![Memory](https://upload.wikimedia.org/wikipedia/commons/5/58/Memory.gif)
Memory is the faculty of the
mind by which
data
In the pursuit of knowledge, data (; ) is a collection of discrete Value_(semiotics), values that convey information, describing quantity, qualitative property, quality, fact, statistics, other basic units of meaning, or simply sequences of sy ...
or
information
Information is an abstract concept that refers to that which has the power to inform. At the most fundamental level information pertains to the interpretation of that which may be sensed. Any natural process that is not completely random ...
is
encoded
In communications and information processing, code is a system of rules to convert information—such as a letter, word, sound, image, or gesture—into another form, sometimes shortened or secret, for communication through a communication ...
, stored, and retrieved when needed. It is the retention of information over time for the purpose of influencing future action. If
past events could not be remembered, it would be impossible for language, relationships, or
personal identity
Personal identity is the unique numerical identity of a person over time. Discussions regarding personal identity typically aim to determine the necessary and sufficient conditions under which a person at one time and a person at another time ca ...
to develop.
Memory loss is usually described as
forgetfulness
Forgetting or disremembering is the apparent loss or modification of information already encoded and stored in an individual's short or long-term memory. It is a spontaneous or gradual process in which old memories are unable to be recalled from ...
or
amnesia.
Memory is often understood as an
informational processing system with explicit and implicit functioning that is made up of a
sensory processor,
short-term (or
working) memory, and
long-term memory
Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long- ...
.
This can be related to the
neuron
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
.
The sensory processor allows information from the outside world to be sensed in the form of chemical and physical stimuli and attended to various levels of focus and intent. Working memory serves as an encoding and retrieval processor. Information in the form of stimuli is encoded in accordance with explicit or implicit functions by the working memory processor. The working memory also retrieves information from previously stored material. Finally, the function of long-term memory is to store through various categorical models or systems.
Declarative, or explicit, memory is the conscious storage and recollection of data.
Under declarative memory resides
semantic and
episodic memory
Episodic memory is the memory of everyday events (such as times, location geography, associated emotions, and other contextual information) that can be explicitly stated or conjured. It is the collection of past personal experiences that occurred ...
. Semantic memory refers to memory that is encoded with specific meaning,
while episodic memory refers to information that is encoded along a spatial and temporal plane.
Declarative memory is usually the primary process thought of when referencing memory.
Non-declarative, or implicit, memory is the unconscious storage and recollection of information.
An example of a non-declarative process would be the unconscious learning or retrieval of information by way of
procedural memory
Procedural memory is a type of implicit memory ( unconscious, long-term memory) which aids the performance of particular types of tasks without conscious awareness of these previous experiences.
Procedural memory guides the processes we perform ...
, or a priming phenomenon.
Priming
Priming may refer to:
* Priming (agriculture), a form of seed planting preparation, in which seeds are soaked before planting
* Priming (immunology), a process occurring when a specific antigen is presented to naive lymphocytes causing them to d ...
is the process of
subliminally arousing specific responses from memory and shows that not all memory is consciously activated,
whereas procedural memory is the slow and gradual learning of skills that often occurs without conscious attention to learning.
Memory is not a perfect processor, and is affected by many factors. The ways by which information is encoded, stored, and retrieved can all be corrupted. Pain, for example, has been identified as a physical condition that impairs memory, and has been noted in animal models as well as chronic pain patients. The amount of attention given new stimuli can diminish the amount of information that becomes encoded for storage.
Also, the storage process can become corrupted by physical damage to areas of the brain that are associated with memory storage, such as the hippocampus.
Finally, the retrieval of information from long-term memory can be disrupted because of decay within long-term memory.
Normal functioning, decay over time, and brain damage all affect the accuracy and capacity of the memory.
Sensory memory
Sensory memory holds information, derived from the senses, less than one second after an item is perceived. The ability to look at an item and remember what it looked like with just a split second of observation, or memorization, is the example of sensory memory. It is out of cognitive control and is an automatic response. With very short presentations, participants often report that they seem to "see" more than they can actually report. The first precise experiments exploring this form of sensory memory were conducted by
George Sperling
George Sperling (born 1934) is an American cognitive psychologist, researcher, and educator. Sperling documented the existence of iconic memory (one of the sensory memory subtypes). Through several experiments, he showed support for his hypothe ...
(1963) using the "partial report paradigm." Subjects were presented with a grid of 12 letters, arranged into three rows of four. After a brief presentation, subjects were then played either a high, medium or low tone, cuing them which of the rows to report. Based on these partial report experiments, Sperling was able to show that the capacity of sensory memory was approximately 12 items, but that it degraded very quickly (within a few hundred milliseconds). Because this form of memory degrades so quickly, participants would see the display but be unable to report all of the items (12 in the "whole report" procedure) before they decayed. This type of memory cannot be prolonged via rehearsal.
Three types of sensory memories exist.
Iconic memory
Iconic memory is the visual sensory memory register pertaining to the visual domain and a fast-decaying store of visual information. It is a component of the visual memory system which also includes visual short-term memory (VSTM) and long-term mem ...
is a fast decaying store of visual information, a type of sensory memory that briefly stores an image that has been perceived for a small duration.
Echoic memory
Echoic memory is the sensory memory that registers specific to auditory information (sounds). Once an auditory stimulus is heard, it is stored in memory so that it can be processed and understood. Unlike most visual memory, where a person can choo ...
is a fast decaying store of auditory information, also a sensory memory that briefly stores sounds that have been perceived for short durations.
Haptic memory
Haptic memory is the form of sensory memory specific to touch stimuli. Haptic memory is used regularly when assessing the necessary forces for gripping and interacting with familiar objects. It may also influence one's interactions with novel obje ...
is a type of sensory memory that represents a database for touch stimuli.
Short-term memory
Short-term memory is also known as working memory. Short-term memory allows recall for a period of several seconds to a minute without rehearsal. Its capacity, however, is very limited. In 1956,
George A. Miller (1920–2012), when working at
Bell Laboratories
Nokia Bell Labs, originally named Bell Telephone Laboratories (1925–1984),
then AT&T Bell Laboratories (1984–1996)
and Bell Labs Innovations (1996–2007),
is an American industrial research and scientific development company owned by mult ...
, conducted experiments showing that the store of short-term memory was 7±2 items. (Hence, the title of his famous paper,
"The Magical Number 7±2.") Modern perspectives estimate the capacity of short-term memory to be lower, typically on the order of 4–5 items,
or argue for a more flexible limit based on information instead of items.
Memory capacity can be increased through a process called
chunking.
For example, in recalling a ten-digit
telephone number
A telephone number is a sequence of digits assigned to a landline telephone subscriber station connected to a telephone line or to a wireless electronic telephony device, such as a radio telephone or a mobile telephone, or to other devices f ...
, a person could chunk the digits into three groups: first, the area code (such as 123), then a three-digit chunk (456), and, last, a four-digit chunk (7890). This method of remembering telephone numbers is far more effective than attempting to remember a string of 10 digits; this is because we are able to chunk the information into meaningful groups of numbers. This is reflected in some countries' tendencies to display telephone numbers as several chunks of two to four numbers.
Short-term memory is believed to rely mostly on an acoustic code for storing information, and to a lesser extent on a visual code. Conrad (1964)
found that test subjects had more difficulty recalling collections of letters that were acoustically similar, e.g., E, P, D. Confusion with recalling acoustically similar letters rather than visually similar letters implies that the letters were encoded acoustically. Conrad's (1964) study, however, deals with the encoding of written text; thus, while memory of written language may rely on acoustic components, generalizations to all forms of memory cannot be made.
Long-term memory
![Memory-Warner-Highsmith](https://upload.wikimedia.org/wikipedia/commons/7/7e/Memory-Warner-Highsmith.jpeg)
The storage in sensory memory and short-term memory generally has a strictly limited capacity and duration, which means that information is not retained indefinitely. By contrast, while the total capacity of long-term memory has yet to be established, it can store much larger quantities of information. Furthermore, it can store this information for a much longer duration, potentially for a whole life span. For example, given a random seven-digit number, one may remember it for only a few seconds before forgetting, suggesting it was stored in short-term memory. On the other hand, one can remember telephone numbers for many years through repetition; this information is said to be stored in long-term memory.
While short-term memory encodes information acoustically, long-term memory encodes it semantically: Baddeley (1966)
discovered that, after 20 minutes, test subjects had the most difficulty recalling a collection of words that had similar meanings (e.g. big, large, great, huge) long-term. Another part of long-term memory is episodic memory, "which attempts to capture information such as 'what', 'when' and 'where.
With episodic memory, individuals are able to recall specific events such as birthday parties and weddings.
Short-term memory is supported by transient patterns of
neuronal communication
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. No ...
, dependent on regions of the
frontal lobe
The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a groove be ...
(especially
dorsolateral prefrontal cortex
The dorsolateral prefrontal cortex (DLPFC or DL-PFC) is an area in the prefrontal cortex of the primate brain. It is one of the most recently derived parts of the human brain. It undergoes a prolonged period of maturation which lasts until adultho ...
) and the
parietal lobe
The parietal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The parietal lobe is positioned above the temporal lobe and behind the frontal lobe and central sulcus.
The parietal lobe integrates sensory informa ...
. Long-term memory, on the other hand, is maintained by more stable and permanent changes in neural connections widely spread throughout the brain. The
hippocampus
The hippocampus (via Latin from Greek , ' seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, ...
is essential (for learning new information) to the consolidation of information from short-term to long-term memory, although it does not seem to store information itself. It was thought that without the hippocampus new memories were unable to be stored into long-term memory and that there would be a very short
attention span, as first gleaned from patient
Henry Molaison after what was thought to be the full removal of both his hippocampi. More recent examination of his brain, post-mortem, shows that the hippocampus was more intact than first thought, throwing theories drawn from the initial data into question. The hippocampus may be involved in changing neural connections for a period of three months or more after the initial learning.
Research has suggested that long-term memory storage in humans may be maintained by
DNA methylation,
and the
'prion' gene.
Further research investigated the molecular basis for
long-term memory
Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long- ...
. By 2015 it had become clear that long-term memory requires gene transcription activation and
de novo protein synthesis.
Long-term memory formation depends on both the activation of memory promoting genes and the inhibition of memory suppressor genes, and
DNA methylation/
DNA demethylation
For molecular biology in mammals, DNA demethylation causes replacement of 5-methylcytosine (5mC) in a DNA sequence by cytosine (C) (see figure of 5mC and C). DNA demethylation can occur by an active process at the site of a 5mC in a DNA sequenc ...
was found to be a major mechanism for achieving this dual regulation.
Rats with a new, strong long-term memory due to
contextual fear conditioning have reduced expression of about 1,000 genes and increased expression of about 500 genes in the hippocampus 24 hours after training, thus exhibiting modified expression of 9.17% of the rat hippocampal genome. Reduced gene expressions were associated with methylations of those genes.
Considerable further research into long-term memory has illuminated the molecular mechanisms by which methylations are established or removed, as reviewed in 2022.
These mechanisms include, for instance, signal-responsive
TOP2B
DNA topoisomerase 2-beta is an enzyme that in humans is encoded by the ''TOP2B'' gene.
Function
This gene encodes a DNA topoisomerase, an enzyme that controls and alters the topologic states of DNA during transcription. This nuclear enzyme is i ...
-induced double-strand breaks in
immediate early genes
Immediate early genes (IEGs) are genes which are activated transiently and rapidly in response to a wide variety of cellular stimuli. They represent a standing response mechanism that is activated at the transcription level in the first round of ...
. Also the
messenger RNAs of many genes that had been subjected to methylation-controlled increases or decreases are transported by neural granules (
messenger RNP) to the
dendritic spine
A dendritic spine (or spine) is a small membranous protrusion from a neuron's dendrite that typically receives input from a single axon at the synapse. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical s ...
s. At these locations the messenger RNAs can be
translated
Translation is the communication of the meaning of a source-language text by means of an equivalent target-language text. The English language draws a terminological distinction (which does not exist in every language) between ''transla ...
into the proteins that control signaling at
neuron
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
al
synapses.
Multi-store model
![multistore model](https://upload.wikimedia.org/wikipedia/commons/b/bf/Multistore_model.png)
The multi-store model (also known as
Atkinson–Shiffrin memory model) was first described in 1968 by
Atkinson Atkinson may refer to:
Places
*Atkinson, Nova Scotia, Canada
* Atkinson, Dominica, a village in Dominica
*Atkinson, Illinois, U.S.
* Atkinson, Indiana, U.S.
*Atkinson, Maine, U.S.
*Atkinson Lake, a lake in Minnesota, U.S.
*Atkinson, Nebraska, U. ...
and
Shiffrin.
The multi-store model has been criticised for being too simplistic. For instance, long-term memory is believed to be actually made up of multiple subcomponents, such as episodic and
procedural memory
Procedural memory is a type of implicit memory ( unconscious, long-term memory) which aids the performance of particular types of tasks without conscious awareness of these previous experiences.
Procedural memory guides the processes we perform ...
. It also proposes that rehearsal is the only mechanism by which information eventually reaches long-term storage, but evidence shows us capable of remembering things without rehearsal.
The model also shows all the memory stores as being a single unit whereas research into this shows differently. For example, short-term memory can be broken up into different units such as visual information and acoustic information. In a study by Zlonoga and Gerber (1986), patient 'KF' demonstrated certain deviations from the Atkinson–Shiffrin model. Patient KF was
brain damaged, displaying difficulties regarding short-term memory. Recognition of sounds such as spoken numbers, letters, words and easily identifiable noises (such as doorbells and cats meowing) were all impacted. Visual short-term memory was unaffected, suggesting a dichotomy between visual and audial memory.
Working memory
![Working memory model](https://upload.wikimedia.org/wikipedia/commons/e/ed/Working_memory_model.svg)
In 1974 Baddeley and Hitch proposed a "working memory model" that replaced the general concept of short-term memory with an active maintenance of information in the short-term storage. In this model, working memory consists of three basic stores: the central executive, the phonological loop and the visuo-spatial sketchpad. In 2000 this model was expanded with the multimodal episodic buffer (
Baddeley's model of working memory
Baddeley's model of working memory is a model of human memory proposed by Alan Baddeley and Graham Hitch in 1974, in an attempt to present a more accurate model of primary memory (often referred to as short-term memory). Working memory splits pri ...
).
The central executive essentially acts as an attention sensory store. It channels information to the three component processes: the phonological loop, the visuo-spatial sketchpad, and the episodic buffer.
The phonological loop stores auditory information by silently rehearsing sounds or words in a continuous loop: the articulatory process (for example the repetition of a telephone number over and over again). A short list of data is easier to remember. The phonological loop is occasionally disrupted.
Irrelevant speech or background noise can impede the phonological loop.
Articulatory suppression Articulatory suppression is the process of inhibiting memory performance by speaking while being presented with an item to remember. Most research demonstrates articulatory suppression by requiring an individual to repeatedly say an irrelevant speec ...
can also confuse encoding and words that sound similar can be switched or misremembered through the phonological similarity effect. the phonological loop also has a limit to how much it can hold at once which means that it is easier to remember a lot of short words rather than a lot of long words, according to the word length effect.
The
visuospatial sketchpad stores visual and spatial information. It is engaged when performing spatial tasks (such as judging distances) or visual ones (such as counting the windows on a house or imagining images). Those with
Aphantasia
Aphantasia ( , ) is the inability to create mental imagery.
The phenomenon was first described by Francis Galton in 1880 but has since remained relatively unstudied. Interest in the phenomenon renewed after the publication of a study in 2015 ...
will not be able to engage the visuospatial sketchpad.
The episodic buffer is dedicated to linking information across domains to form integrated units of visual, spatial, and verbal information and chronological ordering (e.g., the memory of a story or a movie scene). The episodic buffer is also assumed to have links to long-term memory and semantical meaning.
The working memory model explains many practical observations, such as why it is easier to do two different tasks (one verbal and one visual) than two similar tasks (e.g., two visual), and the aforementioned word-length effect. Working memory is also the premise for what allows us to do everyday activities involving thought. It is the section of memory where we carry out thought processes and use them to learn and reason about topics.
Types
Researchers distinguish between recognition and recall memory. Recognition memory tasks require individuals to indicate whether they have encountered a stimulus (such as a picture or a word) before. Recall memory tasks require participants to retrieve previously learned information. For example, individuals might be asked to produce a series of actions they have seen before or to say a list of words they have heard before.
By information type
Topographical memory involves the ability to orient oneself in space, to recognize and follow an itinerary, or to recognize familiar places. Getting lost when traveling alone is an example of the failure of topographic memory.
Flashbulb memories are clear
episodic memories
Episodic memory is the memory of everyday events (such as times, location geography, associated emotions, and other contextual information) that can be explicitly stated or conjured. It is the collection of past personal experiences that occurred ...
of unique and highly emotional events. People remembering where they were or what they were doing when they first heard the news of
President Kennedy
John Fitzgerald Kennedy (May 29, 1917 – November 22, 1963), often referred to by his initials JFK and the nickname Jack, was an American politician who served as the 35th president of the United States from 1961 until assassination of Joh ...
's
assassination,
the
Sydney Siege or of
9/11 are examples of flashbulb memories.
Anderson
Anderson or Andersson may refer to:
Companies
* Anderson (Carriage), a company that manufactured automobiles from 1907 to 1910
* Anderson Electric, an early 20th-century electric car
* Anderson Greenwood, an industrial manufacturer
* Anderson ...
(1976)
divides long-term memory into ''
declarative (explicit)'' and ''
procedural (implicit)'' memories.
Declarative
Declarative memory
Explicit memory (or declarative memory) is one of the two main types of long-term human memory, the other of which is implicit memory. Explicit memory is the conscious, intentional recollection of factual information, previous experiences, and c ...
requires
conscious
Consciousness, at its simplest, is sentience and awareness of internal and external existence. However, the lack of definitions has led to millennia of analyses, explanations and debates by philosophers, theologians, linguisticians, and scien ...
recall, in that some conscious process must call back the information. It is sometimes called ''
explicit memory
Explicit memory (or declarative memory) is one of the two main types of long-term human memory, the other of which is implicit memory. Explicit memory is the conscious, intentional recollection of factual information, previous experiences, and con ...
'', since it consists of information that is explicitly stored and retrieved. Declarative memory can be further sub-divided into
semantic memory
Semantic memory refers to general world knowledge that humans have accumulated throughout their lives. This general knowledge (word meanings, concepts, facts, and ideas) is intertwined in experience and dependent on culture. We can learn about n ...
, concerning principles and facts taken independent of context; and
episodic memory
Episodic memory is the memory of everyday events (such as times, location geography, associated emotions, and other contextual information) that can be explicitly stated or conjured. It is the collection of past personal experiences that occurred ...
, concerning information specific to a particular context, such as a time and place. Semantic memory allows the encoding of abstract
knowledge
Knowledge can be defined as Descriptive knowledge, awareness of facts or as Procedural knowledge, practical skills, and may also refer to Knowledge by acquaintance, familiarity with objects or situations. Knowledge of facts, also called pro ...
about the world, such as "Paris is the capital of France". Episodic memory, on the other hand, is used for more personal memories, such as the sensations, emotions, and personal associations of a particular place or time. Episodic memories often reflect the "firsts" in life such as a first kiss, first day of school or first time winning a championship. These are key events in one's life that can be remembered clearly.
Research suggests that declarative memory is supported by several functions of the medial temporal lobe system which includes the hippocampus.
Autobiographical memory
Autobiographical memory is a memory system consisting of episodes recollected from an individual's life, based on a combination of episodic (personal experiences and specific objects, people and events experienced at particular time and place) a ...
– memory for particular events within one's own life – is generally viewed as either equivalent to, or a subset of, episodic memory.
Visual memory
Visual memory describes the relationship between perceptual processing and the encoding, storage and retrieval of the resulting neural representations. Visual memory occurs over a broad time range spanning from eye movements to years in order ...
is part of memory preserving some characteristics of our senses pertaining to visual experience. One is able to place in memory information that resembles objects, places, animals or people in sort of a
mental image
A mental image is an experience that, on most occasions, significantly resembles the experience of 'perceiving' some object, event, or scene, but occurs when the relevant object, event, or scene is not actually present to the senses. There are ...
. Visual memory can result in
priming
Priming may refer to:
* Priming (agriculture), a form of seed planting preparation, in which seeds are soaked before planting
* Priming (immunology), a process occurring when a specific antigen is presented to naive lymphocytes causing them to d ...
and it is assumed some kind of perceptual representational system underlies this phenomenon.
Procedural
In contrast,
procedural memory
Procedural memory is a type of implicit memory ( unconscious, long-term memory) which aids the performance of particular types of tasks without conscious awareness of these previous experiences.
Procedural memory guides the processes we perform ...
(or ''
implicit memory
In psychology, implicit memory is one of the two main types of long-term human memory. It is acquired and used unconsciously, and can affect thoughts and behaviours. One of its most common forms is procedural memory, which allows people to perfo ...
'') is not based on the conscious recall of information, but on
implicit learning. It can best be summarized as remembering how to do something. Procedural memory is primarily used in learning
motor skill
A motor skill is a function that involves specific movements of the body's muscles to perform a certain task. These tasks could include walking, running, or riding a bike. In order to perform this skill, the body's nervous system, muscles, and br ...
s and can be considered a subset of implicit memory. It is revealed when one does better in a given task due only to repetition – no new explicit memories have been formed, but one is
unconsciously
The unconscious mind (or the unconscious) consists of the processes in the mind which occur automatically and are not available to introspection and include thought processes, memories, interests, and motivations.
Even though these processes exist ...
accessing aspects of those previous experiences. Procedural memory involved in
motor learning
Motor learning refers broadly to changes in an organism's movements that reflect changes in the structure and function of the nervous system. Motor learning occurs over varying timescales and degrees of complexity: humans learn to walk or talk over ...
depends on the
cerebellum and
basal ganglia
The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an exter ...
.
A characteristic of procedural memory is that the things remembered are automatically translated into actions, and thus sometimes difficult to describe. Some examples of procedural memory include the ability to ride a bike or tie shoelaces.
By temporal direction
Another major way to distinguish different memory functions is whether the content to be remembered is in the past,
retrospective memory, or in the future,
prospective memory Prospective memory is a form of memory that involves remembering to perform a planned action or recall a planned intention at some future point in time.McDaniel, M. A., & Einstein, G. O. (2007). ''Prospective memory: An overview and synthesis of an ...
. John Meacham introduced this distinction in a paper presented at the 1975
American Psychological Association
The American Psychological Association (APA) is the largest scientific and professional organization of psychologists in the United States, with over 133,000 members, including scientists, educators, clinicians, consultants, and students. It ha ...
annual meeting and subsequently included by
Ulric Neisser
Ulric Richard Gustav Neisser (December 8, 1928 – February 17, 2012) was a German-American psychologist, Cornell University professor, and member of the US National Academy of Sciences. He has been referred to as the "father of cognitive p ...
in his 1982 edited volume, ''Memory Observed: Remembering in Natural Contexts''. Thus, retrospective memory as a category includes semantic, episodic and autobiographical memory. In contrast, prospective memory is memory for future intentions, or ''remembering to remember'' (Winograd, 1988). Prospective memory can be further broken down into event- and time-based prospective remembering. Time-based prospective memories are triggered by a time-cue, such as going to the doctor (action) at 4pm (cue). Event-based prospective memories are intentions triggered by cues, such as remembering to post a letter (action) after seeing a mailbox (cue). Cues do not need to be related to the action (as the mailbox/letter example), and lists, sticky-notes, knotted handkerchiefs, or string around the finger all exemplify cues that people use as strategies to enhance prospective memory.
Study techniques
To assess infants
Infants do not have the language ability to report on their memories and so verbal reports cannot be used to assess very young children's memory. Throughout the years, however, researchers have adapted and developed a number of measures for assessing both infants' recognition memory and their recall memory.
Habituation
Habituation is a form of non-associative learning in which an innate (non-reinforced) response to a stimulus decreases after repeated or prolonged presentations of that stimulus. Responses that habituate include those that involve the intact org ...
and
operant conditioning techniques have been used to assess infants' recognition memory and the deferred and elicited imitation techniques have been used to assess infants' recall memory.
Techniques used to assess infants' recognition memory include the following:
* Visual paired comparison procedure (relies on habituation): infants are first presented with pairs of visual stimuli, such as two black-and-white photos of human faces, for a fixed amount of time; then, after being familiarized with the two photos, they are presented with the "familiar" photo and a new photo. The time spent looking at each photo is recorded. Looking longer at the new photo indicates that they remember the "familiar" one. Studies using this procedure have found that 5- to 6-month-olds can retain information for as long as fourteen days.
* Operant conditioning technique: infants are placed in a crib and a ribbon that is connected to a mobile overhead is tied to one of their feet. Infants notice that when they kick their foot the mobile moves – the rate of kicking increases dramatically within minutes. Studies using this technique have revealed that infants' memory substantially improves over the first 18-months. Whereas 2- to 3-month-olds can retain an operant response (such as activating the mobile by kicking their foot) for a week, 6-month-olds can retain it for two weeks, and 18-month-olds can retain a similar operant response for as long as 13 weeks.
Techniques used to assess infants' recall memory include the following:
* Deferred imitation technique: an experimenter shows infants a unique sequence of actions (such as using a stick to push a button on a box) and then, after a delay, asks the infants to imitate the actions. Studies using deferred imitation have shown that 14-month-olds' memories for the sequence of actions can last for as long as four months.
* Elicited imitation technique: is very similar to the deferred imitation technique; the difference is that infants are allowed to imitate the actions before the delay. Studies using the elicited imitation technique have shown that 20-month-olds can recall the action sequences twelve months later.
To assess children and older adults
Researchers use a variety of tasks to assess older children and adults' memory. Some examples are:
* Paired associate learning – when one learns to associate one specific word with another. For example, when given a word such as "safe" one must learn to say another specific word, such as "green". This is stimulus and response.
* Free recall – during this task a subject would be asked to study a list of words and then later they will be asked to recall or write down as many words that they can remember, similar to free response questions.
Earlier items are affected by retroactive interference (RI), which means the longer the list, the greater the interference, and the less likelihood that they are recalled. On the other hand, items that have been presented lastly suffer little RI, but suffer a great deal from proactive interference (PI), which means the longer the delay in recall, the more likely that the items will be lost.
[Baddeley, Alan D., "The Psychology of Memory", pp. 131–132, Basic Books, Inc., Publishers, New York, 1976, ]
* Cued recall – one is given a significant hints to help retrieve information that has been previously encoded into the person's memory; typically this can involve a word relating to the information being asked to remember. This is similar to fill in the blank assessments used in classrooms.
* Recognition – subjects are asked to remember a list of words or pictures, after which point they are asked to identify the previously presented words or pictures from among a list of alternatives that were not presented in the original list.
This is similar to multiple choice assessments.
* Detection paradigm – individuals are shown a number of objects and color samples during a certain period of time. They are then tested on their visual ability to remember as much as they can by looking at testers and pointing out whether the testers are similar to the sample, or if any change is present.
* Savings method – compares the speed of originally learning to the speed of relearning it. The amount of time saved measures memory.
* Implicit-memory tasks – information is drawn from memory without conscious realization.
Failures
* Transience – memories degrade with the passing of time. This occurs in the storage stage of memory, after the information has been stored and before it is retrieved. This can happen in sensory, short-term, and long-term storage. It follows a general pattern where the information is rapidly forgotten during the first couple of days or years, followed by small losses in later days or years.
* Absent-mindedness – Memory failure due to the lack of
attention. Attention plays a key role in storing information into long-term memory; without proper attention, the information might not be stored, making it impossible to be retrieved later.
Physiology
Brain areas involved in the
neuroanatomy of memory such as the
hippocampus
The hippocampus (via Latin from Greek , ' seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, ...
, the
amygdala
The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verte ...
, the
striatum, or the
mammillary bodies
The mammillary bodies are a pair of small round bodies, located on the undersurface of the brain that, as part of the diencephalon, form part of the limbic system. They are located at the ends of the anterior arches of the fornix. They consist o ...
are thought to be involved in specific types of memory. For example, the hippocampus is believed to be involved in spatial learning and
declarative learning
Declarative learning is acquiring information that one can speak about (contrast with motor learning). The capital of a state is a declarative piece of information, while knowing how to ride a bike is not. Episodic memory and semantic memory a ...
, while the amygdala is thought to be involved in
emotional memory
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 ...
.
Damage to certain areas in patients and animal models and subsequent memory deficits is a primary source of information. However, rather than implicating a specific area, it could be that damage to adjacent areas, or to a pathway ''traveling'' through the area is actually responsible for the observed deficit. Further, it is not sufficient to describe memory, and its counterpart,
learning, as solely dependent on specific brain regions. Learning and memory are usually attributed to changes in neuronal
synapses, thought to be mediated by
long-term potentiation
In neuroscience, long-term potentiation (LTP) is a persistent strengthening of synapses based on recent patterns of activity. These are patterns of synaptic activity that produce a long-lasting increase in signal transmission between two neurons ...
and
long-term depression
In neurophysiology, long-term depression (LTD) is an activity-dependent reduction in the efficacy of neuronal synapses lasting hours or longer following a long patterned stimulus. LTD occurs in many areas of the CNS with varying mechanisms depen ...
.
In general, the more emotionally charged an event or experience is, the better it is remembered; this phenomenon is known as the memory enhancement effect. Patients with amygdala damage, however, do not show a memory enhancement effect.
Hebb distinguished between short-term and long-term memory. He postulated that any memory that stayed in short-term storage for a long enough time would be consolidated into a long-term memory. Later research showed this to be false. Research has shown that direct injections of
cortisol or
epinephrine help the storage of recent experiences. This is also true for stimulation of the amygdala. This proves that excitement enhances memory by the stimulation of hormones that affect the amygdala. Excessive or prolonged stress (with prolonged cortisol) may hurt memory storage. Patients with amygdalar damage are no more likely to remember emotionally charged words than nonemotionally charged ones. The hippocampus is important for explicit memory. The hippocampus is also important for memory consolidation. The hippocampus receives input from different parts of the cortex and sends its output out to different parts of the brain also. The input comes from secondary and tertiary sensory areas that have processed the information a lot already. Hippocampal damage may also cause
memory loss
Amnesia is a deficit in memory caused by brain damage or disease,Gazzaniga, M., Ivry, R., & Mangun, G. (2009) Cognitive Neuroscience: The biology of the mind. New York: W.W. Norton & Company. but it can also be caused temporarily by the use o ...
and problems with memory storage. This memory loss includes
retrograde amnesia
In neurology, retrograde amnesia (RA) is a loss of memory-access to events that occurred or information that was learned in the past. It is caused by an injury or the onset of a disease. It tends to negatively affect episodic, autobiographical, ...
which is the loss of memory for events that occurred shortly before the time of brain damage.
Cognitive neuroscience
Cognitive neuroscientists consider memory as the retention, reactivation, and reconstruction of the experience-independent internal representation. The term of internal
representation implies that such a definition of memory contains two components: the expression of memory at the behavioral or conscious level, and the underpinning physical neural changes (Dudai 2007). The latter component is also called
engram or memory traces (Semon 1904). Some neuroscientists and psychologists mistakenly equate the concept of engram and memory, broadly conceiving all persisting after-effects of experiences as memory; others argue against this notion that memory does not exist until it is revealed in behavior or thought (Moscovitch 2007).
One question that is crucial in cognitive neuroscience is how information and mental experiences are coded and represented in the brain. Scientists have gained much knowledge about the neuronal codes from the studies of plasticity, but most of such research has been focused on simple learning in simple neuronal circuits; it is considerably less clear about the neuronal changes involved in more complex examples of memory, particularly declarative memory that requires the storage of facts and events (Byrne 2007).
Convergence-divergence zone The theory of convergence-divergence zones Antonio Damasio, Time-locked multiregional retroactivation: A systems-level proposal for the neural substrates of recall and recognition, Cognition, 33 (1989) 25-62 was proposed by Antonio Damasio, in 1989, ...
s might be the neural networks where memories are stored and retrieved. Considering that there are several kinds of memory, depending on types of represented knowledge, underlying mechanisms, processes functions and modes of acquisition, it is likely that different brain areas support different memory systems and that they are in mutual relationships in neuronal networks: "components of memory representation are distributed widely across different parts of the brain as mediated by multiple neocortical circuits".
*
Encoding
In communications and information processing, code is a system of rules to convert information—such as a letter, word, sound, image, or gesture—into another form, sometimes shortened or secret, for communication through a communication ...
. Encoding of
working memory
Working memory is a cognitive system with a limited capacity that can hold information temporarily. It is important for reasoning and the guidance of decision-making and behavior. Working memory is often used synonymously with short-term memory, ...
involves the spiking of individual neurons induced by sensory input, which persists even after the sensory input disappears (Jensen and Lisman 2005; Fransen et al. 2002). Encoding of
episodic memory
Episodic memory is the memory of everyday events (such as times, location geography, associated emotions, and other contextual information) that can be explicitly stated or conjured. It is the collection of past personal experiences that occurred ...
involves persistent changes in molecular structures that alter
synaptic transmission between neurons. Examples of such structural changes include
long-term potentiation
In neuroscience, long-term potentiation (LTP) is a persistent strengthening of synapses based on recent patterns of activity. These are patterns of synaptic activity that produce a long-lasting increase in signal transmission between two neurons ...
(LTP) or
spike-timing-dependent plasticity
Spike-timing-dependent plasticity (STDP) is a biological process that adjusts the strength of connections between neurons in the brain. The process adjusts the connection strengths based on the relative timing of a particular neuron's output and in ...
(STDP). The persistent spiking in working memory can enhance the synaptic and cellular changes in the encoding of episodic memory (Jensen and Lisman 2005).
* Working memory. Recent functional imaging studies detected working memory signals in both
medial temporal lobe
The temporal lobe is one of the four major lobes of the cerebral cortex in the brain of mammals. The temporal lobe is located beneath the lateral fissure on both cerebral hemispheres of the mammalian brain.
The temporal lobe is involved in pro ...
(MTL), a brain area strongly associated with
long-term memory
Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long- ...
, and
prefrontal cortex
In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA46 ...
(Ranganath et al. 2005), suggesting a strong relationship between working memory and long-term memory. However, the substantially more working memory signals seen in the prefrontal lobe suggest that this area plays a more important role in working memory than MTL (Suzuki 2007).
*
Consolidation and
reconsolidation.
Short-term memory (STM) is temporary and subject to disruption, while long-term memory (LTM), once consolidated, is persistent and stable. Consolidation of STM into LTM at the molecular level presumably involves two processes: synaptic consolidation and system consolidation. The former involves a protein synthesis process in the medial temporal lobe (MTL), whereas the latter transforms the MTL-dependent memory into an MTL-independent memory over months to years (Ledoux 2007). In recent years, such traditional consolidation dogma has been re-evaluated as a result of the studies on reconsolidation. These studies showed that prevention after
retrieval affects subsequent retrieval of the memory (Sara 2000). New studies have shown that post-retrieval treatment with protein synthesis inhibitors and many other compounds can lead to an amnestic state (Nadel et al. 2000b; Alberini 2005; Dudai 2006). These findings on reconsolidation fit with the behavioral evidence that retrieved memory is not a carbon copy of the initial experiences, and memories are updated during retrieval.
Genetics
Study of the genetics of human memory is in its infancy though many genes have been investigated for their association to memory in humans and non-human animals. A notable initial success was the association of
APOE
Apolipoprotein E (APOE) is a protein involved in the metabolism of fats in the body of mammals. A subtype is implicated in Alzheimer's disease and cardiovascular disease.
APOE belongs to a family of fat-binding proteins called apolipoproteins. ...
with memory dysfunction in
Alzheimer's disease. The search for genes associated with normally varying memory continues. One of the first candidates for normal variation in memory is the protein ''
KIBRA,'' which appears to be associated with the rate at which material is forgotten over a delay period. There has been some evidence that memories are stored in the nucleus of neurons.
Genetic underpinnings
Several genes, proteins and enzymes have been extensively researched for their association with memory. Long-term memory, unlike short-term memory, is dependent upon the synthesis of new proteins. This occurs within the cellular body, and concerns the particular transmitters, receptors, and new synapse pathways that reinforce the communicative strength between neurons. The production of new proteins devoted to synapse reinforcement is triggered after the release of certain signaling substances (such as calcium within hippocampal neurons) in the cell. In the case of hippocampal cells, this release is dependent upon the expulsion of magnesium (a binding molecule) that is expelled after significant and repetitive synaptic signaling. The temporary expulsion of magnesium frees NMDA receptors to release calcium in the cell, a signal that leads to gene transcription and the construction of reinforcing proteins. For more information, see
long-term potentiation
In neuroscience, long-term potentiation (LTP) is a persistent strengthening of synapses based on recent patterns of activity. These are patterns of synaptic activity that produce a long-lasting increase in signal transmission between two neurons ...
(LTP).
One of the newly synthesized proteins in LTP is also critical for maintaining long-term memory. This protein is an autonomously active form of the enzyme
protein kinase C (PKC), known as
PKMζ. PKMζ maintains the activity-dependent enhancement of synaptic strength and inhibiting
PKMζ erases established long-term memories, without affecting short-term memory or, once the inhibitor is eliminated, the ability to encode and store new long-term memories is restored. Also,
BDNF
Brain-derived neurotrophic factor (BDNF), or abrineurin, is a protein found in the and the periphery. that, in humans, is encoded by the ''BDNF'' gene. BDNF is a member of the neurotrophin family of growth factors, which are related to the canon ...
is important for the persistence of long-term memories.
The long-term stabilization of synaptic changes is also determined by a parallel increase of pre- and postsynaptic structures such as
axonal bouton,
dendritic spine
A dendritic spine (or spine) is a small membranous protrusion from a neuron's dendrite that typically receives input from a single axon at the synapse. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical s ...
and
postsynaptic density
The postsynaptic density (PSD) is a protein dense ''specialization'' attached to the postsynaptic membrane. PSDs were originally identified by electron microscopy as an electron-dense region at the membrane of a postsynaptic neuron. The PSD is in ...
.
[
]
On the molecular level, an increase of the postsynaptic scaffolding proteins
PSD-95
PSD-95 (postsynaptic density protein 95) also known as SAP-90 (synapse-associated protein 90) is a protein that in humans is encoded by the ''DLG4'' (discs large homolog 4) gene.
PSD-95 is a member of the membrane-associated guanylate kinase (MA ...
and
HOMER1
Homer protein homolog 1 or Homer1 is a neuronal protein that in humans is encoded by the ''HOMER1'' gene. Other names are Vesl and PSD-Zip45.
Structure
Homer1 protein has an N-terminal EVH1 domain, involved in protein interaction, and a C-te ...
c has been shown to correlate with the stabilization of synaptic enlargement.
The cAMP response element-binding protein (
CREB
CREB-TF (CREB, cAMP response element-binding protein) is a cellular transcription factor. It binds to certain DNA sequences called cAMP response elements (CRE), thereby increasing or decreasing the transcription of the genes. CREB was first de ...
) is a
transcription factor
In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The f ...
which is believed to be important in consolidating short-term to long-term memories, and which is believed to be downregulated in
Alzheimer's disease.
DNA methylation and
demethylation Demethylation is the chemical process resulting in the removal of a methyl group (CH3) from a molecule. A common way of demethylation is the replacement of a methyl group by a hydrogen atom, resulting in a net loss of one carbon and two hydrogen at ...
Rats exposed to an intense
learning event may retain a life-long memory of the event, even after a single training session. The long-term memory of such an event appears to be initially stored in the
hippocampus
The hippocampus (via Latin from Greek , ' seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, ...
, but this storage is transient. Much of the long-term storage of the memory seems to take place in the
anterior cingulate cortex
In the human brain
The human brain is the central organ (anatomy), organ of the human nervous system, and with the spinal cord makes up the central nervous system. The brain consists of the cerebrum, the brainstem and the cerebellum. It c ...
. When such an exposure was experimentally applied, more than 5,000 differently methylated DNA regions appeared in the
hippocampus
The hippocampus (via Latin from Greek , ' seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, ...
neuron
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
al
genome
In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...
of the rats at one and at 24 hours after training. These alterations in methylation pattern occurred at many
gene
In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s that were
down-regulated, often due to the formation of new
5-methylcytosine sites in CpG rich regions of the genome. Furthermore, many other genes were
upregulated
In the biological context of organisms' production of gene products, downregulation is the process by which a cell decreases the quantity of a cellular component, such as RNA or protein, in response to an external stimulus. The complementary pro ...
, likely often due to hypomethylation. Hypomethylation often results from the removal of methyl groups from previously existing
5-methylcytosines in DNA. Demethylation is carried out by several proteins acting in concert, including the
TET enzymes
The TET enzymes are a family of ten-eleven translocation (TET) methylcytosine dioxygenases. They are instrumental in DNA demethylation. 5-Methylcytosine (see first Figure) is a methylated form of the DNA base cytosine (C) that often regulates ge ...
as well as enzymes of the DNA
base excision repair
Base excision repair (BER) is a cellular mechanism, studied in the fields of biochemistry and genetics, that repairs damaged DNA throughout the cell cycle. It is responsible primarily for removing small, non-helix-distorting base lesions from t ...
pathway (see
Epigenetics in learning and memory
While the cellular and molecular mechanisms of learning and memory have long been a central focus of neuroscience, it is only in recent years that attention has turned to the epigenetic mechanisms behind the dynamic changes in gene transcription r ...
). The pattern of induced and repressed genes in brain neurons subsequent to an intense learning event likely provides the molecular basis for a long-term memory of the event.
Epigenetics
Studies of the molecular basis for memory formation indicate that
epigenetic mechanisms operating in
brain
A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...
neuron
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
s play a central role in determining this capability. Key epigenetic mechanisms involved in memory include the
methylation and
demethylation Demethylation is the chemical process resulting in the removal of a methyl group (CH3) from a molecule. A common way of demethylation is the replacement of a methyl group by a hydrogen atom, resulting in a net loss of one carbon and two hydrogen at ...
of neuronal DNA, as well as modifications of
histone
In biology, histones are highly basic proteins abundant in lysine and arginine residues that are found in eukaryotic cell nuclei. They act as spools around which DNA winds to create structural units called nucleosomes. Nucleosomes in turn a ...
proteins including
methylations,
acetylations and deacetylations.
Stimulation of brain activity in memory formation is often accompanied by the generation of
damage in neuronal DNA that is followed by repair associated with persistent epigenetic alterations. In particular the DNA repair processes of
non-homologous end joining
Non-homologous end joining (NHEJ) is a pathway that repairs double-strand breaks in DNA. NHEJ is referred to as "non-homologous" because the break ends are directly ligated without the need for a homologous template, in contrast to homology direc ...
and
base excision repair
Base excision repair (BER) is a cellular mechanism, studied in the fields of biochemistry and genetics, that repairs damaged DNA throughout the cell cycle. It is responsible primarily for removing small, non-helix-distorting base lesions from t ...
are employed in memory formation.
Role of DNA Topoisomerase II Beta in learning and memory
During a new
learning experience, a set of
gene
In biology, the word gene (from , ; "... Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a b ...
s is rapidly expressed in the
brain
A brain is an organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as vision. It is the most complex organ in a ve ...
. This induced
gene expression is considered to be essential for processing the information being learned. Such genes are referred to as
immediate early gene
Immediate early genes (IEGs) are genes which are activated transiently and rapidly in response to a wide variety of cellular stimuli. They represent a standing response mechanism that is activated at the transcription level in the first round of ...
s (IEGs).
DNA Topoisomerase II Beta (TOP2B) activity is essential for the expression of IEGs in a type of learning experience in mice termed associative fear memory.
[ ] Such a learning experience appears to rapidly trigger TOP2B to induce double-strand breaks in the
promoter DNA of IEG genes that function in
neuroplasticity
Neuroplasticity, also known as neural plasticity, or brain plasticity, is the ability of neural networks in the brain to change through growth and reorganization. It is when the brain is rewired to function in some way that differs from how it p ...
.
Repair
The technical meaning of maintenance involves functional checks, servicing, repairing or replacing of necessary devices, equipment, machinery, building infrastructure, and supporting utilities in industrial, business, and residential installa ...
of these induced breaks is associated with
DNA demethylation
For molecular biology in mammals, DNA demethylation causes replacement of 5-methylcytosine (5mC) in a DNA sequence by cytosine (C) (see figure of 5mC and C). DNA demethylation can occur by an active process at the site of a 5mC in a DNA sequenc ...
of IEG gene promoters allowing immediate expression of these IEG genes.
[
The double-strand breaks that are induced during a learning experience are not immediately repaired. About 600 regulatory sequences in promoters and about 800 regulatory sequences in ]enhancers
In genetics, an enhancer is a short (50–1500 bp) region of DNA that can be bound by proteins ( activators) to increase the likelihood that transcription of a particular gene will occur. These proteins are usually referred to as transcriptio ...
appear to depend on double strand breaks initiated by topoisomerase 2-beta (TOP2B) for activation. The induction of particular double-strand breaks are specific with respect to their inducing signal. When neurons are activated ''in vitro'', just 22 of TOP2B-induced double-strand breaks occur in their genomes.
Such TOP2B-induced double-strand breaks are accompanied by at least four enzymes of the non-homologous end joining (NHEJ) DNA repair pathway (DNA-PKcs, KU70, KU80, and DNA LIGASE IV) (see Figure). These enzymes repair the double-strand breaks within about 15 minutes to two hours. The double-strand breaks in the promoter are thus associated with TOP2B and at least these four repair enzymes. These proteins are present simultaneously on a single promoter nucleosome (there are about 147 nucleotides in the DNA sequence wrapped around a single nucleosome) located near the transcription start site of their target gene.
The double-strand break introduced by TOP2B apparently frees the part of the promoter at an RNA polymerase-bound transcription start site
Transcription is the process of copying a segment of DNA into RNA. The segments of DNA transcribed into RNA molecules that can encode proteins are said to produce messenger RNA (mRNA). Other segments of DNA are copied into RNA molecules calle ...
to physically move to its associated enhancer (see regulatory sequence
A regulatory sequence is a segment of a nucleic acid molecule which is capable of increasing or decreasing the expression of specific genes within an organism. Regulation of gene expression is an essential feature of all living organisms and v ...
). This allows the enhancer, with its bound transcription factor
In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The f ...
s and mediator proteins, to directly interact with the RNA polymerase paused at the transcription start site
Transcription is the process of copying a segment of DNA into RNA. The segments of DNA transcribed into RNA molecules that can encode proteins are said to produce messenger RNA (mRNA). Other segments of DNA are copied into RNA molecules calle ...
to start transcription
Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including:
Genetics
* Transcription (biology), the copying of DNA into RNA, the fir ...
.
Contextual fear conditioning in the mouse causes the mouse to have a long-term memory and fear of the location in which it occurred. Contextual fear conditioning causes hundreds of DSBs in mouse brain medial prefrontal cortex (mPFC) and hippocampus neurons (see Figure: Brain regions involved in memory formation). These DSBs predominately activate genes involved in synaptic processes, that are important for learning and memory.
In infancy
Up until the mid-1980s it was assumed that infants could not encode, retain, and retrieve information. A growing body of research now indicates that infants as young as 6-months can recall information after a 24-hour delay. Furthermore, research has revealed that as infants grow older they can store information for longer periods of time; 6-month-olds can recall information after a 24-hour period, 9-month-olds after up to five weeks, and 20-month-olds after as long as twelve months. In addition, studies have shown that with age, infants can store information faster. Whereas 14-month-olds can recall a three-step sequence after being exposed to it once, 6-month-olds need approximately six exposures in order to be able to remember it.
Although 6-month-olds can recall information over the short-term, they have difficulty recalling the temporal order of information. It is only by 9 months of age that infants can recall the actions of a two-step sequence in the correct temporal order – that is, recalling step 1 and then step 2. In other words, when asked to imitate a two-step action sequence (such as putting a toy car in the base and pushing in the plunger to make the toy roll to the other end), 9-month-olds tend to imitate the actions of the sequence in the correct order (step 1 and then step 2). Younger infants (6-month-olds) can only recall one step of a two-step sequence. Researchers have suggested that these age differences are probably due to the fact that the dentate gyrus
The dentate gyrus (DG) is part of the hippocampal formation in the temporal lobe of the brain, which also includes the hippocampus and the subiculum. The dentate gyrus is part of the hippocampal trisynaptic circuit and is thought to contribute t ...
of the hippocampus and the frontal components of the neural network are not fully developed at the age of 6-months.
In fact, the term 'infantile amnesia' refers to the phenomenon of accelerated forgetting during infancy. Importantly, infantile amnesia is not unique to humans, and preclinical research (using rodent models) provides insight into the precise neurobiology of this phenomenon. A review of the literature from behavioral neuroscientist Dr Jee Hyun Kim suggests that accelerated forgetting during early life is at least partly due to rapid growth of the brain during this period.
Aging
One of the key concerns of older adults is the experience of memory loss
Amnesia is a deficit in memory caused by brain damage or disease,Gazzaniga, M., Ivry, R., & Mangun, G. (2009) Cognitive Neuroscience: The biology of the mind. New York: W.W. Norton & Company. but it can also be caused temporarily by the use o ...
, especially as it is one of the hallmark symptoms of Alzheimer's disease. However, memory loss is qualitatively different in normal aging from the kind of memory loss associated with a diagnosis of Alzheimer's (Budson & Price, 2005). Research has revealed that individuals' performance on memory tasks that rely on frontal regions declines with age. Older adults tend to exhibit deficits on tasks that involve knowing the temporal order in which they learned information; source memory tasks that require them to remember the specific circumstances or context in which they learned information; and prospective memory tasks that involve remembering to perform an act at a future time. Older adults can manage their problems with prospective memory by using appointment books, for example.
Gene transcription
Transcription refers to the process of converting sounds (voice, music etc.) into letters or musical notes, or producing a copy of something in another medium, including:
Genetics
* Transcription (biology), the copying of DNA into RNA, the fir ...
profiles were determined for the human frontal cortex of individuals from age 26 to 106 years. Numerous genes were identified with reduced expression after age 40, and especially after age 70. Genes that play central roles in memory and learning were among those showing the most significant reduction with age. There was also a marked increase in DNA damage
DNA repair is a collection of processes by which a cell identifies and corrects damage to the DNA molecules that encode its genome. In human cells, both normal metabolic activities and environmental factors such as radiation can cause DNA d ...
, likely oxidative damage
Oxidative stress reflects an imbalance between the systemic manifestation of reactive oxygen species and a biological system's ability to readily detoxify the reactive intermediates or to repair the resulting damage. Disturbances in the normal r ...
, in the promoters of those genes with reduced expression. It was suggested that DNA damage may reduce the expression of selectively vulnerable genes involved in memory and learning.
Disorders
Much of the current knowledge of memory has come from studying ''memory disorders'', particularly loss of memory, known as amnesia. Amnesia can result from extensive damage to: (a) the regions of the medial temporal lobe, such as the hippocampus, dentate gyrus, subiculum, amygdala, the parahippocampal, entorhinal, and perirhinal cortices or the (b) midline diencephalic region, specifically the dorsomedial nucleus of the thalamus and the mammillary bodies of the hypothalamus. There are many sorts of amnesia, and by studying their different forms, it has become possible to observe apparent defects in individual sub-systems of the brain's memory systems, and thus hypothesize their function in the normally working brain. Other neurological
Neurology (from el, νεῦρον (neûron), "string, nerve" and the suffix -logia, "study of") is the branch of medicine dealing with the diagnosis and treatment of all categories of conditions and disease involving the brain, the spinal c ...
disorders such as Alzheimer's disease and Parkinson's disease
Parkinson's disease (PD), or simply Parkinson's, is a long-term degenerative disorder of the central nervous system that mainly affects the motor system. The symptoms usually emerge slowly, and as the disease worsens, non-motor symptoms becom ...
can also affect memory and cognition.
Hyperthymesia
Hyperthymesia, also known as hyperthymestic syndrome or highly superior autobiographical memory (HSAM), is a condition that leads people to be able to remember an abnormally large number of their life experiences in vivid detail. It is extraordi ...
, or hyperthymesic syndrome, is a disorder that affects an individual's autobiographical memory, essentially meaning that they cannot forget small details that otherwise would not be stored. Korsakoff's syndrome, also known as Korsakoff's psychosis, amnesic-confabulatory syndrome, is an organic brain disease that adversely affects memory by widespread loss or shrinkage of neurons within the prefrontal cortex.
While not a disorder, a common ''temporary'' failure of word retrieval from memory is the tip-of-the-tongue
Tip of the tongue (also known as ''lethologica'') is the phenomenon of failing to retrieve a word or term from memory, combined with partial recall and the feeling that retrieval is imminent. The phenomenon's name comes from the saying, "It's on ...
phenomenon
A phenomenon ( : phenomena) is an observable event. The term came into its modern philosophical usage through Immanuel Kant, who contrasted it with the noumenon, which ''cannot'' be directly observed. Kant was heavily influenced by Gottfried ...
. Those with Anomic aphasia
Anomic aphasia (also known as dysnomia, nominal aphasia, and amnesic aphasia) is a mild, fluent type of aphasia where individuals have word retrieval failures and cannot express the words they want to say (particularly nouns and verbs). By contra ...
(also called Nominal aphasia or Anomia), however, do experience the tip-of-the-tongue phenomenon on an ongoing basis due to damage to the frontal and parietal lobes of the brain
The lobes of the brain are the major identifiable zones of the cerebral cortex, and they comprise the surface of each hemisphere of the cerebrum. The two hemispheres are roughly symmetrical in structure, and are connected by the corpus callosum. ...
.
Memory dysfunction can also occur after viral infections. Many patients recovering from COVID-19 experience memory lapses. Other viruses can also elicit memory dysfunction, including SARS-CoV-1
Severe acute respiratory syndrome coronavirus 1 (SARS-CoV-1; or Severe acute respiratory syndrome coronavirus, SARS-CoV) is a strain of coronavirus that causes severe acute respiratory syndrome (SARS), the respiratory illness responsible for ...
, MERS-CoV
''Middle East respiratory syndrome–related coronavirus'' (''MERS-CoV''), or EMC/2012 ( HCoV-EMC/2012), is the virus that causes Middle East respiratory syndrome (MERS). It is a species of coronavirus which infects humans, bats, and camels. Th ...
, Ebola virus
''Zaire ebolavirus'', more commonly known as Ebola virus (; EBOV), is one of six known species within the genus '' Ebolavirus''. Four of the six known ebolaviruses, including EBOV, cause a severe and often fatal hemorrhagic fever in humans and o ...
and even influenza virus
''Orthomyxoviridae'' (from Greek ὀρθός, ''orthós'' 'straight' + μύξα, ''mýxa'' 'mucus') is a family of negative-sense RNA viruses. It includes seven genera: ''Alphainfluenzavirus'', ''Betainfluenzavirus'', '' Gammainfluenzavirus'', ' ...
.
Influencing factors
Interference can hamper memorization and retrieval. There is retroactive interference, when learning new information makes it harder to recall old information and proactive interference, where prior learning disrupts recall of new information. Although interference can lead to forgetting, it is important to keep in mind that there are situations when old information can facilitate learning of new information. Knowing Latin, for instance, can help an individual learn a related language such as French – this phenomenon is known as positive transfer.
Stress
Stress has a significant effect on memory formation and learning. In response to stressful situations, the brain releases hormones and neurotransmitters (ex. glucocorticoids and catecholamines) which affect memory encoding processes in the hippocampus. Behavioural research on animals shows that chronic stress produces adrenal hormones which impact the hippocampal structure in the brains of rats. An experimental study by German cognitive psychologists L. Schwabe and O. Wolf demonstrates how learning under stress also decreases memory recall in humans. In this study, 48 healthy female and male university students participated in either a stress test or a control group. Those randomly assigned to the stress test group had a hand immersed in ice cold water (the reputable SECPT or 'Socially Evaluated Cold Pressor Test') for up to three minutes, while being monitored and videotaped. Both the stress and control groups were then presented with 32 words to memorize. Twenty-four hours later, both groups were tested to see how many words they could remember (free recall) as well as how many they could recognize from a larger list of words (recognition performance). The results showed a clear impairment of memory performance in the stress test group, who recalled 30% fewer words than the control group. The researchers suggest that stress experienced during learning distracts people by diverting their attention during the memory encoding process.
However, memory performance can be enhanced when material is linked to the learning context, even when learning occurs under stress. A separate study by cognitive psychologists Schwabe and Wolf shows that when retention testing is done in a context similar to or congruent with the original learning task (i.e., in the same room), memory impairment and the detrimental effects of stress on learning can be attenuated. Seventy-two healthy female and male university students, randomly assigned to the SECPT stress test or to a control group, were asked to remember the locations of 15 pairs of picture cards – a computerized version of the card game "Concentration" or "Memory". The room in which the experiment took place was infused with the scent of vanilla, as odour is a strong cue for memory. Retention testing took place the following day, either in the same room with the vanilla scent again present, or in a different room without the fragrance. The memory performance of subjects who experienced stress during the object-location task decreased significantly when they were tested in an unfamiliar room without the vanilla scent (an incongruent context); however, the memory performance of stressed subjects showed no impairment when they were tested in the original room with the vanilla scent (a congruent context). All participants in the experiment, both stressed and unstressed, performed faster when the learning and retrieval contexts were similar.
This research on the effects of stress on memory may have practical implications for education, for eyewitness testimony
Eyewitness testimony is the account a bystander or victim gives in the courtroom, describing what that person observed that occurred during the specific incident under investigation. Ideally this recollection of events is detailed; however, this is ...
and for psychotherapy: students may perform better when tested in their regular classroom rather than an exam room, eyewitnesses may recall details better at the scene of an event than in a courtroom, and persons with post-traumatic stress
Post-traumatic stress disorder (PTSD) is a mental and behavioral disorder that can develop because of exposure to a traumatic event, such as sexual assault, warfare, traffic collisions, child abuse, domestic violence, or other threats on a ...
may improve when helped to situate their memories of a traumatic event in an appropriate context.
Stressful life experiences may be a cause of memory loss as a person ages. Glucocorticoids
Glucocorticoids (or, less commonly, glucocorticosteroids) are a class of corticosteroids, which are a class of steroid hormones. Glucocorticoids are corticosteroids that bind to the glucocorticoid receptor that is present in almost every vertebr ...
that are released during stress cause damage to neurons that are located in the hippocampal
The hippocampus (via Latin from Greek , 'seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, an ...
region of the brain. Therefore, the more stressful situations that someone encounters, the more susceptible they are to memory loss later on. The CA1 neurons found in the hippocampus are destroyed due to glucocorticoids decreasing the release of glucose and the reuptake of glutamate. This high level of extracellular glutamate allows calcium to enter NMDA receptors
The ''N''-methyl-D-aspartate receptor (also known as the NMDA receptor or NMDAR), is a glutamate receptor and ion channel found in neurons. The NMDA receptor is one of three types of ionotropic glutamate receptors, the other two being AMPA and ...
which in return kills neurons. Stressful life experiences can also cause repression of memories where a person moves an unbearable memory to the unconscious mind. This directly relates to traumatic events in one's past such as kidnappings, being prisoners of war or sexual abuse as a child.
The more long term the exposure to stress is, the more impact it may have. However, short term exposure to stress also causes impairment in memory by interfering with the function of the hippocampus. Research shows that subjects placed in a stressful situation for a short amount of time still have blood glucocorticoid levels that have increased drastically when measured after the exposure is completed. When subjects are asked to complete a learning task after short term exposure they often have difficulties. Prenatal stress also hinders the ability to learn and memorize by disrupting the development of the hippocampus and can lead to unestablished long term potentiation in the offspring of severely stressed parents. Although the stress is applied prenatally, the offspring show increased levels of glucocorticoids when they are subjected to stress later on in life. One explanation for why children from lower socioeconomic backgrounds tend to display poorer memory performance than their higher-income peers is the effects of stress accumulated over the course of the lifetime. The effects of low income on the developing hippocampus is also thought be mediated by chronic stress responses which may explain why children from lower and higher-income backgrounds differ in terms of memory performance.
Sleep
Making memories occurs through a three-step process, which can be enhanced by sleep
Sleep is a sedentary state of mind and body. It is characterized by altered consciousness, relatively inhibited sensory activity, reduced muscle activity and reduced interactions with surroundings. It is distinguished from wakefulness by a de ...
. The three steps are as follows:
# Acquisition which is the process of storage and retrieval of new information in memory
# Consolidation
# Recall
Sleep affects memory consolidation. During sleep, the neural connections in the brain are strengthened. This enhances the brain's abilities to stabilize and retain memories. There have been several studies which show that sleep improves the retention of memory, as memories are enhanced through active consolidation. System consolidation takes place during slow-wave sleep (SWS). This process implicates that memories are reactivated during sleep, but that the process doesn't enhance every memory. It also implicates that qualitative changes are made to the memories when they are transferred to long-term store during sleep. During sleep, the hippocampus replays the events of the day for the neocortex. The neocortex then reviews and processes memories, which moves them into long-term memory. When one does not get enough sleep it makes it more difficult to learn as these neural connections are not as strong, resulting in a lower retention rate of memories. Sleep deprivation makes it harder to focus, resulting in inefficient learning. Furthermore, some studies have shown that sleep deprivation can lead to false memories as the memories are not properly transferred to long-term memory.
One of the primary functions of sleep is thought to be the improvement of the consolidation of information, as several studies have demonstrated that memory depends on getting sufficient sleep between training and test. Additionally, data obtained from neuroimaging studies have shown activation patterns in the sleeping brain that mirror those recorded during the learning of tasks from the previous day, suggesting that new memories may be solidified through such rehearsal.
Construction for general manipulation
Although people often think that memory operates like recording equipment, this is not the case. The molecular mechanisms underlying the induction and maintenance of memory are very dynamic and comprise distinct phases covering a time window from seconds to even a lifetime. In fact, research has revealed that our memories are constructed: "current hypotheses suggest that constructive processes allow individuals to simulate and imagine future episodes, happenings, and scenarios. Since the future is not an exact repetition of the past, simulation of future episodes requires a complex system that can draw on the past in a manner that flexibly extracts and recombines elements of previous experiences – a constructive rather than a reproductive system." People can construct their memories when they encode them and/or when they recall them. To illustrate, consider a classic study conducted by Elizabeth Loftus and John Palmer (1974) in which people were instructed to watch a film of a traffic accident and then asked about what they saw. The researchers found that the people who were asked, "How fast were the cars going when they ''smashed'' into each other?" gave higher estimates than those who were asked, "How fast were the cars going when they ''hit'' each other?" Furthermore, when asked a week later whether they had seen broken glass in the film, those who had been asked the question with ''smashed'' were twice more likely to report that they had seen broken glass than those who had been asked the question with ''hit'' (there was no broken glass depicted in the film). Thus, the wording of the questions distorted viewers' memories of the event. Importantly, the wording of the question led people to construct different memories of the event – those who were asked the question with ''smashed'' recalled a more serious car accident than they had actually seen. The findings of this experiment were replicated around the world, and researchers consistently demonstrated that when people were provided with misleading information they tended to misremember, a phenomenon known as the misinformation effect
The misinformation effect occurs when a person's recall of episodic memories becomes less accurate because of post-event information. The misinformation effect has been studied since the mid-1970s. Elizabeth Loftus is one of the most influential re ...
.
Research has revealed that asking individuals to repeatedly imagine actions that they have never performed or events that they have never experienced could result in false memories. For instance, Goff and Roediger (1998) asked participants to imagine that they performed an act (e.g., break a toothpick) and then later asked them whether they had done such a thing. Findings revealed that those participants who repeatedly imagined performing such an act were more likely to think that they had actually performed that act during the first session of the experiment. Similarly, Garry and her colleagues (1996) asked college students to report how certain they were that they experienced a number of events as children (e.g., broke a window with their hand) and then two weeks later asked them to imagine four of those events. The researchers found that one-fourth of the students asked to imagine the four events reported that they had actually experienced such events as children. That is, when asked to imagine the events they were more confident that they experienced the events.
Research reported in 2013 revealed that it is possible to artificially stimulate prior memories and artificially implant false memories in mice. Using optogenetics
Optogenetics is a biological technique to control the activity of neurons or other cell types with light. This is achieved by expression of light-sensitive ion channels, pumps or enzymes specifically in the target cells. On the level of individ ...
, a team of RIKEN-MIT scientists caused the mice to incorrectly associate a benign environment with a prior unpleasant experience from different surroundings. Some scientists believe that the study may have implications in studying false memory formation in humans, and in treating PTSD and schizophrenia
Schizophrenia is a mental disorder characterized by continuous or relapsing episodes of psychosis. Major symptoms include hallucinations (typically hearing voices), delusions, and disorganized thinking. Other symptoms include social wit ...
.
Memory reconsolidation is when previously consolidated memories are recalled or retrieved from long-term memory to your active consciousness. During this process, memories can be further strengthened and added to but there is also risk of manipulation involved. We like to think of our memories as something stable and constant when they are stored in long-term memory but this isn't the case. There are a large number of studies that found that consolidation of memories is not a singular event but are put through the process again, known as reconsolidation. This is when a memory is recalled or retrieved and placed back into your working memory. The memory is now open to manipulation from outside sources and the misinformation effect which could be due to misattributing the source of the inconsistent information, with or without an intact original memory trace (Lindsay and Johnson, 1989). One thing that can be sure is that memory is malleable.
This new research into the concept of reconsolidation has opened the door to methods to help those with unpleasant memories or those that struggle with memories. An example of this is if you had a truly frightening experience and recall that memory in a less arousing environment, the memory will be weaken the next time it is retrieved. "Some studies suggest that over-trained or strongly reinforced memories do not undergo reconsolidation if reactivated the first few days after training, but do become sensitive to reconsolidation interference with time." This, however does not mean that all memory is susceptible to reconsolidation. There is evidence to suggest that memory that has undergone strong training and whether or not is it intentional is less likely to undergo reconsolidation. There was further testing done with rats and mazes that showed that reactivated memories were more susceptible to manipulation, in both good and bad ways, than newly formed memories. It is still not known whether or not these are new memories formed and it's an inability to retrieve the proper one for the situation or if it's a reconsolidated memory. Because the study of reconsolidation is still a newer concept, there is still debate on whether it should be considered scientifically sound.
Improving
A UCLA research study published in the June 2008 issue of the ''American Journal of Geriatric Psychiatry'' found that people can improve cognitive function
Cognitive skills, also called cognitive functions, cognitive abilities or cognitive capacities, are brain-based skills which are needed in acquisition of knowledge, manipulation of information and reasoning. They have more to do with the mechanisms ...
and brain efficiency through simple lifestyle changes such as incorporating memory exercises, healthy eating
A healthy diet is a diet that maintains or improves overall health. A healthy diet provides the body with essential nutrition: fluid, macronutrients such as protein, micronutrients such as vitamins, and adequate fibre and food energy.
A h ...
, physical fitness and stress reduction into their daily lives. This study examined 17 subjects, (average age 53) with normal memory performance.
Eight subjects were asked to follow a "brain healthy" diet, relaxation, physical, and mental exercise (brain teasers and verbal memory training techniques). After 14 days, they showed greater word fluency (not memory) compared to their baseline performance. No long-term follow-up was conducted; it is therefore unclear if this intervention has lasting effects on memory.
There are a loosely associated group of mnemonic principles and techniques that can be used to vastly improve memory known as the art of memory
The art of memory (Latin: ''ars memoriae'') is any of a number of loosely associated mnemonic principles and techniques used to organize memory impressions, improve recall, and assist in the combination and 'invention' of ideas. An alternative ...
.
The International Longevity Center
The Joseph L. Mailman School of Public Health is the public health graduate school of Columbia University. Located on the Columbia University Medical Center campus in the Washington Heights neighborhood of Manhattan, New York City, the school ...
released in 2001 a report which includes in pages 14–16 recommendations for keeping the mind in good functionality until advanced age. Some of the recommendations are:
* to stay intellectually active through learning
* training or reading
* to keep physically active so to promote blood circulation to the brain
* to socialize
* to reduce stress
* to keep sleep time regular
* to avoid depression or emotional instability
* to observe good nutrition.
Memorization
Memorization is the process of committing something to memory. It is a mental process undertaken in order to store in memory for later recall visual, auditory, or tactical information.
The scientific study of memory is part of cognitive neuros ...
is a method of learning that allows an individual to recall information verbatim. Rote learning
Rote learning is a memorization technique based on repetition. The method rests on the premise that the recall of repeated material becomes faster the more one repeats it. Some of the alternatives to rote learning include meaningful learning, ...
is the method most often used. Methods of memorizing things have been the subject of much discussion over the years with some writers, such as Cosmos Rossellius using visual alphabets
An alphabet is a standardized set of basic written graphemes (called letters) that represent the phonemes of certain spoken languages. Not all writing systems represent language in this way; in a syllabary, each character represents a syllab ...
. The spacing effect
The spacing effect demonstrates that learning is more effective when study sessions are spaced out. This effect shows that more information is encoded into long-term memory by spaced study sessions, also known as ''spaced repetition'' or ''spac ...
shows that an individual is more likely to remember a list of items when rehearsal is spaced over an extended period of time. In contrast to this is cramming: an intensive memorization in a short period of time. the spacing effect is exploited to improve memory in spaced repetition
Spaced repetition is an evidence-based learning technique that is usually performed with flashcards. Newly introduced and more difficult flashcards are shown more frequently, while older and less difficult flashcards are shown less frequently i ...
flashcard training. Also relevant is the Zeigarnik effect
Named after Lithuanian-Soviet psychologist Bluma Zeigarnik, in psychology the Zeigarnik effect occurs when an activity that has been interrupted may be more readily recalled. It postulates that people remember unfinished or interrupted tasks bett ...
which states that people remember uncompleted or interrupted tasks better than completed ones. The so-called Method of loci
The method of loci is a strategy for memory enhancement, which uses visualizations of familiar spatial environments in order to enhance the recall of information. The method of loci is also known as the memory journey, memory palace, journey m ...
uses spatial memory to memorize non-spatial information.
In plants
Plants lack a specialized organ devoted to memory retention, so plant memory has been a controversial topic in recent years. New advances in the field have identified the presence of neurotransmitters in plants, adding to the hypothesis that plants are capable of remembering. Action potential
An action potential occurs when the membrane potential of a specific cell location rapidly rises and falls. This depolarization then causes adjacent locations to similarly depolarize. Action potentials occur in several types of animal cells, ...
s, a physiological response characteristic of neuron
A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. ...
s, have been shown to have an influence on plants as well, including in wound responses and photosynthesis
Photosynthesis is a process used by plants and other organisms to convert light energy into chemical energy that, through cellular respiration, can later be released to fuel the organism's activities. Some of this chemical energy is stored i ...
. In addition to these homologous features of memory systems in both plants and animals, plants have also been observed to encode, store and retrieve basic short-term memories.
One of the most well-studied plants to show rudimentary memory is the Venus flytrap
The Venus flytrap (''Dionaea muscipula'') is a carnivorous plant native to subtropical wetlands on the East Coast of the United States in North Carolina and South Carolina. It catches its prey—chiefly insects and arachnids—with a trapping ...
. Native to the subtropical wetlands of the eastern United States, Venus Fly Traps have evolved the ability to obtain meat for sustenance, likely due to the lack of nitrogen in the soil. This is done by two trap-forming leaf tips that snap shut once triggered by a potential prey. On each lobe, three triggers hairs await stimulation. In order to maximize the benefit to cost ratio, the plant enables a rudimentary form of memory in which two trigger hairs must be stimulated within 30 seconds in order to result in trap closure. This system ensures that the trap only closes when potential prey is within grasp.
The time lapse between trigger hair stimulations suggests that the plant can remember an initial stimulus long enough for a second stimulus to initiate trap closure. This memory isn't encoded in a brain, as plants lack this specialized organ. Rather, information is stored in the form of cytoplasmic calcium levels. The first trigger causes a subthreshold cytoplasmic calcium influx. This initial trigger isn't enough to activate trap closure, so a subsequent stimulus allows for a secondary influx of calcium. The latter calcium rise superimposes on the initial one, creating an action potential that passes threshold, resulting in trap closure. Researchers, to prove that an electrical threshold must be met to stimulate trap closure, excited a single trigger hair with a constant mechanical stimulus using Ag/AgCl electrodes. The trap closed after only a few seconds. This experiment gave evidence to demonstrate that the electrical threshold, not necessarily the number of trigger hair stimulations, was the contributing factor in Venus Fly Trap memory. It has been shown that trap closure can be blocked using uncouplers and inhibitors of voltage-gated channels. After trap closure, these electrical signals stimulate glandular production of jasmonic acid and hydrolase
Hydrolase is a class of enzyme that commonly perform as biochemical catalysts that use water to break a chemical bond, which typically results in dividing a larger molecule into smaller molecules. Some common examples of hydrolase enzymes are este ...
s, allowing for digestion of the prey.
The field of plant neurobiology has gained a large amount of interest over the past decade, leading to an influx of research regarding plant memory. Although the Venus flytrap is one of the more highly studied, many other plants exhibit the capacity to remember, including the ''Mimosa pudica
''Mimosa pudica'' (from la, pudica "shy, bashful or shrinking"; also called sensitive plant, sleepy plant, action plant, touch-me-not, shameplant) is a creeping annual or perennial flowering plant of the pea/legume family Fabaceae. It is often g ...
'' through an experiment conducted by Monica Gagliano and colleagues in 2013. To study the ''Mimosa pudica
''Mimosa pudica'' (from la, pudica "shy, bashful or shrinking"; also called sensitive plant, sleepy plant, action plant, touch-me-not, shameplant) is a creeping annual or perennial flowering plant of the pea/legume family Fabaceae. It is often g ...
,'' Gagliano designed an apparatus with which potted mimosa plants could be repeatedly dropped the same distance and at the same speed. It was observed that the plants defensive response of curling up its leaves decreased over the 60 times the experiment was repeated per plant. To confirm that this was a mechanism of memory rather than exhaustion, some of the plants were shaken post experiment and displayed normal defensive responses of leaf curling. This experiment also demonstrated long-term memory in the plants, as it was repeated a month later and the plants were observed to remain unfazed by the dropping.
See also
* Adaptive memory
Adaptive memory is the study of memory systems that have evolved to help retain survival- and fitness-related information, i.e., that are geared toward helping an organism enhance its reproductive fitness and chances of surviving. One key elemen ...
, memory systems that have evolved to help retain survival-and-fitness information
* Animal memory
* Body memory, hypothetical memory function of individual body parts or cells
* Collective memory
Collective memory refers to the shared pool of memories, knowledge and information of a social group that is significantly associated with the group's identity. The English phrase "collective memory" and the equivalent French phrase "la mémoire c ...
, memory that is shared, passed on, and constructed by a group
*Explicit memory
Explicit memory (or declarative memory) is one of the two main types of long-term human memory, the other of which is implicit memory. Explicit memory is the conscious, intentional recollection of factual information, previous experiences, and con ...
* False memory
In psychology, a false memory is a phenomenon where someone recalls something that did not happen or recalls it differently from the way it actually happened. Suggestibility, activation of associated information, the incorporation of misinformat ...
* Immunological memory
Immunological memory is the ability of the immune system to quickly and specifically recognize an antigen that the body has previously encountered and initiate a corresponding immune response. Generally, these are secondary, tertiary and other subs ...
, a characteristic of adaptive immunity
* Implicit memory
In psychology, implicit memory is one of the two main types of long-term human memory. It is acquired and used unconsciously, and can affect thoughts and behaviours. One of its most common forms is procedural memory, which allows people to perfo ...
, previous experiences help to perform a task with no awareness of those experiences
* Intermediate-term memory Intermediate-term memory (ITM) is a stage of memory distinct from sensory memory, working memory/short-term memory, and long-term memory. While sensory memory persists for several milliseconds, working memory persists for up to thirty seconds, and l ...
* Involuntary memory
Involuntary memory, also known as involuntary explicit memory, involuntary conscious memory, involuntary aware memory, madeleine moment, mind pops and most commonly, involuntary autobiographical memory, is a sub-component of memory that occurs when ...
* Long memory, a statistical property in which intertemporal dependence decays only slowly
* Long-term memory
Long-term memory (LTM) is the stage of the Atkinson–Shiffrin memory model in which informative knowledge is held indefinitely. It is defined in contrast to short-term and working memory, which persist for only about 18 to 30 seconds. Long- ...
, the ability of the brain to store and recover memories
* Method of loci
The method of loci is a strategy for memory enhancement, which uses visualizations of familiar spatial environments in order to enhance the recall of information. The method of loci is also known as the memory journey, memory palace, journey m ...
* Mnemonic major system
The major system (also called the phonetic number system, phonetic mnemonic system, or Herigone's mnemonic system) is a mnemonic technique used to aid in memorizing numbers.
The system works by converting numbers into consonants, then into words ...
* Photographic memory
Eidetic memory ( ; more commonly called photographic memory or total recall) is the ability to recall an image from memory with high precision—at least for a brief period of time—after seeing it only onceThe terms ''eidetic memory'' and ''pho ...
* Politics of memory Politics of memory is the organisation of collective memory by political agents; the political means by which events are remembered and recorded, or discarded. Eventually, politics of memory may determine the way history is written and passed on, he ...
* Prenatal memory
Prenatal memory, also called fetal memory, is important for the development of memory in humans. Many factors can impair fetal memory and its functions, primarily maternal actions. There are multiple techniques available not only to demonstrate th ...
* Procedural memory
Procedural memory is a type of implicit memory ( unconscious, long-term memory) which aids the performance of particular types of tasks without conscious awareness of these previous experiences.
Procedural memory guides the processes we perform ...
, a type most frequently below conscious awareness that helps perform particular types of action
* Sensory memory
During every moment of an organism's life, sensory information is being taken in by sensory receptors and processed by the nervous system. Sensory information is stored in sensory memory just long enough to be transferred to short-term memory. Hu ...
* Short-term memory
* Working memory
Working memory is a cognitive system with a limited capacity that can hold information temporarily. It is important for reasoning and the guidance of decision-making and behavior. Working memory is often used synonymously with short-term memory, ...
Notes
Sources
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* Eysenck, M.W. (2012). Fundamentals of cognition. New York: Psychology Press.
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*
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* Suzuki, W.A. (2007). "Working memory: Signals in the brain". In: Roediger, H.L., Dudai, Y. and Fitzpatrick S.M., eds. ''Science of Memory: Concepts''. New York: Oxford University Press, pp. 147–150.
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Further reading
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External links
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Memory-related resources
from the National Institutes of Health
The National Institutes of Health, commonly referred to as NIH (with each letter pronounced individually), is the primary agency of the United States government responsible for biomedical and public health research. It was founded in the late ...
{{Authority control
Neuropsychological assessment
Sources of knowledge
Mental processes