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onflict processing 5 5 role in social memory 5 6 role in olfaction 6 physiology toggle physiology subsection 6 1 hippocampal theta rhythm 6 2 sharp waves 6 3 long term potentiation 7 research 8 clinical significance toggle clinical significance subsection 8 1 aging 8 2 alzheimer s disease 8 3 stress 8 3 1 ptsd 8 3 2 transient global amnesia 8 4 epilepsy 8 5 schizophrenia 8 6 microcephaly 9 other animals toggle other animals subsection 9 1 other vertebrates 9 1 1 fish 9 1 2 birds 9 2 insects and molluscs 10 see also 11 additional images 12 references 13 external links toggle the table of contents hippocampus 59 languages العربية مصرى azərbaycanca български বাংলা bosanski català čeština dansk deutsch ελληνικά esperanto español eesti euskara فارسی suomi français galego עברית हिन्दी hrvatski magyar հայերեն bahasa indonesia íslenska italiano 日本語 ქართული қазақша 한국어 кыргызча latina lietuvių македонски മലയാളം bahasa melayu nederlands norsk bokmål polski português română русский simple english slovenčina slovenščina српски srpski svenska తెలుగు ไทย tagalog türkçe українська اردو tiếng việt winaray ⵜⴰⵎⴰⵣⵉⵖⵜ ⵜⴰⵏⴰⵡⴰⵢⵜ 粵語 中文 edit links article talk english read edit view history tools tools move to sidebar hide actions read edit view history general what links here related changes upload file permanent link page information cite this page get shortened url print export download as pdf printable version in other projects wikimedia commons wikidata item appearance move to sidebar hide from wikipedia the free encyclopedia vertebrate brain region this article is about the section in the brain for the fish genus see seahorse for the mythological creature see hippocampus mythology for other uses see hippocampus disambiguation hippocampus 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 region as part of the limbic system details part of temporal lobe identifiers latin hippocampus mesh d006624 neuronames 3157 neurolex id birnlex_721 ta98 a14 1 09 321 ta2 5518 fma 275020 anatomical terms of neuroanatomy edit on wikidata the hippocampus pl hippocampi also hippocampus proper is a major component of the brain of humans and many other vertebrates in the human brain the hippocampus the dentate gyrus and the subiculum are components of the hippocampal formation located in the limbic system the hippocampus plays important roles in the consolidation of information from short term memory to long term memory and in spatial memory that enables navigation in humans and other primates the hippocampus is located in the archicortex one of the three regions of allocortex in each hemisphere with direct neural projections to and reciprocal indirect projections from the neocortex the hippocampus as the medial pallium is a structure found in all vertebrates in alzheimer s disease and other forms of dementia the hippocampus is one of the first regions of the brain to be damaged 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 using 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 hippocampi 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 name edit the human hippocampus and fornix left compared with a seahorse right 1 the earliest description of the ridge running along the floor of the inferior horn of the lateral ventricle comes from the venetian anatomist julius caesar aranzi 1587 who likened it first to a silkworm and then to a seahorse latin hippocampus from greek ἱππόκαμπος latinised ìppókampos from ἵππος ippos horse κάμπος kámpos sea monster 2 the german anatomist johann georg duvernoy 1729 the first to illustrate the structure also wavered between seahorse and silkworm ram s horn was proposed by the danish anatomist jacob winsløw in 1732 and a decade later his fellow parisian the surgeon de garengeot used cornu ammonis horn of amun 3 after the ancient egyptian god who was often represented as having a ram s head 4 ammon is the greek name for amun 5 the head region of the hippocampus is enlarged and presents two or three rounded elevations or foot like digitations and hence it was named the pes hippocampi pes meaning foot 6 7 later this part was described as pes hippocampi major with an adjacent bulge in the occipital horn of the lateral ventricle described as pes hippocampi minor later renamed as the calcar avis 3 8 in 1786 félix vicq d azyr published an authoritative description naming just the hippocampus but the term remained largely unused with no description of any function proposed until in the middle of the 20th century it was associated with memory 2 mayer mistakenly used the term hippopotamus in 1779 and was followed by some other authors until karl friedrich burdach resolved this error in 1829 in 1861 the hippocampus minor became the center of a dispute over human evolution between thomas henry huxley and richard owen satirized as the great hippocampus question the term hippocampus minor fell from use in anatomy textbooks and was officially removed in the nomina anatomica of 1895 9 the modern term for the structure is just the hippocampus with the term cornu ammonis that is ammon s horn surviving in the names of the hippocampal subfields ca1 ca4 10 11 12 in the limbic system edit hippocampus in the limbic system the hippocampus is one of the structures of the limbic lobe first described by broca in 1878 as the cortical areas that line the deep edge of the cerebral cortex 13 the limbic lobe is the main component of the limbic system 14 the cingulate gyrus and the parahippocampal gyrus are the two main parts of the described lobe which had been largely associated with olfaction 13 many studies later culminating in work by papez and maclean the involvement of other interacting brain regions associated with emotion was recognized 14 the hippocampus is anatomically connected to parts of the brain that are involved with emotional behavior including the septal area the hypothalamic mammillary bodies and the anterior nuclear complex in the thalamus maclean proposed that the associated structures of the limbic lobe be included in what he termed as the limbic system 14 anatomy edit main article hippocampus anatomy cross section of cerebral hemisphere showing structure and location of hippocampus coronal section of the brain of a macaque monkey showing hippocampal formation circled the hippocampus is a five centimeter long ridge of gray matter tissue within the parahippocampal gyrus that can only be seen when the gyrus is opened 15 16 the hippocampus is an inward fold of three layered archicortex one of three regions of the allocortex into the medial temporal lobe of the brain where it elevates into the floor of each lateral ventricle inferior horn 17 18 19 20 the hippocampus stretches along its anterior posterior axis from the amygdala to the splenium of the corpus callosum with the head body and tail as subregions of this axis 21 15 the dentate gyrus ca subfields fimbria and subiculum are divisions across the short axis the proximal distal axis 21 the hippocampal formation refers to the hippocampus and its related adjoining parts to include the dentate gyrus the subiculum the presubiculum parasubiculum and the entorhinal cortex 17 sometimes the subiculum presubiculum and parasubiculum are grouped together as the subicular complex but the regions are neuroanatomically distinct some sources may only include the hippocampus dentate gyrus and subiculum being regions of the hippocampal three layered archicortex 17 but the six regions are linked together serially by almost unidirectional neural pathways 17 other sources include the indusium griseum gyrus fasciolaris the medial and longitudinal striae and uncus and exclude subicular regions 22 23 the neural layout and pathways within the hippocampal formation are very similar in all mammals 19 the hippocampus has a generally similar appearance across the range of mammals from egg laying mammals such as the echidna to humans and other primates 24 the hippocampal size to body size ratio broadly increases being about twice as large for primates as for the echidna 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 0 to 3 5 cm 3 as compared to 320 to 420 cm 3 for the volume of the neocortex 25 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 26 neuroanatomy edit the hippocampus and dentate gyrus that is folded into the hippocampal archicortex have the shape of a curved rolled up tube the curve of the hippocampus known as cornu ammonis uses the initial letters ca to name the hippocampal subfields ca1 ca4 ca4 is in fact the polymorphic layer or hilus of the dentate gyrus but ca4 is still sometimes in use to describe the part of ca3 that inserts between the dentate gyrus regions or blades 17 27 it can be distinguished as an area where the cortex narrows into a single layer of densely packed pyramidal neurons which curl into a tight u shape one edge of the u is ca4 the hilus of the dentate gyrus this is embedded into the backward facing flexed dentate gyrus in humans the hippocampus is described as having an anterior and posterior part in other primates they are termed rostral and caudal and in rodents they are termed ventral and dorsal 28 both parts are of similar composition but belong to different neural circuits 28 the dentate gyrus combined with other hippocampal regions form a banana like structure with the two hippocampi joined at the stems by the commissure of fornix also called the hippocampal commissure 19 29 in humans and other 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 30 in a cross section of the hippocampus including the dentate gyrus several layers will be shown the dentate gyrus has three layers of cells the outer molecular layer the middle granular layer and the inner polymorphic layer also known as the hilus 31 the ca3 subfield 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 32 the input to the hippocampus from varying cortical and subcortical structures comes from the entorhinal cortex via the perforant path 33 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 34 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 33 the ec is located in the parahippocampal gyrus a cortical region adjacent to the hippocampus 35 this gyrus conceals the hippocampus the parahippocampal gyrus is adjacent to 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 35 circuitry edit basic circuit of the hippocampus as drawn by ramón y cajal dg dentate gyrus sub subiculum ec entorhinal cortex the major input to the hippocampus is through the entorhinal cortex ec whereas its major output is via ca1 to the subiculum 36 information reaches ca1 via two main pathways direct and indirect 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 cell fibers 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 36 axons from ca1 then project back to the entorhinal cortex completing the circuit 37 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 give a recurrent excitation which is an important mechanism found in some memory processing microcircuits 38 several other connections play important roles in hippocampal function 19 beyond the output to the ec additional output pathways go to other cortical areas including the prefrontal cortex 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 16 the hippocampus receives modulatory input from the serotonin norepinephrine and dopamine system...
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