認知症の病因「タウタンパク質」の病変を抑制 〜Mark4遺伝子欠損がマウスのタウオパチー症状を改善~
アルツハイマー病など認知症の原因となる多くの疾患では、脳にタウと呼ばれるタンパク質が凝集して蓄積し(タウオパチー)、それが神経細胞を殺し、脳を萎縮させると考えられています。しかし、これらの疾患でどうしてタウタンパク質が凝集、蓄積するのかは、よくわかっていません。タウは、疾患脳では過剰な「リン酸化」という修飾を受けており、それが蓄積を引き起こすのではないかと考えられています。また、このタウのリン酸化を媒介する酵素の一つ、MARK4は、アルツハイマー病のリスクの増加とも関連することが知られています。東京都立大学大学院理学研究科生命科学専攻のGrigorii Sultanakhmetov大学院生(当時)、斎藤太郎助教、Adam Weitemier准教授、安藤香奈絵准教授らは、MARK4という酵素を欠損したマウスでは、タウの凝集が減少することを発見しました。ヒトの疾患の原因となるタウを神経細胞に発現したマウスでは、神経細胞のネットワークが壊れ、脳内で炎症が起き、認知機能が低下し、寿命が短縮しますが、MARK4を持たないマウスでは、タウによって引き起こされるこれらの症状が改善されました。本研究から、MARK4がタウの異常な蓄積とそれによる脳機能低下に関わることがわかり、MARK4の阻害がアルツハイマー病など認知症の原因となる疾患の治療に役立つ可能性が示されました。
2. ポイント
・脳の神経細胞におけるタウの凝集と蓄積(タウオパチー)は、アルツハイマー病やその他認知症の原因となる疾患を引き起こす。
・MARK4という酵素を欠損したマウスは、タウの凝集が起きにくいことがわかった。
・MARK4欠損によって、タウによる認知機能低下や寿命短縮が改善した。
・MARK4の阻害は、タウの凝集と蓄積を緩和し、認知症の治療に役立つ可能性がある。
3. 研究の背景
日本を含む多くの国が高齢化社会を迎え、アルツハイマー病など加齢に伴う神経変性疾患の発症率も増加しています。これらの神経変性疾患では、脳の機能を担っている神経細胞が次第に死んでいくため、記憶障害や生活機能障害が引き起こされます。また、その特徴的な病変として、脳の神経細胞の中にタウと呼ばれるタンパク質が凝集して沈着している現象が見られます(タウオパチー)。異常に凝集したタウの量は疾患の進行に伴って増加し、この凝集したタウは神経毒性を持つことがわかっています。そのため、タウの蓄積の抑制は、タウオパチーを伴う認知症の治療戦略として検討されていますが、どのようにタウの凝集と蓄積が起きるのかははっきりとわかっていません。
発症過程で起きるタウの変化の一つに、過剰なリン酸化があります。タウは多くの部位で、様々な酵素によってリン酸化を受けますが、なかでもMicrotubule affinity regulating kinase 4 (MARK4)は、タウ病変と共局在し、疾患脳で活性が増加していることや、その変異がアルツハイマー病のリスクを増加させることなどから、アルツハイマー病でのタウ病変に関与するのではないかと考えられてきました。また、ショウジョウバエを使った実験では、MARK4の活性が増加すると、タウによる神経細胞死が悪化することが報告されていました。そこで今回の研究では、ヒトと同じ哺乳類であるマウスで、MARK4とタウ病変の関係を調べました。
4.研究の詳細
ヒトで神経変性疾患の原因となる変異をもつタウを神経細胞に発現したマウスでは、認知機能が低下し、寿命が短縮します。また脳の神経細胞では、タウの凝集、リン酸化タウの蓄積が見られ、神経細胞同士のコネクション(シナプス)が失われるのが観察されます。このタウオパチーマウスを、MARK4をコードする遺伝子Mark4を欠損したマウスと掛け合わせ、MARK4の有無でタウオパチーマウスの症状が変化するかを、行動、生化学的、組織学的解析で調べました。
その結果、Mark4欠損マウスでは、Mark4遺伝子を持つマウスと比べて、死亡率と認知機能が改善していることがわかりました。脳を生化学的に調べると、Mark4遺伝子欠損マウスではリン酸化タウが減少し、また凝集したタウを標識するチオフラビンSのシグナルが低下していました。Mark4遺伝子欠損マウスの神経細胞は、タウを発現させてもより多くのシナプスを維持していました。また、タウの発現によって、脳の免疫細胞であるグリアが活性化し炎症を起こしますが、Mark4遺伝子欠損マウスでは、グリアの一種アストロサイトの活性化が抑制されていました。アストロサイトの活性化は加齢によっても起きますが、Mark4遺伝子欠損マウスの脳では、老化によるアストロサイトの活性化も抑制されていました。
【画像:https://kyodonewsprwire.jp/img/202404199722-O1-8fI7VZyV】
5.研究の意義と波及効果
これらの結果から、MARK4がタウオパチーの発症機構に寄与していることが明らかになり、またMARK4の阻害がタウオパチーを伴う認知症の治療戦略となる可能性が示されました。Mark4遺伝子欠損マウスは正常に発育していたため、その阻害によって甚大な副作用は起きないことが示唆されます。MARK4を阻害する薬剤の開発により、アルツハイマー病やその他のタウ病変を伴う疾患の、根本的な治療薬の開発が期待できます。
(論文情報)
タイトル:Mark4 ablation attenuates pathological phenotypes in a mouse model of tauopathy
著者:Grigorii Sultanakhmetov, Sophia Jobien M. Limlingan, Aoi Fukuchi, Keisuke Tsuda, Hirokazu Suzuki, Iori Kato, Taro Saito, Adam Z. Weitemier and Kanae Ando
掲載誌:Brain Communications
DOI:10.1093/braincomms/fcae136
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File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/d064ec4523cd2f49c67136b7b84705846d19c90b_0.file.side.tpl.php
Line: 43
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b47775ef5_14228348
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 417
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 248
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 184
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 186
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 156
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/fbdaa42e51518f8881faeabdc1199f6dd175d1ce_0.file.base_index_layout.tpl.php
Line: 88
Function: instanceBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3cb92fa0_94546305
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 116
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 71
Function: endChild
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_671b089d2e5dc3_21870328
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 238
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 116
Function: _execute
File: /www/wwwroot/homev2.kingsoft.jp/application/libraries/SmartyLibrary.php
Line: 45
Function: fetch
File: /www/wwwroot/homev2.kingsoft.jp/application/controllers/ArticleController.php
Line: 32
Function: view
File: /www/wwwroot/homev2.kingsoft.jp/public/index.php
Line: 292
Function: require_once
A PHP Error was encountered
Severity: Notice
Message: Undefined offset: 1
Filename: smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line Number: 41
Backtrace:
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line: 41
Function: _error_handler
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d14aa11_83151422
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/d064ec4523cd2f49c67136b7b84705846d19c90b_0.file.side.tpl.php
Line: 43
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b47775ef5_14228348
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 417
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 248
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 184
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 186
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 156
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/fbdaa42e51518f8881faeabdc1199f6dd175d1ce_0.file.base_index_layout.tpl.php
Line: 88
Function: instanceBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3cb92fa0_94546305
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 116
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 71
Function: endChild
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_671b089d2e5dc3_21870328
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 238
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 116
Function: _execute
File: /www/wwwroot/homev2.kingsoft.jp/application/libraries/SmartyLibrary.php
Line: 45
Function: fetch
File: /www/wwwroot/homev2.kingsoft.jp/application/controllers/ArticleController.php
Line: 32
Function: view
File: /www/wwwroot/homev2.kingsoft.jp/public/index.php
Line: 292
Function: require_once
A PHP Error was encountered
Severity: Notice
Message: Trying to access array offset on value of type null
Filename: smarty/49292b3df7e5bd60f7381ff16c5ca4532cf9476f_0.file.card.tpl.php
Line Number: 24
Backtrace:
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/49292b3df7e5bd60f7381ff16c5ca4532cf9476f_0.file.card.tpl.php
Line: 24
Function: _error_handler
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d136f97_11902452
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 384
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line: 41
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d14aa11_83151422
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/d064ec4523cd2f49c67136b7b84705846d19c90b_0.file.side.tpl.php
Line: 43
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b47775ef5_14228348
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 417
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 248
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 184
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 186
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 156
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/fbdaa42e51518f8881faeabdc1199f6dd175d1ce_0.file.base_index_layout.tpl.php
Line: 88
Function: instanceBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3cb92fa0_94546305
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 116
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 71
Function: endChild
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_671b089d2e5dc3_21870328
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 238
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 116
Function: _execute
File: /www/wwwroot/homev2.kingsoft.jp/application/libraries/SmartyLibrary.php
Line: 45
Function: fetch
File: /www/wwwroot/homev2.kingsoft.jp/application/controllers/ArticleController.php
Line: 32
Function: view
File: /www/wwwroot/homev2.kingsoft.jp/public/index.php
Line: 292
Function: require_once
A PHP Error was encountered
Severity: Notice
Message: Trying to access array offset on value of type null
Filename: smarty/49292b3df7e5bd60f7381ff16c5ca4532cf9476f_0.file.card.tpl.php
Line Number: 29
Backtrace:
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/49292b3df7e5bd60f7381ff16c5ca4532cf9476f_0.file.card.tpl.php
Line: 29
Function: _error_handler
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d136f97_11902452
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 384
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line: 41
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d14aa11_83151422
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/d064ec4523cd2f49c67136b7b84705846d19c90b_0.file.side.tpl.php
Line: 43
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b47775ef5_14228348
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 417
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 248
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 184
Function: callBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 186
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 156
Function: process
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/fbdaa42e51518f8881faeabdc1199f6dd175d1ce_0.file.base_index_layout.tpl.php
Line: 88
Function: instanceBlock
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3cb92fa0_94546305
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
Line: 116
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 71
Function: endChild
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_671b089d2e5dc3_21870328
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 238
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_templatebase.php
Line: 116
Function: _execute
File: /www/wwwroot/homev2.kingsoft.jp/application/libraries/SmartyLibrary.php
Line: 45
Function: fetch
File: /www/wwwroot/homev2.kingsoft.jp/application/controllers/ArticleController.php
Line: 32
Function: view
File: /www/wwwroot/homev2.kingsoft.jp/public/index.php
Line: 292
Function: require_once
A PHP Error was encountered
Severity: Notice
Message: Undefined offset: 2
Filename: smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line Number: 41
Backtrace:
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/08d1cc69876d91563afb9c1f4d21cc8b7b96caa4_0.file.tips.tpl.php
Line: 41
Function: _error_handler
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b3d14aa11_83151422
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/d064ec4523cd2f49c67136b7b84705846d19c90b_0.file.side.tpl.php
Line: 43
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_resource_base.php
Line: 123
Function: content_66f26b47775ef5_14228348
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_template_compiled.php
Line: 114
Function: getRenderedTemplateCode
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 217
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_template.php
Line: 386
Function: render
File: /www/wwwroot/homev2.kingsoft.jp/application/cache/smarty/597cefa22c7600eaa32b117569a9685c70819274_0.file.show.tpl.php
Line: 417
Function: _subTemplateRender
File: /www/wwwroot/homev2.kingsoft.jp/application/third_party/smarty/libs/sysplugins/smarty_internal_runtime_inheritance.php
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Function: callBlock
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