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ハイパーテキスト転送プロトコル RFC 2616 WebブラウザとWebサーバの間でHTMLなどのコンテンツの送受信に用いられる通信プロトコル リクエスト-レスポンス型 トランスポート・プロトコルとして通常TCPを使用 基本的な考え方は非常に単純であり「何を」「どうして」ほしいのかを相手に要求する。「何を」に当たるのがURL、「どうして」がメソッドにあたる。 World Wide WebにおけるWebページなどのリソースは、Uniform Resource Identifierによって指定される。 ポート番号80をデフォルトとして使用する（送信時は8080）。 TLSで暗号化され、セキュリティを確保したHTTPは、HTTPSと呼ばれる（httpsは実際にはURIスキームの1つであり、実際のプロトコルにはHTTP over SSL/TLSが用いられる）。 HTTP は基本的にサーバが状態を保持しない (stateless) プロトコルだが、データベースなどを使用するWebアプリケーションにおいては状態保持が必要だったため、そのためにいわゆる Cookie とよばれる機構が Netscape Communications Corporation によって導入された。Cookie を使用することによって状態を管理し、"セッション" を維持することが可能になる。 HTTPの拡張プロトコルとしてWebDAVがある。 UPnPでは、HTTPをUDP上で使用するHTTPUや、マルチキャストで使用するHTTPMUが規定された。 HTTP/0.9 URLのみの簡単なやりとり HTTP/1.0 NNTPやSMTPのような各種ヘッダが定義 HTTP_Cookieなどの利用 HTTP/1.1 複数データを転送するためのキープアライブ（keep-alive）機能やプロキシなどの利用も想定された仕様 バーチャルホストをサポートした。インターネット人気に伴い多くの企業がWebサイトを持ち始めたが、当時ではまだまだ企業が自前のWebサーバを運用するのは人員、効率の問題で難しかったためISPのサーバでホスティングをしていた。当時はまだ一社ごとに専用サーバを用意するほどのことでもないため一台のサーバで複数のWebサイトを運用していた。 しかしバーチャルホストには問題がある。例えばある1台のサーバに foo.example.com と bar.example.com という二つの仮想Webサーバがあるとする。ここではクライアントは http //foo.example.com/index.html にアクセスしたいとする。そのためにはまず foo.example.com をIPアドレスに解決するためDNSサーバに問い合わせ、そのサーバにアクセスし GET index.html を要求する。しかしサーバ側のIPアドレスは foo.example.com と bar.example.com 共におなじIPアドレスである。もし foo.example.com にも bar.example.com にも index.html というファイルが存在すればクライアントはどちらのサーバにアクセスしたのかわかるすべがない。 これを解決するにはそれぞれにIPアドレスを付与することで解決できるが、IPv4の資源を無駄にすることになる。 HTTP/1.1ではこれを解決するためにHostヘッダを追加した。 HTTP/1.0のヘッダ GET /index.html HTTP/1.0 HTTP/1.1のヘッダ GET /index.html HTTP/1.1 Host foo.example.com 動作 通信の開始 他のプロトコル同様クライアント側とサーバ側ではHTTPの役割が大きく異なる。HTTP通信を開始できるのはクライアント側のみである。 クライアント側はサーバにリクエストを送り、サーバはクライアントにレスポンスを返すのが最も典型的なHTTPのやりとりである。 接続 システム間でメッセージをやりとりするにはTCP接続を確立させる必要がある。 HTTP/0.9ではクライアントのリクエストごとにTCP接続を確立させる必要があったが、これは当時のWebサイトがシンプルなテキストベースであることが多かったためである。近年ではJavaScriptやアニメーション画像など、多数のオブジェクトが埋め込まれたWebサイトが一般的となってきているが、これら全てのオブジェクトを取得するたびにTCP接続を確立するのはサーバやネットワークに大きな負担を強いるため、HTTP/1.1では持続的接続がサポートされることとなった。ただしこの機能が利用できるのはサーバ側がその要求を許可した場合のみである。 パイプライン クライアントは前のリクエストに対するサーバの応答を待たずに別のリクエストを発行できる。 メソッド HTTPでは8つのメソッドが定義されている。ただし実際のHTTP通信ではGETとPOSTメソッドだけで殆どを占める。 HTTPメソッドの一覧 メソッド HTTP/0.9 HTTP/1.0 HTTP/1.1 GET ○ ○ ○ POST ○ ○ PUT △ ○ HEAD ○ ○ DELETE △ ○ OPTION ○ TRACE ○ CONNECT ○ GET 指定されたURIのリソースを取り出す。HTTPの最も基本的な動作で、HTTP/0.9では唯一のメソッド。 POST GETとは反対にクライアントがサーバにデータを送信するメソッドである。Webフォームや電子掲示板、Wikiなどに投稿する。GETの場合と同じくサーバはクライアントにデータを返すことができる。 PUT 指定したURIにリソースを保存する。URIが指し示すリソースが存在しない場合は、サーバはそのURIにリソースを作成する。画像のアップロードなどが代表的。 DELETE 指定したURIのリソースを削除する。 OPTION サーバを調査するメソッド。例えばサーバがサポートしているHTTPのバージョンなどを調査できる。 HEAD GETと似ているがサーバはHTTPヘッダのみ返す。クライアントはWebページを取得せずともそのWebページが存在するかどうかを知ることが出来る。例えばWebページのリンク先が生きているか検証するときなどにリンク先のデータを全て取得することなく調査することが出来る。 TRACE サーバまでのネットワーク経路をチェックできる。サーバは受け取ったメッセージのそれ自体をレスポンスのデータにコピーして応答する。WindowsのTracertやUNIXのTracerouteとよく似た動作。 CONNECT 暗号化したメッセージをプロキシで転送する際に用いる。 サーバの連携 バーチャルホスト リダイレクト 301 MovedというステータスコードとURIを受け取りクライアントはこの受け取ったURIに再度GETを送る。 クッキー(HTTP_Cookie) HTTPメッセージ クライアントからのHTTPリクエストは3つの要素から構成される。それぞれメソッド、URI、HTTPのバージョンでありスペースで区切られている。 下にもっとも単純な、クライアントとサーバ(www.google.co.jp 80)とのHTTPプロトコルのやり取りの例を挙げる。 クライアントのリクエスト GET / HTTP/1.0 GETがメソッド、URIは / 、バージョンはHTTP/1.0であることを示す。 URIは/でルートリソースを対象にしたリクエストであることを示している。TRACEなど特定のサーバを対象としないリクエストの場合には*が表示される。 サーバのレスポンス HTTP/1.0 200 OK Cache-Control private Content-Type text/html Set-Cookie PREF=ID=72c1ca72230dea65 LD=ja TM=1113132863 LM=111 3132863 S=nNO7MIp W2o7Cqeu_; expires=Sun, 17-Jan-2038 19 14 07 GMT; path=/; domain=.google.co.jp Server GWS/2.1 Date Sun, 10 Apr 2005 11 34 23 GMT Connection Close html head meta http-equiv="content-type" content="text/html; charset=Shift_JI S" title Google /title style !-- ・・・以下省略 上のリクエストのGETにあたる部分をメソッドといい、 HTTP/1.0では、GET, HEAD, PUT, POST, DELETE, LINK, UNLINK、 HTTP/1.1ではさらに、OPTIONS, TRACEがある。 GETメソッドのレスポンスにはヘッダ情報のあとに改行が挟まれ、コンテンツ本体が送られる。 HEADメソッドのレスポンスにはコンテンツサイズや更新日時などの情報を含むヘッダのみが送られる。 また、リクエストの2行目以降はヘッダを送る。 HTTPヘッダフィールド ヘッダの各要素は フィールド名 内容 のペアで構成される。 ブラウザの情報を表すUser-Agent、使用候補言語を表すAccept-Language、他ページへのリンクを辿った場合にそのリンク元ページのURLを表すRefererなどが代表的なフィールドである。 なお、リクエスト時のHostヘッダはHTTP/1.1では必須であるが、HTTP/1.0では無くても良い。 但し、サーバがバーチャルホストを利用している場合は、Hostヘッダが無いとリソース取得に失敗するので、たとえHTTP/1.0を使用していてもHostヘッダを付加しなければならない。 HTTPヘッダフィールドの一覧 リクエストヘッダ ヘッダ概要HTTP/0.9HTTP/1.0HTTP/1.1 Acceptクライアントの受け入れ可能コンテンツタイプを示す○○ Accept-Charsetクライアントの受け入れ可能文字セットを示す○○ Accept-Encodingクライアントの受け入れ可能文字エンコーディングを示す○○ Accept-Languageクライアントの受け入れ可能言語を示す○○ Authorizationクライアントの認証情報を示す○○ Cookieクライアントの状態管理情報をサーバに返す Cookie2HTTP/1.1のSet-Cookie2ヘッダの受け入れ可能をサーバに知らせる Expectクライアントがサーバに期待する動作を示す○ Fromリクエスト発行者個人の情報を示す。一般的に電子メールアドレスを使用する○○ Host要求しているオブジェクトがあるホストを示す○ If-Matchif文を用い条件が真の場合のみリクエストを処理するようサーバに要求する○ If-Modified-Since指定日及び指定時刻以降にオブジェクトが変更されている場合のみリクエストを処理するよう要求する○○ If-None-MatchIf-Matchの逆で条件が真でない場合のみリクエストを処理する要求○ If-Range条件が真の場合のみ指定したオブジェクトの範囲を返すようサーバに要求する○ If-Unmodified-SinceIf-Modified-Sinceの逆で真でないときのみ実行する○ Max-Forwardsリクエストの中間システム経由数を最大いくつまでかを指定する○ Proxy-Authorizationクライアントがプロキシサーバに対して自身の認証を行う○ Rangeオブジェクト全体でなくリソースの一部を要求する○ Refererリクエストの出所を示す。一般的にはユーザの辿ったWebページのURLが用いられる。○○ TEレスポンスの受け入れ可能転送エンコーディングを示す○ レスポンスヘッダ ヘッダ概要HTTP/0.9HTTP/1.0HTTP/1.1 Accept-Rangesオブジェクトの一部に対するリクエストをサーバが受け入れ可能か示す○ Ageオブジェクトの経過時間を秒単位で返す○ AllowオブジェクトがサポートするHTTPメソッドを示す○○ ETagオブジェクトのエンティティタグ値を示す○ Locationオブジェクトの場所を示す○○ Proxy-Authenticateプロキシサーバがクライアントに認証を要求するときに用いる○ Retry-Afterリクエストの再試行をいつ行うかをクライアントに通知する○○ Serverサーバのベンダー名、バージョン番号を占めす○○ Set-Cookie2サーバがクライアントにCookieを送信するときに用いる Varyサーバのレスポンス内容を決定する際にリクエストURI以外に使用したHTTPヘッダのリストを示す○ WWW-Authenticateクライアントに対してリクエストの再発行を要求する。認証情報も含まれる○○ 一般ヘッダ ヘッダ概要HTTP/0.9HTTP/1.0HTTP/1.1 Cache-Controlメッセージの経由する中間キャッシュの動作を指示する○ Connection中間システムが転送すべきでないヘッダのリストを示す○○ Dateメッセージの作成日時を示す○○ Pragmaメッセージに関する追加情報を示す○○ Trailerメッセージボディの後に追加のヘッダーが表れることを示す○ Transfer-Encodingクライアントの転送を目的としたオブジェクトのエンコーディングを示す○ Upgrade通信相手に別のプロトコルにアップデートするよう要求する○ User-AgentクライアントのWebブラウザなどの情報を示す○○ Warningメッセージに関する追加情報を示す。通常はキャッシュの問題を警告するときに使われる○ エンティティヘッダ ヘッダ概要HTTP/0.9HTTP/1.0HTTP/1.1 Content-Encodingオブジェクトのエンコーディングを示す○○ Content-Languageオブジェクトの言語（人間の言語）を示す○○ Content-Lengthオブジェクトのサイズをバイト単位で示す○○ Content-Locationオブジェクトの場所を示す○ Content-MD5オブジェクトのメッセージダイジェストを運ぶ○ Content-Rangeメッセージボディで運ばれるオブジェクトの範囲を示す○ Content-Typeオブジェクトのタイプを示す○○ Expiresオブジェクトの有効期限の日時を示す○○ Last-Modifiedオブジェクトが最後に変更された日時を示す○○ Accept サーバのレスポンスに含まれるメッセージボディで受け入れることが出来るコンテンツタイプと各コンテンツタイプの相対的な優先度を指定するリクエストヘッダ。指定できるコンテンツタイプはIANAによって定義されている。 Accept text/plain; q=0.5, text/html, text/x-dvi; q=0.8, text/x-c 上記のようにAcceptヘッダには行をわけて複数のコンテンツタイプを指定できる。上記の例はいずれの4のコンテンツタイプのいずれも受け入れ可能であることを示す。0.5や0.8といった数字は品質係数で0～1の範囲の数値である。数値の指定がなければ1.0となる。 text/plain; q=0.5 text/html text/x-dvi; q=0.8 text/x-c Accept-Charset レスポンスで返されるメッセージボディの文字コードを指定するリクエストヘッダ。Acceptと同じく複数指定でき品質係数も設定できる。定義済み文字セットはIANAが管理している。 Accept-Charset unicode, *; q=0.8 この例だとクライアントはUnicode文字セットを優先的に希望しているが他の文字セットとの相対優先度0.8で受け入れている。ただしサーバからのレスポンスのHTTPヘッダそのものの文字コードは常にISO-8859-1である。 Accept-Encoding Accept-Language レスポンスの言語（人間の言語）に対する優先度を指定する。言語コードはISO-639の2文字の省略コードを用いる。書き方は他のAccept-群と変わらず。 Accept-Language en-gb, en; q=0.8 上記の例はまずイギリス英語を要求し、利用できない場合はその他の英語を要求する。 Accept-Ranges Acceptで始まる他のヘッダフィールドと違いレスポンスヘッダーである。現在の仕様では2つの指定方法しかない。 Age リソースの推定経過時間を表示するレスポンスヘッダ。キャッシュサーバーはAgeヘッダの値からキャッシュしたリソースが有効かどうかを判定する。 Allow Authentication-info ユーザ認証のやりとりの最後で用いられる、成功したレスポンスのサーバが含めることの出来るレスポンスヘッダー。 Authorization サーバに対するクライアント自身の認証を行うことが出来る。 Cache-Control キャッシングの動作を指定するためのマスターヘッダ。 Connection Content-Encoding Content-Language リソースを英語などの自然言語で示すのに使われる。言語の指定はAccept-Languageヘッダと同じ。 Content-Length Content-Location Content-MD5 メッセージボディが変更されず宛先に届くことを保証する。MD5アルゴリズムを実行する。ただし悪意の改ざんに対しては当然MD5も改ざんされるのであまり機能はしない。どちらかといえば偶発的な変更の保証をしている。 Content-Range ダウンロードの再開に用いられる。 Content-Type メッセージボディに含まれるオブジェクトタイプを示す。次の例はリソースがテキストファイル、文字セットはISO-8859-4を使用していることを示している。 Content-Type text/plain; Charset=ISO-8859-4 Cookie クライアントがHTTP状態管理を望む場合にサーバから受け取ったクッキーを以後のリクエストに次の例のようなヘッダーを付加する。 Cookie $Version="1"; NAME="VALUE"; $Path="/shopping"; $domain="www.shop.com"+ $Port="80" $VersionはHTTPのバージョン、NAMEはクッキーの名前である。$から始まるクッキー名は使用が禁止されている。 Cookie2 基本的にCookieヘッダーとCookie2ヘッダーは別物である。 Date サーバがメッセージを生成した日時を示す。リソースの時間を示すLast-Modifiedヘッダーとは区別する必要がある。 HTTP/1.1では次のような形式を用いるようRFC1123で定義されている。 Date Sun, 06, Nov 1994 08 49 37 GMT HTTP仕様ではレスポンスにDateヘッダーを含めることを求めている。ただしレスポンスのステータスがサーバエラーの場合にはDateヘッダーは返らない。 ETag 主にキャッシングのパフォーマンスを向上する目的で使われる。 Expect サーバに対して特定の動作の期待を知らせる。用途としてはクライアントがサーバに対して100 Continueステータスを返すことを期待する場合に使われる。 Expect 100-continue サーバが期待に応じれない場合は417 Expectation Failedを返す。クライアントがいくつかのプロキシ経由で通信している場合、各プロキシサーバはExpectヘッダの一切の修正を許されない。 Expires オブジェクトの有効期限を示す。このヘッダで指定された日時までキャッシュはレスポンスのコピーを保持し、リクエストに対するレスポンスとして返すことが出来る。サーバがオブジェクトのキャッシュを望まない場合にはExpiresヘッダに過去の日時を設定することが多い。また、HTTP仕様では1年以上先の日時は設定できない。 Expires Thu, 28 Aug 2010 16 00 00 GMT Cache-Controlヘッダのmax-ageディレクティブはExpiresヘッダより優先されるため注意が必要である。 From リクエストを発行したユーザを特定することが出来る。1990年代では電子メールアドレスを設定することが多かったが、迷惑メールの問題もあり現在では殆ど使われていない。 From hoge@hogehoge.com Host 主にレンタルサーバのサポートを目的としてHTTP/1.1で導入された。現在ではHostヘッダを利用できない場合レンタルサーバのウェブサイトとまともな通信が出来ないと言ってよい（詳細はHTTP#歴史を参照）。 If-Match クライアントのリクエストを条件付きのリクエストにするために使われる。サーバは一定の条件が真であった場合のみリクエストを受け入れることが出来る。例えばウィキペディアを編集する際、記事のソースを取得し、書き換える際の間に別のユーザが既に編集していないかを判断するときなどに用いられる。 「if文」も参照 利用者：HogeがHTTPの記事を取得。ETagは1234 利用者：HageがHTTPの記事を取得。ETagは1234 利用者：HogeがHTTPのETagを再度取得。先ほど取得したETag 1234と現在のETag 1234が一致。 利用者：HogeがHTTPの記事を編集。ETagは1256になる。 利用者：HageがHTTPのETagを再度取得。先ほど取得したETagと現在のETagはマッチせず。 サーバは利用者：Hageの書き込みを拒否。 If-Modified-Since このヘッダーで指定された日時以降にオブジェクトが変更されている場合のみリクエストに応答するようサーバに要求する。リソースの削減に効果がある。 If-None-Match If-Matchと逆で条件が真でない場合のみリクエストを処理するよう要求する。 If-Range クライアントがキャッシュにオブジェクトの一部分を持っている場合にパフォーマンスを向上できる。 If-Unmodified-Since If-Modified-Sinceの逆の働きをする Last-Modified サーバオブジェクトの最終更新日時を示す。クライアントはこのヘッダを利用しIf-Modified-Sinceヘッダ等と組み合わせることによって効果を発揮する。 Location サーバがクライアントにリダイレクト先URLを知らせる際に用いられる。一般的にステータスコードが3xx代のレスポンスと共に使われるが201 Createdのレスポンスでも使うことが出来る。Content-Locationヘッダと名前が似ているが全く関係のない別のヘッダであるため注意。 Max-Forwards プロキシサーバ等を経由する際の最大ホップ数を指定する。二重ループなどでサーバから応答が得られない場合の問題解決の際、OPTIONメソッドやTRACEメソッドと共に用いられる。 HTTPステータスコード ステータスコードはクライアントのリクエストが成功したかどうかを示した上で追加情報を提供するいずれも3桁の数字から成る。具体的には100-199が情報提供、200-299が成功を示す。300-399はリダイレクト、400-499はエラーを示す。 セキュリティ技術 Basic認証 HTTP/1.1でBasic認証が定義されており最も単純なセキュリティ技術である。しかし仕様書を読むと定義を書いた著者自身が認証技術に疎いことがよくわかる。『HTTPプロトコル セキュア＆スケーラブルなWeb開発』の著者は「基本認証を用いるくらいならなにも使わない方がまし」と著書に書いている。通常サーバは401ステータスコードで応答する。 行末文字はWindowsと同じCRLF。 RFC 2818 - HTTP Over TLS RFC 2817 - Upgrading to TLS Within HTTP/1.1 RFC 2616 - HTTP/1.1 ハイパーテキスト転送プロトコル -- HTTP/1.1 RFC 2068 - HTTP/1.1（初版，RFC 2616 によって obsolete） TS X 0085 2004 - ハイパテキスト転送プロトコル HTTP/1.1 標準仕様書(TS) RFC 1945 - HTTP/1.0 HttpTea Freeware HTTP Logger Studying HTTP
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Ultimate Spider-Man (Ultimate Spider-Man (Graphic Novels)) Marvel Enterprises?Brian Michael Bendis?Bill Jemas?Mark Bagley? The Complete Frank Miller Spider-Man Marvel Enterprises?Frank Miller? The Best of Spider Man Marvel Enterprises?J. Michael Straczynski?John Romita?Greg Rucka?Peter Milligan?Eduardo Risso?Duncan Fegredo?Paul Jenkins?Staz Johnson?Brian Michael Bendis? G.I. Joe A Real American Hero Marvel Enterprises?Larry Hama?Herb Trimpe?Steven Grand? Spider Man The Movie (Marvel Graphic Novels) Marvel Enterprises?Stan Lee?Alan Davis? The Incredible Hulk Boiling Point (Incredible Hulk) Marvel Enterprises?Bruce Jones? Blaze of Glory Marvel Enterprises?John Ostrander?Leonardo Manco? G.I. Joe (G.I. Joe) Marvel Enterprises?Larry Hama?Herb Trimpe? Cable The Shining Path (Cable) Marvel Enterprises?David Tischman?Igor Kordey? Ghost Rider The Hammer Lane (Ghost Rider) Marvel Enterprises?Devin Grayson? Elektra Wolverine The Redeemer Marvel Enterprises?Greg Rucka?Yoshitaka Amano? The Further Adventures of Cyclops and Phoenix Marvel Enterprises?Peter Milligan?John Paul Leon? The Clandestine Vs. the X-Men Marvel Enterprises?Alan Davis? X-Men Visionaries (X-Men Visionaries) Marvel Enterprises?Jim Lee?Ann Nocenti? The Punisher War Zone Marvel Enterprises?Chuck Dixon? Marvel Masterworks Presents the Amazing Spider-Man (Marvel Masterworks (Numbered)) Marvel Enterprises?Stan Lee? Thor The Death of Odin (Mighty Thor Lord of Asgard) Marvel Enterprises?Dan Jurgens? Daredevil The Man Without Fear! (Daredevil) Marvel Enterprises?Brian Michael Bendis?Alexander Maleev? 2000-2001 Year in Review Fanboys and Badgirls Bill Joe's Marvelous Adventure (Marvel Comics) Marvel Enterprises?Jim McLaughlin? G.I. Joe A Real American Hero (G.I. Joe) Marvel Enterprises?Larry Hama? Howard the Duck Marvel Enterprises?Phil Winslade?Glenn Fabry?Garry Leach?Steve Gerber? Union Jack Marvel Enterprises?Ben Raab?John Cassady? Son of Origins of Marvel Comics Marvel Enterprises?Stan Lee?Jack Kirby?John Buscema?Don Heck?Bill Everett?Gene Colan? X-Men Evolution (X-Men) Marvel Enterprises?Devin Grayson? The Incredible Hulk Return of the Monster (Incredible Hulk) Marvel Enterprises?Bruce Jones?John Romita? Marvel Visionaries (Marvel Visionaries) Marvel Enterprises?Stan Lee?Arnold Drake?Jim Steranko? Daredevil Wake Up (Daredevil) Marvel Enterprises?Brian Michael Bendis? Essential Daredevil The Man Without Fear (Essential Daredevil) Marvel Enterprises?Stan Lee?Wally Wood?John Romita?Gene Colan? Contest of Champions Marvel Enterprises?Bill Mantlo? Marvel Legends Wolverine (Wolverine) Marvel Enterprises?Larry Hama?Marc Silvestri?Dan Green?Hilary Barta?Tom Palmer? Avengers The Kang Dynasty (Avengers) Marvel Enterprises?Kurt Busiek? Daredevil The Movie (Daredevil) Marvel Enterprises?Mark Stephen Johnson?Bruce Jones?Manuel Garcia? The Ultimates (Ultimates) Marvel Enterprises?Mark Millar?Bryan Hitch?Andrew Currie? Spider-Man Revelations (Spider-Man) Marvel Enterprises?Todd Dezago?Tom DeFalco?Howard MacKie?J. M. Dematteis?Luke Ross?Steve Skroce?Mike Wieringo?Bud Larosa?Scott Hanna?John Romita? Cable The End (Cable) Marvel Enterprises?David Tischman?Darko Macan?Igor Kordey? New X-Men (New X-Men) Marvel Enterprises?Grant Morrison?Frank Quitely? Spider-Man Marvel ComicsTodd Dezago? Cage Marvel Enterprises?Brian Azzarello? Ultimate Spider-Man Legacy (Ultimate Spider-Man (Graphic Novels)) Marvel Enterprises?Brian Michael Bendis?Mark Bagley? Daredevil Yellow (Daredevil) Marvel Enterprises?Jeph Loeb?Tim Sale?
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Market Analysis The global Online Sports Betting Market is predicted to increase at a cagr of 13.6%, from USD 24,988.4 million in 2019 to USD 59,530.7 million by the forecast period. The global online sports betting market is expanding due to increased demand for sports, which is essential for reaching development goals. In a number of geographical, cultural, and political contexts, sport can play an important role in promoting societal integration and economic prosperity. Sports impact on social capital, culture, trust, and crime can help with increasing capital accumulation, thereby improving and strengthening market functioning. All these outcomes further reinforce each other to grow the market. Furthermore, participating in sports allows players to live longer and healthier lives. The global E-Sports market has been expanding in recent years, with an increasing number of players participating as competitors or spectators. The modern sports-only websites are now growing in popularity just like the traditional sports betting website, now offer chances to bet on events and tournaments, similar to how many people bet on E-Sports. The major growth factor of such market share would be the increasing sports demand and rising demand for e-Sports activities. The online sports betting market is seeing an increase in growth due to rising digital innovation. However, due to Government prohibitions, some nations are expected to put some restrain in the global market expansion. Then again, the enhancement of digital innovation through advanced programming and technological use can bring more opportunities in the global market due to high-quality games. COVID-19 Impact on the Worldwide Online Sports Betting Market The spread of new coronavirus has impacted numerous industries, including manufacturing, from raw material production to final product distribution. On the other hand, in order to track the spread of a deadly virus, governments around the world imposed restrictions on a variety of sporting activities. Scheduled events including the English Premier League, Serie A in Italy, La Liga in Spain, Super Lig in Turkey, and Major League Soccer in the United States were canceled or postponed due to a rise in the number of COVID-19 cases globally. Some organizations prefer to reschedule events in order to decrease expenditure losses, such as the Belgian Cup final, which was initially slated for March 2020 but was moved to August 1st. Another case in point is the African Nations Championship 2020 competition, which was scheduled to take place in Cameroon in April 2020 but has been postponed until further notice. Since governments imposed lockdowns to contain and monitor the spread of the fatal virus, a sizable percentage of the workforce has been instructed to remain indoors, resulting in a surge in demand for virtual entertainment and media. Microsoft, for instance, announced a large rise in multiplayer participation in March and April 2020. Furthermore, when physical venues stayed closed to prevent the spread of the virus, online casino gaming and fantasy sports betting grew by more than 30%. Request a Free Sample @ https //www.marketresearchfuture.com/sample_request/8641 Market Segmentation The global online sports betting market is classified into various segments based on sports type, by type, by application. Moreover, on the basis of sports type, the online sports betting market is bifurcated into football, cricket, tennis, baseball, volleyball, basketball, hockey, horse riding, boxing, golf, racing, and many others. Further, the football segment is sub-segments into FIFA, Premier League, UEA/EURO, and others. The market analysis report suggests that the FIFA segment has registered for acquiring the largest market value in the preview period and increasing at a CAGR of 125 in the forecast period. According to the by Type segment, the online sports betting market is classified into line-in-play, fixed old betting, daily fantasy, e-sports, exchange betting, spread betting, and others. The online sports betting market is again segregated into web-based and mobile-based based on the application segment. Regional Analysis According to the global market analysis report, the regional bifurcation of the online sports betting market includes North America, Europe, Asia-Pacific, and the rest of the world. Moreover, the market study also states that Europe has dominated the online sports betting market in the preview years and is expected to grow at a CAGR of 12.1%. The driving factor would be the increasing zeal of watching sports with the advancement of innovative game formats and gaming modes. However, during the forecast period, the North American market is predicted to grow at the fastest rate of 15.8%. Key Players Notable players in the worldwide online sports betting market include 888 Holdings PLC (UK), Flutter Entertainment PLC (UK), Bet365 Group Ltd (UK), Fortuna Entertainment Group (Netherlands), Churchill Downs Incorporated (US), Entain PLC (UK), DraftKings (US), Betsson AB (Sweden), Betfred Ltd (UK), Webis Holdings PLC (Isle of Man), Kindred Group PLC (Malta), Sportech PLC (UK), BetAmerica (US), HKJC Football Betting Limited (Hong Kong), and Megapari (Cyprus). Industry Updates In April 2019, Betfred made the debut of its American Sports betting platform at Betting on Sports America 2019 at Meadowlands, New Jersey. In January 2022, Inspired, one of the leading B2B gaming content providers, announced to have acquired Sportech Lotteries, Inc., a subsidiary of Sportech PLC. Browse Full Report Details @ https //www.marketresearchfuture.com/reports/online-gambling-market-8641 Table of Contents 1 Executive Summary 2 Scope of The Report 2.1 Market Definition 2.2 Scope of The Study 2.2.1 Research Objectives 2.2.2 Assumptions Limitations 2.3 Market Structure Continued… Similar Report** Open Source Intelligence (OSINT) Market By Security Type (Human Intelligence, Content Intelligence, Dark Web Analysis, Link/Network Analysis, Data Analytics, Text Analytics, Artificial Intelligence, Big Data, Others), Technology (Bid Data Software, Video Analytics, Text Analytics, Visualization Tool, Cyber Security, Web Analysis, Social Media Analysis, Others), Application (Military Defense, Homeland Security, Private Sector, Public Sector, National Security, Others) Voice Assistant Market** https //mrfrinformation.tistory.com/615 Network-Attached Storage Market** https //telescope.ac/information-technology-Lc1XMRDqL/5WlKY9Kx4 About Market Research Future At Market Research Future (MRFR), we enable our customers to unravel the complexity of various industries through our Cooked Research Report (CRR), Half-Cooked Research Reports (HCRR), Raw Research Reports (3R), Continuous-Feed Research (CFR), and Market Research Consulting Services. Contact Market Research Future (Part of Wantstats Research and Media Private Limited) 99 Hudson Street, 5Th Floor New York, NY 10013 United States of America 1 628 258 0071 (US) 44 2035 002 764 (UK) Email sales@marketresearchfuture.com Website https //www.marketresearchfuture.com
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JDorama.com http //www.jdorama.com/ ドメイン管理者の国はシンガポール Ｑ：誰がウェブサイトを所有しているか？ Ａ：シンガポールの日本のドラマ愛好家。 Kegareta Shita [汚れた舌] 部分転載：http //www.jdorama.com/drama.936.htm "Naoko s Filthy Tongue has fans licking lips" Yu Yamada gets leg over actor Shun Oguri to get leg up in career http //www.jdorama.com/viewtopic.php?p=766917#766917 The Nakamura-Takeuchi split she gets the kid, alimony and a new career, he gets hosed 部分転載：http //www.jdorama.com/viewtopic.php?p=748428#748428 Sexy thespian Tomoko Yamaguchi uses booby trap to snare audience for TV show http //www.jdorama.com/viewtopic.php?p=727787#727787 Offbeat panty bar offers uninhibited lust with the improper stranger http //www.jdorama.com/viewtopic.php?p=695445#695445 How low can they go? 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Pseudo pedo-porn peddlers loll in lascivious limbo Latest Akihabara geek fetish? One-eyed virginal maid mummies Offbeat 'panty bar' offers uninhibited lust with the improper stranger One, Two, Poo in Your Shoe; Three, Four, Wee on the Floor Pesky perverts pry on otaku's dolled-up maids Sex workers on the ball with World Cup fever Stressed teachers indulging their wild sides with coworkers, parents -- and Top baseball commentator the latest to strike out in NTVs string of sex scandals WaiWaiの記事を転載した英語サイト：J 毎日新聞謝罪記事の問題点
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Spectacular Spider-Man The Hunger (Spectacular Spider-Man) Marvel Enterprises?Paul Jenkins?Humberto Ramos?John Miesegaes? The Punisher (Punisher) Marvel Enterprises?Garth Ennis?Steve Dillon?Darick Robertson? Peter Parker, Spider-Man Senseless Violence (Peter Parker, Spider-Man) Marvel Enterprises?Zeb Wells? Thor Vikings (Max) Marvel Enterprises?Garth Ennis? Marvel Masterworks Presents The Fantastic Four 1 (Marvel Masterworks (Numbered)) Marvel Enterprises?Stan Lee?Jack Kirby? Stan Lee Meets Marvel Enterprises?Stan Lee?Brian Michael Bendis?Joss Whedon?Jeph Loeb?Roy Thomas? Thor Blood Oath (Thor (Graphic Novels)) Marvel Enterprises?Michael Avon Oeming?Scott Kolins?Tom Brevoort? Fantastic Four 2 (Fantastic Four (Graphic Novels)) Marvel Enterprises?J. Michael Straczynski? Marvel Masterworks The Fantastic Four Nos. 21 - 30 (Marvel Masterworks (Numbered)) Marvel Enterprises?Stan Lee?Jack Kirby? Nextwave Agents of H.a.t.e. 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Vampire Survivors 項目数：211(140+2+17+3+21+1+4+1+1+6+8+7) 総ポイント：1575(1200+15+95+20+105+5+20+5+5+30+40+35) 難易度：★★☆☆☆ 配信日：2022/10/20 ストア：https //www.xbox.com/ja-jp/games/store/vampire-survivors/9pd5bm2z8c4l GamePass対応 実績表記は英語だがゲーム自体は日本語にも対応。ただ実績内容には誤りが多い ゲーム内「アンロック」の日本語で記載されている条件を満たすと解除される キャラやステージの指定が多いもののキャラ毎の武器、アイテムの特徴をある程度把握できれば未プレイでも十分コンプ可能 コンプまで4～50時間前後か 話題になった作品のため、参考になる攻略情報や攻略動画は少し探すと簡単に見つかる Vampire Survivors 攻略Wiki https //gamerch.com/vampiresurvivors/ + DLC未購入の場合に取れる実績とスコア 実績数 171/211 ゲーマースコア 1375/1575 Wings Reach Level 5.レベル5に到達する 5 Crown Reach Level 10.レベル10に到達する 5 Hollow Heart Survive 1 minute with any character.いずれかのキャラクターで1分間生き残る 5 Runetracer Survive 5 minutes with Pasqualina.パスカリーナで5分間生き残る 5 Peachone Survive 10 minutes with any character.いずれかのキャラクターで10分間生き残る 5 Arca Get Fire Wand to Level 4.炎の杖をレベル4に上げる 5 Bracer Get King Bible To Level 4.王の聖書をレベル4に上げる 5 Candelabrador Get Santa Water To Level 4.聖水をレベル4に上げる 5 Porta Get Lightning Ring To Level 4.雷の指輪をレベル4に上げる 5 Duplicator Get Magic Wand To Level 7.魔法の杖をレベル7に上げる 5 Ebony WIngs Get Peachone To Level 7.純白の翼をレベル7に上げる 5 Spellbinder Get Runetracer To Level 7.軌跡の魔弾をレベル7に上げる 5 Empty Tome Hold 6 different weapons at once.6つの武器を一度に持つ 5 Fire Wand Destroy 20 light sources.光源を20個破壊する 5 Garlic Find 5 Floor Chickens.トリ肉を5つ見つける 5 Clover Find a Little Clover.小さなクローバーを見つける 5 Magnet Find a Vacuum.バキュームを見つける 5 Clock Lancet Find an Orologion.懐中時計を見つける 5 Cross Find a Rosary.ロザリオを見つける 5 Lightning Ring Defeat a total of 5000 enemies.敵を合計で5,000体倒す 5 Mortaccio Defeat a total of 3000 Skeletons.スケルトンを3,000体倒す 10 Pentagram Survive 20 minutes with any character.いずれかのキャラクターで20分間生き残る 5 Hyper Mad Forest Defeat the giant Blue Venus in the Mad Forest.狂乱の森で巨大なブルー・ヴィーナスを倒す 20 Hyper Inlaid Library Defeat the Hag in the Inlaid Library.象眼の図書館でハグを倒す 20 Inlaid Library Reach Level 20 in the Mad Forest.狂乱の森でレベル20に到達する 5 Pummarola Survive 5 minutes with Gennaro.ジェンナーロで5分間生き残る 5 Stone Mask Find a Stone Mask.石の仮面を見つける 10 Bloody Tear Evolve the Whip.ムチを進化させる 5 Holy Wand Evolve the Magic Wand.魔法の杖を進化させる 5 Thousand Edge Evolve the Knife.ナイフを進化させる 5 Death Spiral Evolve the Axe.オノを進化させる 5 Heaven Sword Evolve the Cross.十字架を進化させる 5 Unholy Vespers Evolve the King Bible.王の聖書を進化させる 5 Hellfire Evolve the Fire Wand.炎の杖を進化させる 5 Poe Ratcho Get Garlic to level 7.ニンニクをレベル7に上げる 5 Soul Eater Evolve the Garlic.ニンニクを進化させる 5 Vandalier Unite Ebony Wings and Peachone.漆黒の翼 純白の翼を合体させる 5 Green Acres Unlock Hyper mode for 2 normal stages.ハイパーモードを2ステージ分アンロックする 5 Suor Clerici Recover a total of 1000 HP.HPを合計1,000回復する 5 Dommario Earn 5000 coins in a single run.1回のプレイで5,000枚のコインを獲得する 10 Krochi Defeat a total of 100000 enemies.敵を合計で100,000体倒す 10 Tiragisú Survive 20 minutes with Krochi.クロチで20分間生存する 5 La Borra Evolve the Santa Water.聖水を進化させる 5 Thunder Loop Evolve the Lightning Ring.雷の指輪を進化させる 5 Reroll 1 Reach Level 80 with Mortaccio.モルタッチオでレベル80になる 5 Skip 2 Survive 30 minutes in Green Acres.緑の地で30分間生存する 10 Reroll 2 Reach Level 80 with Yatta Cavallo.ヤッタ・カヴァロでレベル80に到達する 5 Yatta Cavallo Defeat a total of 3000 Lion Heads.ライオンヘッドを合計で3,000体倒す 10 Lama Survive 20 minutes with at least +10% Curse.呪い+10%以上で20分間生存する 5 Skull O'Maniac Survive 30 minutes with Lama.ラマで30分間生存する 5 Dairy Plant Reach Level 40 in the Inlaid Library.象眼の図書館でレベル40に到達する 5 Hyper Dairy Plant Defeat the Sword Guardian in the Dairy Plant.酪農場にいるソードガーディアンを倒す 20 Milky Way Map Find the Milky Way Map.天の川の地図を見つける 5 Coffin Dairy Plant Find and open the coffin in the Dairy Plant.酪農場にある棺桶を探して開ける 20 Song Of Mana Survive 15 minutes with Poppea.ポピアで15分間生き残る 5 Mannajja Evolve the Song Of Mana.マナの歌を進化させる 5 Christine Get Pentagram to Level 7.五芒星の魔法陣をレベル7に上げる 5 Il Molise Unlock Hyper mode for any normal stage.任意のステージのハイパーモードをアンロックする 5 Skip 1 Survive 15 minutes in Molise.地図にない場所で15分間生存する 5 Gorgeous Moon Evolve the Pentagram.五芒星の魔法陣を進化させる 5 Mindbender Fill 50 entries in the Collection.コレクションを50項目満たす 5 Coffin Mad Forest Find and open the coffin in the Mad Forest.狂乱の森にある棺桶を探して開ける 20 Phiera Der Tuphello Survive 10 minutes with Pugnala.プニャーラで10分間生き残る 5 Eight The Sparrow Survive 15 minutes with Pugnala.プニャーラで15分間生き残る 5 Phieraggi Evolve and unite Phiera Der Tuphello and Eight The Sparrow.拳銃『フィエラ』と拳銃『エイト』を進化、合体させる 5 NO FUTURE Evolve the Runetracer.軌跡の魔弾を進化させる 5 Reroll 3 Reach Level 80 with Bianca Ramba.ビアンカ ランバでレベル80に到達する 5 Banish 1 Fill 60 entries in the Collection.コレクションを60項目満たす 5 Hyper Gallo Tower Defeat the Giant Enemy Crab in Gallo Tower.ガロの塔で巨大なエネミー・クラブを倒す 20 Sorceress' Tears Find the Sorceress' Tears.魔女の涙を見つける 10 Gallo Tower Reach Level 60 in Dairy Plant.酪農場でレベル60に到達する 5 Bianca Ramba Defeat a total of 3000 Milk Elementals.ミルクエレメンタルを合計で3,000体倒す 10 Banish 2 Fill 70 entries in the Collection.コレクションを70項目満たす 5 Randomazzo Find the Randomazzo.Randomazzoを見つける 10 Gatti Amari Survive 15 minutes with Giovanna.ジオヴァーナで15分間生き残る 5 Coffin Inlaid Library Find and open the coffin in the Inlaid Library.象眼の図書館にある棺桶を探して開ける 20 V - Chaos in the Dark Night Reach Level 50 with Giovanna.ジアヴァーナでレベル50に到達する 5 IV - Awake Reach Level 50 with Krochi.クロチでレベル50に到達する 5 VI - Sarabande of Healing Find the Randomazzo.Randomazzoを見つける 5 XVI - Slash Reach Level 50 with Lama.ラマでレベル50に到達する 5 XVII - Lost And Found Painting Reach Level 50 with Poppea.ポピアでレベル50に到達する 5 XIX - Heart of Fire Reach Level 50 with Arca.アルカでレベル50に到達する 5 O'Sole Meeo Defeat a total of 3000 Dragon Shrimps.ドラゴンシュリンプを合計で3,000体倒す 10 The Bone Zone Unlock Hyper mode for 3 normal stages.ハイパーモードを3ステージ分アンロックする 5 Skip 3 Survive 30 minutes in The Bone Zone.遺骸地帯で30分間生存する 10 Reroll 4 Reach Level 80 with O'Sole Meeo.オ・ソーレ ミーオでレベル80に到達する 5 XI - Waltz of Pearls Reach Level 50 with Imelda.イメルダでレベル50に到達する 5 VII - Iron Blue Will Reach Level 50 with Gennaro.ジェンナーロでレベル50に到達する 5 Coffin Gallo Tower Find and open the coffin in the Gallo Tower.ガロの塔にある棺桶を探して開ける 20 XVIII - Boogaloo of Illusions Reach Level 50 with Concetta.コンチェッタでレベル50に到達する 5 Vicious Hunger Evolve the Gatti Amari.獰猛な猫を進化させる 5 Valkyrie Turner Evolve the Shadow Pinion.黒色のドリルを進化させる 5 XV - Disco of Gold Reach minute 31 in the Inlaid Library.象眼の図書館で31分生存する 5 Banish 3 Fill 80 entries in the Collection.コレクションを80項目満たす 5 Magic Banger Find the Magic Banger.魔法の楽器を見つける 5 Shadow Pinion Survive 15 minutes with Concetta.コンチェッタで15分間生き残る 5 Moongolow Unlock Hyper mode for 4 normal stages.ハイパーモードを4ステージ分アンロックする 5 Skip 4 Survive 15 minutes in Moongolow.骸骨地帯で30分間生存する 5 Glass Vizard Find and buy the Glass Vizard.Glass Vizardを見つけて買う 5 XIV - Jail of Crystal Reach Level 50 with Pasqualina.パスカリーナでレベル50に到達する 5 XII - Out of Bounds Reach minute 31 in the Gallo Tower.ガロの塔で31分生存する 5 Banish 4 Fill 90 entries in the Collection.コレクションを90項目満たす 5 Yellow Sign Find the Yellow Sign.黄色い看板を見つける 50 Seeker of the Infinite Corridor Obtain the Infinite Corridor.無限回廊を入手する 10 Seeker of the Crimson Shroud Obtain the Crimson Shroud.深紅の聖骸衣を入手する 10 X - Beginning Reach Level 50 with Antonio.アントニオでレベル50に到達する 5 VIII - Mad Groove Reach minute 31 in the Mad Forest.狂乱の森で31分生存する 5 Torrona's Box Hold 6 different weapon evolutions at once.1回で6つの異なる武器を進化させる 10 Omni Get Torrona's Box to Level 9.トローナの箱をレベル9に上げる 10 Cappella Magna Reach Level 80 in the Gallo Tower.ガロの塔でレベル80に到達する 5 Coffin Cappella Magna Find and open the coffin in the Cappella Magna.マグナ礼拝堂にある棺桶を探して開ける 20 Vento Sacro Survive 15 minutes with Zi'Assunta.ジ・アスンタで15分間生き残る 5 Fuwalafuwaloo Unite Vento Sacro and Bloody Tear.聖なる風と血染めの鞭を合体させる 5 Hyper Cappella Magna Defeat the Trinacria in the Cappella Magna.マグナ礼拝堂でトリナクリアを倒す 20 Sir Ambrojoe Defeat a total of 6000 Stage Killers.ステージ・キラーを合計で6,000体倒す 10 III - Tragic Princess Reach Level 50 with Porta.ポルタでレベル50に到達する 5 XX - Silent Old Sanctuary Reach minute 31 in the Dairy Plant.酪農場で31分生存する 5 Banish 5 Fill 100 entries in the Collection.コレクションを100項目満たす 5 Reroll 5 Reach Level 80 with Sir Ambrojoe.サー アンブロジョーでレベル80に到達する 5 Grim Grimoire Find the Grim Grimoire.Grim Grimoireを見つける 5 Ars Gouda Find the Ars Gouda.Ars Goudaを見つける 5 Great Gospel Find the Great Gospel.大いなる福音を見つける 20 Game Killer Defeat the final enemy in the Cappella Magna.マグナ礼拝堂で最後の敵を倒す 10 Boss Rash Unlock Hyper mode for all 5 normal stages.ハイパーモードを5ステージ分アンロックする 5 Skip 5 Survive 15 minutes in Boss Rash.ボスラッシュで15分間生存する 10 II - Twilight Requiem Reach Level 50 with Dommario.ドンマリオでレベル50に到達する 5 I - Gemini Reach Level 50 with Pugnala.プニャーラでレベル50に到達する 5 Forbidden Scrolls Find the Forbidden Scrolls of Morbane.モルベインの禁じられた巻物を見つける 50 XIII - Wicked Season Reach Level 50 with Christine.クリスティーンでレベル50に到達する 5 Queen Sigma Complete the Collection.コレクションをコンプリートする 100 Victory Sword Defeat 100000 enemies in a single run with Queen Sigma.シグマを使用して1回の戦闘で敵を100,000体倒す 10 IX - Divine Bloodline Reach Level 50 with Suor Clerici.クレーリチでレベル50に到達する 5 XXI - Blood Astronomia Reach Level 50 with Poe.ポーでレベル50に到達する 5 Candybox Discover every standard evolution and union.すべての一般の進化と融合を発見する 5 Bracelet Survive 30 minutes with either Gallo or Divano.ガロかディヴァーノのどちらかで30分間生き残る 5 Tri-Bracelet Evolve the Bracelet and then the Bi-Bracelet.ブレスレットを進化させ、さらにバイブレスレットを進化させる 5 The Eudaimonia Machine Obtain all standard relics from all stages.全ステージですべての一般レリックを入手する 5 Seventh Trumpet Obtain the Seventh Trumpet.第七のラッパ吹きを入手する 10 Gracia's Mirror Obtain Gracia's Mirror.グラシアの鏡を入手する 10 Greatest Jubilee See the final fireworks.最後の花火を見る 50 Tiny Update(Update) 15G EXTRA Seal Banish 10 or more weapons in a single run.1回の戦闘で10個以上の武器を消去する 5 Tiny Bridge Reach Level 80 in Inverse Gallo Tower.反転のガロの塔でレベル80に到達する 10 Legacy of the Moonspell(DLC) 95G Miang Find and open the coffin in Mt.Moonspell.ムーンスペル山にある棺桶を探して開ける 5 Silver Wind Survive 15 minutes with Miang Moonspell.ミアン・ムーンスペルで15分間生き残る 5 Menya Evolve the Silver Wind.銀の風を進化させる 5 Four Seasons Survive 15 minutes with Menya Moonspell.メンヤ・ムーンスペルで15分間生き残る 5 Syuuto Evolve the Four Seasons.春夏秋冬を進化させる 5 Summon Night Survive 15 minutes with Syuuto Moonspell.シュウト・ムーンスペルで15分間生き残る 5 Babi-Onna Evolve the Summon Night.夜の召喚を進化させる 5 Mirage Robe Survive 15 minutes with Babi-Onna.バビ・オーナで15分間生き残る 5 McCoy-Oni Evolve the Mirage Robe.幻惑の着物を進化させる 5 108 Bocce Survive 15 minutes with McCoy-Oni.真鬼で15分間生き残る 5 Megalo Menya Defeat 100000 enemies in a single run with Menya Moonspell.メンヤ・ムーンスペルを使用して1回の戦闘で敵を100,000体倒す 10 Megalo Syuuto Defeat 100000 enemies in a sinle run with Syuuto Moonspell.シュウト・ムーンスペルを使用して1回の戦闘で敵を100,000体倒す 10 Gav'Et-Oni Defeat 6000 Kappa.6,000匹の河童を倒す 5 Night Sword Find a Night Sword.夜の剣を探す 5 Muramasa Evolve the Night Sword.夜の剣を進化させる 5 Hyper Mt.Moonspell Defeat the Orochimario in Mt.Moonspell.ムーンスペル山でオロチマリオを倒す 5 Boo Roo Boolle Evolve the Mille Bolle Blu.藍の泡を進化させる 5 The Chaotic One(Update) 20G EXTRA Bat Country Reach Level 80 in Inverse Mad Forest.反転の狂乱の森でレベル80に到達する 10 EXTRA Apoplexy Find the Apoplexy at minute 09 00 in Bat Country.バットカントリーで09 00分に脳卒中を見つける 5 EXTRA Chaos Malachite Find the Chaos Malachite at minute 18 00 in Bat Country.バットカントリーで18 00分にカオスマラカイトを見つける 5 Tides of the Foscari(DLC) 105G Eleanor Uziron Find and open the coffin in Lake Foscari.フォスカリ湖にある棺桶を探して開ける 5 SpellString Survive 15 minutes with Eleanor Uziron.エレノール・ウジロンで15分生き残る 5 SpellStream Get SpellStream to Level 6.スペルストリームをレベル6に上げる 5 SpellStrike Get SpellStrike to Level 6.スペルストライクをレベル6に上げる 5 Maruto Cuts Unite SpellString, SpellStream, and SpellStrike.スペルストリング、スペルストリーム、スペルストライクを合体させる 5 Eskizzibur Survive 15 minutes with Maruto Cuts.マルト・カッツで15分生き残る 5 Keitha Muort Evolve the Eskizzibur.エスキッジブールを進化させる 5 Flash Arrow Survive 15 minutes with Keitha Muort.キーサ・ムオルトで15分生き残る 5 Millionaire Evolve the Flash Arrow.閃光の矢を進化させる 5 Abyss Foscari With Keitha, break the Seal of the Lake.キーサで湖の封印を解く 5 Luminaire Foscari With Maruto, break the Seal of the Abyss.マルトでアビスの封印を解く 5 Genevieve Gruyère With Eleanor, break the Seal of the Banished.エレノールで追放者の封印を解く 5 Prismatic Missile Survive 15 minutes with Luminaire Foscari.ルミネール・フォスカリで15分生き残る 5 Luminaire Evolve the Prismatic Missile.プリズムミサイルを進化させる 5 Shadow Servant Survive 15 minutes with Genevieve Gruyère.ジュヌヴィーヴ・グリュイエールで15分生き残る 5 Ophion Evolve the Shadow Servant.シャドー・サーヴァントを進化させる 5 Je-Ne-Viv Defeat 100000 enemies in a single run with Genevieve Gruyère.ジュヌヴィーヴ・グリュイエールで死なずに敵を100,000体倒す 5 Happy Birthday Defeat a total of 6000 Sammies.サミーを合計で6,000体倒す 5 Rottin'Ghoul Defeat a total of 6000 Rotting Ghouls.ロッティン・グールを合計で6,000体倒す 5 Hyper Lake Foscari Defeat the Avatar of Gaea in Lake Foscari.フォスカリ湖でガイア・アバターを倒す 5 Hyper Abyss Foscari Defeat Je-Ne-Viv in Abyss Foscari.アビス・フォスカリでジュ・ヌ・ヴィーヴを倒す 5 The Whimsy One(Update) 5G EXTRA Seal II Banish 20 or more weapons in a single run.1回の戦闘で20個以上の武器を消去する 5 Overwhelming Update(Update) 20G EXTRA Astral Stair Reach Level 80 in Inverse Inlaid Library.反転の象眼の図書館でレベル80に到達する 5 EXTRA Chaos Rosalia Find the Chaos Rosalia in Astral Stair.アストラル階段でカオス・ロザリアを見つける 5 EXTRA Trisection Find the Trisection in Astral Stair.アストラル階段でトライセクションを見つける 5 EXTRA Astral Stair Map Find the Astral Stair Map.アストラル階段の地図を見つける 5 Mt.Moonspell Map(DLC) 5G Mt.Moonspell Map Find the Mt.Moonspell Map.ムーンスペル山の地図を見つける 5 Lake Foscari Map(DLC) 5G Lake Foscari Map Find the Lake Foscari Map.フォスカリ湖の地図を見つける 5 Whiteout(Update) 30G EXTRA Whiteout Find 20 Orologions.懐中時計を20個見つける 5 EXTRA Glass Fandango Get Glass Fandango to Level 7.グラス・ファンダンゴをレベル7に上げる 5 EXTRA Celestial Voulge Evolve the Glass Fandango.グラス・ファンダンゴを進化させる 5 EXTRA She-Moon Eeta Survive 20 minutes in Whiteout.ホワイトアウトで20分間生き残る 5 EXTRA Antidote Find the Antidote in Whiteout.ホワイトアウトで解毒薬を見つける 5 EXTRA Chaos Altemanna Find the Chaos Altemanna in Tiny Bridge.小さな橋でカオス・アルテマンナを見つける 5 Space 54(Update) 40G EXTRA Adventures Obtain the Atlas Gate in Boss Rash.ボスラッシュでアトラスゲートを見つける 2 EXTRA Space 54 Find 5 Golden Fingers.黄金の指を5本見つける 4 EXTRA Phas3r Get Phas3r to level 7.フェイザーをレベル7に上げる 6 EXTRA Photonstorm Evolve the Phas3r.フェイザーを進化させる 8 EXTRA Space Dude Survive 20 minutes in Space 54.宇宙54で20分間生き残る 10 EXTRA Brave Story Find the Brave Story at minute 18 in Space 54.宇宙54の18分に勇敢な物語を見つける 15 EXTRA Pako Battiliar Defeat a total of 161616 bats.コウモリを合計161,616匹倒す 1 EXTRA Bat Robbert Evolve the Pako Battiliar.パコ・バッティリアを進化させる 2 Laborratory(Update) 35G EXTRA Seal III Banish 40 or more weapons in a single run.1回の戦闘で40個以上の武器を消去する 2 EXTRA Laborratory Find 33 Rosaries.ロザリオを33個見つける 4 EXTRA Santa Javelin Get Santa Javelin to level 7.聖ジャヴェリンをレベル7に上げる 6 EXTRA Seraphic Cry Evolve the Santa Javelin.聖ジャヴェリンを進化させる 8 EXTRA Santa Ladonna Survive 20 minutes in Laborratory.ラボラトリーで20分間生き残る 10 EXTRA Arma Dio Find an Arma Dio in Laborratory.ラボラトリーでアルマ・ジロを見つける 15 EXTRA Carlo Cart Deal a total of 25120 damage with the Laborratory Train.ラボラトリーのトレインで合計25,120ダメージを与える 1 ※全体的な流れ ゲームシステムは1プレイ最大30分程度を繰り返す形 基本的には色々なキャラを使い、武器レベルを上げながら生存し続ければ何かしらはされていく どうしてもクリア出来ないならば、シークレットから「クイーン シグマ」 DLCを所有している場合は「メガロ メイヤ」を先にアンロックすると多少楽になる ■レベル50に到達する関連、31分生存する関連、Game Killer ガロの塔でRandomazzoのロックを解除した後に達成することで解除される。 ■キャラクター解放関連 キャラの開放が実績の解除に影響していないキャラもいる。 上記Wikiのキャラクター一覧だと「Gyorunton」から「Red Death」までのキャラが該当。 中には強力なキャラもいるが、癖の強いものが多いので実績目的なら後回しでも問題ない。 ■シークレットコード 「Forbidden Scrolls of Morbane」入手後か、「コレクション」の背景が薄紫になっているアイテム欄の7～8番目を 一定回数クリックすることで「シークレット」が解放される。 ここで特定のコードを入力するとキャラやステージ等が解放されるが、実績は解除されない点に注意。 (シークレットによる解放後でも条件を満たせば実績を解除可能) シークレット一覧 https //gamerch.com/vampiresurvivors/entry/380604 ■Gav'Et-Oni 河童はステージ「ムーンスペル山」の水のある辺りで出現する傾向にある。 ステージ「ムーンスペル山」開始地点から西へ進んでから北の方にある坂道を上っていくと、マップ北西端辺りにある凍った湖のような場所につく。 この凍った湖では河童が大量にポップするので条件を達成しやすい。 ■EXTRA Chaos Malachite バットカンパニーで18分になるとカオスマラカイトを示す緑の矢印が表示される。 19分までに取りに行かないと消えてしまうため注意。 ■Abyss Foscari キーサでフォスカリ湖をスタートすると地図に？マークが表示される。 ？マークの場所に行くと封印があり、封印を解くにはキーサの武器を進化させる必要があるため、進化させてから行くとよい。 (閃光の矢+腕甲+クローバー) 封印を解いた後はアビス・フォスカリに移動するが、すぐに終了してOK。 ■Luminaire Foscari マルトでアビス・フォスカリをスタートすると？マークが表示される。 ？マークの場所に行くと封印があり、封印を解くにはマルトの武器を進化させる必要があるため、進化させてから行くとよい。 (エスキッジブール+鎧) ■Genevieve Gruyère、Hyper Abyss Foscari ※Luminaire Foscariを解除しておく必要あり。 エレノールでアビス・フォスカリをスタートすると？マークが表示される。 ？マークの場所に行くと封印があり、封印を解くにはエレノールの武器を進化させる必要があるため、進化させてから行くとよい。 (スペルストリング+スペルストリーム+スペルストライク) また、道中にある王冠を拾いレベルMAXまで上げておく。 封印を解くとしばらく経ったのち戦闘が始まるが、武器が全てなくなる。 床に武器が落ちているのでそれらを拾いつつボスの体に触れないよう立ち回る。 プリズムミサイルが出始めたら進化できるまで拾うと宝箱が出てくるので進化させる。 そうするとルミネール・フォスカリが登場し、しばらく攻撃していると倒すことができる。 なお一度封印を解けば次回以降は最初から出現しており武器の没収もされないため、上記の方法で倒せないなら強いキャラで挑むのもアリ。 ■Happy Birthday https //www.youtube.com/watch?v=9XgXJZ-qIm8 ■Hyper Lake Foscari フォスカリ湖で25分経つと出現する。 出現場所近くだと回復してしまい倒せない。 少し離れたところにいくと回復しなくなるため、その間に倒す。 ■EXTRA Astral Stair、EXTRA Chaos Rosalia、EXTRA Trisection、EXTRA Astral Stair Map、Mt.Moonspell Map、Lake Foscari Map https //kakinblog.com/vampire-survivors1-5-0/ ■Whiteoutでの追加分 https //kakinblog.com/vampire-survivors1-7-0/ ■Space 54での追加分 https //kakinblog.com/vampire-survivors-p190/ ■Laborratoryでの追加分 https //kakinblog.com/vampire-survivors-p110/
https://w.atwiki.jp/tiger/pages/4.html
TEC-/- BTK-/- double mutant T cells exhibit severely impaired T cell activitity. RLK-/-ITK-/- double mutant celles exhibit severely imparired Th2 responses. Grb2(+/-) mice disrupt T cell signaling networks and development. Dendric cells and macrophages of MEK3 deficient mice have impaired IL12 production. Bam32(-/-) B cell develop normally but have impaired T-independent antibody responses in vivo. T-cell and B-cell of RAP1A deficient mice impair integrin-mediated cell adhesion. T-cell of WASP deficient mice impair the proliferaction and antigen receptor cap formation in response to anti-CD3zeta stimulation. T-cell of SHB defective mice impair the phosphorylation of LAT and consequently the activation of MAP kinase pathways. B-cell of 3BP2 (-/-) deficient mice have defective in proliferation, cell cycle progression, PLC-gamma2 phosphorylation, calcium mobilization, NF-ATp dephosphorylation, and Erk and Jnk activation in response to BCR ligation. B-cell of Vav2(-/-) deficient mice are defective in the ability to switch immunoglobulin class. T-cell of Vav1(-/-) deficient mice exhibit impaired antigen receptor signaling. Vav1(-/-)Vav2(-/-) mice exhibit greatly reduced the mature B-cells. Vav-1-/-Vav-2-/- B cells were unresponsive to BCR-driven proliferation in vitro and to thymus-indepen-dent antigen in vivo. Fyn-deficient mice exhibit a remarkably specific lymphoid defect thymocytes are refractile to stimulation through the TCR with mitogen or antigen. Lck-deficient mice show a pronounced thymic atrophy, with a dramatic reduction in the double-positive (CD4+CD8+) thymocyte population. T cell from mice deficient in LCK is required for normal signal transduction through the TCR. T cells from mice deficient in SLAP-130/Fyb show markedly impaired proliferation. B cell of chicken deficient ITK reduce IP3 generation and phospholipase C gamma 2 tyrosine phosphorylation. T cell of mice deficient ITK reduce IP3 generation and phospholipase C gamma 1 tyrosine phosphorylation. T cell of mice deficient ITK have failure of Th2 development. Mice deficient in ITK have reduced proliferative responses to MHC stimulation and to anti-TCR cross-linking Mutations in Btk cause X-linked immunodeficiency. Gads(GRAP2) has a role in thymocyte proliferaction for maturation of T-cells. Gads(GRAP2) has a role for homeostatic proliferaction in B cells. Grap negatively regulates T-cell proliferation. Gab2 is a substrate of ZAP-70 and functions as a switch molecule toward inhibition of TCR signal transuduction. B cell signaling causes tyrosine phosphorylation of Gab1, and in turn SHP2 bind to Gab1 Gab1 phosophorylation potentiate the phosphorylation of Akt, PI3K-dependent response. RasGRP1 mediates Ras activation following TCR stimulatioin. RasGRP1 and RasGRP3 induces RAS activation in B-cell to response to T-cell stimulation. Grb2-hSos1-PLCgamma1-p36/p38-ZAP70 complexes localize in the vicinity of TCR-zeta Gads(Grap2) plays an important role in T-cell signaling via its association with SLP-76 and LAT. Lck is required for normal signal transduction through the TCR. ZAP-70 plays crucial roles in T-cell activation and development. Syk triggers cellular activation in T-cell. TEC-/- BTK-/- double mutant T cells exhibit severely impaired T cell activitity. 1 J Exp Med. 2000 Dec 4;192(11) 1611-24. Severe B cell deficiency in mice lacking the tec kinase family members Tec and Btk. Ellmeier W, Jung S, Sunshine MJ, Hatam F, Xu Y, Baltimore D, Mano H, Littman DR. Molecular Pathogenesis Program, Skirball Institute of Biomolecular Medicine. wilfried.ellmeier@univie.ac.at The cytoplasmic protein tyrosine kinase Tec has been proposed to have important functions in hematopoiesis and lymphocyte signal transduction. Here we show that Tec-deficient mice developed normally and had no major phenotypic alterations of the immune system. To reveal potential compensatory roles of other Tec kinases such as Bruton s tyrosine kinase (Btk), Tec/Btk double-deficient mice were generated. These mice exhibited a block at the B220(+)CD43(+) stage of B cell development and displayed a severe reduction of peripheral B cell numbers, particularly immunoglobulin (Ig)M(lo)IgD(hi) B cells. Although Tec/Btk(null) mice were able to form germinal centers, the response to T cell-dependent antigens was impaired. Thus, Tec and Btk together have an important role both during B cell development and in the generation and/or function of the peripheral B cell pool. The ability of Tec to compensate for Btk may also explain phenotypic differences in X-linked immunodeficiency (xid) mice compared with human X-linked agammaglobulinemia (XLA) patients. Publication Types Research Support, Non-U.S. Gov t PMID 11104803 [PubMed - indexed for MEDLINE] RLK-/-ITK-/- double mutant celles exhibit severely imparired Th2 responses. 1 Nat Immunol. 2001 Dec;2(12) 1183-8. Mutation of Tec family kinases alters T helper cell differentiation. Schaeffer EM, Yap GS, Lewis CM, Czar MJ, McVicar DW, Cheever AW, Sher A, Schwartzberg PL. National Human Genome Research Institute, National Institutes of Health, Bethesda, MD 20892, USA. The Tec kinases Rlk and Itk are critical for full T cell receptor (TCR)-induced activation of phospholipase C-gamma and mitogen-activated protein kinase. We show here that the mutation of Rlk and Itk impaired activation of the transcription factors NFAT and AP-1 and production of both T helper type 1 (TH1) and TH2 cytokines. Consistent with these biochemical defects, Itk-/- mice did not generate effective TH2 responses when challenged with Schistosoma mansoni eggs. Paradoxically, the more severely impaired Rlk-/-Itk-/- mice were able to mount a TH2 response and produced TH2 cytokines in response to this challenge. In addition, Rlk-/-Itk-/- cells showed impaired TCR-induced repression of the TH2-inducing transcription factor GATA-3, suggesting a potential mechanism for TH2 development in these hyporesponsive cells. Thus, mutations that affect Tec kinases lead to complex alterations in CD4+ TH cell differentiation. Publication Types Research Support, Non-U.S. Gov t Research Support, U.S. Gov t, P.H.S. PMID 11702066 [PubMed - indexed for MEDLINE] Grb2(+/-) mice disrupt T cell signaling networks and development. 1 Nat Immunol. 2001 Jan;2(1) 29-36. Disruption of T cell signaling networks and development by Grb2 haploid insufficiency. Gong Q, Cheng AM, Akk AM, Alberola-Ila J, Gong G, Pawson T, Chan AC. Howard Hughes Medical Institute, Washington University School of Medicine, St. Louis, MO 63110, USA. The developmental processes of positive and negative selection in the thymus shape the T cell antigen receptor (TCR) repertoire and require the integration of multiple signaling networks. These networks involve the efficient assembly of macromolecular complexes and are mediated by multimodular adaptor proteins that permit the functional integration of distinct signaling molecules. We show here that decreased expression of the adaptor protein Grb2 in Grb2+/- mice weakens TCR-induced c-Jun N-terminal kinase (JNK) and p38, but not extracellular signal-regulated kinase (ERK), activation. In turn, this selective effect decreases the ability of thymocytes to undergo negative, but not positive, selection. We also show that there are differences in the signaling thresholds of the three mitogen-activated protein kinase (MAPK) families. These differences may provide a mechanism by which quantitative differences in signal strength can alter the balance of downstream signaling pathways to induce the qualitatively distinct biological outcomes of proliferation, differentiation or apoptosis. PMID 11135575 [PubMed - indexed for MEDLINE] Dendric cells and macrophages of MEK3 deficient mice have impaired IL12 production. 1 EMBO J. 1999 Apr 1;18(7) 1845-57. Defective IL-12 production in mitogen-activated protein (MAP) kinase kinase 3 (Mkk3)-deficient mice. Lu HT, Yang DD, Wysk M, Gatti E, Mellman I, Davis RJ, Flavell RA. Howard Hughes Medical Institute and Section of Immunobiology, Yale University School of Medicine, New Haven, CT 06520, USA. The p38 mitogen-activated protein kinase (MAPK) pathway, like the c-Jun N-terminal kinase (JNK) MAPK pathway, is activated in response to cellular stress and inflammation and is involved in many fundamental biological processes. To study the role of the p38 MAPK pathway in vivo, we have used homologous recombination in mice to inactivate the Mkk3 gene, one of the two specific MAPK kinases (MAPKKs) that activate p38 MAPK. Mkk3(-/-) mice were viable and fertile; however, they were defective in interleukin-12 (IL-12) production by macrophages and dendritic cells. Interferon-gamma production following immunization with protein antigens and in vitro differentiation of naive T cells is greatly reduced, suggesting an impaired type I cytokine immune response. The effect of the p38 MAPK pathway on IL-12 expression is at least partly transcriptional, since inhibition of this pathway blocks IL-12 p40 promoter activity in macrophage cell lines and IL-12 p40 mRNA is reduced in MKK3-deficient mice. We conclude that the p38 MAP kinase, activated through MKK3, is required for the production of inflammatory cytokines by both antigen-presenting cells and CD4(+) T cells. PMID 10202148 [PubMed - indexed for MEDLINE] Bam32(-/-) B cell develop normally but have impaired T-independent antibody responses in vivo. 1 Immunity. 2003 Oct;19(4) 621-32. Bam32 links the B cell receptor to ERK and JNK and mediates B cell proliferation but not survival. Han A, Saijo K, Mecklenbrauker I, Tarakhovsky A, Nussenzweig MC. Laboratory of Molecular Immunology, The Rockefeller University, New York, NY 10021, USA. Bam32 is an adaptor protein recruited to the plasma membrane upon B cell receptor (BCR) crosslinking in a phosphoinositol 3-kinase (PI3K)-dependent manner; however, its physiologic function is unclear. To determine its physiologic function, we produced Bam32-deficient mice. Bam32(-/-) B cells develop normally but have impaired T-independent antibody responses in vivo and diminished responses to BCR crosslinking in vitro. Biochemical analysis revealed that Bam32 acts in a novel pathway leading from the BCR to MAPK/ERK Kinases (MEK1/2), MAPK/ERK Kinase Kinase-1 (MEKK1), extracellular signal-regulated kinase (ERK), and c-jun NH2-terminal kinase (JNK), but not p38 mitogen-activated protein kinase (p38). This pathway appears to be initiated by hematopoietic progenitor kinase-1 (HPK1), which interacts directly with Bam32, and differs from all previously characterized BCR signaling pathways in that it is required for normal BCR-mediated proliferation but not for B cell survival. PMID 14563325 [PubMed - indexed for MEDLINE] T-cell and B-cell of RAP1A deficient mice impair integrin-mediated cell adhesion. 1 Mol Cell Biol. 2006 Jan;26(2) 643-53. Rap1A-deficient T and B cells show impaired integrin-mediated cell adhesion. Duchniewicz M, Zemojtel T, Kolanczyk M, Grossmann S, Scheele JS, Zwartkruis FJ. Department of Computational Molecular Biology, Max Planck Institute for Molecular Genetics, Ihnestrasse 73, D-14195 Berlin, Germany. Studies in tissue culture cells have demonstrated a role for the Ras-like GTPase Rap1 in the regulation of integrin-mediated cell-matrix and cadherin-mediated cell-cell contacts. To analyze the function of Rap1 in vivo, we have disrupted the Rap1A gene by homologous recombination. Mice homozygous for the deletion allele are viable and fertile. However, primary hematopoietic cells isolated from spleen or thymus have a diminished adhesive capacity on ICAM and fibronectin substrates. In addition, polarization of T cells from Rap1-/- cells after CD3 stimulation was impaired compared to that of wild-type cells. Despite this, these defects did not result in hematopoietic or cell homing abnormalities. Although it is possible that the relatively mild phenotype is a consequence of functional complementation by the Rap1B gene, our genetic studies confirm a role for Rap1A in the regulation of integrins. PMID 16382154 [PubMed - indexed for MEDLINE] T-cell of WASP deficient mice impair the proliferaction and antigen receptor cap formation in response to anti-CD3zeta stimulation. 1 Immunity. 1998 Jul;9(1) 81-91. Wiskott-Aldrich syndrome protein-deficient mice reveal a role for WASP in T but not B cell activation. Snapper SB, Rosen FS, Mizoguchi E, Cohen P, Khan W, Liu CH, Hagemann TL, Kwan SP, Ferrini R, Davidson L, Bhan AK, Alt FW. Howard Hughes Medical Institute, Children s Hospital, Boston, Massachusetts 02115, USA. The Wiskott-Aldrich syndrome (WAS) is a human X-linked immunodeficiency resulting from mutations in a gene (WASP) encoding a cytoplasmic protein implicated in regulating the actin cytoskeleton. To elucidate WASP function, we disrupted the WASP gene in mice by gene-targeted mutation. WASP-deficient mice showed apparently normal lymphocyte development, normal serum immunoglobulin levels, and the capacity to respond to both T-dependent and T-independent type II antigens. However, these mice did have decreased peripheral blood lymphocyte and platelet numbers and developed chronic colitis. Moreover, purified WASP-deficient T cells showed markedly impaired proliferation and antigen receptor cap formation in response to anti-CD3epsilon stimulation. Yet, purified WASP-deficient B cells showed normal responses to anti-Ig stimulation. We discuss the implications of our findings regarding WASP function in receptor signaling and cytoskeletal reorganization in T and B cells and compare the effects of WASP deficiency in mice and humans. PMID 9697838 [PubMed - indexed for MEDLINE] T-cell of SHB defective mice impair the phosphorylation of LAT and consequently the activation of MAP kinase pathways. 9 Sep 24;274(39) 28050-7. Requirement of the Src homology 2 domain protein Shb for T cell receptor-dependent activation of the interleukin-2 gene nuclear factor for activation of T cells element in Jurkat T cells. Lindholm CK, Gylfe E, Zhang W, Samelson LE, Welsh M. Department of Medical Cell Biology, Box 571, Biomedicum, Uppsala University, S-75123 Uppsala, Sweden. Stimulation of the T cell antigen receptor (TCR) induces tyrosine phosphorylation of numerous intracellular proteins. We have recently investigated the role of the adaptor protein Shb in the early events of T cell signaling and observed that Shb associates with Grb2, linker for activation of T cells (LAT) and the TCR zeta-chain in Jurkat cells. We now report that Shb also associates with phospholipase C-gamma1 (PLC-gamma1) in these cells. Overexpression of Src homology 2 domain defective Shb caused diminished phosphorylation of LAT and consequently the activation of mitogen-activated protein kinases was decreased upon TCR stimulation. In addition, the Shb mutant also blocked phosphorylation of PLC-gamma1 and the increase in cytoplasmic Ca(2+) following TCR stimulation. Nuclear factor for activation of T cells is a major target for Ras and calcium signaling pathways in T cells following TCR stimulation, and the overexpression of the mutant Shb prevented TCR-dependent activation of the nuclear factor for activation of T cells. Consequently, endogenous interleukin-2 production was decreased under these conditions. The results indicate a role for Shb as a link between the TCR and downstream signaling events involving LAT and PLC-gamma1 and resulting in the activation of transcription of the interleukin-2 gene. PMID 10488157 [PubMed - indexed for MEDLINE] B-cell of 3BP2 (-/-) deficient mice have defective in proliferation, cell cycle progression, PLC-gamma2 phosphorylation, calcium mobilization, NF-ATp dephosphorylation, and Erk and Jnk activation in response to BCR ligation. 1 Mol Cell Biol. 2006 Jul;26(14) 5214-25. 3BP2 deficiency impairs the response of B cells, but not T cells, to antigen receptor ligation. de la Fuente MA, Kumar L, Lu B, Geha RS. Division of Immunology, Children s Hospital, 300 Longwood Ave., Boston, MA 02115, USA. The adapter protein 3BP2 is expressed in lymphocytes; binds to Syk/ZAP-70, Vav, and phospholipase C-gamma (PLC-gamma); and is thought to be important for interleukin-2 gene transcription in T cells. To define the role of 3BP2 in lymphocyte development and function, we generated 3BP2-deficient mice. T-cell development, proliferation, cytokine secretion, and signaling in response to T-cell receptor (TCR) ligation were all normal in 3BP2(-/-) mice. 3BP2(-/-) mice had increased accumulation of pre-B cells in the bone marrow and a block in the progression of transitional B cells in the spleen from the T1 to the T2 stage, but normal numbers of mature B cells. B-cell proliferation, cell cycle progression, PLC-gamma2 phosphorylation, calcium mobilization, NF-ATp dephosphorylation, and Erk and Jnk activation in response to B-cell receptor (BCR) ligation were all impaired. These results suggest that 3BP2 is important for BCR, but not for TCR signaling. PMID 16809760 [PubMed - indexed for MEDLINE] B-cell of Vav2(-/-) deficient mice are defective in the ability to switch immunoglobulin class. 1 Nat Immunol. 2001 Jun;2(6) 542-7. Comment in Nat Immunol. 2001 Jun;2(6) 482-4. Signal transduction through Vav-2 participates in humoral immune responses and B cell maturation. Doody GM, Bell SE, Vigorito E, Clayton E, McAdam S, Tooze R, Fernandez C, Lee IJ, Turner M. Laboratory of Lymphocyte Signaling and Development, Molecular Immunology Programme, The Babraham Institute, Babraham, Cambridge CB2 4AT, UK. B and T lymphocytes develop normally in mice lacking the guanine nucleotide exchange factor Vav-2. However, the immune responses to type II thymus-independent antigen as well as the primary response to thymus-dependent (TD) antigen are defective. Vav-2-deficient mice are also defective in their ability to switch immunoglobulin class, form germinal centers and generate secondary immune responses to TD antigens. Mice lacking both Vav-1 and Vav-2 contain reduced numbers of B lymphocytes and display a maturational block in the development of mature B cells. B cells from Vav-1(-/-)Vav-2(-/-) mice respond poorly to antigen receptor triggering, both in terms of proliferation and calcium release. These studies show the importance of Vav-2 in humoral immune responses and B cell maturation. PMID 11376342 [PubMed - indexed for MEDLINE] T-cell of Vav1(-/-) deficient mice exhibit impaired antigen receptor signaling. 1 Nature. 1995 Mar 30;374(6521) 474-7. Defective T-cell receptor signalling and positive selection of Vav-deficient CD4+ CD8+ thymocytes. Fischer KD, Zmuldzinas A, Gardner S, Barbacid M, Bernstein A, Guidos C. Program in Molecular Biology and Cancer, Samuel Lunenfeld Research Institute, Mount Sinai Hospital, Toronto, Ontario, Canada. During lymphocyte development, cellular proliferation and positive and negative selection events ensure the production of T and B lymphocytes bearing highly diverse, but self-tolerant, repertoires of antigen receptors. These processes are initiated when engagement of growth-factor receptors, or the T and B lymphocyte antigen receptors, induces tyrosine phosphorylation of specific SH2- and SH3-domain-containing cytoplasmic proteins, including Vav. Here we show that vav-/- embryonic stem cells generate only limited numbers of immature and mature T and B lymphocytes in the RAG-2 blastocyst complementation assay. Furthermore, Vav-deficient T lymphocytes showed severely impaired antigen receptor signalling. Finally, we demonstrate that Vav-dependent signalling pathways regulate maturation, but not CD4/CD8 lineage commitment, during T-cell-receptor-mediated positive selection of immature CD4+ CD8+ precursors into mature CD4+ CD8- or CD4- CD8+ T cells. PMID 7700360 [PubMed - indexed for MEDLINE] Vav1(-/-)Vav2(-/-) mice exhibit greatly reduced the mature B-cells. Vav-1-/-Vav-2-/- B cells were unresponsive to BCR-driven proliferation in vitro and to thymus-indepen-dent antigen in vivo. 1 Nat Immunol. 2001 Jun;2(6) 548-55. Comment in Nat Immunol. 2001 Jun;2(6) 482-4. Compensation between Vav-1 and Vav-2 in B cell development and antigen receptor signaling. Tedford K, Nitschke L, Girkontaite I, Charlesworth A, Chan G, Sakk V, Barbacid M, Fischer KD. Abteilung Physiologische Chemie, Universitat Ulm, Albert-Einstein-Allee 11, D-89069 Ulm, Germany. Vav-1 and Vav-2 are closely related Dbl-homology GTP exchange factors (GEFs) for Rho GTPases. Mutation of Vav-1 disrupts T cell development and T cell antigen receptor-induced activation, but has comparatively little effect on B cells. We found that combined deletion of both Vav-1 and Vav-2 in mice resulted in a marked reduction in mature B lymphocyte numbers. Vav-1(-/-)Vav-2(-/-) B cells were unresponsive to B cell antigen receptor (BCR)-driven proliferation in vitro and to thymus-independent antigen in vivo. BCR-stimulated intracellular calcium mobilization was greatly impaired in Vav-1(-/-)Vav-2(-/-) B cells. These findings establish a role for Vav-2 in BCR calcium signaling and reveal that the Vav family of GEFs is critical to B cell development and function. PMID 11376343 [PubMed - indexed for MEDLINE] Fyn-deficient mice exhibit a remarkably specific lymphoid defect thymocytes are refractile to stimulation through the TCR with mitogen or antigen. 1 Cell. 1992 Sep 4;70(5) 751-63. Defective T cell receptor signaling in mice lacking the thymic isoform of p59fyn. Appleby MW, Gross JA, Cooke MP, Levin SD, Qian X, Perlmutter RM. Howard Hughes Medical Institute, Department of Immunology, University of Washington, Seattle 98195. Considerable evidence supports the hypothesis that the nonreceptor protein tyrosine kinase p59fyn participates in signal transduction from the T cell receptor (TCR). To examine this hypothesis in detail, we have produced mice that lack the thymic isoform of p59fyn but retain expression of the brain isoform of the protein. fynTnull mice exhibit a remarkably specific lymphoid defect thymocytes are refractile to stimulation through the TCR with mitogen or antigen, while peripheral T cells, following what appears to be a normal maturation sequence, reacquire significant signaling capabilities. These data confirm that p59fynT plays a pivotal role in TCR signal transduction and demonstrate that additional developmentally regulated signaling components also contribute to TCR-induced lymphocyte activation. PMID 1516132 [PubMed - indexed for MEDLINE] Lck-deficient mice show a pronounced thymic atrophy, with a dramatic reduction in the double-positive (CD4+CD8+) thymocyte population. 1 Nature. 1992 May 14;357(6374) 161-4. Comment in Nature. 1993 Jan 21;361(6409) 213. Profound block in thymocyte development in mice lacking p56lck. Molina TJ, Kishihara K, Siderovski DP, van Ewijk W, Narendran A, Timms E, Wakeham A, Paige CJ, Hartmann KU, Veillette A, et al. Ontario Cancer Institute, University of Toronto, Canada. The protein Lck (p56lck) has a relative molecular mass of 56,000 and belongs to the Src family of tyrosine kinases. It is expressed exclusively in lymphoid cells, predominantly in thymocytes and peripheral T cells. Lck associates specifically with the cytoplasmic domains of both CD4 and CD8 T-cell surface glycoproteins and interacts with the beta-chain of the interleukin-2 receptor, which implicates Lck activity in signal transduction during thymocyte ontogeny and activation of mature T cells. Here we generate an lck null mutation by homologous recombination in embryonic stem cells to evaluate the role of p56lck in T-cell development and activation. Lck-deficient mice show a pronounced thymic atrophy, with a dramatic reduction in the double-positive (CD4+CD8+) thymocyte population. Mature, single-positive thymocytes are not detectable in these mice and there are only very few peripheral T cells. These results illustrate the crucial role of this T-cell-specific tyrosine kinase in the thymocyte development. PMID 1579166 [PubMed - indexed for MEDLINE] T cell from mice deficient in LCK is required for normal signal transduction through the TCR. 1 Cell. 1992 Aug 21;70(4) 585-93. Genetic evidence for the involvement of the lck tyrosine kinase in signal transduction through the T cell antigen receptor. Straus DB, Weiss A. Howard Hughes Medical Institute, Department of Medicine, University of California, San Francisco 94143. Signaling through the T cell antigen receptor (TCR) results both in rapid increases in tyrosine phosphorylation on a number of proteins and in the activation of the phosphatidylinositol pathway. It is not clear how stimulation of the TCR leads to these signaling events. Mutants of the Jurkat T cell line have been previously isolated that fail to show increases in calcium following receptor stimulation. Analysis of one of these mutants, JCaM1, which is defective in the induction of tyrosine phosphorylation, revealed a defect in the expression of functional lck tyrosine kinase. The lack of lck activity was caused in part by a splicing defect. Expression of the lck cDNA in JCaM1 restores the ability of the cell to respond to TCR stimulation. These results indicate that lck is required for normal signal transduction through the TCR. PMID 1505025 [PubMed - indexed for MEDLINE] T cells from mice deficient in SLAP-130/Fyb show markedly impaired proliferation. 1 Science. 2001 Sep 21;293(5538) 2263-5. Coupling of the TCR to integrin activation by Slap-130/Fyb. Peterson EJ, Woods ML, Dmowski SA, Derimanov G, Jordan MS, Wu JN, Myung PS, Liu QH, Pribila JT, Freedman BD, Shimizu Y, Koretzky GA. The Abramson Family Cancer Research Institute, Department of Medicine, School of Medicine, University of Pennsylvania, Philadelphia, PA, 19104, USA. SLAP-130/Fyb (SLP-76-associated phosphoprotein or Fyn-binding protein; also known as Fyb/Slap) is a hematopoietic-specific adapter, which associates with and modulates function of SH2-containing leukocyte phosphoprotein of 76 kilodaltons (SLP-76). T cells from mice lacking SLAP-130/Fyb show markedly impaired proliferation following CD3 engagement. In addition, the T cell receptor (TCR) in SLAP-130/Fyb mutant cells fails to enhance integrin-dependent adhesion. Although TCR-induced actin polymerization is normal, TCR-stimulated clustering of the integrin LFA-1 is defective in SLAP-130/Fyb-deficient cells. These data indicate that SLAP-130/Fyb is important for coupling TCR-mediated actin cytoskeletal rearrangement with activation of integrin function, and for T cells to respond fully to activating signals. PMID 11567141 [PubMed - indexed for MEDLINE] B cell of chicken deficient ITK reduce IP3 generation and phospholipase C gamma 2 tyrosine phosphorylation. 1 J Exp Med. 1996 Jul 1;184(1) 31-40. A role for Bruton s tyrosine kinase in B cell antigen receptor-mediated activation of phospholipase C-gamma 2. Takata M, Kurosaki T. Department of Oncology and Immunology, Wyeth-Ayerst Research, Pearl River, New York 10965, USA. Defects in the gene encoding Bruton s tyrosine kinase (Btk) result in a disease called X-linked agammaglobulinemia, in which there is a profound decrease of mature B cells due to a block in B cell development. Recent studies have shown that Btk is tyrosine phosphorylated and activated upon B cell antigen receptor (BCR) stimulation. To elucidate the functions of this kinase, we examined BCR signaling of DT40 B cells deficient in Btk. Tyrosine phosphorylation of phospholipase C (PLC)-gamma 2 upon receptor stimulation was significantly reduced in the mutant cells, leading to the loss of both BCR-coupled phosphatidylinositol hydrolysis and calcium mobilization. Pleckstrin homology and Src-homology 2 domains of Btk were required for PLC-gamma 2 activation. Since Syk is also required for the BCR-induced PLC-gamma 2 activation, our findings indicate that PLC-gamma 2 activation is regulated by Btk and Syk through their concerted actions. PMID 8691147 [PubMed - indexed for MEDLINE] T cell of mice deficient ITK reduce IP3 generation and phospholipase C gamma 1 tyrosine phosphorylation. 1 J Exp Med. 1998 May 18;187(10) 1721-7. T cell receptor-initiated calcium release is uncoupled from capacitative calcium entry in Itk-deficient T cells. Liu KQ, Bunnell SC, Gurniak CB, Berg LJ. Program of Immunology, Division of Medical Sciences, Harvard University, Boston, Massachusetts 02115, USA. Itk, a Tec family tyrosine kinase, plays an important but as yet undefined role in T cell receptor (TCR) signaling. Here we show that T cells from Itk-deficient mice have a TCR-proximal signaling defect, resulting in defective interleukin 2 secretion. Upon TCR stimulation, Itk-/- T cells release normal amounts of calcium from intracellular stores, but fail to open plasma membrane calcium channels. Since thapsigargin-induced store depletion triggers normal calcium entry in Itk-/- T cells, an impaired biochemical link between store depletion and channel opening is unlikely to be responsible for this defect. Biochemical studies indicate that TCR-induced inositol 1,4,5 tris-phosphate (IP3) generation and phospholipase C gamma1 tyrosine phosphorylation are substantially reduced in Itk-/- T cells. In contrast, TCR-zeta and ZAP-70 are phosphorylated normally, suggesting that Itk functions downstream of, or in parallel to, ZAP-70 to facilitate TCR-induced IP3 production. These findings support a model in which quantitative differences in cytosolic IP3 trigger distinct responses, and in which only high concentrations of IP3 trigger the influx of extracellular calcium. PMID 9584150 [PubMed - indexed for MEDLINE] T cell of mice deficient ITK have failure of Th2 development. 1 Immunity. 1999 Oct;11(4) 399-409. Impaired NFATc translocation and failure of Th2 development in Itk-deficient CD4+ T cells. Fowell DJ, Shinkai K, Liao XC, Beebe AM, Coffman RL, Littman DR, Locksley RM. Department of Medicine, University of California San Francisco 94143, USA. Naive Itk-deficient CD4+ T cells were unable to establish stable IL-4 production, even when primed in Th2-inducing conditions. In contrast, IFNgamma production was little affected. Failure to express IL-4 occurred even among cells that had gone through multiple cell divisions and was associated with a delay in the kinetics and magnitude of NFATc nuclear localization. IL-4 production was restored genetically by retroviral reconstitution of Itk or biochemically by augmenting the calcium flux with ionomycin. In vivo, Itk-deficient mice were unable to establish functional Th2 cells. Development of protective Th1 cells was unimpeded. These data define a nonredundant role for Itk in modulating signals from the TCR/CD28 pathways that are specific for the establishment of stable IL-4 but not IFNgamma expression. PMID 10549622 [PubMed - indexed for MEDLINE] Mice deficient in ITK have reduced proliferative responses to MHC stimulation and to anti-TCR cross-linking 1 Immunity. 1995 Dec;3(6) 757-69. Altered T cell receptor signaling and disrupted T cell development in mice lacking Itk. Liao XC, Littman DR. Department of Microbiology and Immunology, University of California, San Francisco 94143-0414, USA. Itk is a T cell protein tyrosine kinase (PTK) that, along with Btk and Tec, belongs to a family of cytoplasmic PTKs with N-terminal pleckstrin homology domains. Btk plays a critical role in B lymphocyte development. To determine whether Itk has an analogous role in T lymphocytes, we used gene targeting to prepare mice lacking expression of Itk. Such animals had decreased numbers of mature thymocytes, an effect most clearly observed in mice expressing T cell receptor (TCR) transgenes. Mature T cells from Itk-deficient mice had reduced proliferative responses to allogeneic MHC stimulation and to anti-TCR cross-linking, but responded normally to stimulation with phorbol ester plus ionomycin or with IL-2. These results provide genetic evidence that Itk is involved in T cell development and also suggest that Itk has an important role in proximal events in TCR-mediated signaling pathways. PMID 8777721 [PubMed - indexed for MEDLINE] Mutations in Btk cause X-linked immunodeficiency. 1 Semin Immunol. 1998 Aug;10(4) 309-16. Btk function in B cell development and response. Satterthwaite AB, Li Z, Witte ON. Department of Microbiology and Molecular Genetics, University of California, Los Angeles 90095-1662, USA. Mutations in Bruton s tyrosine kinase (Btk) result in the B cell immunodeficiencies XLA in humans and Xid in mice. Both the maintenance of peripheral B cell numbers and their response to B cell antigen receptor (BCR) crosslinking depend on Btk. Btk integrates signals from multiple cell surface receptors, including BCR and G-protein coupled receptors. These Btk dependent signals control B cell proliferation and survival by mediating Ca2+ flux, activating JNK and p38 and inducing cell cycle regulatory genes. Publication Types Review PMID 9695187 [PubMed - indexed for MEDLINE] Gads(GRAP2) has a role in thymocyte proliferaction for maturation of T-cells. 1 Science. 2001 Mar 9;291(5510) 1987-91. Requirement for the SLP-76 adaptor GADS in T cell development. Yoder J, Pham C, Iizuka YM, Kanagawa O, Liu SK, McGlade J, Cheng AM. Medical Scientist Training Program, Washington University School of Medicine, St. Louis, MO 63110, USA. GADS is an adaptor protein implicated in CD3 signaling because of its ability to link SLP-76 to LAT. A GADS-deficient mouse was generated by gene targeting, and the function of GADS in T cell development and activation was examined. GADS- CD4-CD8- thymocytes exhibited a severe block in proliferation but still differentiated into mature T cells. GADS- thymocytes failed to respond to CD3 cross-linking in vivo and were impaired in positive and negative selection. Immunoprecipitation experiments revealed that the association between SLP-76 and LAT was uncoupled in GADS- thymocytes. These observations indicate that GADS is a critical adaptor for CD3 signaling. PMID 11239162 [PubMed - indexed for MEDLINE] Gads(GRAP2) has a role for homeostatic proliferaction in B cells. 1 Eur J Immunol. 2005 Apr;35(4) 1184-92. Expression and function of the adaptor protein Gads in murine B cells. Yankee TM, Draves KE, Clark EA. Department of Immunology, University of Washington, Seattle, USA. tyankee@kumc.edu Nearly all hematopoietic receptors are dependent on adaptor proteins for the activation of downstream signaling pathways. The Gads adaptor protein is expressed in many hematopoietic tissues, including bone marrow, lymph node, and spleen. Using intracellular staining, we detected Gads protein in a number cells, including B cells, T cells, NK cells, monocytes, and plasmacytoid DC, but not in macrophages, neutrophils, or monocyte-derived DC. In the B cell compartment, Gads was first expressed after immature B cells leave the bone marrow and was down-regulated after B cell antigen receptor (BCR) ligation. Female Gads(-/-) mice had increased numbers of splenic B cells, as compared to female Gads(+/+) mice, suggesting a role for Gads in B cell homeostasis. Although B cell production and turnover of splenic B cell subsets appeared normal in Gads(-/-) mice, homeostatic proliferation was significantly impaired in Gads(-/-) B cells. Whereas BCR ligation can induce apoptosis in wild-type transitional stage 1 (T1) B cells, Gads(-/-) T1 B cells were resistant to BCR-induced apoptosis. Gads(-/-) B cells also showed increased BCR-mediated calcium mobilization. We conclude that Gads may have a negative regulatory role in signaling through survival pathways, and is necessary for normal homeostatic proliferation in B cells. PMID 15761845 [PubMed - indexed for MEDLINE] Grap negatively regulates T-cell proliferation. 1 Mol Cell Biol. 2002 May;22(10) 3230-6. Grap negatively regulates T-cell receptor-elicited lymphocyte proliferation and interleukin-2 induction. Shen R, Ouyang YB, Qu CK, Alonso A, Sperzel L, Mustelin T, Kaplan MH, Feng GS. Program in Signal Transduction Research, The Burnham Institute, La Jolla, California 92037, USA. Grb-2-related adaptor protein (Grap) is a Grb2-like SH3-SH2-SH3 adaptor protein with expression restricted to lymphoid tissues. Grap(-/-) lymphocytes isolated from targeted Grap-deficient mice exhibited enhanced proliferation, interleukin-2 production, and c-fos induction in response to mitogenic T-cell receptor (TCR) stimulation, compared to wild-type cells. Ectopic expression of Grap led to a suppression of Elk-1-directed transcription induced by the Ras/Erk pathway, without having effects on gene expression mediated by Jnk and p38 mitogen-activated protein kinases. Together, these data suggest that Grap, unlike Grb2, acts as a negative regulator of TCR-stimulated intracellular signaling by downregulating signal relay through the Ras/Erk pathway. PMID 11971956 [PubMed - indexed for MEDLINE] Gab2 is a substrate of ZAP-70 and functions as a switch molecule toward inhibition of TCR signal transuduction. 1 J Biol Chem. 2001 Nov 30;276(48) 45175-83. Epub 2001 Sep 25. Docking protein Gab2 is phosphorylated by ZAP-70 and negatively regulates T cell receptor signaling by recruitment of inhibitory molecules. Yamasaki S, Nishida K, Hibi M, Sakuma M, Shiina R, Takeuchi A, Ohnishi H, Hirano T, Saito T. Molecular Genetics, Chiba University Graduate School of Medicine, Chiba 260-8670, Japan. To maintain various T cell responses and immune equilibrium, activation signals triggered by T cell antigen receptor (TCR) must be regulated by inhibitory signals. Gab2, an adaptor protein of the insulin receptor substrate-1 family, has been shown to be involved in the downstream signaling from cytokine receptors. We investigated the functional role of Gab2 in TCR-mediated signal transduction. Gab2 was phosphorylated by ZAP-70 and co-precipitated with phosphoproteins, such as ZAP-70, LAT, and CD3zeta, upon TCR stimulation. Overexpression of Gab2 in Jurkat cells or antigen-specific T cell hybridomas resulted in the inhibition of NF-AT activation, interleukin-2 production, and tyrosine phosphorylation. The structure-function relationship of Gab2 was analyzed by mutants of Gab2. The Gab2 mutants lacking SHP-2-binding sites mostly abrogated the inhibitory activity of Gab2, but its inhibitory function was restored by fusing to active SHP-2 as a chimeric protein. A mutant with defective phosphatidylinositol 3-kinase binding capacity also impaired the inhibitory activity, and the pleckstrin homology domain-deletion mutant revealed a crucial function of the pleckstrin homology domain for localization to the plasma membrane. These results suggest that Gab2 is a substrate of ZAP-70 and functions as a switch molecule toward inhibition of TCR signal transduction by mediating the recruitment of inhibitory molecules to the TCR signaling complex. PMID 11572860 [PubMed - indexed for MEDLINE] B cell signaling causes tyrosine phosphorylation of Gab1, and in turn SHP2 bind to Gab1 Gab1 phosophorylation potentiate the phosphorylation of Akt, PI3K-dependent response. 1 J Biol Chem. 2001 Apr 13;276(15) 12257-65. Epub 2001 Jan 22. The Gab1 docking protein links the b cell antigen receptor to the phosphatidylinositol 3-kinase/Akt signaling pathway and to the SHP2 tyrosine phosphatase. Ingham RJ, Santos L, Dang-Lawson M, Holgado-Madruga M, Dudek P, Maroun CR, Wong AJ, Matsuuchi L, Gold MR. Departments of Microbiology and Immunology and Zoology, University of British Columbia, Vancouver, British Columbia V6T 1Z3, Canada. B cell antigen receptor (BCR) signaling causes tyrosine phosphorylation of the Gab1 docking protein. This allows phosphatidylinositol 3-kinase (PI3K) and the SHP2 tyrosine phosphatase to bind to Gab1. In this report, we tested the hypothesis that Gab1 acts as an amplifier of PI3K- and SHP2-dependent signaling in B lymphocytes. By overexpressing Gab1 in the WEHI-231 B cell line, we found that Gab1 can potentiate BCR-induced phosphorylation of Akt, a PI3K-dependent response. Gab1 expression also increased BCR-induced tyrosine phosphorylation of SHP2 as well as the binding of Grb2 to SHP2. We show that the pleckstrin homology (PH) domain of Gab1 is required for BCR-induced phosphorylation of Gab1 and for Gab1 participation in BCR signaling. Moreover, using confocal microscopy, we show that BCR ligation can induce the translocation of Gab1 from the cytosol to the plasma membrane and that this requires the Gab1 PH domain as well as PI3K activity. These findings are consistent with a model in which the binding of the Gab1 PH domain to PI3K-derived lipids brings Gab1 to the plasma membrane, where it can be tyrosine-phosphorylated and then act as an amplifier of BCR signaling. PMID 11278704 [PubMed - indexed for MEDLINE] RasGRP1 mediates Ras activation following TCR stimulatioin. RasGRP1 and RasGRP3 induces RAS activation in B-cell to response to T-cell stimulation. 1 Immunol Lett. 2006 May 15;105(1) 77-82. Epub 2006 Feb 20. The role of RasGRPs in regulation of lymphocyte proliferation. Coughlin JJ, Stang SL, Dower NA, Stone JC. Department of Biochemistry, University of Alberta, Edmonton, Alta., Canada T6G 2H7. RasGRP1 links TCR signaling to Ras in T cells, while both RasGRP1 and RasGRP3 link BCR signaling to Ras in B cells. T cells deficient in RasGRP1 have defective proliferative responses as do B cells deficient in both RasGRP1 and RasGRP3, confirming the importance of Ras activation in lymphocyte proliferation. While aged Rasgrp1-/- mice develop late-onset autoimmunity characterized by splenomegaly and the presence of anti-nuclear antibodies (ANA), the additional loss of RasGRP3 expression inhibits this phenotype. We show here that the autoimmunity in Rasgrp1-/- mice is T cell dependent. Compared to wildtype, Rasgrp1-/- T cells induce greater in vitro B cell proliferation that is due, at least in part, to increased production of interleukin-4 (IL-4). Rasgrp1 Rasgrp3 double mutant B cells are less responsive to this T cell stimulation. The reduced double mutant B cell proliferative response was paralleled by decreased induction of cyclin D2 upon stimulation with IL-4 and anti-IgM. Taken together these results suggest that double mutant mice fail to generate autoimmunity due to their decreased B cell cyclin D2 accumulation, and thus proliferation, in response to the elevated levels of IL-4 produced by mutant T cells. PMID 16530850 [PubMed - indexed for MEDLINE] Grb2-hSos1-PLCgamma1-p36/p38-ZAP70 complexes localize in the vicinity of TCR-zeta 1 J Biol Chem. 1995 Aug 4;270(31) 18428-36. Ligation of the T-cell antigen receptor (TCR) induces association of hSos1, ZAP-70, phospholipase C-gamma 1, and other phosphoproteins with Grb2 and the zeta-chain of the TCR. Nel AE, Gupta S, Lee L, Ledbetter JA, Kanner SB. Department of Medicine, UCLA School of Medicine 90024, USA. Signaling by the T-cell antigen receptor (TCR) involves both phospholipase C (PLC)-gamma 1 and p21ras activation. While failing to induce Shc/Grb2 association, ligation of the TCR/CD3 receptor in Jurkat T-cells induced hSos1-Grb2 complexes. In addition to hSos1, Grb2 participates in the formation of a tyrosine phosphoprotein complex that includes 145-, 95-, 70-, 54-, and 36-38-kDa proteins. p145 was identified as PLC-gamma 1 and p70 as the protein tyrosine kinase, ZAP-70. Although of the same molecular weight, p95 was not recognized by an anti-serum to p95 Vav. The SH2 domains of Grb2 and PLC-gamma 1 were required for the formation of this protein complex. In anti-CD3-treated cells, Grb2 redistributed from the cytosol to a particulate cell compartment along with p36/p38, ZAP-70, and PLC-gamma 1. Part of the Grb2 complex associated with the particulate compartment could be extracted with Nonidet P-40, while the rest was Nonidet P-40 insoluble. In both the detergent-soluble and -insoluble fractions, Grb2 coimmunoprecipitated with the zeta-chain of the TCR. Taken together, these results indicate that anti-CD3 induces Grb2-hSos1-PLC-gamma 1-p36/p38-ZAP70 complexes, which localize in the vicinity of TCR-zeta. PMID 7629168 [PubMed - indexed for MEDLINE] Gads(Grap2) plays an important role in T-cell signaling via its association with SLP-76 and LAT. 1 Curr Biol. 1999 Jan 28;9(2) 67-75. The hematopoietic-specific adaptor protein gads functions in T-cell signaling via interactions with the SLP-76 and LAT adaptors. Liu SK, Fang N, Koretzky GA, McGlade CJ. Department of Medical Biophysics, University of Toronto, The Arthur and Sonia Labatt Brain Tumour Research Centre, Hospital for Sick Children, Research Institute, 555 University Ave, Toronto, Ontario M5G 1X8, Canada. BACKGROUND The adaptor protein Gads is a Grb2-related protein originally identified on the basis of its interaction with the tyrosine-phosphorylated form of the docking protein Shc. Gads protein expression is restricted to hematopoietic tissues and cell lines. Gads contains a Src homology 2 (SH2) domain, which has previously been shown to have a similar binding specificity to that of Grb2. Gads also possesses two SH3 domains, but these have a distinct binding specificity to those of Grb2, as Gads does not bind to known Grb2 SH3 domain targets. Here, we investigated whether Gads is involved in T-cell signaling. RESULTS We found that Gads is highly expressed in T cells and that the SLP-76 adaptor protein is a major Gads-associated protein in vivo. The constitutive interaction between Gads and SLP-76 was mediated by the carboxy-terminal SH3 domain of Gads and a 20 amino-acid proline-rich region in SLP-76. Gads also coimmunoprecipitated the tyrosine-phosphorylated form of the linker for activated T cells (LAT) adaptor protein following cross-linking of the T-cell receptor; this interaction was mediated by the Gads SH2 domain. Overexpression of Gads and SLP-76 resulted in a synergistic augmentation of T-cell signaling, as measured by activation of nuclear factor of activated T cells (NFAT), and this cooperation required a functional Gads SH2 domain. CONCLUSIONS These results demonstrate that Gads plays an important role in T-cell signaling via its association with SLP-76 and LAT. Gads may promote cross-talk between the LAT and SLP-76 signaling complexes, thereby coupling membrane-proximal events to downstream signaling pathways. PMID 10021361 [PubMed - indexed for MEDLINE] Lck is required for normal signal transduction through the TCR. 1 Cell. 1992 Aug 21;70(4) 585-93. Genetic evidence for the involvement of the lck tyrosine kinase in signal transduction through the T cell antigen receptor. Straus DB, Weiss A. Howard Hughes Medical Institute, Department of Medicine, University of California, San Francisco 94143. Signaling through the T cell antigen receptor (TCR) results both in rapid increases in tyrosine phosphorylation on a number of proteins and in the activation of the phosphatidylinositol pathway. It is not clear how stimulation of the TCR leads to these signaling events. Mutants of the Jurkat T cell line have been previously isolated that fail to show increases in calcium following receptor stimulation. Analysis of one of these mutants, JCaM1, which is defective in the induction of tyrosine phosphorylation, revealed a defect in the expression of functional lck tyrosine kinase. The lack of lck activity was caused in part by a splicing defect. Expression of the lck cDNA in JCaM1 restores the ability of the cell to respond to TCR stimulation. These results indicate that lck is required for normal signal transduction through the TCR. PMID 1505025 [PubMed - indexed for MEDLINE] ZAP-70 plays crucial roles in T-cell activation and development. Syk triggers cellular activation in T-cell. 1 Mol Cell Biol. 1998 Mar;18(3) 1388-99. Genetic evidence for differential coupling of Syk family kinases to the T-cell receptor reconstitution studies in a ZAP-70-deficient Jurkat T-cell line. Williams BL, Schreiber KL, Zhang W, Wange RL, Samelson LE, Leibson PJ, Abraham RT. Department of Immunology, Mayo Clinic, Rochester, Minnesota 55905, USA. T-cell antigen receptor (TCR) engagement activates multiple protein tyrosine kinases (PTKs), including the Src family member, Lck, and the Syk-related PTK, ZAP-70. Studies in ZAP-70-deficient humans have demonstrated that ZAP-70 plays crucial roles in T-cell activation and development. However, progress toward a detailed understanding of the regulation and function of ZAP-70 during TCR signaling has been hampered by the lack of a suitable T-cell model for biochemical and genetic analyses. In this report, we describe the isolation and phenotypic characterization of a Syk- and ZAP-70-negative somatic mutant derived from the Jurkat T-cell line. The P116 cell line displays severe defects in TCR-induced signaling functions, including protein tyrosine phosphorylation, intracellular Ca2+ mobilization, and interleukin-2 promoter-driven transcription. These signaling defects were fully reversed by reintroduction of catalytically active versions of either Syk or ZAP-70 into the P116 cells. However, in contrast to ZAP-70 expression, Syk expression triggered a significant degree of cellular activation in the absence of TCR ligation. Transfection experiments with ZAP-70-Syk chimeric proteins indicated that both the amino-terminal regulatory regions and the carboxy-terminal catalytic domains of Syk and ZAP-70 contribute to the distinctive functional properties of these PTKs. These studies underscore the crucial role of ZAP-70 in TCR signaling and offer a powerful genetic model for further analyses of ZAP-70 regulation and function in T cells. PMID 9488454 [PubMed - indexed for MEDLINE]
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【Tags Miku Owata-P tO K】 Original Music title 片想いサンバ English music title One-sided Love Samba Romaji music title Kataomoi Sanba Music Lyrics written, Voice edited by オワタP (Owata-P) Music arranged by オワタP (Owata-P) Singer(s) 初音ミク (Hatsune Miku) Click here for the original Japanese Lyrics English Lyrics (translated by blacksaingrain)： I love you. I love you. My heart is about to burst. I love you. I love you. I can t stop these feelings. So I want you to know how I m feeling now. I love you. I love you. Get the message of mine at once. I ve been in love with you for a long time. You re unreliable though, I love you all the more for it. I love you. I m in love. But you put me behind you. I ve been in love with you for a long time. I m watching you anywhere anytime. I like you. I love you. But you re too insensible. I ll get a hustle on. I won t give up. I m gonna make you happy. I won t cry. I ll keep my chin up. I m gonna do everything I can do for you. I m sure you d say we re just friends. You re not interested in me, are you? Just get it. It pains me a lot. I can t fight back my tears any more. Get the message of mine. I can t bear one-sided love any more. Don t tease me like this. At heart you already knew it, didn t you? Anyway just pick-me-up. I ll never say "No". If only you d hold me in your arms. Even though, I like the way we re going now. Well, I m so pleased with it. I talk with you. I laugh with you. Those make me really happy. I won t tell you how I feel. I don t wanna spoil the relationship between us. I talk with you. I laugh with you. I m scared to go further. I won t go. I won t do it. I won t tell you how I feel. I don t know. I won t stop it. Stay just the way you are. I m sure you d say we re just friends. You ll never take an interest in me, will you? Just get it. I m so sad. I can t always smile. Get the message of mine. I can t bear one-sided love any more. Don t tease me like this. At heart you already knew it, didn t you? Maybe I d get tired of you by now. If you need me, just hold me back. I just wanna hear you say "I love you" Am I not doing it right? Am I getting the short end of the stick? I wonder what you re thinking now. I wanna see through you. I go out with you. And at the end, we just say "Bye- bye" Doesn t that strike you as weird? So, please snog me at once! Get the message of mine. I can t bear one-sided love any more. Don t tease me like this. At heart you already knew it, didn t you? As you are about to leave, I jump at you. And I snatch a kiss from you. You get into a flurry. And I grin at you gleefully. I love you. I love you. My heart is about to burst. I love you. I love you. I can t stop these feelings. Now I wanna share the feelings with you at once. I love you. I love you. I don t wanna forget how I m feeling now. English Lyrics (translated by motokokusanagi2009)： I love you so much I love you so much That my heart is about to burst out I love you I really love you I can t stop it So, I want you to know How I m feeling about you I love you I really love you Why don t you notice my feelings? I ve been in love with you for such a long time I love you even tho you re not dependable enough I love you, I m in love with you But I m not the one in your eyes I ve been in love with you for such a long time You re in my eyes wherever and whenever I love you, I love you from the bottom of my heart But you are really insensitive I ll do my best, I m not gonna lose this love I ll make you happy I won t cry, I ll look forward I ll do anything for you You think I m just your friend No more and no less Please get wise, It s so painful I m about to cry Realize how I ve been feeling about you I m fed up with this one-sided love Don t tease me You know my feelings at heart, don t you? Just ask me out I won t let you down Hold me tight That s more than enough Actually, I like this relationship I m satisfied enough I ll have a chat with you, I ll laugh with you That makes me happy I won t declare my love to you Cuz I don t wanna break this relationship I ll have a chat with you, I ll laugh with you I can t go forward any further I won t go, I won t do I won t declare my love to you I don t know, I won t stop You can stay as you are I m just your friend And we won t go any further, right? Get wise, I m feeling sad Even I cry once in a while Realize how I ve been feeling about you I m fed up with this one-sided love Don t tease me You know my feelings at heart, don t you? I m about give up If you don t want it, please stop me I want you to say just one this "I love you." Am I approaching in a wrong way? Am I in a no-win situation? What are you thinking? I wanna take a peek at your feelings Me and you go out together And say goodbye at the end like always It s clumsy, isn t it? So, please hold me tight right now Realize how I ve been feeling about you I m fed up with this one-sided love Don t tease me You know my feelings at heart, don t you? I jumped at you as you were about to leave me And kissed on your mouth I smirked at you panicking I love you so much I love you so much That my heart is about to burst out I love you I really love you I can t stop it What I want now is An access to your heart I love you I really love you I don t wanna forget this feelings Romaji lyrics (transliterated by blacksaingrain)： kimi ga sukide kimi ga sukide kono kokoro ga hajikesouda yo sukida kimi ga sukida kono kimochi wa tomerarenai yo dakara hayaku kimi ni kono kokoro wo shittehoshii yo sukida kimi ga sukida kono kimochi ni hayaku kizuite yo kimi no koto ga mukashi kara sukida yo kimi no tayorinai toko ga sukida yo kimi ga sukida yo koishiteiru yo dakedo kimi wa watashi wo mitenai yo watashi mukashi kara sukida yo kimi wo itsudemo dokodemo miteru yo kimi ga sukida yo aishiteiru yo dakedo kimi wa totemo nibui yo ganbaru yo makenai yo kimi wo shiawase ni shiteyan yo nakanai yo mae muku yo watashi ga nandemo shiteyan yo watashi no koto wa shosen otomodachi de nannimo kimi wa kangaetenai desho kizuite yo kurushii yo sorosoro watashi mo naichau yo watashi no kimochi ni kizuite yo kataomoi wa mou iyanan yo jirashitari nanka shinaide yo honto wa kizuiteirun desho iikara watashi wo sasotte yo kotowattari nanka shinai yo dakishimetekureru dake demo iinda yo dakedo ima no kankei wa sukida yo betsuni koredemo manzokushiteru yo kimi to hanasu yo kimi to warau yo sorede watashi wa shiawase nanda yo watashi kokuhaku wa shinai yo ima no kankei kowasu no iyada yo watashi hanasu yo watashi warau yo kore ijou susumu no kowai yo ikanai yo yaranai yo watashi no kimochi wa iwanai yo shiranai yo tomenai yo kimi wa konomama de iinda yo watashi no koto wa shosen otomodachi de korekara saki ni susumu toka nai desho kizuite yo kanashii yo watashi mo tamaniwa nakunda yo watashi no kimochi ni kizuite yo kataomoi wa mou iyanan yo jirashitari nanka shinaide yo honto wa kizuiteirun desho watashi mo sorosoro akichau yo iya nara hikitometemisete yo hitokoto sukida to ittehoshiinda yo watashi yarikata machigatteru ka na watashi sonna ikikatashiteru ka na kimi wa nani kangaeteiru no ka na kimi no kimochi wo nozokikondemitainda futari de asobi ni dekakeru soshite sonomama saigo wa baibai sore wa yappari nanka henda yo ne dakara hayaku daitekudasai watashi no kimochi nikizuiteyo kataomoi wamouiya nanyo jirashitari nanka shinaide yo honto wa kizuiteirun desho wakaregiwa ni tobitsuitemiru sonomama kuchibiru oshiateru awateru kimi wo niyari to mitsumetemiru kimi ga sukide kimi ga sukide kono kokoro ga hajikesouda yo sukida kimi ga sukida kono kimochi wa tomerarenai yo ima wa hayakukimi to kono kokoro wo kayowasetakute sukida kimi ga sukida kono kimochi wo wasuretakunai yo [Owata-P, OwataP, Garuna]