IDBObjectStoreでのエラーの原因がどうしても分かりません。

クローム開発者ツールのエラーメッセージは以下の通りです。
Uncaught DOMException: Failed to execute 'put' on 'IDBObjectStore':
Evaluating the object store's key path did not yield a value.
at IDBOpenDBRequest.request.onupgradeneeded

よろしくお願い致します。

function indexDataBase(){
  var indexedDB           = window.indexedDB;
  var request             = null;
  var db                  = null;
  var store_01            = null;
  var transaction_01      = null;
  var titleIndex_01       = null;
  var authorIndex_01      = null;

  request                 = indexedDB.open('books_01', 1);
  request.onupgradeneeded = function(){
    db             = request.result;
    store_01       = db.createObjectStore('book_01', {keyPath:'isbn_01'});
    titleIndex_01  = store_01.createIndex('by_title_01', 'title_01', {unique:false});
    authorIndex_01 = store_01.createIndex('by_author_01', 'author_01');
    store_01.put({title_01:'Quarry Memories_01', author_01:'Fred_01', isbn:123456_01}); // エラーが発生
  };

  request.onsuccess = function(){
    db             = request.result;
    transaction_01 = db.transaction('book_01', 'readwrite');
    store_01       = transaction_01.objectStore('book_01');
    store_01.put({title_01:'Quarry Memories_01', author_01:'Fred_01', isbn:234567_02});
  };
}

indexDataBase();