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hebb in 1949 140 but for many years remained unexplained in 1973 tim bliss and terje lømo described a phenomenon in the rabbit hippocampus that appeared to meet hebb s specifications a change in synaptic responsiveness induced by brief strong activation and lasting for hours or days or longer 141 this phenomenon was soon referred to as long term potentiation ltp 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 signaling cascades that can trigger ltp is acknowledged as preventing a more complete understanding 142 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 143 the best studied form of ltp has been seen in ca1 of the hippocampus and occurs at synapses that terminate on dendritic spines and use the neurotransmitter glutamate 142 the synaptic changes depend on the n methyl d aspartate nmda receptor a cell surface receptor that allows calcium to enter the postsynaptic spine when glutamate and glycine are bound and postsynaptic depolarization occur at the same time 144 drugs that interfere with nmda receptors block ltp and have major effects on some types of memory especially spatial memory genetically modified mice that are modified to disable the ltp mechanism also generally show severe memory deficits 144 research edit a brain implant for use as a hippocampal prosthesis has been the subject of research since the early 2000s 145 it was reported in 2018 that a nonlinear multi input multi output model mimo had been developed that in some studies had been shown to restore and improve memory function 146 this has been followed by a modified version known as the memory decoding model mdm 147 this model has been shown to have the potential use in significant modification of memory a study concluded that further research could be pointed towards an evaluation of both models in particular focusing on the hippocampal theta wave input 148 clinical significance edit aging edit see also neurobiological effects of physical exercise structural growth aging brain and memory and aging 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 plays a central role in memory there has been considerable interest in the possibility that age related declines could be caused by hippocampal deterioration 149 studies show that hippocampal volume loss is a noted feature from middle age more so in women than in men 150 where there is age related shrinkage memory performance will be impaired 149 there are also reports that memory tasks tend to produce less hippocampal activation in the elderly than in the young 149 furthermore a randomized control trial published in 2011 found that aerobic exercise could increase the size of the hippocampus in adults aged 55 to 80 and also improve spatial memory 151 alzheimer s disease edit in alzheimer s disease and other forms of dementia the hippocampus is one of the first regions of the brain to be damaged 152 short term memory loss and disorientation are included among the early symptoms amyloid beta deposits begin in the frontal lobes before the signs of neurofibrillary tangles are seen in the hippocampus 153 damage to the hippocampus can also result from oxygen starvation hypoxia encephalitis or medial temporal lobe epilepsy extensive bilateral hippocampal damage may cause anterograde amnesia the inability to form and retain new memories dementia is very often caused by cerebral ischemia that is believed to trigger changes in the hippocampus changes in ca1 the hippocampal area that underlies episodic memory cause episodic memory impairment the earliest symptom of post ischemic dementia 154 stress edit the hippocampus contains high levels of glucocorticoid receptors which make it more vulnerable to long term stress than most other brain areas 155 there is evidence that humans having experienced severe long lasting traumatic stress show atrophy of the hippocampus more than of other parts of the brain 156 these effects show up in post traumatic stress disorder 157 and they may contribute to the hippocampal atrophy reported in schizophrenia 158 and severe depression 159 anterior hippocampal volume in children is positively correlated with parental family income and this correlation is thought to be mediated by income related stress 160 a study has revealed atrophy as a result of depression but this can be stopped with anti depressants even if they are not effective in relieving other symptoms 161 chronic stress resulting in elevated levels of glucocorticoids notably of cortisol is seen to be a cause of neuronal atrophy in the hippocampus this atrophy results in a smaller hippocampal volume which is also seen in cushing s syndrome the higher levels of cortisol in cushing s syndrome is usually the result of medications taken for other conditions 162 163 neuronal loss also occurs as a result of impaired neurogenesis 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 163 after treatment with medication to reduce cortisol in cushing s syndrome the hippocampal volume is seen to be restored by as much as 10 162 this change is seen to be due to the reforming of the dendrites 163 this dendritic restoration can also happen when stress is removed 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 164 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 165 166 in rats dna damage increases in the hippocampus under conditions of stress 167 ptsd edit some studies shows correlation of reduced hippocampus volume and post traumatic stress disorder ptsd 168 169 a study of vietnam war combat veterans with ptsd showed a 20 reduction in the volume of their hippocampus compared with veterans with no such symptoms 170 this finding was not replicated in those with chronic ptsd traumatized at an air show plane crash in 1988 ramstein germany 171 it is also the case that non combat twin brothers of vietnam veterans with ptsd also had smaller hippocampi than other controls raising questions about the nature of the correlation 172 a 2016 study strengthened the theory that a smaller hippocampus increases the risk for post traumatic stress disorder and a larger hippocampus increases the likelihood of efficacious treatment 173 transient global amnesia edit transient global amnesia is a syndrome of unknown cause that results in a sudden and temporary anterograde amnesia and variable past memory loss an attack usually lasts for up to 24 hours during which time the memory loss is experienced the only diagnostic evidence of tga is given by dwi mri that shows lesions as small dots in the ca1 subfield 174 175 the lesions are detectable from around 24 to 96 hours after symptoms onset and can be seen to have been resolved at a six month follow up 176 the lesions are between 1 and 5 mm and can be single or multiple and may be confined to either hemisphere 174 175 lesions in the dominant hemisphere affect episodic verbal memory and those in the non dominant hemisphere affect visuospatial memory 176 ca1 lesions shows selective affect over other ca subfields 177 the selective vulnerability of ca1 neurons suggests a cause of metabolic stress that could result from emotional or behavioral stress 177 178 other possible causes have been debated including cerebral venous reflex arterial ischema epilepsy and migraine 175 tga as a network disease has also been put forward dti studies show a decreased connectivity between brain regions that may impact the hippocampus 175 epilepsy edit an eeg showing epilepsy right hippocampal seizure onset an eeg showing epilepsy left hippocampal seizure onset the hippocampus is one of the few brain regions where new neurons are generated this process of neurogenesis is confined to the dentate gyrus 179 neurogenesis can be positively affected by exercise or negatively affected by epileptic seizures 179 seizures in temporal lobe epilepsy can affect the normal development of new neurons and can cause tissue damage hippocampal sclerosis specific to the mesial temporal lobe is the most common type of such tissue damage 180 181 it is not yet clear however whether the epilepsy is usually caused by hippocampal abnormalities or whether the hippocampus is damaged by cumulative effects of seizures 182 however in experimental settings where repetitive seizures are artificially induced in animals hippocampal damage is a frequent result this may be a consequence of the concentration of excitable glutamate receptors in the hippocampus hyperexcitability can lead to cytotoxicity and cell death 163 it may also have something to do with the hippocampus being a site of continuous neurogenesis and to abnormalities in this process 179 163 schizophrenia edit a reduction in hippocampal volume is well reported in those with schizophrenia 183 67 the left hippocampus seems to be affected more than the right 184 the volume reduction is small but consistent it can be first seen in the prodromal stage and can show modest progression as the disease advances 67 the reduced volume has been shown to be independent of treatment with antipsychotics some studies suggest that hippocampal alterations play a role in causing the psychotic symptoms of schizophrenia 185 186 it has been suggested that hippocampal dysfunction might produce an alteration of dopamine release in the basal ganglia thereby indirectly affecting the integration of information in the prefrontal cortex 187 in those with psychosis the greatest reduction is seen to be in ca3 and ca2 subfields and a correlation has been made between the reduced volume and memory dysfunction 67 a reduced hippocampal volume has been shown to result in a decreased connectivity between the hippocampus and the prefrontal cortex with the declarative memory characteristic of hippocampal function becoming selectively impaired 183 the impaired connectivity is evident both at rest and during task activity 67 evidence of dysfunctional gaba inhibitory transmission has been shown in studies of the whole hippocampus 67 188 another noted change in the hippocampus is a reduced m4 muscarinic cholinergic receptor but an unaffected m1 receptor 67 reduced neurogenesis has been shown in the dentate gyrus with a noted reduction in the density of neurons and oligodendrocytes 67 hyperactivity in the hippocampus has consistently been linked to schizophrenia using blood oxygenation level dependent imaging in functional mri and in cerebral blood volume studies 67 189 the ca1 subfield is mostly indicated to be affected and hyperactivity almost exclusively found in the anterior hippocampus 190 67 hippocampal hyperactivity has been suggested to be a result of dysfunctional gabaergic inhibition 188 it has been suggested that hippocampal hyperactivity at early stages of schizophrenia may contribute by excitotoxic mechanisms to hippocampal atrophy and volume reduction at later stages of the disorder 191 hippocampal hyperactivity has been suggested to contribute to symptoms of schizophrenia including those of psychosis 192 193 188 microcephaly edit hippocampus atrophy has been characterized in those with microcephaly 194 mouse models with wdr62 mutations which recapitulate human point mutations show a deficiency in hippocampal development and neurogenesis 195 other animals edit other vertebrates edit non mammalian vertebrates lack a brain structure that looks like the mammalian hippocampus but they have one that is considered homologous to it the hippocampus 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 196 197 the pallium is usually divided into three zones medial lateral and dorsal 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 evidence that these hippocampal like structures are involved in spatial cognition in reptiles and in fish 198 fish edit in teleost fish the forebrain is everted like an inside out sock with structures that lie on the outside as contrasted with other vertebrate structures that lie in the interior next to the ventricles 199 one of the consequences of this is that the medial pallium the hippocampal zone of a typical vertebrate is thought to correspond to the lateral pallium of a typical fish 200 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 26 studies in goldfish show that damage to both the lateral pallium and the medial pallium impairs spatial memory coding 201 202 it is not clear if the medial pallium plays a similar role in basal vertebrates such as sharks and rays or even lampreys and hagfish 203 the dorsolateral pallium of the teleost is considered as homologous to the hippocampus in terrestrial vertebrates 204 in 2023 the goldfish brain was mapped by molecular parcellization showing that its telencephalon subregions were homogeneous to the hippocampal subfields in the mouse 201 birds edit see also avian pallium in birds the correspondence is sufficiently well established that most anatomists refer to the medial pallial zone as the avian hippocampus 205 numerous species of birds have strong spatial skills in particular those that cache store food 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 206 insects and molluscs edit some types of insects such as cockroaches 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 suggesting that there is no common evolutionary origin mushroom bodies in insect brains are associated with learning and memory carried out in the mammalian hippocampus 207 the brain of th...
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