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=== 情動や意欲、意思決定 ===
<ref name=Krapp1996><pubmed>8861960</pubmed></ref>
　ドーパミンやオピオイドの作用を変化させると、強迫的な選択行動等がひきおこされることから側坐核がhedonic（快楽的）な感情のプロセスに関与していることが示された。Berridgeらは、げっ歯類の側坐核の中で吻側―尾側軸の特定の異なる領域が、「快楽や報酬」領域と「恐怖や嫌悪」領域という異なる情動と対応していることを明らかにした<ref name=Berridge2008><pubmed>18311558</pubmed></ref>。
<ref name=Knoefler1996><pubmed>8703005</pubmed></ref>
 
<ref name=Rose2001><pubmed>11562365</pubmed></ref>
　[[サル]]でも[[ビククリン]]をごく少量注入して異なる腹側線条体領域の活動を局所的に障害すると、行動の抑制と意欲の低下・[[性行動の神経回路|性的行動]]の亢進・異常な運動を繰り返す[[不安様行動]]が障害個所に特徴的に表れた<ref name=Worbe2009><pubmed>19068490</pubmed></ref> 。Shellの内側部は外側視床下部を抑制していて、これを障害すると[[摂食行動]]が引き起こされる<ref name=Kelley2005><pubmed>16289609</pubmed></ref> 。
<ref name=Krapp1998><pubmed>9851981</pubmed></ref>
<ref name=Kawaguchi2002><pubmed>12185368</pubmed></ref>
　線条体には、報酬と関連した感覚刺激や報酬そのものを予測的に期待する発火と、これらの後に反応する発火とが見られる。サルの電気生理学的実験では、背側線条体では課題の比較的前半つまり感覚刺激やその予測に関連した神経発火が見られるのに対し、腹側線条体では、課題の後半つまり報酬の予測や報酬を得た後に発火するものが多く観察されている<ref name=Tremblay2009><pubmed></pubmed></ref><ref name=Nakamura2012><pubmed>23136434</pubmed></ref> 。従って、腹側線条体は報酬を得るというゴールに達するため行動を起こす意欲driving forceの源となっている可能性がある。
<ref name=Sellick2004><pubmed>15543146</pubmed></ref>
 
<ref name=Sellick2003><pubmed>14514650</pubmed></ref>
　[[ヒト]]の非侵襲的イメージングでは腹側線条体が報酬の予測・評価・報酬予測誤差の表現や、動機に基づいた学習に関与していることが示されたが、報酬の時間的予測については結論に差異がある<ref name=Knutson2001><pubmed>11459880</pubmed></ref><ref name=Pagnoni2002><pubmed>11802175</pubmed></ref><ref name=Tanaka2004><pubmed>15235607</pubmed></ref><ref name=ODoherty2004><pubmed>15087550</pubmed></ref><ref name=Kuhnen2005><pubmed>16129404</pubmed></ref> 。
<ref name=Hoveyda1999><pubmed>10507728</pubmed></ref>
 
<ref name=Masui2008><pubmed>18606784</pubmed></ref>
　Nicolaらは腹側被蓋野・内側前頭葉・扁桃体基底外側複合体の、側坐核単一細胞の発火への影響を調べた。[[ラット]]が音を弁別してレバー押しやnose pokeで反応することを学習すると、側坐核ニューロンは音に反応するが、腹側被蓋野・内側前頭葉・扁桃体からの入力をブロックすると側坐核ニューロンの発火が弱まり、弁別反応の正解率も低下する<ref name=Yun2004><pubmed>15044531</pubmed></ref><ref name=Ishikawa2008><pubmed>18463262</pubmed></ref><ref name=Ambroggi2008><pubmed>18760700</pubmed></ref> 。したがって、これらの領域からの情報が側坐核で統合することが刺激―行動に必須であると結論付けられた。
<ref name=Pan2013><pubmed>23325761</pubmed></ref>
 
<ref name=Burlison2008><pubmed>18294628</pubmed></ref>
　一方、側坐核は報酬などの目的達成のためのオペラント条件付けそのものというより、現在進行中の行動から、一定の時間を経て別の行動に変化する過程に重要であるという意見がある<ref name=Cardinal2001><pubmed>11375482</pubmed></ref><ref name=Cardinal2002><pubmed>12034134</pubmed></ref><ref name=Nicola2007><pubmed>16983543</pubmed></ref> 。さらに、サルの腹側線条体細胞の特徴として、例えば視覚刺激→行動という単一試行の課題では課題に反応する細胞の割合は10%前後だが、複数のステップを経て報酬を得るような[[多試行報酬スケジュール課題]]では反応する細胞が60%前後と非常に多い。これらはスケジュールのうち特定の段階で、視覚手がかりへの応答や運動への応答報酬投与に応答する<ref name=Shidara1998><pubmed>9502820</pubmed></ref> 。
<ref name=AlShammari2011><pubmed>21749365</pubmed></ref>
 
<ref name=Adell2000><pubmed>10768861</pubmed></ref>
　側坐核におけるドーパミンの作用については多くの知見がある。報酬を得られたらその行動を学習し、報酬を得られなかったら柔軟性を発揮して別の行動を選択する。この相反する行動決定の切り替えのメカニズムの少なくとも一部に、側坐核におけるphasicまたはtonicなドーパミンの作用のバランスが関与している。Phasicな作用は辺縁系（海馬）からの入力で主にドーパミン[[D1受容体]]を介する調節を受けている。Tonicな作用は前頭葉からの入力で主にドーパミン[[D2受容体]]を介する調節を受けている。腹側淡蒼球 (ventral pallidum, VP)は通常ドーパミン系をtonicに抑制している。海馬から興奮性の入力を受けると、側坐核は抑制性の投射をこのVPに送り、[[脱抑制]]機構によりドーパミン細胞をtonicに興奮させる。このtonicなドーパミン投射はD2受容体を介して前頭葉からの入力を抑制する。一方、phasicな作用は脚橋被蓋核からドーパミン細胞への入力による。これらの入力の側坐核でのバランスによって適切な行動の選択が可能となる<ref name=Goto2008><pubmed>18786735</pubmed></ref> 。
<ref name=Masui2007><pubmed>17938243</pubmed></ref>
 
<ref name=Magnuson2013><pubmed>23823474</pubmed></ref>
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2025年10月5日 (日) 12:12時点における最新版

[1] [2] [3] [4] [5] [6] [7] [8] [9] [10] [11] [12] [13] [14] [15] [16] [17] [18] [19] [20] [21] [22] [23] [24] [25] [26] [27] [28] [29] [30] [31] [32] [33] [34] [35] [36] [37] [38] [39] [40] [41] [42] [43] [44] [45] [46] [47] [48] [49] [50] [51] [52] [53] [54] [55] [56] [57] [58] [59] [60] [61] [62] [63] [64] [65] [66] [67] [68] [69] [70] [71] [72] [73] [74] [75] [76] [77] [78] [79] [80] [81] [82] [83]

  1. ↑ Krapp, A., Knöfler, M., Frutiger, S., Hughes, G.J., Hagenbüchle, O., & Wellauer, P.K. (1996).
    The p48 DNA-binding subunit of transcription factor PTF1 is a new exocrine pancreas-specific basic helix-loop-helix protein. The EMBO journal, 15(16), 4317-29. [PubMed:8861960] [PMC] [WorldCat]
  2. ↑ Knöfler, M., Krapp, A., Hagenbüchle, O., & Wellauer, P.K. (1996).
    Constitutive expression of the gene for the cell-specific p48 DNA-binding subunit of pancreas transcription factor 1 in cultured cells is under control of binding sites for transcription factors Sp1 and alphaCbf. The Journal of biological chemistry, 271(36), 21993-2002. [PubMed:8703005] [WorldCat] [DOI]
  3. ↑ Rose, S.D., Swift, G.H., Peyton, M.J., Hammer, R.E., & MacDonald, R.J. (2001).
    The role of PTF1-P48 in pancreatic acinar gene expression. The Journal of biological chemistry, 276(47), 44018-26. [PubMed:11562365] [WorldCat] [DOI]
  4. ↑ Krapp, A., Knöfler, M., Ledermann, B., Bürki, K., Berney, C., Zoerkler, N., ..., & Wellauer, P.K. (1998).
    The bHLH protein PTF1-p48 is essential for the formation of the exocrine and the correct spatial organization of the endocrine pancreas. Genes & development, 12(23), 3752-63. [PubMed:9851981] [PMC] [WorldCat] [DOI]
  5. ↑ Kawaguchi, Y., Cooper, B., Gannon, M., Ray, M., MacDonald, R.J., & Wright, C.V. (2002).
    The role of the transcriptional regulator Ptf1a in converting intestinal to pancreatic progenitors. Nature genetics, 32(1), 128-34. [PubMed:12185368] [WorldCat] [DOI]
  6. ↑ Sellick, G.S., Barker, K.T., Stolte-Dijkstra, I., Fleischmann, C., Coleman, R.J., Garrett, C., ..., & Houlston, R.S. (2004).
    Mutations in PTF1A cause pancreatic and cerebellar agenesis. Nature genetics, 36(12), 1301-5. [PubMed:15543146] [WorldCat] [DOI]
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    A novel gene for neonatal diabetes maps to chromosome 10p12.1-p13. Diabetes, 52(10), 2636-8. [PubMed:14514650] [WorldCat] [DOI]
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    Neonatal diabetes mellitus and cerebellar hypoplasia/agenesis: report of a new recessive syndrome. Journal of medical genetics, 36(9), 700-4. [PubMed:10507728] [PMC] [WorldCat]
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    Transcriptional autoregulation controls pancreatic Ptf1a expression during development and adulthood. Molecular and cellular biology, 28(17), 5458-68. [PubMed:18606784] [PMC] [WorldCat] [DOI]
  10. ↑ Pan, F.C., Bankaitis, E.D., Boyer, D., Xu, X., Van de Casteele, M., Magnuson, M.A., ..., & Wright, C.V. (2013).
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    Pdx-1 and Ptf1a concurrently determine fate specification of pancreatic multipotent progenitor cells. Developmental biology, 316(1), 74-86. [PubMed:18294628] [PMC] [WorldCat] [DOI]
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    A novel PTF1A mutation in a patient with severe pancreatic and cerebellar involvement. Clinical genetics, 80(2), 196-8. [PubMed:21749365] [WorldCat] [DOI]
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    Role of the basic helix-loop-helix transcription factor p48 in the differentiation phenotype of exocrine pancreas cancer cells. Cell growth & differentiation : the molecular biology journal of the American Association for Cancer Research, 11(3), 137-47. [PubMed:10768861] [WorldCat]
  14. ↑ Masui, T., Long, Q., Beres, T.M., Magnuson, M.A., & MacDonald, R.J. (2007).
    Early pancreatic development requires the vertebrate Suppressor of Hairless (RBPJ) in the PTF1 bHLH complex. Genes & development, 21(20), 2629-43. [PubMed:17938243] [PMC] [WorldCat] [DOI]
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    Pancreas-specific Cre driver lines and considerations for their prudent use. Cell metabolism, 18(1), 9-20. [PubMed:23823474] [PMC] [WorldCat] [DOI]
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    Transcriptional regulation of pancreas development and β-cell function [Review]. Endocrine journal, 64(5), 477-486. [PubMed:28420858] [WorldCat] [DOI]
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    HMGA1 expression levels are elevated in pancreatic intraepithelial neoplasia cells in the Ptf1a-Cre; LSL-KrasG12D transgenic mouse model of pancreatic cancer. British journal of cancer, 117(5), 639-647. [PubMed:28697176] [PMC] [WorldCat] [DOI]
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    p48 subunit of mouse PTF1 binds to RBP-Jkappa/CBF-1, the intracellular mediator of Notch signalling, and is expressed in the neural tube of early stage embryos. Genes to cells : devoted to molecular & cellular mechanisms, 6(4), 345-60. [PubMed:11318877] [WorldCat] [DOI]
  20. ↑ Hoshino, M., Nakamura, S., Mori, K., Kawauchi, T., Terao, M., Nishimura, Y.V., ..., & Nabeshima, Y. (2005).
    Ptf1a, a bHLH transcriptional gene, defines GABAergic neuronal fates in cerebellum. Neuron, 47(2), 201-13. [PubMed:16039563] [WorldCat] [DOI]
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    Molecular machinery governing GABAergic neuron specification in the cerebellum. Cerebellum (London, England), 5(3), 193-8. [PubMed:16997750] [WorldCat] [DOI]
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    The long adventurous journey of rhombic lip cells in jawed vertebrates: a comparative developmental analysis. Frontiers in neuroanatomy, 5, 27. [PubMed:21559349] [PMC] [WorldCat] [DOI]
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    Specification of spatial identities of cerebellar neuron progenitors by ptf1a and atoh1 for proper production of GABAergic and glutamatergic neurons. The Journal of neuroscience : the official journal of the Society for Neuroscience, 34(14), 4786-800. [PubMed:24695699] [PMC] [WorldCat] [DOI]
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    Temporal identity transition from Purkinje cell progenitors to GABAergic interneuron progenitors in the cerebellum. Nature communications, 5, 3337. [PubMed:24535035] [PMC] [WorldCat] [DOI]
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    Cerebellar GABAergic progenitors adopt an external granule cell-like phenotype in the absence of Ptf1a transcription factor expression. Proceedings of the National Academy of Sciences of the United States of America, 104(12), 5193-8. [PubMed:17360405] [PMC] [WorldCat] [DOI]
  26. ↑ Millen, K.J., & Gleeson, J.G. (2008).
    Cerebellar development and disease. Current opinion in neurobiology, 18(1), 12-9. [PubMed:18513948] [PMC] [WorldCat] [DOI]
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    Regulation of early cerebellar development. The FEBS journal, 290(11), 2786-2804. [PubMed:35262281] [WorldCat] [DOI]
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    Math1 is essential for genesis of cerebellar granule neurons. Nature, 390(6656), 169-72. [PubMed:9367153] [WorldCat] [DOI]
  30. ↑ Machold, R., & Fishell, G. (2005).
    Math1 is expressed in temporally discrete pools of cerebellar rhombic-lip neural progenitors. Neuron, 48(1), 17-24. [PubMed:16202705] [WorldCat] [DOI]
  31. ↑ Wang, V.Y., Rose, M.F., & Zoghbi, H.Y. (2005).
    Math1 expression redefines the rhombic lip derivatives and reveals novel lineages within the brainstem and cerebellum. Neuron, 48(1), 31-43. [PubMed:16202707] [WorldCat] [DOI]
  32. ↑ Glasgow, S.M., Henke, R.M., Macdonald, R.J., Wright, C.V., & Johnson, J.E. (2005).
    Ptf1a determines GABAergic over glutamatergic neuronal cell fate in the spinal cord dorsal horn. Development (Cambridge, England), 132(24), 5461-9. [PubMed:16291784] [WorldCat] [DOI]
  33. ↑ Hori, K., & Hoshino, M. (2012).
    GABAergic neuron specification in the spinal cord, the cerebellum, and the cochlear nucleus. Neural plasticity, 2012, 921732. [PubMed:22830054] [PMC] [WorldCat] [DOI]
  34. ↑ Fujitani, Y., Fujitani, S., Luo, H., Qiu, F., Burlison, J., Long, Q., ..., & Wright, C.V. (2006).
    Ptf1a determines horizontal and amacrine cell fates during mouse retinal development. Development (Cambridge, England), 133(22), 4439-50. [PubMed:17075007] [WorldCat] [DOI]
  35. ↑ Nakhai, H., Sel, S., Favor, J., Mendoza-Torres, L., Paulsen, F., Duncker, G.I., & Schmid, R.M. (2007).
    Ptf1a is essential for the differentiation of GABAergic and glycinergic amacrine cells and horizontal cells in the mouse retina. Development (Cambridge, England), 134(6), 1151-60. [PubMed:17301087] [WorldCat] [DOI]
  36. ↑ Dullin, J.P., Locker, M., Robach, M., Henningfeld, K.A., Parain, K., Afelik, S., ..., & Perron, M. (2007).
    Ptf1a triggers GABAergic neuronal cell fates in the retina. BMC developmental biology, 7, 110. [PubMed:17910758] [PMC] [WorldCat] [DOI]
  37. ↑ Fujiyama, T., Yamada, M., Terao, M., Terashima, T., Hioki, H., Inoue, Y.U., ..., & Hoshino, M. (2009).
    Inhibitory and excitatory subtypes of cochlear nucleus neurons are defined by distinct bHLH transcription factors, Ptf1a and Atoh1. Development (Cambridge, England), 136(12), 2049-58. [PubMed:19439493] [WorldCat] [DOI]
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