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It is important to note that this study involved only male subjects, which may be significant as sex steroid hormones may have different effects in response to cortisol administration.

Men and women also respond to emotional stimuli differently and this may affect cortisol levels. This was also the first Functional magnetic resonance imaging fMRI study done utilising glucocorticoids, therefore more research is necessary to further substantiate these findings.

It is believed that sleep plays an active role in consolidation of declarative memory. Specifically, sleep's unique properties enhance memory consolidation , such as the reactivation of newly learned memories during sleep.

For example, it has been suggested that the central mechanism for consolidation of declarative memory during sleep is the reactivation of hippocampal memory representations.

This reactivation transfers information to neocortical networks where it is integrated into long-term representations. Slow-wave sleep , often referred to as deep sleep, plays the most important role in consolidation of declarative memory and there is a large amount of evidence to support this claim.

One study found that the first 3. Additional hours of sleep do not add to the initial level of performance. Thus this study suggests that full sleep may not be important for optimal performance of memory.

However this is not the case for subjects who were tested for the second half of their sleep cycle, as they experience less SWS. Another key piece of evidence regarding SWS's involvement in declarative memory consolidation is a finding that people with pathological conditions of sleep, such as insomnia, exhibit both reduction in Slow-Wave Sleep and also have impaired consolidation of declarative memory during sleep.

This in turn indicated that SWS is associated with poor declarative memory consolidation but not with age itself.

Some researchers suggest that sleep spindle , a burst of brain activity occurring during stage 2 sleep, plays a role in boosting consolidation of declarative memories.

This supports the hypothesis that sleep spindle helps to consolidate recent memory traces but not memory performance in general.

There is a relatively small body of evidence that supports the idea that REM sleep helps consolidate highly emotional declarative memories.

For instance Wagner, et al. The view that sleep plays an active role in declarative memory consolidation is not shared by all researchers.

For instance Ellenbogen, et al. However, during sleep, when interference is minimal, memories can be consolidated without associative interference.

More research is needed to make a definite statement whether sleep creates favourable conditions for consolidation or it actively enhances declarative memory consolidation.

The encoding of explicit memory depends on conceptually driven, top-down processing, in which a subject reorganizes the data to store it.

The later recall of information is thus greatly influenced by the way in which the information was originally processed.

The depth-of-processing effect is the improvement in subsequent recall of an object about which a person has given thought to its meaning or shape.

Simply put: To create explicit memories, you have to do something with your experiences: think about them, talk about them, write them down, study them, etc.

The more you do, the better you will remember. Testing of information while learning has also shown to improve encoding in explicit memory.

If a student reads a text book and then tests themselves afterward, their semantic memory of what was read is improved.

This study — test method improves encoding of information. This Phenomenon is referred to as the Testing Effect. Retrieval : Because a person has played an active role in processing explicit information, the internal cues that were used in processing it can also be used to initiate spontaneous recall.

The conditions in which information is memorized can affect recall. If a person has the same surroundings or cues when the original information is presented, they are more likely to remember it.

This is referred to as encoding specificity and it also applies to explicit memory. In a study where subjects were asked to perform a cued recall task participants with a high working memory did better than participants with a low working memory when the conditions were maintained.

When the conditions were changed for recall both groups dropped. The subjects with higher working memory declined more.

Several neural structures are proposed to be involved in explicit memory. Most are in the temporal lobe or closely related to it, such as the amygdala , the hippocampus , the rhinal cortex in the temporal lobe, and the prefrontal cortex.

While the human brain is certainly regarded for its plasticity, there is some evidence that shows traumatic brain injury TBI in young children can have negative effects on explicit memory.

Researchers have looked at children with TBI in early childhood i. Findings showed that children with severe TBI in late childhood experienced impaired explicit memory while still maintaining implicit memory formation.

Researchers also found that children with severe TBI in early childhood had both increased chance of having both impaired explicit memory and implicit memory.

While children with severe TBI are at risk for impaired explicit memory, the chances of impaired explicit memory in adults with severe TBI is much greater.

Alzheimer's disease has a profound effect on explicit memory. Mild cognitive impairment is an early sign of Alzheimer's disease. People with memory conditions often receive cognitive training.

When an fMRI was used to view brain activity after training, it found increased activation in various neural systems that are involved with explicit memory.

However, if the task is presented repeatedly they can learn and retain some new knowledge of the task. This effect is more apparent if the information is familiar.

The person with Alzheimer's must also be guided through the task and prevented from making errors. This means that people with Alzheimer's have difficulty remembering where items are placed in unfamiliar environments.

The effects of Alzheimer's disease are seen in the episodic part of explicit memory. This can lead to problems with communication.

A study was conducted where Alzheimer's patients were asked to name a variety of objects from different periods. The results shown that their ability to name the object depended on frequency of use of the item and when the item was first acquired.

Alzheimer's patients have difficulty distinguishing between different melodies they have never heard before. People with Alzheimer's also have issues with picturing future events.

This is due to a deficit in episodic future thinking. Amnesiacs are frequently portrayed in television and movies. Some of the better-known examples include:.

Having lost her short term memory in a car crash, Lucy can only remember the current day's events until she falls asleep. When she wakes up the next morning, she has no recollection of the previous day's experiences.

Although this movie is not the most accurate representation of a true amnesic patient, it is useful for informing viewers of the detrimental effects of amnesia.

Memento a film inspired by the case of Henry Molaison H. Unlike most amnesiacs, Leonard retains his identity and the memories of events that occurred before the injury, but loses all ability to form new memories.

This loss of ability to form new memories indicates that the head injury affected the medial temporal lobe of the brain resulting in the inability for Leonard to form declarative memory.

Finding Nemo features a reef fish named Dory with an inability to develop declarative memory. This prevents her from learning or retaining any new information such as names or directions.

The exact origin of Dory's impairment is not mentioned in the film, but her memory loss accurately portrays the difficulties facing amnesiacs.

Submit link. Submit link Login Sign up Imprint. Long-Term Memory Subtype Description Example Declarative explicit Conscious memory of facts and events Semantic Factual information The capital of Germany is Berlin Episodic Specific personal experiences Your 10th birthday Non-declarative implicit Modes of learning that are nonconscious - one learning to perform a sequence of actions that do not necessarily invoke knowledge Priming also known as "pattern completion," where one has the ability to complete a pattern they have once seen before.

This priming differs from Psychology priming. If you were given a picture of half of a letter from the alphabet and you recognized which letter it is, you would be able to complete the letter.

Perceptual learning Perceptual ability to differentiate sensories through experience of stimuli Differentiating between categories such as smells, colors, tastes Category learning " The model of language Declarative and procedural memory fall into two categories of human language.

Procedural memory builds rule-governed structure merging or series of forms and representations into complex structures such as: phonology inflectional and derivational morphology compositional semantics the meaning of composition of words into complex structures syntax Broca and Wernicke's Brain Region Broca's area is important to procedural memory, for, "Broca's area is involved in the expressive aspects of spoken and written language production of sentences constrained by the rules of grammar and syntax.

History The study of human memory stretches back over the last years. Al , and proposes that the hippocampus does three things with episodic memory: Mediates the recording of episodic memories Identifies common features between episodes Links these common episodes in a memory space.

Amygdala as seen in red Amygdala The amygdala is believed to be involved in the encoding and retrieval of emotionally charged memories.

Hippocampal lesion studies The Morris water maze The Morris water navigation task tests spatial learning in rats. Amygdala lesion studies Adolph, Cahill and Schul completed a study showing that emotional arousal facilitates the encoding of material into long term declarative memory.

Episodic and semantic memory. In Organization of Memory, ed. E Tulving, W Donaldson, pp. Retrieved Reisberg, Daniel ed. The Oxford Handbook of Cognitive Psychology.

These reflect subthreshold membrane potentials and strongly modulate the spiking of hippocampal neurons and synchronise across the hippocampus in a travelling wave pattern.

From rodent studies it has been proposed that the trisynaptic circuit generates the hippocampal theta rhythm.

Theta rhythmicity is very obvious in rabbits and rodents and also clearly present in cats and dogs. Whether theta can be seen in primates is not yet clear.

An example would be the phase with which theta rhythms, at the time of stimulation of a neuron, shape the effect of that stimulation upon its synapses.

What is meant here is that theta rhythms may affect those aspects of learning and memory that are dependent upon synaptic plasticity.

During sleep or during resting, when an animal is not engaged with its surroundings, the hippocampal EEG shows a pattern of irregular slow waves, somewhat larger in amplitude than theta waves.

This pattern is occasionally interrupted by large surges called sharp waves. Sharp waves are most frequent during sleep when they occur at an average rate of around 1 per second in rats but in a very irregular temporal pattern.

Sharp waves are less frequent during inactive waking states and are usually smaller. Sharp waves have also been observed in humans and monkeys.

In macaques, sharp waves are robust but do not occur as frequently as in rats. One of the most interesting aspects of sharp waves is that they appear to be associated with memory.

Wilson and McNaughton , [88] and numerous later studies, reported that when hippocampal place cells have overlapping spatial firing fields and therefore often fire in near-simultaneity , they tend to show correlated activity during sleep following the behavioral session.

This enhancement of correlation, commonly known as reactivation , has been found to occur mainly during sharp waves.

Sharp waves in Hebbian theory are seen as persistently repeated stimulations by presynaptic cells, of postsynaptic cells that are suggested to drive synaptic changes in the cortical targets of hippocampal output pathways.

Since at least the time of Ramon y Cajal , psychologists have speculated that the brain stores memory by altering the strength of connections between neurons that are simultaneously active.

As a candidate mechanism for long-term memory , LTP has since been studied intensively, and a great deal has been learned about it.

However, the complexity and variety of the intracellular signalling cascades that can trigger LTP is acknowledged as preventing a more complete understanding.

The hippocampus is a particularly favorable site for studying LTP because of its densely packed and sharply defined layers of neurons, but similar types of activity-dependent synaptic change have also been observed in many other brain areas.

Genetically modified mice that are modified to disable the LTP mechanism, also generally show severe memory deficits. Age-related conditions such as Alzheimer's disease and other forms of dementia for which hippocampal disruption is one of the earliest signs [] have a severe impact on many types of cognition including memory.

Even normal aging is associated with a gradual decline in some types of memory, including episodic memory and working memory or short-term memory.

Because the hippocampus is thought to play a central role in memory, there has been considerable interest in the possibility that age-related declines could be caused by hippocampal deterioration.

There is, however, a reliable relationship between the size of the hippocampus and memory performance; so that where there is age-related shrinkage, memory performance will be impaired.

The hippocampus contains high levels of glucocorticoid receptors , which make it more vulnerable to long-term stress than most other brain areas.

Chronic stress resulting in elevated levels of glucocorticoids , notably of cortisol , is seen to be a cause of neuronal atrophy in the hippocampus.

Another factor that contributes to a smaller hippocampal volume is that of dendritic retraction where dendrites are shortened in length and reduced in number, in response to increased glucocorticoids.

This dendritic retraction is reversible. There is, however, evidence derived mainly from studies using rats that stress occurring shortly after birth can affect hippocampal function in ways that persist throughout life.

Sex-specific responses to stress have also been demonstrated in the rat to have an effect on the hippocampus.

Chronic stress in the male rat showed dendritic retraction and cell loss in the CA3 region but this was not shown in the female.

This was thought to be due to neuroprotective ovarian hormones. The hippocampus is one of the few brain regions where new neurons are generated.

This process of neurogenesis is confined to the dentate gyrus. Seizures in temporal lobe epilepsy can affect the normal development of new neurons and can cause tissue damage.

Hippocampal sclerosis is the most common type of such tissue damage. This may be a consequence of the concentration of excitable glutamate receptors in the hippocampus.

Hyperexcitability can lead to cytotoxicity and cell death. The causes of schizophrenia are not well understood, but numerous abnormalities of brain structure have been reported.

The most thoroughly investigated alterations involve the cerebral cortex, but effects on the hippocampus have also been described.

Many reports have found reductions in the size of the hippocampus in people with schizophrenia. It is unclear whether hippocampal alterations play any role in causing the psychotic symptoms that are the most important feature of schizophrenia.

It has been suggested that on the basis of experimental work using animals, hippocampal dysfunction might produce an alteration of dopamine release in the basal ganglia , thereby indirectly affecting the integration of information in the prefrontal cortex.

MRI studies have found a smaller brain volume and larger ventricles in people with schizophrenia — however researchers do not know if the shrinkage is from the schizophrenia or from the medication.

Cortical patterns are altered, and a reduction in the volume and thickness of the cortex particularly in the frontal and temporal lobes has been noted.

It has further been proposed that many of the changes seen are present at the start of the disorder which gives weight to the theory that there is abnormal neurodevelopment.

The hippocampus has been seen as central to the pathology of schizophrenia, both in the neural and physiological effects.

Several lines of evidence implicate changes in the synaptic organization and connectivity, in and from the hippocampus [] Many studies have found dysfunction in the synaptic circuitry within the hippocampus and its activity on the prefrontal cortex.

The glutamatergic pathways have been seen to be largely affected. The subfield CA1 is seen to be the least involved of the other subfields, [] [] and CA4 and the subiculum have been reported elsewhere as being the most implicated areas.

It was further concluded that schizophrenia is not due to any known neurodegenerative disorder.

Transient global amnesia is a dramatic, sudden, temporary, near-total loss of short-term memory. Various causes have been hypothesized including ischemia, epilepsy, migraine [] and disturbance of cerebral venous blood flow, [] leading to ischemia of structures such as the hippocampus that are involved in memory.

There has been no scientific proof of any cause. However, diffusion weighted MRI studies taken from 12 to 24 hours following an episode has shown there to be small dot-like lesions in the hippocampus.

These findings have suggested a possible implication of CA1 neurons made vulnerable by metabolic stress. Some studies shows correlation of reduced hippocampus volume and posttraumatic stress disorder PTSD.

The hippocampus has a generally similar appearance across the range of mammals, from monotremes such as the echidna to primates such as humans.

It does not, however, increase at anywhere close to the rate of the neocortex -to-body-size ratio. Therefore, the hippocampus takes up a much larger fraction of the cortical mantle in rodents than in primates.

In adult humans the volume of the hippocampus on each side of the brain is about 3. There is also a general relationship between the size of the hippocampus and spatial memory.

When comparisons are made between similar species, those that have a greater capacity for spatial memory tend to have larger hippocampal volumes.

Non-mammalian species do not have a brain structure that looks like the mammalian hippocampus, but they have one that is considered homologous to it.

The hippocampus, as pointed out above, is in essence part of the allocortex. Only mammals have a fully developed cortex, but the structure it evolved from, called the pallium , is present in all vertebrates, even the most primitive ones such as the lamprey or hagfish.

The medial pallium forms the precursor of the hippocampus. It does not resemble the hippocampus visually because the layers are not warped into an S shape or enfolded by the dentate gyrus, but the homology is indicated by strong chemical and functional affinities.

There is now evidence that these hippocampal-like structures are involved in spatial cognition in birds, reptiles, and fish.

In birds, the correspondence is sufficiently well established that most anatomists refer to the medial pallial zone as the "avian hippocampus".

There is evidence that food-caching birds have a larger hippocampus than other types of birds and that damage to the hippocampus causes impairments in spatial memory.

The story for fish is more complex. In teleost fish which make up the great majority of existing species , the forebrain is distorted in comparison to other types of vertebrates: most neuroanatomists believe that the teleost forebrain is in essence everted, like a sock turned inside-out, so that structures that lie in the interior, next to the ventricles, for most vertebrates, are found on the outside in teleost fish, and vice versa.

Several types of fish particularly goldfish have been shown experimentally to have strong spatial memory abilities, even forming "cognitive maps" of the areas they inhabit.

Some types of insects, and molluscs such as the octopus, also have strong spatial learning and navigation abilities, but these appear to work differently from the mammalian spatial system, so there is as yet no good reason to think that they have a common evolutionary origin; nor is there sufficient similarity in brain structure to enable anything resembling a "hippocampus" to be identified in these species.

Some have proposed, however, that the insect's mushroom bodies may have a function similar to that of the hippocampus. Submit link.

Submit link Login Sign up Imprint. Vertebrate brain region involved in memory consolidation. This article is about the section in the brain.

For the fish genus Hippocampus , see Seahorse. For other uses, see Hippocampus disambiguation. Humans have two hippocampi, one in each hemisphere of the brain.

They are located in the medial temporal lobe of the brain. In this lateral view of the human brain, the frontal lobe is at the left, the occipital lobe at the right, and the temporal and parietal lobes have largely been removed to reveal one of the hippocampi underneath.

Hippocampus lowest pink bulb as part of the limbic system. Main article: Hippocampus anatomy.

See also: Amnesia. Main article: Place cell. Further information: Reward system. Main article: Theta wave. Main article: Sharp waves and ripples.

See also: Long-term potentiation and Sleep and learning. Morphology in four rodent species. Main hippocampal regions in marmoset. Neuronal loss shown in rat CA1 following ischemia.

Left hippocampus along with other subcortical structures, in glass brain. Medicine portal Anatomy portal.

Neuroanatomy: text and atlas third ed. McGraw-Hill Companies. The hippocampus book first ed. New York: Oxford University Press.

Scientific Reports. The Hippocampus Book. Oxford University Press. Dorland's illustrated medical dictionary 32nd ed.

Neuroscience 5th ed. Sunderland, Mass. Neuroscience 5. Archived from the original on Origins of neuroscience: a history of explorations into brain function.

Journal of Comparative Neurology. The Journal of Neuroscience. Behavioural Brain Research. Nature Neuroscience. New York: Kevin Feyen.

Neurological Sciences. September International Journal of Psychophysiology. Bibcode : Sci David Bibcode : PLoSO Journal of Comparative and Physiological Psychology.

Bibcode : PLoSO.. Neuroscience and Biobehavioral Reviews. The American Journal of Psychiatry. Meta-analysis and comparison with bipolar disorder".

Archives of General Psychiatry. Biological Psychiatry. Overview of the Glutamatergic System. National Academies Press US. Reviews in the Neurosciences.

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In Alzheimer's disease and other forms of dementia , the hippocampus is one of the first regions of the brain to suffer damage; [5] short-term memory loss and disorientation are included among the early symptoms.

Damage to the hippocampus can also result from oxygen starvation hypoxia , encephalitis , or medial temporal lobe epilepsy.

People with extensive, bilateral hippocampal damage may experience anterograde amnesia : the inability to form and retain new memories.

Since different neuronal cell types are neatly organized into layers in the hippocampus, it has frequently been used as a model system for studying neurophysiology.

The form of neural plasticity known as long-term potentiation LTP was initially discovered to occur in the hippocampus and has often been studied in this structure.

LTP is widely believed to be one of the main neural mechanisms by which memories are stored in the brain.

In rodents as model organisms , the hippocampus has been studied extensively as part of a brain system responsible for spatial memory and navigation.

Many neurons in the rat and mouse hippocampus respond as place cells : that is, they fire bursts of action potentials when the animal passes through a specific part of its environment.

Hippocampal place cells interact extensively with head direction cells , whose activity acts as an inertial compass, and conjecturally with grid cells in the neighboring entorhinal cortex.

The German anatomist Duvernoy , the first to illustrate the structure, also wavered between "seahorse" and "silkworm". Another reference appeared with the term pes hippocampi , which may date back to Diemerbroeck in , introducing a comparison with the shape of the folded back forelimbs and webbed feet of the mythological hippocampus , a sea-monster with a horse's forequarters and a fish's tail.

The hippocampus was then described as pes hippocampi major , with an adjacent bulge in the occipital horn , described as the pes hippocampi minor and later renamed as the calcar avis.

Mayer mistakenly used the term hippopotamus in , and was followed by some other authors until Karl Friedrich Burdach resolved this error in In the hippocampus minor became the centre of a dispute over human evolution between Thomas Henry Huxley and Richard Owen , satirised as the Great Hippocampus Question.

The term hippocampus minor fell from use in anatomy textbooks, and was officially removed in the Nomina Anatomica of The term limbic system was introduced in by Paul MacLean [12] to describe the set of structures that line the edge of the cortex Latin limbus meaning border : These include the hippocampus, cingulate cortex , olfactory cortex , and amygdala.

Paul MacLean later suggested that the limbic structures comprise the neural basis of emotion. The hippocampus is anatomically connected to parts of the brain that are involved with emotional behavior—the septum , the hypothalamic mammillary body , and the anterior nuclear complex in the thalamus , and is generally accepted to be part of the limbic system.

The hippocampus can be seen as a ridge of gray matter tissue , elevating from the floor of each lateral ventricle in the region of the inferior or temporal horn.

The term hippocampal formation is used to refer to the hippocampus proper and its related parts.

However, there is no consensus as to what parts are included. Sometimes the hippocampus is said to include the dentate gyrus and the subiculum.

Some references include the dentate gyrus and the subiculum in the hippocampal formation, [1] and others also include the presubiculum, parasubiculum , and entorhinal cortex.

The hippocampus, including the dentate gyrus, has the shape of a curved tube, which has been compared to a seahorse, and a ram's horn C ornu A mmonis.

One edge of the "U," — CA4, is embedded into the backward-facing, flexed dentate gyrus. The hippocampus is described as having an anterior and posterior part in primates or a ventral and dorsal part in other animals.

Both parts are of similar composition but belong to different neural circuits. In primates , the part of the hippocampus at the bottom, near the base of the temporal lobe , is much broader than the part at the top.

This means that in cross-section the hippocampus can show a number of different shapes, depending on the angle and location of the cut.

In a cross-section of the hippocampus, including the dentate gyrus, several layers will be shown. The dentate gyrus has three layers of cells or four if the hilus is included.

The layers are from the outer in - the molecular layer , the inner molecular layer , the granular layer , and the hilus.

The CA3 in the hippocampus proper has the following cell layers known as strata: lacunosum-moleculare, radiatum, lucidum, pyramidal, and oriens.

CA2 and CA1 also have these layers except the lucidum stratum. The input to the hippocampus from varying cortical and subcortical structures comes from the entorhinal cortex via the perforant path.

The entorhinal cortex EC is strongly and reciprocally connected with many cortical and subcortical structures as well as with the brainstem.

Different thalamic nuclei , from the anterior and midline groups , the medial septal nucleus , the supramammillary nucleus of the hypothalamus, and the raphe nuclei and locus coeruleus of the brainstem all send axons to the EC, so that it serves as the interface between the neocortex and the other connections, and the hippocampus.

The EC is located in the parahippocampal gyrus , [2] a cortical region adjacent to the hippocampus. The parahippocampal gyrus also includes the perirhinal cortex , which plays an important role in the visual recognition of complex objects.

There is also substantial evidence that it makes a contribution to memory, which can be distinguished from the contribution of the hippocampus.

It is apparent that complete amnesia occurs only when both the hippocampus and the parahippocampus are damaged. The major input to the hippocampus is through the entorhinal cortex EC , whereas its major output is via CA1 to the subiculum.

Axons from the EC that originate in layer III are the origin of the direct perforant pathway and form synapses on the very distal apical dendrites of CA1 neurons.

Conversely, axons originating from layer II are the origin of the indirect pathway, and information reaches CA1 via the trisynaptic circuit.

In the initial part of this pathway, the axons project through the perforant pathway to the granule cells of the dentate gyrus first synapse.

From then, the information follows via the mossy fibres to CA3 second synapse. From there, CA3 axons called Schaffer collaterals leave the deep part of the cell body and loop up to the apical dendrites and then extend to CA1 third synapse.

Basket cells in CA3 receive excitatory input from the pyramidal cells and then give an inhibitory feedback to the pyramidal cells.

This recurrent inhibition is a simple feedback circuit that can dampen excitatory responses in the hippocampus.

The pyramidal cells gives a recurrent excitation which is an important mechanism found in some memory processing microcircuits.

Several other connections play important roles in hippocampal function. A major output goes via the fornix to the lateral septal area and to the mammillary body of the hypothalamus which the fornix interconnects with the hippocampus.

A very important projection comes from the medial septal nucleus, which sends cholinergic , and gamma amino butyric acid GABA stimulating fibers GABAergic fibers to all parts of the hippocampus.

The inputs from the medial septal nucleus play a key role in controlling the physiological state of the hippocampus; destruction of this nucleus abolishes the hippocampal theta rhythm and severely impairs certain types of memory.

Areas of the hippocampus are shown to be functionally and anatomically distinct. The dorsal hippocampus DH , ventral hippocampus VH and intermediate hippocampus serve different functions, project with differing pathways, and have varying degrees of place cells.

Using the radial arm maze , lesions in the DH were shown to cause spatial memory impairment while VH lesions did not. Its projecting pathways include the medial septal nucleus and supramammillary nucleus.

The intermediate hippocampus has overlapping characteristics with both the ventral and dorsal hippocampus.

This region has the smallest number of place cells. The ventral hippocampus functions in fear conditioning and affective processes.

There continues to be some interest in hippocampal olfactory responses, in particular, the role of the hippocampus in memory for odors, but few specialists today believe that olfaction is its primary function.

Over the years, three main ideas of hippocampal function have dominated the literature: response inhibition , episodic memory , and spatial cognition.

It derived much of its justification from two observations: first, that animals with hippocampal damage tend to be hyperactive ; second, that animals with hippocampal damage often have difficulty learning to inhibit responses that they have previously been taught, especially if the response requires remaining quiet as in a passive avoidance test.

British psychologist Jeffrey Gray developed this line of thought into a full-fledged theory of the role of the hippocampus in anxiety.

The second major line of thought relates the hippocampus to memory. Although it had historical precursors, this idea derived its main impetus from a famous report by American neurosurgeon William Beecher Scoville and British-Canadian neuropsychologist Brenda Milner [38] describing the results of surgical destruction of the hippocampi when trying to relieve epileptic seizures in an American man Henry Molaison , [39] known until his death in as "Patient H.

This case attracted such widespread professional interest that Molaison became the most intensively studied subject in medical history.

There is now universal agreement that the hippocampi play some sort of important role in memory; however, the precise nature of this role remains widely debated.

The third important theory of hippocampal function relates the hippocampus to space. The spatial theory was originally championed by O'Keefe and Nadel, who were influenced by American psychologist E.

Tolman's theories about " cognitive maps " in humans and animals. O'Keefe and his student Dostrovsky in discovered neurons in the rat hippocampus that appeared to them to show activity related to the rat's location within its environment.

Later research has focused on trying to bridge the disconnect between the two main views of hippocampal function as being split between memory and spatial cognition.

In some studies, these areas have been expanded to the point of near convergence. In an attempt to reconcile the two disparate views, it is suggested that a broader view of the hippocampal function is taken and seen to have a role that encompasses both the organisation of experience mental mapping , as per Tolman's original concept in and the directional behaviour seen as being involved in all areas of cognition, so that the function of the hippocampus can be viewed as a broader system that incorporates both the memory and the spatial perspectives in its role that involves the use of a wide scope of cognitive maps.

It has also been proposed that the spiking activity of hippocampal neurons is associated spatially, and it was suggested that the mechanisms of memory and planning both evolved from mechanisms of navigation and that their neuronal algorithms were basically the same.

Many studies have made use of neuroimaging techniques such as functional magnetic resonance imaging fMRI , and a functional role in approach-avoidance conflict has been noted.

The anterior hippocampus is seen to be involved in decision-making under approach-avoidance conflict processing. It is suggested that the memory, spatial cognition, and conflict processing functions may be seen as working together and not mutually exclusive.

Psychologists and neuroscientists generally agree that the hippocampus plays an important role in the formation of new memories about experienced events episodic or autobiographical memory.

This is partly why returning to a location where an emotional event occurred may evoke that emotion.

There is a deep emotional connection between episodic memories and places. Due to bilateral symmetry the brain has a hippocampus in each cerebral hemisphere.

If damage to the hippocampus occurs in only one hemisphere, leaving the structure intact in the other hemisphere, the brain can retain near-normal memory functioning.

Although the retrograde effect normally extends many years back before the brain damage, in some cases older memories remain. This retention of older memories leads to the idea that consolidation over time involves the transfer of memories out of the hippocampus to other parts of the brain.

In particular, efficiency of verbal memory retention is related to the anterior parts of the right and left hippocampus.

The right head of the hippocampus is more involved in executive functions and regulation during verbal memory recall. The tail of the left hippocampus tends to be closely related to verbal memory capacity.

Damage to the hippocampus does not affect some types of memory, such as the ability to learn new skills playing a musical instrument or solving certain types of puzzles, for example.

This fact suggests that such abilities depend on different types of memory procedural memory and different brain regions.

Furthermore, amnesic patients frequently show "implicit" memory for experiences even in the absence of conscious knowledge. For example, patients asked to guess which of two faces they have seen most recently may give the correct answer most of the time in spite of stating that they have never seen either of the faces before.

Some researchers distinguish between conscious recollection , which depends on the hippocampus, and familiarity , which depends on portions of the medial temporal lobe.

When rats are exposed to an intense learning event, they may retain a life-long memory of the event even after a single training session.

The memory of such an event appears to be first stored in the hippocampus, but this storage is transient.

Much of the long-term storage of the memory seems to take place in the anterior cingulate cortex. Furthermore, many other genes were upregulated , likely often due to the removal of methyl groups from previously existing 5-methylcytosines 5mCs in DNA.

Demethylation of 5mC can be carried out by several proteins acting in concert, including TET enzymes as well as enzymes of the DNA base excision repair pathway see Epigenetics in learning and memory.

Studies on freely moving rats and mice have shown many hippocampal neurons to act as place cells that cluster in place fields , and these fire bursts of action potentials when the animal passes through a particular location.

This place-related neural activity in the hippocampus has also been reported in monkeys that were moved around a room whilst in a restraint chair.

Other cells in smaller proportion are inhibitory interneurons , and these often show place-related variations in their firing rate that are much weaker.

There is little, if any, spatial topography in the representation; in general, cells lying next to each other in the hippocampus have uncorrelated spatial firing patterns.

Place cells are typically almost silent when a rat is moving around outside the place field but reach sustained rates as high as 40 Hz when the rat is near the center.

Neural activity sampled from 30—40 randomly chosen place cells carries enough information to allow a rat's location to be reconstructed with high confidence.

The size of place fields varies in a gradient along the length of the hippocampus, with cells at the dorsal end showing the smallest fields, cells near the center showing larger fields, and cells at the ventral tip showing fields that cover the entire environment.

In humans, cells with location-specific firing patterns have been reported during a study of patients with drug-resistant epilepsy.

They were undergoing an invasive procedure to localize the source of their seizures , with a view to surgical resection. The patients had diagnostic electrodes implanted in their hippocampus and then used a computer to move around in a virtual reality town.

A study showed that the posterior part of the hippocampus is larger in these drivers than in the general public, and that a positive correlation exists between the length of time served as a driver and the increase in the volume of this part.

It was also found the total volume of the hippocampus was unchanged, as the increase seen in the posterior part was made at the expense of the anterior part, which showed a relative decrease in size.

There have been no reported adverse effects from this disparity in hippocampal proportions. The anterior part of the right hippocampus was larger and the posterior part was smaller, compared with sighted individuals.

There are several navigational cells in the brain that are either in the hippocampus itself or are strongly connected to it, such as the speed cells present in the medial entorhinal cortex.

Together these cells form a network that serves as spatial memory. The first of such cells discovered in the s were the place cells, which led to the idea of the hippocampus acting to give a neural representation of the environment in a cognitive map.

Getting lost is a common symptom of amnesia. These have been assigned as head direction cells , grid cells and boundary cells. Approach-avoidance conflict happens when a situation is presented that can either be rewarding or punishing, and the ensuing decision-making has been associated with anxiety.

Overall findings showed that the anterior hippocampus is sensitive to conflict, and that it may be part of a larger cortical and subcortical network seen to be important in decision making in uncertain conditions.

A review makes reference to a number of studies that show the involvement of the hippocampus in conflict tasks. The authors suggest that a challenge is to understand how conflict processing relates to the functions of spatial navigation and memory and how all of these functions need not be mutually exclusive.

The hippocampus shows two major "modes" of activity, each associated with a distinct pattern of neural population activity and waves of electrical activity as measured by an electroencephalogram EEG.

The main characteristics described below are for the rat, which is the animal most extensively studied. The theta mode appears during states of active, alert behavior especially locomotion , and also during REM dreaming sleep.

An active cell typically stays active for half a second to a few seconds. As the rat behaves, the active cells fall silent and new cells become active, but the overall percentage of active cells remains more or less constant.

In many situations, cell activity is determined largely by the spatial location of the animal, but other behavioral variables also clearly influence it.

The LIA mode appears during slow-wave non-dreaming sleep, and also during states of waking immobility such as resting or eating. Sharp waves are frequently generated in sets, with sets containing up to 5 or more individual sharp waves and lasting up to ms.

The spiking activity of neurons within the hippocampus is highly correlated with sharp wave activity. These two hippocampal activity modes can be seen in primates as well as rats, with the exception that it has been difficult to see robust theta rhythmicity in the primate hippocampus.

Although this movie is not the most accurate representation of a true amnesic patient, it is useful for informing viewers of the detrimental effects of amnesia.

Memento a film inspired by the case of Henry Molaison H. Unlike most amnesiacs, Leonard retains his identity and the memories of events that occurred before the injury, but loses all ability to form new memories.

This loss of ability to form new memories indicates that the head injury affected the medial temporal lobe of the brain resulting in the inability for Leonard to form declarative memory.

Finding Nemo features a reef fish named Dory with an inability to develop declarative memory. This prevents her from learning or retaining any new information such as names or directions.

The exact origin of Dory's impairment is not mentioned in the film, but her memory loss accurately portrays the difficulties facing amnesiacs.

Submit link. Submit link Login Sign up Imprint. Long-Term Memory Subtype Description Example Declarative explicit Conscious memory of facts and events Semantic Factual information The capital of Germany is Berlin Episodic Specific personal experiences Your 10th birthday Non-declarative implicit Modes of learning that are nonconscious - one learning to perform a sequence of actions that do not necessarily invoke knowledge Priming also known as "pattern completion," where one has the ability to complete a pattern they have once seen before.

This priming differs from Psychology priming. If you were given a picture of half of a letter from the alphabet and you recognized which letter it is, you would be able to complete the letter.

Perceptual learning Perceptual ability to differentiate sensories through experience of stimuli Differentiating between categories such as smells, colors, tastes Category learning " The model of language Declarative and procedural memory fall into two categories of human language.

Procedural memory builds rule-governed structure merging or series of forms and representations into complex structures such as: phonology inflectional and derivational morphology compositional semantics the meaning of composition of words into complex structures syntax Broca and Wernicke's Brain Region Broca's area is important to procedural memory, for, "Broca's area is involved in the expressive aspects of spoken and written language production of sentences constrained by the rules of grammar and syntax.

History The study of human memory stretches back over the last years. Al , and proposes that the hippocampus does three things with episodic memory: Mediates the recording of episodic memories Identifies common features between episodes Links these common episodes in a memory space.

Amygdala as seen in red Amygdala The amygdala is believed to be involved in the encoding and retrieval of emotionally charged memories.

Hippocampal lesion studies The Morris water maze The Morris water navigation task tests spatial learning in rats.

Amygdala lesion studies Adolph, Cahill and Schul completed a study showing that emotional arousal facilitates the encoding of material into long term declarative memory.

Episodic and semantic memory. In Organization of Memory, ed. E Tulving, W Donaldson, pp. Retrieved Reisberg, Daniel ed. The Oxford Handbook of Cognitive Psychology.

Annual Review of Psychology. Current Science. Autobiographical memory. Conway Eds. Hove, UK: Psychology Press. Memory: A Contribution to Experimental Psychology.

Behavioural Brain Research. Nature Neuroscience. Proceedings of the National Academy of Sciences. Current Directions in Psychological Science.

Journal of Cognitive Neuroscience. Psychological Bulletin. Nat Neurosci. Behav Neurosci. Nature Reviews Neuroscience.

Brain Cogn. Brain Sci. Learning and Memory. American Journal of Psychiatry. Brain Imaging and Behaviour. Current Opinion in Neurobiology.

Sleep Res. J Cogn Neurosci. J Neurosci. Eur J Neurosci. Learn Mem. Psychol Sci. Current Biology. Annu Rev Psychol. Fundamentals of Human Neuropsychology.

Worth Publishers. Principles of neural science. McGraw Hill Medical. J Verb Learn Verb Behav. Teaching of Psychology.

Cerebral Cortex. Petri and M. Brain Injury. Neurorehabilitation and Neural Repair. Journal of Clinical and Experimental Neuropsychology.

Dementia and Geriatric Cognitive Disorders. Brain and Language. British Medical Journal. Trends in Cognitive Sciences.

Human memory. Amnesia anterograde childhood post-traumatic psychogenic retrograde transient global Decay theory Forgetting curve Interference theory Memory inhibition Motivated forgetting Repressed memory Retrieval-induced forgetting Selective amnesia Weapon focus.

Confabulation False memory Hindsight bias Imagination inflation List of memory biases Memory conformity Mere-exposure effect Misattribution of memory Misinformation effect Source-monitoring error Wernicke—Korsakoff syndrome.

Absent-mindedness Atkinson—Shiffrin memory model Context-dependent memory Childhood memory Cryptomnesia Effects of alcohol Emotion and memory Exosomatic memory Flashbacks Free recall Involuntary memory Levels-of-processing effect Memory and trauma Memory improvement Metamemory Mnemonic Muscle memory Priming Intertrial Prospective memory Recovered-memory therapy Retrospective memory Sleep and memory State-dependent memory Transactive memory.

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