Anne Shirley returns to Avonlea after five years of teaching at an orphanage in Nova Scotia while separated from her fiancé, Gilbert Blythe, who was finishing medical school. After catching up with Diana Barry and other old friends, Anne is reunited with Gilbert on the island. However, instead of settling down immediately, Gilbert asks Anne to move to New York since he has been offered a staff position at a prestigious medical institution. Her reluctant agreement comes with the promise of being published and that they will return to Prince Edward Island to raise their family.
In New York, Anne is employed as a junior editor at a small publishing house, where she meets Jack Garrison, a writer and local sensation who becomes interested in Anne. Seeing her potential as a writer, he offers to become the editor for her new manuscript and use his influence to have it published under both their names. When the manuscript is accepted, Anne is outraged to learn that Jack has presented it as his own; she breaks off contact with Jack, who reveals he has fallen in love with her. Meanwhile, Gilbert is troubled by his work at the hospital - most of the doctors are more interested in prestige than saving lives. Disillusioned by life in New York, Anne and Gilbert return to Prince Edward Island.
Upon returning to Avonlea, Gilbert buys Green Gables from Mr. Harrison and they set about restoring the house as they plan their wedding. However, the couple find the effect of the war in Europe to be much more present; Fred Wright, Diana's husband, eventually enlists in the war as his family life becomes increasingly strained. Increasing pressure results in Gilbert enlisting as medical officer and he convinces Anne to marry him before he leaves. When he is declared missing in action, Anne leaves for Europe as a Red Cross volunteer in hopes of finding him.
Anne's desperate search leads to a chance meeting with Jack Garrison, along with his lover, Colette, and their infant son, Dominic. Jack, involved in dangerous espionage activities, leaves suddenly and asks that Anne take Colette and Dominic to London. However, Colette is killed in an explosion while Anne briefly sees Gilbert as his troupe is forced to abandon their field hospital. Anne promises Colette on her last breath to take care of Dominic and find his father. Left to care for Dominic, Anne finds an injured Fred and his arm must be amputated. They leave for London and reside in Jack's apartment until they can return to Canada; in that time, Anne works for a British newspaper office and becomes acquainted with Fergus Keegan, Jack's editor, and Maude Montrose, one of Jack's contemporaries and a fellow spy. When Jack arrives to make arrangements for his family, he learns of Colette's death and asks for Anne to care for Dominic should anything happen to him. Though Anne intends to return to Canada with Fred and Dominic, her connection to Jack as well as a resurfacing chance of finding Gilbert results in taking Dominic with her to France.
Disguised as a nun and befriending entertainers Margaret Bush and Elsie James, Anne struggles to bring Dominic to Kit Garrison, Jack's aunt who has also become involved in war efforts. Reunited again with Jack, Anne travels with him in hopes for finding her husband. As Anne's search continues to be fruitless, Jack attempts to persuade her to return with him to the United States and form a family with him and Dominic; Anne refuses, determined to continue searching for Gilbert in Germany. When she encounters Margaret and Elsie again, she joins them in entertaining the soldiers; in the crowd is Gilbert and Anne is reunited with him at last. Jack arranges for them to leave immediately as the armistice to end the war approaches. However, Keegan's fear of being exposed as a traitor by Jack's group results in Jack being shot on the train; as he dies, he asks that Anne take care of his son. Once the armistice is declared, Anne and Gilbert attempt to find Dominic, but after finding Kit Garrison's cháteau empty, they are forced to return to Canada in hopes of finding the child from home.
After a year of living back in Avonlea, Anne and Gilbert arrive at the local train station to collect Dominic, whom they are adopting as their own. They decide to leave Green Gables to Diana and Fred while they "make a new life, but built on all the old foundations" and "build a good home, and raise a family, with lots of scope for the imagination."
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