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orant 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 systems and from the nucleus reuniens of the thalamus to field ca1 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 impairs spatial memory 39 subfields edit hippocampal subfields and subregions drawing by italian pathologist camillo golgi of a hippocampus stained using silver nitrate the hippocampal subfields ca1 ca4 and subregions head body and tail are functionally and anatomically differentiated and connect differently to other brain regions 40 41 42 their cells are morphologically different 43 they also have different levels of vulnerability to disease 40 in humans the head of the hippocampus is also termed the anterior hippocampus the body is the intermediate hippocampus and the tail the posterior hippocampus the terms used for corresponding subregions in other primates are anterior or rostral intermediate and posterior or caudal and in rodents ventral or temporal intermediate and dorsal or septal 41 42 the subregions along the longitudinal axis have been linked to different functions project with different neural pathways and have varying numbers of place cells with differentially sized place fields 44 41 for example the posterior hippocampus has particularly been linked to spatial memory 41 in the rat the dorsal hippocampus has more place cells than both the ventral and intermediate hippocampal regions and place cells in the dorsal hippocampal provide a more fine grained representation of the spatial environment than in the intermediate and ventral hippocampus 45 46 the intermediate hippocampus has overlapping characteristics with both the ventral and dorsal hippocampus 44 47 studies in 2002 showed that alterations to the ventral hippocampus reduced the amount of information sent to the amygdala by the dorsal and ventral hippocampus consequently altering fear conditioning in rats 48 in 2007 studies using anterograde tracing methods located the moderate projections to two primary olfactory cortical areas and prelimbic areas of the medial prefrontal cortex this region has the smallest number of place cells the ventral hippocampus has been particularly linked to fear conditioning and affective processes 49 50 function edit theories edit three main theories of hippocampal function have been in dominance response inhibition episodic memory and spatial cognition the response inhibition theory caricatured by john o keefe and lynn nadel as slam on the brakes was very popular up to the 1960s 51 it was based largely on two observations first that animals with hippocampal damage tend to be hyperactive second that animals with hippocampal damage often have difficulty learning to inhibit previously learnt responses especially if the response requires remaining quiet as in a passive avoidance test british psychologist jeffrey gray developed this line of thought into a complete theory of the role of the hippocampus in anxiety called the behavioral inhibition system 52 53 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 54 it described the results of surgical destruction of the hippocampi when trying to relieve epileptic seizures in an american man henry molaison known until his death in 2008 as patient h m 55 56 the unexpected outcome of the surgery was severe anterograde and partial retrograde amnesia molaison was unable to form new episodic memories after his surgery and could not remember any events that occurred just before his surgery but he did retain memories of events that occurred many years earlier extending back into his childhood this case attracted such widespread professional interest that molaison became the most intensively studied subject in medical history 55 rats and cognitive maps the third important theory of hippocampal function relates the hippocampus to space and spatial memory with the idea of a cognitive map first proposed by american psychologist e c tolman this theory was followed further by o keefe and in 1971 he and his student dostrovsky discovered neurons in the rat hippocampus that seemed to show activity related to the rat s location within its environment the neurons were described as place cells 57 a book was later produced in 1978 the hippocampus as a cognitive map written by o keefe and nadel 58 it has been generally agreed that the hippocampus plays a key role in spatial coding but the details are widely debated 59 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 organization of experience mental mapping as per tolman s original concept in 1948 and the directional behavior 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 60 61 62 63 this relates to the purposive behaviorism born of tolman s original goal of identifying the complex cognitive mechanisms and purposes that guided behavior 64 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 65 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 66 role in memory edit see also amnesia and epigenetics in learning and memory the hallmark function of the hippocampus is the formation of explicit memory also known as declarative memory 67 episodic memory and semantic memory are the two components of explicit memory 68 the hippocampus also encodes emotional context from the amygdala 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 69 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 70 severe damage to the hippocampi in both hemispheres results in profound difficulties in forming new memories anterograde amnesia and often also affects memories formed before the damage occurred retrograde amnesia 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 71 ch 1 experiments using intrahippocampal transplantation of hippocampal cells in primates with neurotoxic lesions of the hippocampus have shown that the hippocampus is required for the formation and recall but not the storage of memories 72 it has been shown that a decrease in the volume of various parts of the hippocampus leads to specific memory impairments 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 73 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 such as procedural memory in implicit memory function implicating 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 74 a study claims to have confirmed that the hippocampus is not associated with implicit memory 75 but other sources say the question is still up for debate as of 2024 76 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 77 when such an intense learning event was experimentally applied more than 5 000 differently methylated dna regions appeared in the hippocampus neuronal genome of the rats at one hour and at 24 hours after training 78 these alterations in methylation pattern occurred at many genes that were down regulated often due to the formation of new 5 methylcytosine sites in cpg rich regions of the genome furthermore many other genes were upregulated 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 79 80 as well as enzymes of the dna base excision repair pathway 81 between systems model edit the between systems memory interference model describes the inhibition of non hippocampal systems of memory during concurrent hippocampal activity 82 specifically it was found that when the hippocampus was inactive non hippocampal systems located elsewhere in the brain were found to consolidate memory in its place however when the hippocampus was reactivated memory traces consolidated by non hippocampal systems were not recalled suggesting that the hippocampus interferes with long term memory consolidation in other memory related systems 83 one of the major implications that this model illustrates is the dominant effects of the hippocampus on non hippocampal networks when information is incongruent with this information in mind future directions could lead towards the study of these non hippocampal memory systems through hippocampal inactivation further expanding the labile constructs of memory additionally many theories of memory are holistically based around the hippocampus this model could add beneficial information to hippocampal research and memory theories such as the multiple trace theory 84 85 lastly the between system memory interference model allows researchers to evaluate their results on a multiple systems model suggesting that some effects may not be simply mediated by one portion of the brain 86 role in spatial memory and navigation edit main article place cell spatial firing patterns of eight place cells recorded from the ca1 layer of a rat the rat ran back and forth along an elevated track stopping at each end to eat a small food reward dots indicate positions where action potentials were recorded with color indicating which neuron emitted that action potential hippocampal connections in spatial cognition there are several types of navigational cells in the brain that are either in the hippocampus itself or are strongly connected to it they include the place cells speed cells present in the medial entorhinal cortex head direction cells grid cells and boundary cells 59 87 together these cells form a network that serves as spatial memory the first of these types of cell discovered in the 1970s 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 58 when the hippocampus is dysfunctional orientation is affected people may have difficulty in remembering how they arrived at a location and how to proceed further getting lost is a common symptom of amnesia 88 studies with animals have shown that an intact hippocampus is required for initial learning and long term retention of some spatial memory tasks in particular ones that require finding the way to a hidden goal 89 90 91 92 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 93 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 94 speed cells are thought to provide input to the hippocampal grid cells 95 this place related neural activi...
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