Mekhoa ea boipheliso ea mekhoa ea bohloa boko bo botong bo phelang (2014)

Chia-Chien Chen,1 Ju Lu,2 'me Yi Zuo1,*
  • 1Lefapha la Limolek'hule, Cell le Biology ea Ntshetsopele, Univesithi ea California ho Santa Cruz, Santa Cruz, CA, USA
  • 2Lefapha la Saense ea Tlhaho le Setsi sa James H. Clark, Univesithi ea Stanford, Stanford, CA, USA

Tlhahlobo e Nyane ARTICLE

Ka pele. Neuroanat., 09 May 2014 | Doi: 10.3389 / fnana.2014.00028

inahaneloang

Li-spendritic spine ke libaka tse ngata tse thabisang ka ho fetisisa bokong ba 'mele, kahoo li ka sebetsa e le matšoao a sebopeho se sebetsang. Mesebetsi ea morao-rao e bonts'itse hore neuronal coding of memory e ka amana le liphetoho tse potlakileng ho etsoang lesapo la mokokotlo le ho felisoa. Tsoelo-pele ea theknoloji e thusitse bafuputsi ho ithuta matla a mokokotlo Ka vivo nakong ea nts'etsopele le tlasa maemo a fapaneng a 'mele le a bophelo. Re lumela hore kutlwisiso e ntlafalitsoeng ea methapo ea sepakapaka e tla thusa ho hlakisa melao-motheo ea phetisetso ea potoloho e amanang le boiphihlelo le ts'ebetso ea tlhahisoleseling bokong bo phelang.

Keywords: dendritic spine, Ka vivo, ho nahana ka lifoto tse peli, polasetiki e nang le boiphihlelo, potoloho ea neural, cortex ea mokokotlo

SELELEKELA

Li-spine tsa dendritic li khahlile meloko ea litsebi tsa methapo ea kutlo ho tloha ha litlhaloso tsa tsona tsa pele ke Santiago Ramón y Cajal lilemong tse fetang lekholo tse fetileng (Ramon y Cajal, 1888). Li-protrusions tsena tse bonojoana li tsoa mokokotlong oa dendritic mme li tšoana le "meutloa e hlabang kapa likhahla tse khutšoane" joalo ka ha e hlalositsoe ka ho hlaka ke Cajal. Ke libaka tsa postynaptic tsa bongata bo boholo (> 90%) ba li-synapses tse nyakallisang tsa glutamatergic bokong ba mamalia, 'me li na le likarolo tsa bohlokoa tsa limolek'hule bakeng sa lets'oao la postynaptic le polasetiki. Ka hona, mokokotlo le matla a tsona a sebopeho li ka sebetsa e le litsupa tsa khokahano ea synaptic le liphetoho tsa eona (Segal, 2005; Tada le Sheng, 2006; Likotsi le Dunaevsky, 2007).

Boholo ba lithuto tsa pele ho dendritic spine li ile tsa hlahloba lithane tsa "neural" tse bobebe tse nang le khanya kapa elektrone ea elektrone (Lund et al., 1977; Woolley et al., 1990; Harris le Kater, 1994; Hering le Sheng, 2001; Lippman le Dunaevsky, 2005). Leha ba ne ba fana ka leseli la bohlokoa ka moroko morphology le kabo, litlhahlobo tsena tse tsitsitseng tsa liteko tsa "li-" snapshots "tse ikhethang tsa lesapo la mokokotlo. Kenyelletso ea mekhoa ea ho ngola mabitso a fluorescent le li-microscopy tse ngata-ngata li ile tsa fetola tšimo. Ho 2002, mosebetsi oa bopula-maliboho o tsoang li-laboratoring tse peli (Grutzendler et al., 2002; Trachtenberg et al., 2002) e bonts'itse monyetla oa ho latela mokokotlo o tšoanang bokong bo phelang ka nako e telele (ke ho re, libeke) tsa nako. Ha e le hantle, matla a lesapo la mokokotlo a emela matla a phetoho e fapaneng. Le ha methapo e tsitsitseng e emela mabitso a synaptic, ke karoloana e nyane feela ea mesifa ea nakoana e emelang mabitso a mahlale a nakoana, 'me tse ling kaofela li emela li-synaptogenesis tse sa sebetseng (Trachtenberg et al., 2002; Knott et al., 2006; Cane et al., 2014). Ho tsoa lipatlisisong tse joalo tsa nako e telele, setšoantšo se matla sa mesifa se hlahile: sebopeho sa spines, se holisa, se nyenyefatsa le ho khutla nakong eohle ea bophelo ba phoofolo. Ntle le moo, morphology le matla a tsona a fapana ka mefuta ea li-neuronal, ka methati ea tsoelo-pele, le karabelo liphihlelong tse kang ho susumetsa maikutlo le ho hloka, ho ruisa tikoloho le likarolo tse fapaneng tsa ho ithuta (Holtmaat le Svoboda, 2009; Fu le Zuo, 2011).

Tlhahlobo ena e shebisisa sephetho sa Ka vivo lithuto tsa monahano. Ho tšoantšisa matla a lesapo la mokokotlo, bafuputsi ba shebile haholo likarolo tse peli: liphetoho tse akaretsang letsoalo la lesapo la mokokotlo, le sebaka se ikhethileng haufi le dendrite moo ho thehoang lesapo la mokokotlo le ho felisoa. Le ha letsoalo la lesapo la mokokotlo le fana ka khakanyo e lekantsoeng ea palo e felletseng ea liphihlelo tsa thabo ea "postinaptic neuron", sebaka sa lesapo la mokokotlo se susumetsa tlatsetso ea lipontšo tsa sona tsa motlakase le tsa lik'hemik'hale tse arolelanoang karabong e kopaneng ea soma (Nevian et al., 2007; Spruston, 2008). Ho utloisisa hore na matla a lesapo la mokokotlo a ikamahanya joang le likarolo tsa 'mele le tsa' mele tsa potoloho e itseng ea neural maemong a fapaneng a boitsoaro ho bohlokoa ho tlatsong ea ts'ebetso ea tlhahisoleseling le mekhoa ea ho boloka bokong.

DIPUO TSA BOTLUO NTS'A KHAFA

Boholo ba lesapo la mokokotlo le fapana haholo ho palo ea li-neuron tse fapaneng, mohlomong li bontša phapang ea methapo ea kutlo le ts'ebetso (Nimchinsky et al., 2002; Ballesteros-Yanez et al., 2006). Tekano lipakeng tsa ho thetsoa lesapo la mokokotlo le ho emisoa ho tiisa phetoho ea letsoalo la lesapo la mokokotlo: karolo e eketsehileng ea lesapo la mokokotlo e felisoa ka karolo ea methapo e eketsa letsoalo la mokokotlo ka lona, ​​mme ka lehlakoreng le leng. Ka "cortex" ea "cerebral", le ha makala a dendritic a tsitsitse ha nako e ntse e feta.Trachtenberg et al., 2002; Mizrahi le Katz, 2003; Chow et al., 2009; Mostany le Portera-Cailliau, 2011; Schubert et al., 2013), linoko li thehoa khafetsa le ho felisoa. Sekhahla sa ho thehoa ha lesapo la mokokotlo le ho felisoa hoa fetoha ha nako e ntse e ea, ho fella ka phetoho e seng ea monotonate ho letsoalo la lesapo la mokokotlo (Figure Figure11). Mohlala, spines ho dendrites ea apical dendrites ea layer 2 / 3 pyramidal neurons in rodent barrel cortex shows butle-butle ea fokotseha motility (elongation le khutsufatso ea spines) le sekhahla sa phallo (e hlalositsoeng e le kakaretso ea phaello le tahlehelo ea methapo) pakeng tsa letsatsi la 7 la kamora ho pepa. le 24 (P7-24; Lendvai et al., 2000; Cruz-Martin et al., 2010). Leha ho le joalo, letsoalo la lesapo la mokokotlo le ntse le eketseha nako ena kaofela (Cruz-Martin et al., 2010). Kamora karolo ena ea pele ea phaello ea lesapo la mokokotlo, ho felisoa ha lesapo la mokokotlo ho qala ho hlahisa sebopeho, ho lebisang phokotsong e akaretsang ea letsoalo la mokokotlo (Holtmaat et al., 2005; Zuo et al., 2005b; Yang et al., 2009). P pakeng tsa P28 le P42, 17% ea methapo e felisoa hammoho le li-dendrites tsa apole tsa 5 pyramidal neurons ka har'a "barrel cortex", ha 5% ea methapo e mecha e thehiloe ka nako e ts'oanang.Zuo et al., 2005a, b). Habohlokoa, ha se mefuta eohle ea methapo e ka bang bonolo ho e felisa: e nang le lihlooho tse kholo e tsitsitseng ho feta e mosesaane. Ha boholo ba lesapo la lesapo la mokokotlo le hokahana ka matla a synaptic, ketsahalo ena e fana ka maikutlo a hore li-synapses tse matla li tiile haholoanyane (Holtmaat et al., 2005). Ho feta moo, linoko tse sa tsoa thehoa li na le menyetla ea ho felisoa ho feta mehala e neng e le teng pele (Xu et al., 2009), le boholo ba methapo e tsitsitseng e thehiloeng pele bocha e lula e kenyellelitsoe potolohong ea methapo ea batho ba baholo (Zuo et al., 2005a; Yang et al., 2009; Yu et al., 2013). Qetellong, ho liphoofolo tse kholo tsa ho theha lesapo la mokokotlo le ho felisoa ho fihlela ho lekana; letsoalo la lesapo la mokokotlo le lula le batla le fetoha ho fihlela ho qala ho tsofala (Zuo et al., 2005a; Mostany et al., 2013).

Tšoantšiso 1 

Spine e nchafatsoa maemong a fapaneng a bophelo ba phoofolo. Rapid spinogenesis matsatsing a pele a peleho e lateloa ke ho pongoa ha lesapo la mokokotlo butle butle lilemong tsa bocha. Ha e le motho e moholo, ho thehoa lesapo la mokokotlo le ho felisoa ho fihlela ho lekana, ka karoloana e nyane ea methapo ...

LIJENO TSE BOTLENG KAPA HO BUA LE TLHALOSO EA SESOTHO

Cortex ea "cerebral" e na le bokhoni bo hlollang ba ho hlophisa potoloho ea eona ho latela liphihlelo. Ka hona, tsela eo liphihlelo tsa maikutlo (kapa ho haella ka eona) li amang matla a mokokotlo li khahlang haholo ho litsebi tsa kelello. Ho bontšoa ka matla ho potoloha le ho sa foleng ha maikutlo ho ama matla a mokokotlo, empa phello e tobileng e ipapisitse le phapang ea nako le nako, hammoho le sethala sa phoofolo. Nakong ea bokhachane pele ho nako, litlatsetso tsa maikutlo li bapala karolo ea ho ruta ho tsitsisa le ho holofatsa methapo ea mmele. Ka "visual" cortex, ho amoha pono ea tlhaho ho tloha tsoalong ho thibele ho fokotseha ha letsoalo la mokokotlo le ho emisoa ha lesapo la mokokotlo ((spine morphology)Majewska le Sur, 2003; Tropea et al., 2010). Ho tlosoa hoa liphatsa tsa lefutso tsa PirB ho phethile phello ea ho nyahlatsoa ka kotloloho ho ts'ebetso ea lesapo la mokokotlo (Djurisic et al., 2013). Ho litoeba tse neng li sentsoe ke bofokoli ba pono ba nakong e fetileng, leseli la lesapo la mokokotlo le neng le entsoe ka mokhoa o fokolang le ka ts'oaroa ke ts'ebetso ea pharmacological ea tsamaiso ea GABAergic, e fana ka maikutlo a karolo ea bohlokoa ea lipotoloho tse thibelang phetoho ea maqhubu a thabo (Tropea et al., 2010). Hamorao, boiphihlelo ba maikutlo bo khanna ho pongoa ha lesapo la mokokotlo (le hlalosoang ka tahlehelo ea methapo ea methapo). Ho ts'oara ha makhooa ohle a litoeba ka likhoeli tsa 1 tse nang le matsatsi a 4 kapa matsatsi a 14 ho fokotsitse ho felisoa ha lesapo ka mokokotlong, empa ho ile ha siea sebopeho sa lesapo la mokokotlo le sa senyehe haholo (Zuo et al., 2005b; Yu et al., 2013). Pharmacological blockade ea li-receptor tsa NMDA e etselitse phello ea ho ts'oara ha whisker, ho bonts'a ho kenya letsoho ha tsela ea li-receptor tsa NMDA ts'ebetsong e joalo e itšetlehileng ka lesapo la mokokotlo (Zuo et al., 2005b).

Le ha ho ts'oaroa hoa whisker ho tlosa ho kenella hoa maikutlo lefats'eng ka bophara, ho ts'oara whisker e 'ngoe le e' ngoe ("chessboard trimming") ka mokhoa o hlakileng ho phahamisa phapang efe kapa efe ea maemo le ts'ebetso ea libare tsa boahelani, ka ho etsa joalo ho hlahisa boiphihlelo ba maikutlo a haufinyane. Paradigm e joalo e bontšitsoe ho khothaletsa phetoho ea lesapo la mokokotlo le ho tsitsisa methapo e sa tsoa thehoa ka mokhoa o ikhethileng ka subclass ea li-neuron tsa cortical (Trachtenberg et al., 2002; Holtmaat et al., 2006). Li-spines tse ncha li ile tsa eketsoa ka mokhoa o ikhethileng mokokotlong oa li-neuron tsa 5 tsa piramide tse nang le li-aplic tse thata tsa apical, ho fapana le tse nang le marang-rang a bonolo (Holtmaat et al., 2006). Ho litoeba tse nang le phoso tsa íCaMKII-T286A, chessboard e ile ea sitoa ho boloka botsitso ba methapo e mecha e phehellang moeling pakeng tsa libare tse sa sirelelitsoeng le tse senyehileng (Wilbrecht et al., 2010). Haufinyane tjena, boithuto bo botle bo kopaneng le ts'usumetso ea optogenetic le Ka vivo monahano o bonts'itse hore ke mohlala oa tšebetso ea neural, ho fapana le boholo, e khethollang botsitso ba methapo ea methapo (Wyatt et al., 2012).

Ho ts'oana le ho chechela ha chessboard, ho nyatsuoa ka bokhutšoaane ba bosoasoi (MD) ho eketsa karohano lipakeng tsa likhakanyo tse tsoang mahlong a mabeli. Ka mokhoa o ts'oanang le oa ho chessboard, MD e fumanoe e eketsa ho etsoa ha lesapo la mokokotlo hammoho le li-apical dendritic li-sits tsa "5 pyramidal neurons" sebakeng sa bonono sa "cortex" ea "bonase". Leha ho le joalo phello ena ha ea ka ea bonoa ka li-neuron tsa 2 / 3, kapa tikolohong ea monocular (Hofer et al., 2009), hape e bonts'a mofuta oa sele e ikhethileng e feto-fetohang. Ho khahlisang, MD ea bobeli e ile ea hloleha ho eketsa sebopeho sa lesapo, empa ka mokhoa o khethollang li-spine tse entsoeng nakong ea MD ea pele, e fana ka maikutlo a hore li-spines tse ncha tse entsoeng nakong ea MD ea pele li ne li e-na le li-synapses tse sebetsang hape nakong ea MD ea bobeli (Hofer et al., 2009).

DIPUO TSA BOTLULE HA BA ITHUTA

Sebopeho se matla sa methapo ea dendritic e hlahisa mohopolo o atileng oa hore methapo e ka sebetsa e le sebopeho sa sebopeho sa ho ithuta le ho hopola. Ho 'nile ha fanoa ka maikutlo a hore li-spine tse sa tsoa hlaha (hangata li na le lihlooho tse nyane) tse fumanehang mohopolong, ha libaka tse tsitsitseng (hangata tse nang le lihlooho tse kholo) li sebetsa e le libaka tsa polokelo ea memori (Bourne le Harris, 2007). Ka 'nete, Ka vivo lithuto tsa monahano li bontšitse hore ho "cortex ea" cerebral cortex, matla a lesapo la mokokotlo a amana ka kotloloho le ho ithuta. Ka "motor motor" cortex, ho etsoa ha lesapo la mokokotlo ho qala hang ha phoofolo e ithuta mosebetsi o mocha. Ka mor'a ts'ebetso ena e potlakileng ea letsoai, letsoalo la lesapo la mokokotlo le khutlela boemong ba motheo ka ho felisoa ha lesapo la mokokotlo (Xu et al., 2009; Yu le Zuo, 2011). Ho linonyana tsa lipina, sekhahla se phahameng sa ho fetoloa ha lesapo la mokokotlo pele thuto ea pina e fumanoa e ka ikamahanya le boemo bo eketsehileng ba ho etsisa lipina hamorao (Roberts le al., 2010). Ho litoeba, palo ea methapo e fumanoeng nakong ea ho ithuta ea pele e lumellana hantle le ts'ebetso ea makoloi ea ho fumana thepa (Xu et al., 2009); le ho pholoha ha likholumolumo tse ncha tse amanang le boits'oaro ba motor (3)Yang et al., 2009). Ntle le moo, litsebo tse fapaneng tsa makoloi li kanna tsa akaretsoa ka methapo e fapaneng ea li-motor cortex, ha ho ithuta mofuta oa motor motor ho litoeba tse koetlisitsoeng pele ho tsoela pele ho kenya letsoho phetoho e matla ho motor motor cortex (Xu et al., 2009). Haufinyane, ho boetse ho fumanoe hore boemo ba glucocorticoid bo ama matla a thuto ea mokokotlo oa motor. Ho koetlisoa ke litoeba ho litlhoro tsa glucocorticoid ho felletse ka tekanyo e phahameng ea sebopeho sa lesapo, athe mekotla ea glucocorticoid e latelang thupelo e ne e hlokahala bakeng sa ho tiisa methapo ea methapo e entsoeng nakong ea thupelo le ho hopola nako e telele (Liston et al., 2013). Tlatsetso, e 'nileng ea nkuoa e le thuto ea maqhubu (Hyman, 2005), e etsa hore ho be le liphetoho tse tšoanang tsa nakoana ho matla a lesapo la mokokotlo joalo ka ha ho ithutoa ka makoloi. Sebelisa paradigm e nang le maemo a koae ea koae, phuputso ea morao-rao e bonts'itseng ea hore ho pepesoa ha cocaine ea pele ho ile ha phahamisa sebopeho sa lesapo la mokokotlo ka pele le hore palo ea methapo e mecha e kopaneng e amanang le khetho ea taba ea koae (e koahelang koae).Munoz-Cuevas et al., 2013). Ho khahlisang le ho feta, matla a lesapo la mokokotlo libakeng tse fapaneng tsa cortical a ka fapana nakong ea mosebetsi o tšoanang. Mohlala, mokhoa o tšosang oa paradigm eo li-contex tsa maoto le matsoho a nang le eona e nang le litšisinyeho tsa maoto li bonts'itse litlamorao tse fapaneng ho cortex ea "auditory". Ka "cortex" ea kutlo, ho ile ha fumaneha hore sebopeho se matla sa lesapo la mokokotlo se ne se kopantsoe le maemo a ts'abo a marang-rang, ha maemo a sa lefelloeng a ne a amahanngoa le ho felisoa ha methapo e mengata (Moczulska et al., 2013). Karolong e ka pele ea mokokotlo, ho felisoa ha lesapo la mokokotlo ho fumanoe ho amahanngoa le ho ithuta, ha ho thehoa lesapo la mokokotlo ho ne ho amahanngoa le tšabo, 'me ho khutlisetsa methapo ho felile ha ho entsoe nakong ea ho timela (Lai et al., 2012). Ha li kopantsoe hammoho, liphuputso tsena li senola phapang ea melao ea nakoana e ipapisitseng le matla a susumetsang mokokotlo o ithutileng oona. Hore na methapo ea methapo e thehiloe kapa e tlosoa nakong ea ho ithuta ho ipapisitse le paradigm ea boits'oaro le mefuta e ikhethileng ea li-neuronal le mefuta ea lisele tse nkang karolo molemong oa ho ithuta.

Ho bohlokoa ho hlokomela hore mehlala eohle e tšohliloeng ka holimo e bua ka memori e sa senyeheng, e sa kenyeng khopotso e hlakileng ea nako, sebaka, le boiphihlelo ba liketsahalo (ke hore, memori e hlolohileng). Tlhatlhobo ea Ka vivo matla a lesapo la mokokotlo a amanang le ho hopola lintho tse etsahalang ka nakoana a ipaka ho le thata haholo. Ka lehlakoreng le leng, hippocampus, sebopeho sa bohlokoa bakeng sa mohopolo o fokolisang matla, se patoa ka tlas'a cortex le ka nq'ane ho ts'ebetso ea microscopy e tloaelehileng. Ka lehlakoreng le leng, ho nahanoa hore memori e fokotsehang e bolokiloe ka mokhoa o hlakileng marang-rang a maholo a neocortical, e etsa hore ho be thata ho menahano e lebisitsoeng. Ka hona, nts'etsopele ea mahlale a tebileng a ho nahana ka boko (mohlala, microendoscopy, adapter optics) hammoho le kutlwisiso e ntle ea kabo ea memori ka cortex e tšoere senotlolo sa lipatlisiso tsa nako e tlang tsa matla a lesapo la mokokotlo tse tlasa mohopolo oa morao oa kelello.

LIHLOOHO TSE PHELANG LI-DINAMICS TLHABENG

Liphetoho ho dendritic spine densities li hlokometsoe ho mafu a fapaneng a methapo ea methapo le neuropsychiatric. Boloetse bo bong le bo bong bo hlahisa le litlolo tsa eona tse ikhethang ho matla a lesapo la mokokotlo, bo ntlafatsang le ho feta taba ea hore methapo ea methapo ea kutlo e sebetsa ka mokhoa o hlakileng bakeng sa ts'ebetso e nepahetseng ea kelello. Ho na le tumellano e ntseng e hola ea hore letsoalo la lesapo la mokokotlo le amana le khaello ea boits'oaro le ho fokotseha hoa ts'ebetso ea kelello (bakeng sa lintlha tse ling bona Fiala et al., 2002; Penzes et al., 2011).

Ka mefuta ea stroke, ho bonts'oa hore ischemia e matla e lebisa ho tahlehelo ea lesapo la mokokotlo kapele, e fetoloang kamora ho pheta-pheta haeba pholoso e etsoa nakong e khutšoane ea X (20-60 min; Zhang et al., 2005). Kamora ho otloa, ho thehwa ha lesapo la mokokotlo le ho felisoa hoa morao ho eketsehang sebakeng sa peri-infarct, empa eseng libakeng tse hole le infarct kapa contralateral hemisphere (Brown et al., 2009; Johnston et al., 2013). Plastiki ena e bakiloeng ke likotsi e fihla sehlohlolong sa eona ka mor'a beke ea 1; ho tloha ka nako eo ho ea holimo sekhahla sa ho etsoa ha lesapo la mokokotlo le ho felisoa hoa theoha ka sekhahla. Ketsahalo ena e bonts'a ho ba teng ha nako e mahlonoko eo ho eona likaroloana tsa cortical tse sa pheleng li leng bohlokoa haholo ho kenella lipakeng tsa kalafo (Brown et al., 2007, 2009). Ka mohlala oa mouse bakeng sa bohloko bo sa foleng, ho ruruha ho sa lekanyetsoang ha methapo ea kutlo, ho phahamisa mokokotlo le ho felisoa. Hoa tšoana le mofuta oa stroke, ho phahama ha sekhahla sa ho khothaletsoa ha lesapo ho tlisoa ke ho felisoa, ho lebisang keketseho ea pele ea letsoalo la mokokotlo le lateloang ke phokotso ea lona. Litlamorao tse joalo li ka felisoa ka tetrodotoxin blockade, e bonts'a hore ho nchafatsa lesapo la lesapo la mokokotlo ka mor'a lesapo ho its'etleha mesebetsing (Kim le Nabekura, 2011).

Mefuta e fetotsoeng ea lesapo la mokokotlo e boetse e tlalehiloe mefuteng ea liphoofolo ea maloetse a mpefatsang. Mohlala, tahlehelo ea lesapo la mokokotlo e potlakisoa sebakeng sa haufi le li-plaque tsa β-amyloid sebakeng sa "cortex"Tsai et al., 2004; Lihlapi et al., 2005). Ka mohlala oa phoofolo ea mofuta oa Huntington's lefu la lesapo la mokokotlo le eketseha, empa methapo e sa tsoa thehoa ha e phehelle ho kenyelletsoa potolohong ea lehae, e lebisang ho fokotseha ha palo ea lesapo la mokokotlo (Murmu et al., 2013). Le ha maloetse a neurodegenerative a tloaetse ho amahanngoa le tahlehelo ea lesapo la mokokotlo, mathata a methapo ea methapo a bontša likaroloana tse fapaneng tsa mokokotlo. Ka setšoantšo sa "mouse" sa Fragile X, li-spine li ngata haholo, 'me liperesente tse ngata tsa tsona li hlaha li sa shebahale hantle ha ho hlahlojoa linama tse kholo tsa batho ba baholo (Comery et al., 1997; Irwin et al., 2000). Ka vivo Boithuto bo boetse bo bonts'itse hore liphoofolong tse joalo ho hlaha ha lesapo la mokokotlo le eketsehile libakeng tse fapaneng tsa cortical (Cruz-Martin et al., 2010; Pan et al., 2010; Padmashri et al., 2013), hape ha ho leoa le phoka kapa ho ithuta ka enjine ho ka fetolang matla a mokokotlo (Pan et al., 2010; Padmashri et al., 2013). Ho litoeba tse tebileng tsa MiceP2, e amanang le mofuta oa amanang le Rett Syndrome, ho fumanoe hore melemo le tahlehelo ea mokokotlo li phahame. Leha ho le joalo, methapo e mecha e kotsing ea ho e qobisa ho feta litoeba tsa mofuta o hlaha, e leng se bakang tahlehelo ea methapo (Jiang et al., 2013).

TLHOHONOLOFETSO EA BOTLALO HO LIJO TSA BOTEI

Tsamaiso ea methapo e na le lihlopha tse peli tsa lisele: li-neuron le glia. Karolo e khahlang ka ho fetisisa ea lisele tsa glial ke ho nka karolo ha tsona ho li-synaptic tšebetsong le ho matla. Haufinyane tjena, liphuputso tse 'maloa tse khahlisang li fuputse karolo ea ho supa ha glial ho tlatsana le lesapo la mokokotlo le polasetiki. Mohlala, thibelo ea ho ts'oaroa hoa bonohe ba linaleli ho bontšitsoe ho potlakisa ho felisoa ha lesapo la mokokotlo nakong ea kholo ea kholo (Yu et al., 2013). Mofuta o mong oa lisele tsa glial, microglia, le tsona o fumanoe o le haufi-ufi le li-spendritic spines. Morero oa lits'ebetso tsa microglial le ho ikopanya le lesapo la mokokotlo li laoloa ka mafolofolo ke boiphihlelo ba maikutlo mme o ameha ho feliseng lesapo la mokokotlo (Tremblay et al., 2010). Ntle le moo, ho theoha ha kokoana-hloko ea micoglia ho felletse ka phokotseho e kholo ea thuto ea mokokotlo e kenyellelitseng sebopeho sa lesapo la mokokotlo, le ho tlosoa ka mokhoa o ikhethileng oa "neurotrophic factor" (BDNF) ho microglia ho khutlisa litlamorao tsa ho fokotsoa hoa maqhubu (microglial depletion)Parkhurst et al., 2013).

TLHOKOMELISO E NANG KOTSI EA LIJO TSA BOTEI

Ho nahanisisa ka meetso ea methapo ho khothalelitse hore ho hlaha le ho nyamela ha mesifa ha ho tšoane kapa hona ho ba teng ka mokhoa o tšoanang, empa ho e-ba teng libakeng tse ikhethileng tsa "libaka tse chesang" ka "motor motor" cortex. tloaelo ea ho qhoqha. Ntle le moo, ho eketsoa ha lesapo la bobeli le lecha sehlopheng hangata ho amahanngoa le ho eketsoa ha lesapo la mokokotlo la pele. Ka lehlakoreng le leng, methapo e thehiloeng nakong ea ts'ebetso ea ho etsa mesebetsi e fapaneng ea makoloi kapa nakong ea ho ruisa makoloi ha e kopane (Fu et al., 2012). Ha ho kopantsoe, lintlha tsena li bonts'a hore ts'ebetso ea mokokotlo o mocha o pheta-phetang e hlokahala bakeng sa ho hlaha hoa lesapo la mokokotlo la bobeli. Khethollo e tšoanang ea sepakapaka ea matla a lesapo la mokokotlo e hlokometsoe ho paradigm ea boemo ba tšabo: lesapo la mokokotlo le tlosoang nakong ea boemo ba ts'abo hangata le nkeloa sebaka ke lesapo sebakeng sa lona (kahare ho 2 μm) nakong ea tšabo e felileng (Lai et al., 2012). Ho khahlisang, matla a lesapo la mokokotlo a boetse a susumetsoa ke matla a thibelo ea methapo ea kutlo. Ho felloa ke matla ka kotloloho ho eketsa haholo matla a kopaneng a spine le inhibitory synapses tse haufi le mokato oa 2 / 3 pyramidal neurons (Chen le al., 2012). Liphumano tsena li tšehetsa mofuta o kopaneng oa polasetiki, eo li-synapses tse kopaneng li ka bang le karolo ea ho kenya letsoho tlhahisoleseling e le ngoe ho feta li-synapses tse hasantsoeng ho potoloha dendritic arbor (Govindarajan et al., 2006).

Ho kopanya Ka vivo ho rekota ka seleng ka bongata le ho nahana ka lesapo la mokokotlo o le mong, mosebetsi oa morao-rao o bonts'itse hore lits'oants'o tsa lithapo tse fapaneng tsa maqhubu a fapaneng li kenelletse kahare ho li-dendrites tsa li-neuron tsa "pyramidal"Chen le al., 2011). Ho fumana hona ho hlahisa potso e khahlisang: na li-spine tse kopantsoeng tse ncha li lumellana le se kenyelletsoang ka litšobotsi tse ts'oanang kapa tse fapaneng (mohlala, mekhoa ea tšebetso, thepa ea tokiso)? Bakeng sa ho araba potso ena, ho tla hlokahala hore re fapanyetse mehala sebakeng se pharaletseng sa mokokotlo oa dendritic, ho khetholla "hotspots" ea remine ea lesapo la mokokotlo, le ho kopanya mohopolo oa methapo ea methapo e nang le monahano o sebetsang oa nako. Liteko tse joalo li ke ke tsa thusa feela ho hlakisa mekhoa ea methapo ea kutlo e amanang le lesapo la mokokotlo, empa e tla fana ka leseli ho boemeli ba tlhahisoleseling le polokelo ea methapo ea kutlo.

MAHLOMOLA MAHALA

Sehloohong sena, re shebile lipatlisiso tsa morao-rao mabapi le matla a methapo ea methapo e ka hare bokong bo phelang. Le ha lithuto tsena li hatetse pele kutlwisiso ea rona ea hore na matla a lesapo la mokokotlo a fetoha joang ka nakoana le ka moea, lipotso tse ngata li sala mehloling e fapaneng. Mohlala, na ho na le matšoao a limolek'hule a khethollang mesifa e tsitsitseng ea spine le spine tse sa tsoa etsoa hore li felisoe? Na palo eohle ea methapo e bolokiloe ka mokhoa oa homeostatic, e le hore dendrite e ka boloka tlhokahalo ea metabolic ea phetiso ea synaptic? Na ho kopantsoe ha methapo e mecha ho bonts'a liphetoho matleng a khokahano e teng le axon e le 'ngoe (ha u ntse u boloka topology e tšoanang), kapa na ho supa ho theha likhokahanyo tse eketsehileng ka li-axon tsa nakong e fetileng tse sa amaneng haufi? Ho bohlokoa ho hlokomela hore mesebetsi eohle e tšohliloeng ka holimo e lebisitse ka lehlakoreng la postynaptic, e leng halofo feela ea pale. Morero o mong o ka sehloohong oa ho ajoa ha lesapo la mokokotlo le matla o teng lehlakoreng la mopresidente: boitsebiso le geometry ea li-axons tsa Presynaptic le ho fumaneha ha li-axonal boutons. Ho tseba tlhaiso-leseling e joalo ea bokhachane ho bohlokoa ho rarollleng lipotso tse ngata tse hlahisoang ke ho bonoa ha matla a lesapo la mokokotlo. Leha ho le joalo, ho tsebahala ha mohokahanyi oa lesapo la mokokotlo oa setšoantšo sa dendritic e lula e le phephetso, hobane "axonaptic axon" e ka hlaha mohloling o tsoang mehloling e mengata, 'me hangata e hokahana le lits'ebetso tse ling tse ngata tsa axonal. Ntle le moo, ho sa na le ho hongata ho lokelang ho ithutoa mabapi le tatellano ea moralo o hlophisitsoeng o etsahalang sebakeng sa khokahano lipakeng tsa axonal bouton le lesapo la mokokotlo, le hore na tatellano e joalo e amana joang le ho qaptjoa ha li-synapses. Ho nahana ka nako e le 'ngoe ka li-axonal boutons le likarolo tsa bona tsa tšebelisano molemong oa boitšoaro bo fetisisang ho tla fana ka tlhahisoleseling e ngata ho araba potso ena. Litlhahlobo tse khutlisetsang tse hlahang khafetsa tse kang li-microscopy tsa elektrone (Knott et al., 2009) le Array Tomography (Micheva le Smith, 2007; Michele et al., 2010) e kanna ea tlatselletsa Ka vivo nahana ka ho netefatsa boteng ba li-synapses, le ho senola menoana ea menoana ea meetso ea litšoantšo.

Tatellano ea nakoana ea moea le li-rearrangements tse ikhethileng tsa khokahano ea li-neuronal, le hore na liphetoho tsena ka bongata li kenya letsoho joang phetohong ea boitšoaro ka lebaka la liphihlelo, ke e 'ngoe ea lipotso tsa mantlha mabapi le ts'ebetso ea methapo ea kutlo. Tsoelo-pele mekhoeng ea ho nahana, hammoho le nts'etsopele ea electrophysiology, molek'hule ea liphatsa tsa lefutso le li-optogenetics, e tla thusa ho senola pherekano ea potoloho ea methapo maemong a microscopic, hammoho le mekhoa ea ho kenyelletsa litaba, ho kopanya le ho boloka bokong.

TLHOKOMELISO MOTSAMAI

Chia-Chien Chen ke eena ea entseng setšoantšo. Chia-Chien Chen, Ju Lu le Yi Zuo ba ngotse buka e ngotsoeng ka letsoho.

Tlhōlisano ea Tlhaloso

Bangoli ba bolela hore lipatlisiso li ne li etsoa ka ho se be le likamano leha e le life tsa khoebo kapa tsa lichelete tse ka nkoang e le khohlano e ka 'nang ea e-ba le thahasello.

lumela hore baa fokola

Mosebetsi ona o tšehelitsoe ke chelete ea thuso (R01MH094449) e tsoang Setsing sa Naha sa Bophelo ba Bakuli ho isa Yi Zuo.

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