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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.

There are, however, qualitatively similar sharp waves and similar state-dependent changes in neural population activity. The underlying currents producing the theta wave are generated mainly by densely packed neural layers of the entorhinal cortex, CA3, and the dendrites of pyramidal cells.

The theta wave is one of the largest signals seen on EEG, and is known as the hippocampal theta rhythm. 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.

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