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nal 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 activity in the hippocampus has also been reported in monkeys that were moved around a room whilst in a restraint chair 96 however the place cells may have fired in relation to where the monkey was looking rather than to its actual location in the room 97 over many years many studies have been carried out on place responses in rodents which have given a large amount of information 59 place cell responses are shown by pyramidal cells in the hippocampus and by granule cells in the dentate gyrus 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 to 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 59 in some cases the firing rate of hippocampal cells depends not only on place but also the direction a rat is moving the destination toward which it is traveling or other task related variables 98 the firing of place cells is timed in relation to local theta waves a spatiotemporal process termed phase precession 99 100 cells with location specific firing patterns have been reported during a study of people with drug resistant epilepsy they were undergoing an invasive procedure to localize the source of their seizures with a view to surgical resection they had diagnostic electrodes implanted in their hippocampi and then used a computer to move around in a virtual reality town 101 similar brain imaging studies in navigation have shown the hippocampus to be active 102 a study was carried out on taxi drivers 103 london s black cab drivers need to learn the locations of a large number of places and the fastest routes between them in order to pass a strict test known as the knowledge in order to gain a license to operate 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 104 another study showed opposite findings in blind individuals the anterior part of the right hippocampus was larger and the posterior part was smaller compared with sighted individuals 105 role in approach avoidance conflict processing edit further information reward system 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 106 fmri findings from studies in approach avoidance decision making found evidence for a functional role that is not explained by either long term memory or spatial cognition 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 106 a review makes reference to a number of studies that show the involvement of the hippocampus in conflict tasks the authors suggest that one 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 66 role in social memory edit further information collective memory the hippocampus has received renewed attention for its role in social memory epileptic human subjects with depth electrodes in the left posterior left anterior or right anterior hippocampus demonstrate distinct individual cell responses when presented with faces of presumably recognizable famous people 107 associations among facial and vocal identity were similarly mapped to the hippocampus of rhesus monkeys single neurons in the ca1 and ca3 responded strongly to social stimulus recognition by mri the ca2 was not distinguished and may likely comprise a proportion of the claimed ca1 cells in the study 108 the dorsal ca2 and ventral ca1 subregions of the hippocampus have been implicated in social memory processing genetic inactivation of ca2 pyramidal neurons leads to pronounced loss of social memory while maintaining intact sociability in mice 109 similarly ventral ca1 pyramidal neurons have also been demonstrated as critical for social memory under optogenetic control in mice 110 111 role in olfaction edit in the early 20th century the widely held view was that olfaction was a major hippocampal function 112 this view was argued against pointing out that the hippocampus was present in some animals such as dolphins and whales that did not have a sense of smell and further that lesions in the temporal lobe in dogs had been shown to have no effect on their sense of smell 112 these arguments were concluded in 1947 and held for a few more decades in 1984 and 1987 studies in the rat showed that the entorhinal cortex receives substantial input from the olfactory bulb with part of the ec being directly innervated by the lateral olfactory tract 112 secondary inputs to the ec were also shown to include some from the periamygdaloid and piriform cortices and ca1 in the ventral hippocampus was shown to sends axons to the main olfactory bulb 113 112 it is evident that the hippocampus does have an involvement in memory for odors 114 115 physiology edit examples of rat hippocampal eeg and ca1 neural activity in the theta awake behaving and lia slow wave sleep modes each plot shows 20 seconds of data with a hippocampal eeg trace at the top spike rasters from 40 simultaneously recorded ca1 pyramidal cells in the middle each raster line represents a different cell and a plot of running speed at the bottom the top plot represents a time period during which the rat was actively searching for scattered food pellets for the bottom plot the rat was asleep 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 these modes are named after the eeg patterns associated with them theta and large irregular activity lia the main characteristics described below are for the rat which is the animal most extensively studied 116 the theta mode appears during states of active alert behavior especially locomotion and also during rem sleep dreaming 117 in the theta mode the eeg is dominated by large regular waves with a frequency range of 6 to 9 hz and the main groups of hippocampal neurons pyramidal cells and granule cells show sparse population activity which means that in any short time interval the great majority of cells are silent while the small remaining fraction fire at relatively high rates up to 50 spikes in one second for the most active of them 118 119 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 120 but other behavioral variables also clearly influence it the lia mode appears during slow wave sleep non dreaming and also during states of waking immobility such as resting or eating 117 in the lia mode the eeg is dominated by sharp waves that are randomly timed large deflections of the eeg signal lasting for 25 50 milliseconds sharp waves are frequently generated in sets with sets containing up to 5 or more individual sharp waves and lasting up to 500 ms the spiking activity of neurons within the hippocampus is highly correlated with sharp wave activity most neurons decrease their firing rate between sharp waves however during a sharp wave there is a dramatic increase in firing rate in up to 10 of the hippocampal population 121 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 122 hippocampal theta rhythm edit main article theta wave example of a one second eeg theta wave 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 123 in some situations the eeg is dominated by regular waves at 3 to 10 hz often continuing for many seconds these reflect subthreshold membrane potentials and strongly modulate the spiking of hippocampal neurons and synchronize across the hippocampus in a travelling wave pattern 124 the trisynaptic circuit is a relay of neurotransmission in the hippocampus that interacts with many brain regions from rodent studies it has been proposed that the trisynaptic circuit generates the hippocampal theta rhythm 125 theta rhythmicity previously clearly shown in rabbits and rodents has also been shown in humans 126 in rats the animals that have been the most extensively studied theta is seen mainly in two conditions first when an animal is walking or in some other way actively interacting with its surroundings second during rem sleep 127 the function of theta has not yet been convincingly explained although numerous theories have been proposed 116 the most popular hypothesis has been to relate it to learning and memory 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 128 it is well established that lesions of the medial septum the central node of the theta system cause severe disruptions of memory 129 however the medial septum is more than just the controller of theta it is also the main source of cholinergic projections to the hippocampus 19 it has not been established that septal lesions exert their effects specifically by eliminating the theta rhythm 130 sharp waves edit main article sharp waves and ripples 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 131 these events are associated with bursts of spike activity lasting 50 to 100 milliseconds in pyramidal cells of ca3 and ca1 they are also associated with short lived high frequency eeg oscillations called ripples with frequencies in the range 150 to 200 hz in rats and together they are known as sharp waves and ripples 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 122 sharp waves appear to be associated with memory 132 numerous later studies have 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 133 it has been proposed that sharp waves are in fact reactivations of neural activity patterns that were memorized during behavior driven by strengthening of synaptic connections within the hippocampus 134 this idea forms a key component of the two stage memory theory 135 advocated by buzsáki and others which proposes that memories are stored within the hippocampus during behavior and then later transferred to the neocortex during sleep 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 135 suppression of sharp waves and ripples in sleep or during immobility can interfere with memories expressed at the level of the behavior 136 137 nonetheless the newly formed ca1 place cell code can re emerge even after a sleep with abolished sharp waves and ripples in spatially non demanding tasks 138 long term potentiation edit see also long term potentiation and sleep and learning since at least the time of ramon y cajal 1852 1934 psychologists have speculated that the brain stores memory by altering the strength of connections between neurons that are simultaneously active 139 this idea was formalized by donald 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 lon...
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