The '''hippocampal formation''' is a compound structure in the [[medial temporal lobe]] of the [[brain]]. There is currently no consensus concerning which brain regions are encompassed by the term, with some authors defining it as the [[dentate gyrus]], the [[hippocampus]] proper and the [[subiculum]];<ref name= Martin2003>{{cite book |last1= Martin|first1= JH|title= Neuroanatomy: text and atlas|edition= third|year= 2003|publisher= McGraw-Hill Companies|location= |isbn= 007121237X|page= 382|chapter= Lymbic system and cerebral circuits for emotions, learning, and memory|chapterurl= http://books.google.com.au/books?id=OUC4igr3O4sC&pg=PA382&lpg=PA382&dq=%22the+hippocampal+formation+consists+of+three+components%22&source=bl&ots=gxkaQHHin-&sig=1u-3jI1zqp0VJgVLatYzMqzXaNk&hl=en&ei=EFlPS7fHJ8qHkAXcu-i4Cg&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22the%20hippocampal%20formation%20consists%20of%20three%20components%22&f=false}}</ref> and others including also the presubiculum, [[parasubiculum]], and [[entorhinal cortex]].<ref name= Amaral2007-1a>{{cite book |last1= Amaral|first1= D|last2= Lavenex|first2= P|editor1-first= P|editor1-last= Anderson|editor2-first= R|editor2-last= Morris|editor3-first= D|editor3-last= Amaral|editor4-first= T|editor4-last= Bliss|editor5-first= J|editor5-last= I'Keefe|title= The hippocampus book|edition= first|year= 2007|publisher= Oxford University Press|location= New York|isbn= 0195200271|page= 37|chapter= Hippocampal neuroanatomy|chapterurl= http://books.google.com.au/books?id=zg6oyF1DziQC&pg=PA37&lpg=PA37&dq=%22hippocampal+researchers+tend+to+follow+one+of+several+%22&source=bl&ots=iC5i7KHMoz&sig=D9WaDRnm6nVreL3ePOvANKFwqrE&hl=en&ei=uI1SS8bTFYGg6gPO2qGiBw&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22hippocampal%20researchers%20tend%20to%20follow%20one%20of%20several%20%22&f=true}}</ref> The hippocampal formation is thought to play a role in memory, spatial navigation and control of attention. The neural layout and pathways within the hippocampal formation are very similar in all mammals.<ref name= Amaral2007-1b>{{cite book |last1= Anderson|first1= P|last2= Morris|first2= R|last3= Amaral|first3= D|last4= Bliss|first4=T|last5=O'Keefe|first5=J|editor1-first= P|editor1-last= Anderson|editor2-first= R|editor2-last= Morris|editor3-first= D|editor3-last= Amaral|editor4-first= T|editor4-last= Bliss|editor5-first= J|editor5-last= I'Keefe|title= The hippocampus book|edition= first|year= 2007|publisher= Oxford University Press|location= New York|isbn= 0195200271|page= 3|chapter= The hippocampal formation|chapterurl= http://books.google.com.au/books?id=zg6oyF1DziQC&pg=PA3&lpg=PA3&dq=%22The+basic+layout+of+cells+and+fiber+pathways+of+the+hippocampal+formation+is+much+the+same+in+all+mammals.%22&source=bl&ots=iC5i6PKNqA&sig=z7Up41y-_7sk-TZUGHgxyj94N78&hl=en&ei=mNZRS7T7O4zc7AOVj8yjCg&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22The%20basic%20layout%20of%20cells%20and%20fiber%20pathways%20of%20the%20hippocampal%20formation%20is%20much%20the%20same%20in%20all%20mammals.%22&f=true}}</ref>
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The '''hippocampal formation''' is a compound structure in the [[medial temporal lobe]] of the [[brain]]. There is currently no consensus concerning which brain regions are encompassed by the term, with some authors defining it as the [[dentate gyrus]], the [[hippocampus]] proper and the [[subiculum]];<ref name= Martin2003>{{cite book |last1= Martin|first1= JH|title= Neuroanatomy: text and atlas|edition= third|year= 2003|publisher= McGraw-Hill Companies|location= |isbn= 007121237X|page= 382|chapter= Lymbic system and cerebral circuits for emotions, learning, and memory|chapterurl= http://books.google.com.au/books?id=OUC4igr3O4sC&pg=PA382&lpg=PA382&dq=%22the+hippocampal+formation+consists+of+three+components%22&source=bl&ots=gxkaQHHin-&sig=1u-3jI1zqp0VJgVLatYzMqzXaNk&hl=en&ei=EFlPS7fHJ8qHkAXcu-i4Cg&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22the%20hippocampal%20formation%20consists%20of%20three%20components%22&f=false}}</ref> and others including also the presubiculum, [[parasubiculum]], and [[entorhinal cortex]].<ref name= Amaral2007-1a>{{cite book |last1= Amaral|first1= D|last2= Lavenex|first2= P|editor1-first= P|editor1-last= Anderson|editor2-first= R|editor2-last= Morris|editor3-first= D|editor3-last= Amaral|editor4-first= T|editor4-last= Bliss|editor5-first= J|editor5-last= I'Keefe|title= The hippocampus book|edition= first|year= 2007|publisher= Oxford University Press|location= New York|isbn= 0195200271|page= 37|chapter= Hippocampal neuroanatomy|chapterurl= http://books.google.com.au/books?id=zg6oyF1DziQC&pg=PA37&lpg=PA37&dq=%22hippocampal+researchers+tend+to+follow+one+of+several+%22&source=bl&ots=iC5i7KHMoz&sig=D9WaDRnm6nVreL3ePOvANKFwqrE&hl=en&ei=uI1SS8bTFYGg6gPO2qGiBw&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22hippocampal%20researchers%20tend%20to%20follow%20one%20of%20several%20%22&f=true}}</ref> The hippocampal formation is thought to play a role in [[memory]], spatial [[navigation]] and control of [[attention]]. The neural layout and pathways within the hippocampal formation are very similar in all mammals.<ref name= Amaral2007-1b>{{cite book |last1= Anderson|first1= P|last2= Morris|first2= R|last3= Amaral|first3= D|last4= Bliss|first4=T|last5=O'Keefe|first5=J|editor1-first= P|editor1-last= Anderson|editor2-first= R|editor2-last= Morris|editor3-first= D|editor3-last= Amaral|editor4-first= T|editor4-last= Bliss|editor5-first= J|editor5-last= I'Keefe|title= The hippocampus book|edition= first|year= 2007|publisher= Oxford University Press|location= New York|isbn= 0195200271|page= 3|chapter= The hippocampal formation|chapterurl= http://books.google.com.au/books?id=zg6oyF1DziQC&pg=PA3&lpg=PA3&dq=%22The+basic+layout+of+cells+and+fiber+pathways+of+the+hippocampal+formation+is+much+the+same+in+all+mammals.%22&source=bl&ots=iC5i6PKNqA&sig=z7Up41y-_7sk-TZUGHgxyj94N78&hl=en&ei=mNZRS7T7O4zc7AOVj8yjCg&sa=X&oi=book_result&ct=result&resnum=1&ved=0CAcQ6AEwAA#v=onepage&q=%22The%20basic%20layout%20of%20cells%20and%20fiber%20pathways%20of%20the%20hippocampal%20formation%20is%20much%20the%20same%20in%20all%20mammals.%22&f=true}}</ref>
==Function==
==Function==
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During the nineteenth and early twentieth centuries, based largely on the observation that, between species, the size of the [[olfactory bulb]] varies with the size of the parahippocampal gyrus, the hippocampal formation was thought to be part of the olfactory system.<ref name= Finger2001>{{cite book |last1= Finger|first1= S|title= Origins of neuroscience: a history of explorations into brain function|edition= |year= 2001|publisher= Oxford University Press|location= Oxford/NewYork|isbn= 0195065034|page= 286|chapter= Defining and controlling the circuits of emotion}}</ref>
During the nineteenth and early twentieth centuries, based largely on the observation that, between species, the size of the [[olfactory bulb]] varies with the size of the parahippocampal gyrus, the hippocampal formation was thought to be part of the olfactory system.<ref name= Finger2001>{{cite book |last1= Finger|first1= S|title= Origins of neuroscience: a history of explorations into brain function|edition= |year= 2001|publisher= Oxford University Press|location= Oxford/NewYork|isbn= 0195065034|page= 286|chapter= Defining and controlling the circuits of emotion}}</ref>
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In 1937 [[James Papez|Papez]] theorized that a circuit including the hippocampal formation constitutes the neural substrate of emotional behavior,<ref name= Papez1937>{{cite journal |last1= Papez|first1= JW|title= A proposed mechanism of emotion|journal= Archives of neurology and psychiatry|year= 1937|volume= 38|pages= 725–43}}</ref> and [[Heinrich Klüver|Klüver]] and [[Paul Bucy|Bucy]] reported that, in monkeys, [[resection]] involving the hippocampal formation and the [[amygdala|amygdaloid complex]] has a profound effect on emotional responses.<ref name= Klüver1937>{{cite journal |last1= Klüver|first1= H|last2= Bucy|first2= PC|title= "Psychic blindness" and other symptoms following bilateral temporal lobectomy in Rhesus monkeys|journal= American Journal of Physiology|year= 1937|volume= 119|pages= 352–53|pmid= |doi= |url= }}</ref><ref name= Klüver1939>{{cite journal |last1= Klüver |first1= H|last2= Bucy|first2= PC|title= Preliminary analysis of functions of the temporal lobes in monkeys|journal= Archives of Neurology and psychiatry|year= 1939|volume= 42|pages= 979–1000}}</ref> As a consequence of these publications, the idea that the hippocampal formation is entirely dedicated to [[olfaction]] began to recede.<ref name= Nieuwenhuys2008>{{cite book |last1= Nieuwenhuys|first1= R|last2= Voogd|first2= J|last3= van Huijzen|first3= C|title= The human central nervous system|edition= fourth|year= 2008|publisher= Springer-Verlag|location= Berlin/Heidelberg/New York|isbn= 3540134417|page= 917|chapter= The greater limbic system|chapterurl= http://books.google.com.au/books?id=vAiFe7gZLhoC&pg=PT916&dq=%22from+limbic+lobe+to+limbic+system%22&client=firefox-a&cd=1#v=onepage&q=%22from%20limbic%20lobe%20to%20limbic%20system%22&f=true}}</ref> (Today there is little support for the idea that the hippocampal formation is directly involved in emotion.)
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In 1937 [[James Papez|Papez]] theorized that a circuit including the hippocampal formation constitutes the neural substrate of emotional behavior,<ref name= Papez1937>{{cite journal |last1= Papez|first1= JW|title= A proposed mechanism of emotion|journal= Archives of neurology and psychiatry|year= 1937|volume= 38|pages= 725–43}}</ref> and [[Heinrich Klüver|Klüver]] and [[Paul Bucy|Bucy]] reported that, in monkeys, [[resection]] involving the hippocampal formation and the [[amygdala|amygdaloid complex]] has a profound effect on emotional responses.<ref name= Klüver1937>{{cite journal |last1= Klüver|first1= H|last2= Bucy|first2= PC|title= "Psychic blindness" and other symptoms following bilateral temporal [[lobectomy]] in [[Rhesus monkeys]]|journal= American Journal of Physiology|year= 1937|volume= 119|pages= 352–53|pmid= |doi= |url= }}</ref><ref name= Klüver1939>{{cite journal |last1= Klüver |first1= H|last2= Bucy|first2= PC|title= Preliminary analysis of functions of the temporal lobes in monkeys|journal= Archives of Neurology and psychiatry|year= 1939|volume= 42|pages= 979–1000}}</ref> As a consequence of these publications, the idea that the hippocampal formation is entirely dedicated to [[olfaction]] began to recede.<ref name= Nieuwenhuys2008>{{cite book |last1= Nieuwenhuys|first1= R|last2= Voogd|first2= J|last3= van Huijzen|first3= C|title= The human central nervous system|edition= fourth|year= 2008|publisher= Springer-Verlag|location= Berlin/Heidelberg/New York|isbn= 3540134417|page= 917|chapter= The greater limbic system|chapterurl= http://books.google.com.au/books?id=vAiFe7gZLhoC&pg=PT916&dq=%22from+limbic+lobe+to+limbic+system%22&client=firefox-a&cd=1#v=onepage&q=%22from%20limbic%20lobe%20to%20limbic%20system%22&f=true}}</ref> (Today there is little support for the idea that the hippocampal formation is directly involved in emotion.)
[[Image:CajalHippocampus (modified).png|thumb|300px|left|The hippocampal formation, as drawn by [[Santiago Ramon y Cajal]]. DG: dentate gyrus. Sub: subiculum. EC: entorhinal cortex. CA1-CA3: hippocampus proper]]In an influential 1947 review, [[Alf Brodal]] pointed out that mammal species thought to have no sense of smell nevertheless have fully intact hippocampal formations, that removal of the hippocampal formation did not affect the ability of dogs to perform tasks dependent on olfaction, and that no [[axon|fibers]] were actually known that carry information directly from the olfactory bulb to any part of the hippocampal formation.<ref name= Brodal1947>{{cite journal |last1= Brodal|first1= A|title= Hippocampus and the sense of smell|journal= Brain|year= 1947|volume= 70|pages= 179–222|url= http://brain.oxfordjournals.org/cgi/pdf_extract/70/2/179}}</ref> Though massive direct input from the olfactory bulb to the entorhinal cortex has subsequently been found,<ref name= Shipley1984>{{cite journal |last1= Shipley|first1= MT|last2= Adamek|first2= GD|title= The connections of the mouse olfactory bulb: a study using orthograde and retrograde transport of wheatgerm agglutinin conjugated to horsradish peroxidase|journal= Brain research bulletin|year= 1984|volume= 12|pages= 669–688|pmid= 6206930}}</ref> the current view is that the hippocampal formation is not an integral part of the olfactory system.<ref name= Amaral2007-2/>
[[Image:CajalHippocampus (modified).png|thumb|300px|left|The hippocampal formation, as drawn by [[Santiago Ramon y Cajal]]. DG: dentate gyrus. Sub: subiculum. EC: entorhinal cortex. CA1-CA3: hippocampus proper]]In an influential 1947 review, [[Alf Brodal]] pointed out that mammal species thought to have no sense of smell nevertheless have fully intact hippocampal formations, that removal of the hippocampal formation did not affect the ability of dogs to perform tasks dependent on olfaction, and that no [[axon|fibers]] were actually known that carry information directly from the olfactory bulb to any part of the hippocampal formation.<ref name= Brodal1947>{{cite journal |last1= Brodal|first1= A|title= Hippocampus and the sense of smell|journal= Brain|year= 1947|volume= 70|pages= 179–222|url= http://brain.oxfordjournals.org/cgi/pdf_extract/70/2/179}}</ref> Though massive direct input from the olfactory bulb to the entorhinal cortex has subsequently been found,<ref name= Shipley1984>{{cite journal |last1= Shipley|first1= MT|last2= Adamek|first2= GD|title= The connections of the mouse olfactory bulb: a study using orthograde and retrograde transport of wheatgerm agglutinin conjugated to horsradish peroxidase|journal= Brain research bulletin|year= 1984|volume= 12|pages= 669–688|pmid= 6206930}}</ref> the current view is that the hippocampal formation is not an integral part of the olfactory system.<ref name= Amaral2007-2/>
The hippocampal formation is a compound structure in the medial temporal lobe of the brain. There is currently no consensus concerning which brain regions are encompassed by the term, with some authors defining it as the dentate gyrus, the hippocampus proper and the subiculum;[1] and others including also the presubiculum, parasubiculum, and entorhinal cortex.[2] The hippocampal formation is thought to play a role in memory, spatial navigation and control of attention. The neural layout and pathways within the hippocampal formation are very similar in all mammals.[3]
During the nineteenth and early twentieth centuries, based largely on the observation that, between species, the size of the olfactory bulb varies with the size of the parahippocampal gyrus, the hippocampal formation was thought to be part of the olfactory system.[4]
In 1937 Papez theorized that a circuit including the hippocampal formation constitutes the neural substrate of emotional behavior,[5] and Klüver and Bucy reported that, in monkeys, resection involving the hippocampal formation and the amygdaloid complex has a profound effect on emotional responses.[6][7] As a consequence of these publications, the idea that the hippocampal formation is entirely dedicated to olfaction began to recede.[8] (Today there is little support for the idea that the hippocampal formation is directly involved in emotion.)
File:CajalHippocampus (modified).pngThe hippocampal formation, as drawn by Santiago Ramon y Cajal. DG: dentate gyrus. Sub: subiculum. EC: entorhinal cortex. CA1-CA3: hippocampus properIn an influential 1947 review, Alf Brodal pointed out that mammal species thought to have no sense of smell nevertheless have fully intact hippocampal formations, that removal of the hippocampal formation did not affect the ability of dogs to perform tasks dependent on olfaction, and that no fibers were actually known that carry information directly from the olfactory bulb to any part of the hippocampal formation.[9] Though massive direct input from the olfactory bulb to the entorhinal cortex has subsequently been found,[10] the current view is that the hippocampal formation is not an integral part of the olfactory system.[11]
In 1900, the Russian neurologist Vladimir Bekhterev described two patients with a significant memory deficit who, on autopsy, were found to have softening of hippocampal and adjacent cortical tissue;[12] and in 1957, Scoville and Milner reported memory loss in a series of patients subsequent to their removal of the patients' medial temporal lobes.[13] Thanks to these observations and a great deal of subsequent research, it is now broadly accepted that the hippocampal formation plays a role in some aspects of memory.[11]
EEG evidence from 1938 to the present, stimulation evidence from the 1950's, and modern imaging techniques together suggest a role for some part of the hippocampal formation (in concert with the anterior cingulate cortex) in the control of attention.[11]
In 1971 John O'Keefe and his student Jonathan Dostrovsky discovered neurons in the rat hippocampus that appeared to show activity related to the rat's location within its environment.[14] Despite skepticism from other investigators, O'Keefe and his co-workers, including Lynn Nadel, continued to investigate this question, in a line of work that eventually led to their very influential 1978 book The Hippocampus as a Cognitive Map.[15] As with the memory theory, there is now almost universal agreement that the hippocampal formation plays an important role in spatial coding, but the details are widely debated.[16]
↑ (2001) "Defining and controlling the circuits of emotion" Origins of neuroscience: a history of explorations into brain function, Oxford/NewYork: Oxford University Press.
↑ (1937). A proposed mechanism of emotion. Archives of neurology and psychiatry38: 725–43.
↑ (1937). "Psychic blindness" and other symptoms following bilateral temporal lobectomy in Rhesus monkeys. American Journal of Physiology119: 352–53.
↑ (1939). Preliminary analysis of functions of the temporal lobes in monkeys. Archives of Neurology and psychiatry42: 979–1000.
↑ (2008) "The greater limbic system" The human central nervous system, fourth, Berlin/Heidelberg/New York: Springer-Verlag.
↑ (1984). The connections of the mouse olfactory bulb: a study using orthograde and retrograde transport of wheatgerm agglutinin conjugated to horsradish peroxidase. Brain research bulletin12: 669–688.
↑ (1900). Demonstration eines gehirns mit zerstörung der vorderen und inneren theile der hirnrinde beider schläfenlappen. Neurologische Zeitenblatte19: 990–991.
↑Scoville, WB, Milner B (1957). Loss of Recent Memory After Bilateral Hippocampal Lesions. Journal of neurology, neurosurgery and psychiatry20 (1): 11–21.
↑O'Keefe J, Dostrovsky J (1971). The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. Brain Res34 (1): 171–75.
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