboration may explain the memory enhancement of non-arousing positive or negative items. Studies have shown that dividing attention at encoding decreases an individual's ability to utilize controlled encoding processes, such as autobiographical or semantic elaboration.