Kodi Zithunzi Zolaula Zingagwiritse Ntchito Kukhala Zenizeni Zowononga Ubongo? (2011)

NDEMANGA: Iyi ndi buku la Dr. Hilton lotchedwa Pornography Addiction: A Neuroscience Perspective (2011) , lomwe likupezeka m'gawo lomweli. Iye ali wotsimikiza, monga momwe tilili, kuti mphotho zachilengedwe zimatha kukhala zosokoneza bongo ndipo zimayambitsa kusintha kwa ubongo komweko monga mankhwala osokoneza bongo. Pepala lake laposachedwa lomwe adawunikidwa ndi anzawo ndi Pornography addiction - chilimbikitso chapamwamba chomwe chimaganiziridwa pankhani ya neuroplasticity | Hilton | Socioaffective Neuroscience & Psychology (2013).


January 20, 2011
Donald L. Hilton, Jr. MD, FACS
Pulofesa Wothandizana ndi Chipatala
Dipatimenti ya Neurosurgery
University of Texas Health Sciences Center ku San Antonio

Ubongo wamunthu udapangidwa kuti ukalimbikitse zizolowezi zomwe zimapangitsa kuti munthu akhale ndi moyo. Dongosolo la mesolimbic dopaminergic limapatsa mphotho kudya ndi kugonana ndi zolimbikitsa zamphamvu zosangalatsa. Cocaine, opioid, mowa, ndi mankhwala ena osokoneza bongo amapotoza, kapena kulanda, machitidwe osangalatsawa, ndikupangitsa ubongo kuganiza kuti mankhwala osokoneza bongo ndiofunikira kuti apulumuke. Umboni uli wamphamvu tsopano kuti mphotho zachilengedwe monga chakudya ndi kugonana zimakhudza machitidwe a mphotho momwemonso mankhwalawo amawakhudzira, motero chidwi chomwe chilipo tsopano pa 'chizolowezi chachilengedwe.' Kuledzera, kaya ndi cocaine, chakudya, kapena kugonana kumachitika pamene izi zimaleka kuchititsa homeostasis, ndipo m'malo mwake zimabweretsa zovuta. Mwachitsanzo, pamene kudya kumayambitsa kunenepa kwambiri ndi ochepa omwe anganene kuti chamoyocho chili bwino. Momwemonso, zolaula zimapweteketsa munthu zikawononga kapena kuwononga chidwi chake.

Zaka khumi zapitazo umboni unayamba kusonyeza kuti munthu amamwa mopitirira muyeso makhalidwe achilengedwe omwe amachititsa kuti ubongo ulandire mphotho ya dopaminergic. Mwachitsanzo, Dr. Howard Shaffer, Mtsogoleri wa Kafukufuku wa Zoledzeretsa ku Harvard University, anati mu 2001, “Ndinavutika kwambiri ndi anzanga pamene ndinanena kuti chizolowezi chochuluka chimachokera ku zomwe adakumana nazo … zomwe zimachitika mobwerezabwereza, mokweza, komanso pafupipafupi. Koma zaonekeratu kuti kusintha kwa mitsempha ya ubongo—ndiko kuti, kusintha kwa mitsempha ya ubongo komwe kumathandiza kupititsa patsogolo khalidweli—kumachitika ngakhale kuti palibe mankhwala osokoneza bongo” [1] M’zaka khumi kuchokera pamene ananena izi, wakhala akufufuza kwambiri za zotsatira za ubongo za zizolowezi zachilengedwe monga kutchova juga. Onani zotsatirazi kuchokera mu pepala lomweli la Sayansi la 2001.

Akatswiriwa amakonda kunena kuti chizolowezi chimachitika pamene chizolowezi "chimaba" ubongo womwe unasintha kuti ukhale ndi khalidwe lolimbikitsa kupulumuka monga kudya ndi kugonana. "N'zomveka ngati mungathe kusokoneza ma circuits awa ndi mankhwala, mungathenso kuchita ndi mphotho zachilengedwe," akutero katswiri wa zamaganizo wa ku Stanford University Brian Knutson. Chifukwa chake, mankhwala osokoneza bongo salinso pakati pa nkhaniyi. "Chomwe chikubwera mwachangu ngati nkhani yayikulu ... ndikupitiriza kuchita zinthu zowononga ngakhale zotsatirapo zoyipa," akutero Steven Grant wa NIDA. [2]

M'zaka khumi kuchokera pamene malingaliro osinthikawa adafotokozedwa koyamba, umboni wa lingaliro la kuledzera kwa mphotho zachilengedwe wangokulirakulira. Mu 2005 Dr. Eric Nestler, yemwe tsopano ndi tcheyamani wa sayansi ya ubongo ku Mount Sinai Medical Center ku New York adasindikiza pepala lofunika kwambiri mu Nature Neuroscience lotchedwa "Kodi Pali Njira Yofanana Yoledzera?" Iye anati: ""Umboni wokulirapo ukusonyeza kuti njira ya VTA-NAc ndi madera ena a limbic omwe atchulidwa pamwambapa amagwirizananso, makamaka pang'ono, ndi zotsatira zabwino zamaganizo za mphotho zachilengedwe, monga chakudya, kugonana ndi kuyanjana ndi anthu. Madera omwewa aphatikizidwanso mu zomwe zimatchedwa 'kuledzera kwachilengedwe' (ndiko kuti, kudya mokakamiza mphotho zachilengedwe) monga kudya mopitirira muyeso, kutchova juga kwa matenda ndi zizolowezi zogonana. Zomwe zapezeka poyamba zikusonyeza kuti njira zomwe zimagawidwa zitha kukhalapo: [chitsanzo ndi] kukhudzidwa komwe kumachitika pakati pa mphotho zachilengedwe ndi mankhwala osokoneza bongo." [3]

Mu 2002 kafukufuku wokhudza kuledzera kwa cocaine adasindikizidwa womwe udawonetsa kutayika kwa voliyumu komwe kumayesedwa m'malo angapo a ubongo, kuphatikiza ma lobes akutsogolo. [4] Njirayi inali kugwiritsa ntchito njira yochokera ku MRI yotchedwa voxel-based morphometry (VBM), komwe ma milimita imodzi ya ubongo amayesedwa ndikuyerekezeredwa. Kafukufuku wina wa VBM adasindikizidwa mu 2004 pa methamphetamine yokhala ndi zotsatira zofanana kwambiri. [5] Ngakhale ndizosangalatsa, zomwe zapezekazi sizodabwitsa kwa wasayansi kapena munthu wamba, chifukwa izi ndi "mankhwala enieni."

Nkhaniyi imakhala yosangalatsa kwambiri tikayang'ana chizolowezi chachilengedwe monga kudya mopitirira muyeso komwe kumabweretsa kunenepa kwambiri. Mu 2006 kafukufuku wa VBM adasindikizidwa akuyang'ana makamaka kunenepa kwambiri, ndipo zotsatira zake zinali zofanana kwambiri ndi kafukufuku wa cocaine ndi methamphetamine. [6] Kafukufuku wa kunenepa kwambiri adawonetsa madera angapo a kutayika kwa voliyumu, makamaka m'malo ozungulira, madera okhudzana ndi chiweruzo ndi kuwongolera. Ngakhale kuti kafukufukuyu ndi wofunikira pakuwonetsa kuwonongeka kooneka mu chizolowezi chachilengedwe, mosiyana ndi chizolowezi cha mankhwala osokoneza bongo, zimakhala zosavuta kuvomereza mwachibadwa chifukwa timatha kuwona zotsatira za kudya mopitirira muyeso mwa munthu wonenepa kwambiri.

Nanga bwanji za chizolowezi chogonana? Mu 2007 kafukufuku wa VBM wochokera ku Germany adayang'ana makamaka za pedophilia, ndipo adawonetsa zomwe zapezeka pafupifupi zofanana ndi kafukufuku wa cocaine, methamphetamine, ndi kunenepa kwambiri. [7] Kufunika kwa kafukufukuyu pokhudzana ndi nkhaniyi ndikofunikira kwambiri chifukwa Kukuwonetsa kuti chilakolako chogonana chingayambitse kusintha kwa thupi, komwe kumakhudza ubongo, mwachitsanzo, kuvulaza. Chochititsa chidwi n'chakuti, pepala laposachedwa lapeza mgwirizano waukulu pakati pa zolaula zogonana ndi ana ndi nkhanza zogonana ndi ana. [8] Izi zati, pepalalo lidayang'ana kwambiri gulu laling'ono lomwe, pakati pa mavuto ena, chizolowezi chachikulu cha zolaula. Ngakhale titha kusiyanitsa pakati pa zolaula za ana ndi akuluakulu, ubongo sungakhale ndi nthawi yofanana ndi yaukalamba pankhani yochepetsa dopaminergic ndi kuchepa kwa voliyumu chifukwa cha chizolowezi. Kodi ubongo umasamala ngati munthuyo akuchita zogonana, kapena akuchita izi kudzera mu kugonana, mwachitsanzo, zolaula. Machitidwe agalasi a ubongo amasintha zomwe zimachitika pa zolaula kukhala zenizeni, malinga ndi ubongo. Izi zikuthandizidwa ndi kafukufuku waposachedwa wochokera ku France wosonyeza kuyambika kwa madera okhudzana ndi ma neuron agalasi muubongo wa munthu mwa amuna omwe amaonera zolaula. Olembawo akuti, "tikuganiza kuti ... dongosolo la ma neuron agalasi limalimbikitsa owonera kuti agwirizane ndi momwe anthu ena amaonekera m'mawonekedwe owonetsa za kugonana." [9] Kafukufuku woyambirira amathandizira kuwonongeka kwa kutsogolo makamaka kwa odwala omwe sangathe kulamulira khalidwe lawo logonana. [10] Kafukufukuyu adagwiritsa ntchito MRI yofalitsa kuti aone momwe mitsempha imafalikira kudzera mu zinthu zoyera, komwe kuli ma axon, kapena mawaya olumikizira maselo amitsempha. Adawonetsa kusagwira ntchito bwino m'dera lapamwamba la kutsogolo, dera logwirizana ndi kukakamizidwa, chizindikiro cha chizolowezi.

Kafukufuku wambiri akuwonetsa kusintha kwa kagayidwe kachakudya m'mitsempha ya ubongo pamene ubongo "umaphunzira" kukhala woledzera. Kusintha kumeneku komwe kumachitika mu dopamine reward system kumathanso kuyesedwa ndi ma brain scan monga ma MRI, PET, ndi SPECT scan. Ngakhale tikuyembekezera kuti kafukufuku wa ubongo wowunikira ubongo awonetse zolakwika mu kagayidwe ka dopamine mu cocaine, [11] tingadabwe kupeza kuti kafukufuku waposachedwa akuwonetsanso kusagwira ntchito bwino kwa malo osangalatsa omwewa ndi kutchova juga kwa matenda. [12] Kudya mopitirira muyeso komwe kumabweretsa kunenepa kwambiri, chizolowezi china chachilengedwe, kukuwonetsanso matenda ofanana. [13]

Nkhani ina yofunikira ndi pepala lochokera ku Mayo Clinic lonena za chithandizo cha chizolowezi cha zolaula pa intaneti ndi naltrexone, mankhwala oletsa opioid receptors. [14] Madokotala Bostsick ndi Bucci ku Mayo Clinic adachiza wodwala yemwe anali ndi vuto lolephera kuwongolera kugwiritsa ntchito kwake zolaula pa intaneti.

Anayikidwa pa naltrexone, mankhwala omwe amagwiritsidwa ntchito pa opioid system kuti achepetse kutaya kwa dopamine kuti ipangitse maselo mu nucleus accumbens. Ndi mankhwalawa adatha kuwongolera moyo wake wogonana.

Olemba analemba kuti:

Mwachidule, mawonekedwe a makina a addict's PFC amachititsa kuwonjezeka kwakukulu kwa mankhwala okhudzana ndi mankhwala osokoneza bongo, kuchepetsa mphamvu ya mankhwala osokoneza bongo, komanso kuchepa chidwi chotsatira zolinga zomwe zatsogoleredwa kuti zikhale zofunika kwambiri. Kuwonjezera pa kuvomereza kwa naltrexone kuchokera ku Food and Drug Administration kuti athetse uchidakwa, zochitika zambiri zofalitsidwa m'mabukuzi zasonyeza kuti zingatheke kuthetsa njuga, kudzivulaza, kleptomania, ndi chilakolako chogonana. Tikukhulupirira kuti ichi ndi kufotokoza koyambirira kwa kagwiritsidwe kake kolimbana ndi kugonana kwa pa Intaneti.

Bungwe lodziwika bwino la Royal Society of London linakhazikitsidwa m'ma 1660, ndipo limafalitsa magazini yakale kwambiri ya sayansi padziko lonse lapansi. Mu nkhani yaposachedwa ya Philosophical Transactions of the Royal Society , momwe kumvetsetsa kwa chizolowezi choledzera kunanenedwera pamene kunakambidwa ndi asayansi ena otsogola padziko lonse lapansi pamsonkhano wa Society. Mutu wa nkhani ya magazini yomwe inanena za msonkhanowo unali "The neurobiology of chizolowezi - new vistas." Chochititsa chidwi n'chakuti, mwa nkhani 17, ziwiri zinali zokhudzana ndi chizolowezi chachilengedwe: kutchova juga kwa matenda [15] ndi pepala lolembedwa ndi Dr. Nora Volkow lonena za kufanana kwa ubongo ndi chizolowezi cha mankhwala osokoneza bongo komanso kudya mopitirira muyeso [16] . Pepala lachitatu lolembedwa ndi Dr. Nestler linalankhula za zitsanzo za nyama zomwe chizolowezi chachilengedwe chimakhudzanso DFosB. [17]

DFosB ndi mankhwala omwe Dr. Nestler waphunzira, ndipo akuwoneka kuti amapezeka m'mitsempha ya anthu omwe ali ndi vuto la kugwiritsa ntchito mankhwala osokoneza bongo. Zikuwoneka kuti zili ndi gawo la thupi, koma zimakhudza kwambiri kuledzera. Chochititsa chidwi n'chakuti, idapezeka koyamba m'maselo a ubongo a nyama zomwe zidaphunziridwa mu kuledzera kwa mankhwala osokoneza bongo, koma tsopano yapezeka m'maselo a ubongo omwe ali mu nucleus accumbens okhudzana ndi kugwiritsa ntchito mopitirira muyeso mphotho zachilengedwe. [i] Pepala laposachedwa lofufuza za DFosB ndi gawo lake pakugwiritsa ntchito mopitirira muyeso mphotho ziwiri zachilengedwe, kudya ndi kugonana, likumaliza kuti:

Mwachidule, ntchito yomwe yaperekedwa pano ikupereka umboni wakuti, kuwonjezera pa kugwiritsa ntchito mankhwala osokoneza bongo, mphotho zachilengedwe zimapangitsa kuti milingo ya DFB mu Nac ikhale yochepa… zotsatira zathu zikuwonjezera kuthekera kwakuti kulowetsedwa kwa DFosB mu NAc kungathandize osati kokha pazinthu zofunika kwambiri zokhudzana ndi kugwiritsa ntchito mankhwala osokoneza bongo, komanso mbali za zomwe zimatchedwa zizolowezi zachilengedwe zomwe zimaphatikizapo kumwa mopitirira muyeso mphotho zachilengedwe. [18]

Dr. Nora Volkow ndi mtsogoleri wa National Institute on Drug Abuse (NIDA), ndipo ndi m'modzi mwa asayansi odziwika bwino pankhani ya chizolowezi choledzera padziko lonse lapansi. Iye wazindikira kusintha kumeneku pakumvetsetsa chizolowezi chachilengedwe ndipo walimbikitsa kusintha dzina la NIDA kukhala National Institute on Diseases of Addiction. Magazini ya Science inanena kuti: “Mtsogoleri wa NIDA Nora Volkow nayenso adaganiza kuti dzina la bungwe lake liyenera kuphatikizapo zizolowezi monga zolaula , kutchova juga, ndi chakudya, akutero mlangizi wa NIDA Glen Hanson. ‘Akufuna kutumiza uthenga wakuti [tiyenera] kuyang'ana gawo lonselo.’” [19] (kugogomezedwa kwina).

Mwachidule, m'zaka 10 zapitazi umboni tsopano ukuchirikiza kwambiri chibadwa cha mphotho zachilengedwe zomwe zimawapangitsa kukhala oledzera. Madokotala Malenka ndi Kauer, m'pepala lawo lodziwika bwino lonena za momwe kusintha kwa mankhwala kumachitika m'maselo a ubongo a anthu omwe ali ndi chizolowezi choledzera amati, "chizolowezi chimayimira njira yophunzirira komanso kukumbukira komwe kumachitika m'maselo a ubongo a anthu omwe ali ndi chizolowezi choledzera." [20] Tsopano timatcha kusintha kumeneku m'maselo a ubongo kuti "mphamvu ya nthawi yayitali" komanso "kuvutika maganizo kwa nthawi yayitali," ndipo timalankhula za ubongo ngati pulasitiki, kapena womwe ungasinthe ndikubwezeretsanso mawaya. Dr. Norman Doidge, katswiri wa mitsempha ku Columbia, m'buku lake lakuti The Brain That Changes Itself akufotokoza momwe zolaula zimayambitsa kubwezeretsanso mawaya a mitsempha. Akufotokoza kafukufuku wokhudza amuna omwe amaonera zolaula pa intaneti momwe amaoneka "mosayembekezereka" ngati makoswe omwe amakankhira lever kuti alandire cocaine m'mabokosi oyesera a Skinner. Monga makoswe omwe ali ndi chizolowezi choledzera, akufunafuna njira yotsatira yothetsera vutoli, akudina mbewa pomwe khoswe akukankhira lever. Kusuta zolaula ndi kuphunzira mwachangu , ndipo mwina ichi ndichifukwa chake ambiri omwe alimbana ndi chizolowezi choledzera kangapo amanena kuti chinali chizolowezi chovuta kwambiri kwa iwo kuti achithetse. Ngakhale kuti zizolowezi za mankhwala osokoneza bongo zili ndi mphamvu, zimakhala zopanda ntchito kwenikweni, pomwe kuonera zolaula, makamaka pa intaneti, ndi njira yogwira ntchito kwambiri ya ubongo. Kufufuza nthawi zonse ndikuwunika chithunzi chilichonse kapena kanema wopangidwa kuti aone mphamvu ndi zotsatira zake ndi ntchito yophunzirira ndi kulumikiza mawaya a mitsempha.

Kufika pachimake pa kugonana kwa anthu kumagwiritsa ntchito njira zomwezo zopezera mphotho monga momwe zimakhalira ndi anthu omwe amamwa mowa wa heroin. [21] Ngati sitikumvetsa tanthauzo la luso la zolaula lokonzanso ubongo m'mapangidwe ake, m'mitsempha, komanso m'thupi, tidzadzipangitsa kuti tipitirize kulephera pochiza matenda oopsawa. Komabe, ngati tipereka mphotho yachilengedwe yamphamvuyi cholinga choyenera komanso kugogomezera, tingathandize ambiri omwe ali mu chizolowezi choledzera ndi kutaya mtima kupeza mtendere ndi chiyembekezo.


[1] Constance Holden, “Zizolowezi Zokhudza Khalidwe: Kodi Zilipo? Sayansi , 294 (5544) 2 Novembala 2001, 980.

[2] Chidule.

[3] Eric J. Nestler, “Kodi pali njira yodziwika bwino ya molekyulu yoledzera?” Nature Neuroscience 9(11):1445-9, Nov 2005

[4] Teresa R. Franklin, Paul D. Acton, Joseph A Maldjian, Jason D. Gray, Jason R. Croft, Charles A. Dackis, Charles P. O'Brien, ndi Anna Rose Childress, “Kuchepa kwa Kuganizira Kwambiri kwa Imvi mu Insular, Orbitofrontal, Cingulate, ndi Temporal Cortices ya Odwala a Cocaine,” Biological Psychiatry (51)2, Januwale 15, 2002, 134-142.

[5] Paul M. Thompson, Kikralee M. Hayashi, Sara L. Simon, Jennifer A. Geaga, Michael S. Hong, Yihong Sui, Jessica Y. Lee, Arthur W. Toga, Walter Ling, ndi Edythe D. London, “Zolakwika Za Kapangidwe ka Ubongo wa Anthu Omwe Amagwiritsa Ntchito Methamphetamine,” The Journal of Neuroscience , 24(26) June 30 2004;6028-6036.

[6] Nicola Pannacciulli, Angelo Del Parigi, Kewei Chen, Dec Son NT Le, Eric M. Reiman ndi Pietro A. Tataranni, "Kusokonezeka kwa ubongo mu kunenepa kwambiri kwaumunthu: kafukufuku wa voxel-based morphometry." Neuroimage 31 (4) July 15 2006, 1419-1425.

[7] Boris Schiffer, Thomas Peschel, Thomas Paul, Elke Gizewshi, Michael Forshing, Norbert Leygraf, Manfred Schedlowske, ndi Tillmann HC Krueger, “Kusakhazikika kwa Ubongo mu Frontostriatal System ndi Cerebellum mu Pedophilia,” Journal of Psychiatric Research (41)9, Novembala 2007, 754-762.

[8] M. Bourke, A. Hernandez, 'Butner Study' Redux: Lipoti la Kuchitika kwa Kuzunzidwa kwa Ana ndi Ochita Zolaula za Ana. Journal of Family Violence 24(3) 2009, 183-191.

[9] H. Mouras, S. Stole4ru, V. Moulier, M Pelegrini-Issac, R. Rouxel, B Grandjean, D. Glutron, J Bittoun, Kuyambitsa kwa dongosolo la mirror-neuron pogwiritsa ntchito makanema olaula kumaneneratu kuchuluka kwa kukhwima komwe kwachitika: kafukufuku wa fMRI. NeuroImage 42 (2008) 1142-1150.

[10] Michael H. Miner, Nancy Raymond, Bryon A. Meuller, Martin Lloyd, Kelvin Ol Lim, “Kufufuza koyambirira kwa makhalidwe opupuluma komanso a neuroanatomical a khalidwe lokakamiza kugonana.” Psychiatry Research Neuroimaging Volume 174, Nkhani 2, Novembala 30 2009, Masamba 146-151.

[11] Bruce E. Wexler, Christopher H. Gottschalk, Robert K. Fulbright, Isak Prohovnik, Cheryl M. Lacadie, Bruce J. Rounsaville, ndi John C. Gore, “Kujambula kwa Magnetic Resonance Imaging of Cocaine Craving,” American Journal of Psychiatry , 158, 2001, 86-95.

[12] Jan Reuter, Thomas Raedler, Michael Rose, Iver Hand, Jan Glascher, ndi Christian Buchel, “Kutchova juga kwa matenda kumalumikizidwa ndi kuchepa kwa kuyambitsa kwa dongosolo la mphotho ya mesolimbic,” Nature Neuroscience 8, Januwale 2005, 147-148.

[13] Gene-Jack Wang, Nora D. Volkow, Jean Logan, Naomi R. Pappas, Christopher T. Wong, Wei Zhu, Noelwah Netusil, Joanna S Fowler, “Dopamine ya ubongo ndi kunenepa kwambiri,” Lancet 357(9253) February 3 2001, 354-357.

[14] J. Michael Bostwick ndi Jeffrey A. Bucci, “Kugwiritsa Ntchito Naltrexone Pa Intaneti Pochiza Chizolowezi Chogonana.” Mayo Clinic Proceedings , 2008, 83(2):226-230.

[15] Marc N. Potenza, “The neurobiology of pathologic juga and drug addiction: a view and new revenues,” Philosophical Transactions of the Royal Society , 363, 2008, 3181-3190..

[16] Nora D. Volkow, Gene-Jack Wang, Joanna S. Fowler, Frank Telang, “Ma neural circuits ophatikizana mu chizolowezi ndi kunenepa kwambiri: umboni wa matenda a machitidwe,” Philosophical Transactions of the Royal Society , 363, 2008, 3191-3200.

[16] Eric J. Nestler, “Njira zolembera za kuledzera: udindo wa DFosB,” Philosophical Transactions of the Royal Society , 363, 2008, 3245-3256.

[18] DL Wallace, ndi ena, Mphamvu ya DFosB mu Nucleus Accumberns pa Machitidwe Okhudzana ndi Mphoto Zachilengedwe, The Journal of Neuroscience , 28(4): Okutobala 8, 2008, 10272-10277,

[19] Sayansi 6 Julayi 2007:?Vol. 317. nambala 5834, tsamba 23

[20] Julie A. Kauer, Robert C. Malenka, “Synaptic Plasticity and Addiction,” Nature Reviews Neuroscience, 8, 8440858 Novembala 2007, 844-858.

[21] Gert Holstege, Janniko R. Georgiadis, Anne MJ Paans, Linda C. Meiners, Ferdinand HCE van der Graaf, ndi AAT Simone Reinders, "Kuyambitsa ubongo pa nthawi ya umuna wamwamuna," Journal of Neuroscience 23 (27), 2003, 9185-9193