document_title: Agents of S.H.I.E.L.D. (season 5)

question_text: when does episode 11 of agents of shield come out

root

text: Agents of S.H.I.E.L.D. (season 5)

text: Episodes[edit]

text:
No.
overall
No. in
season
Title
Directed by
Written by
Original air date
U.S. viewers
(millions)

89

90
1

2
"Orientation"
Jesse Bochco

David Solomon
Jed Whedon & Maurissa Tancharoen

DJ Doyle
December 1, 2017 (2017-12-01)
2.54[1]

An unknown group seizes Phil Coulson, Melinda May, Daisy Johnson, Jemma Simmons, Alphonso "Mack" MacKenzie, and Yo-Yo Rodriguez and transports them to a space station through a monolith; Leo Fitz is left behind. Upon arrival, the team is heralded by Virgil, who dies soon after. They quickly regroup and meet Deke, who reveals that the space station, known as the Lighthouse, was built nearly ninety years prior to his knowledge. Hoping to send a message back to Earth, May and Simmons discover it is no longer habitable and appears to have been torn apart by a cataclysmic event. At the same time, Coulson discovers that the monolith transported them not through space, but through time. Elsewhere, Mack and Yo-Yo are held captive by the Kree, who maintain control of the station by encouraging bloodshed among its inhabitants. Back in the station, the team attempts to blend in by agreeing to work for Grill and meets Tess, who saves them during the Renewal. Coulson goes with Tess to learn more about Virgil's message and finds a hidden notebook. Meanwhile, Simmons is taken to meet Kasius, the Kree leader of the station, and is inducted as one of his servants. Later on, Johnson confronts Deke in his framework simulation, where he reveals she is the reason the Earth was torn apart.

91
3
"A Life Spent"
Kevin Hooks
Nora Zuckerman & Lilla Zuckerman
December 8, 2017 (2017-12-08)
1.93[2]

In preparation for Lady Basha's arrival, Simmons is given the task of helping an Inhuman champion control her abilities for her exhibition ceremony. Simmons is successful but becomes distraught when Kasius sells his champion to Lady Basha. While the team adapt to their new jobs, Grill becomes suspicious of the new arrivals and sends one of his men to spy on them. During a space expedition, Coulson, May, Mack, and Tess struggle to make sense of Virgil's notebook and discover a radio transmission supposedly originating from the Earth's surface. Grill's spy reports back, only to be framed as a traitor and exiled to die on Earth. Meanwhile, Yo-Yo helps Johnson get ahold of a Kree tablet to try and save Simmons. However, her mission is thwarted by Deke, who reports her to Kasius. Johnson is captured and taken to become his new champion.

92
4
"A Life Earned"
Stan Brooks
Drew Z. Greenberg
December 15, 2017 (2017-12-15)
1.84[3]

Johnson meets Ben, one of Kasius' other champions, capable of telepathy. Using Ben to corroborate their story, Kasius interrogates Johnson and Simmons about the others. They are able to convince Kasius they came alone, though he remains suspicious. During the interrogation, Ben learns that Kasius intends to destroy the station and everyone on it once he acquires enough profits to leave. Deke returns to the others to gather new information as they learn about the mysterious Level 35. Recognizing his father's voice on the radio transmission, Deke agrees to help them gain access to the restricted floor. There, they discover that Kasius is attempting to breed Inhumans. May also discovers Deke lied about Johnson's whereabouts, but get interrupted by Sinara's arrival. Coulson leaves with Deke, while May fights her alone. Meanwhile, Grill sends Mack to confront someone who owes him payment. Later, Fitz is revealed to be one of the bidders who have arrived for Johnson's exhibition.

93
5
"Rewind"
Jesse Bochco
Craig Titley
December 22, 2017 (2017-12-22)
2.40[4]

Shortly after the others are taken from the diner, Fitz is taken into military custody and interrogated by General Hale and her subordinates about the others' whereabouts, as well as their involvement in the assassination attempt on General Talbot and deaths of Jeffrey Mace and Holden Radcliffe. For the next six months, Fitz is given special privileges to allow him to locate the team, but he fails. With help from Lance Hunter, Fitz escapes the military facility and tracks down Enoch, the man who took the others. Enoch explains that a prophecy foretold these events and takes them to meet Robin, the daughter of Charles Hinton, who expresses her prophetic abilities through drawings. Tracked by the military, Enoch helps them escape to a secret bunker, where Fitz learns that he was left behind so he could save the team. Determined to help them, Fitz and Hunter break back into the same military facility to obtain a cryogenic pod; they find all their old S.H.I.E.L.D. tech, including the Zephyr, which they use to escape. Using the pod, Fitz is put to sleep for the next 74 years. He is awoken by Enoch, who prepares him for their arrival at the Lighthouse.

94
6
"Fun & Games"
Clark Gregg
Brent Fletcher
January 5, 2018 (2018-01-05)
2.49[5]

Down in the station, the Kree have harvested some of the youth for an unscheduled Terrigenesis ceremony. Flint, a friend of Tess, experiences a successful Terrigenesis and is taken into hiding with the team by Yo-Yo. Tess is killed and hanged for Flint's disappearance. During a dinner with the other bidders, Fitz quickly proves himself as a formidable character, earning Kasius' respect. The first exhibition is then held between Ben and May, resulting in May being sent to the surface, while Ben is killed for lying during Johnson's interrogation. Meanwhile, Grill finds the team and holds them captive, until Flint saves them using geokinesis against Grill, killing him. The unexpected arrival of Kasius' brother prompts Sinara to battle Johnson for her exhibition. During which, Johnson, Fitz, and Simmons are able to subdue the others and escape. During the escape, Simmons proposes to Fitz.

95
7
"Together or Not at All"
Brad Turner
Matt Owens
January 12, 2018 (2018-01-12)
2.31[6]

Johnson, Fitz, and Simmons are hunted throughout the station and discover the station is being sustained by gravitonium. Fitz is shot during the chase. Deke finds them and takes them to the rest of the team. Reunited, the team plot their escape to the Earth's surface by using a trawler and Deke's antigravity device. After learning about Tess' death, Flint decides to stay on the station to protect the remaining inhabitants. Mack and Yo-Yo agree to stay behind and help him. Coulson, Johnson, Fitz, Simmons, and Deke fly to the surface, but get caught in the gravity storm. Meanwhile, Kasius kills his brother and plots to regain his father's acceptance by recapturing Johnson with help from Sinara. On the surface, Enoch finds May, but their meeting is cut short by the approaching gravity storm. A mysterious figure takes them to safety, where they are greeted by an elderly Robin.

96
8
"The Last Day"
Nina Lopez-Corrado
James C. Oliver & Sharla Oliver
January 19, 2018 (2018-01-19)
2.41[7]

Coulson, Johnson, Fitz, Simmons, and Deke survive the crash and reunite with May in the Zephyr. They meet Samuel Voss, an acquaintance of Deke's father. Up on the Lighthouse, Kasius punishes the inhabitants by cutting off their resources. Flint, Mack, and Yo-Yo go to retrieve the weapons hidden by Fitz and discover Kasius has released the Vrellnexians onto the lower levels. They defeat the Vrellnexians and rescue the remaining inhabitants. On the surface, the team struggle to get answers from Robin and find a machine built into the Zephyr. While Fitz and Simmons attempt to remove Johnson's inhibitor, May and Coulson find a shard of the time monolith in Voss' locker. Deke questions Voss about the shard, which belonged to his parents, and gets knocked out. Voss then tries to kill Johnson and impales Robin to prevent her from giving the team answers. Before dying, Robin is comforted by May as she reveals their history and finally tells May how to save the world, which requires Flint. In the past, May is shown taking care of Robin and encouraging Fitz to build his machine so they can travel through time.

97
9
"Best Laid Plans"
Garry A. Brown
George Kitson
January 26, 2018 (2018-01-26)
2.27[8]

As the gravity storm worsens, the team attempt to relaunch the Zephyr. May reveals that using the shard, Flint can recreate the monolith allowing the team to travel back through time. Similar to the Lighthouse, Fitz and Simmons find gravitonium on the Zephyr and realize the designs were manufactured by Fitz himself. On the station, Flint, Mack, and Yo-Yo successfully gain control of the lower levels. Tess is resurrected by Kasius and sent to relay demands for him, who threatens to detonate explosives killing everyone if they are not met. Mack and Yo-Yo retaliate by placing the explosives on Level 35, the medical floor, which would prevent Kasius from breeding more Inhumans. During a standoff with Kasius, Flint evacuates the inhabitants to the upper levels and uses the remaining explosives to separate them from the lower levels, out of Kasius' reach. Using the gravity storm to her advantage, May navigates the Zephyr into space as Johnson battles Sinara again, this time killing her. Johnson then contacts Mack while the team are heading for the Lighthouse. Kasius reveals he has his own seer telling him what will happen.

98
10
"Past Life"
Eric Laneuville
DJ Doyle
February 2, 2018 (2018-02-02)
2.22[9]

While Enoch stays on the Zephyr, the team focus on getting to Flint and rescuing the other Inhumans from captivity. Distraught over Sinara's death, Kasius infects Tye, the Inhuman trainer, with odium, a substance that excites maniacal behavior and sends him after the team. During a fight with Tye, Coulson seemingly gets infected. Kasius' seer is revealed to be Yo-Yo from the future, one who has lived through the time loop and endured endless torture. She reveals that the team's ultimate return will bring the end of the world and that the only solution is to let Coulson succumb to his infection. As Flint recreates the monolith, Mack leaves to find Yo-Yo and witnesses her future self's death by Kasius, who consumes the remaining odium and fights him. With help from Simmons, Mack kills Kasius and reunites with Yo-Yo, who is still alive in their timeline. Deke returns to the Zephyr in time to save Enoch and fix the machine, although seemingly at the cost of both of their lives, as the team returns to the present.

99
11
"All the Comforts of Home"
Kate Woods
Drew Z. Greenberg
March 2, 2018 (2018-03-02)
1.90[10]

Back in their time, the team meets Noah, Enoch's successor, who has been monitoring the world from the Lighthouse in their absence. Coulson, May, Fitz, Simmons, Mack, and Yo-Yo investigate a beacon emitting from Earth, which fits Voss' description of the first signs of the end. Reluctant to join the team, Johnson stays behind and discovers that Deke survived the explosion. After he is arrested, Johnson is forced to leave the bunker to save him and prevent him from exposing the team, who are now most wanted. Upon finding the beacon, the team reunites with Agent Piper, who is revealed to be working for General Hale. They are then ambushed by a group of Hale's robotic soldiers, led by an assassin. During the fight, the assassin attempts to kill Mack with their chakram, but Yo-Yo blocks the attack and loses both her arms. Piper realigns with the team as they retreat to the Lighthouse with the beacon, which detonates as Noah sacrifices himself to save Johnson and Fitz. Later, General Hale criticizes the assassin, revealed to be her daughter, Ruby, for jeopardizing the operation. In Philadelphia, General Hale recruits Carl Creel to join her team, to which he reluctantly agrees.

100
12
"The Real Deal"
Kevin Tancharoen
Jed Whedon & Maurissa Tancharoen & Jeffrey Bell
March 9, 2018 (2018-03-09)
2.04[11]

The destruction of three monoliths caused by the beacon's explosion creates a dimensional rift that manifests the fears of the team. Using gravitonium, Fitz creates a device capable of sealing the rift; Coulson volunteers to deliver the device before he collapses. The team finds out about his infection, which is a result of the Ghost Rider burning off the life-sustaining GH.325 drug within him. The team is forced to accelerate their plan when Yo-Yo is attacked by a manifestation of a Simmons LMD. Coulson sends Deke to the surface to call for backup, then proceeds alone to seal the rift. He encounters a manifestation of Mike Peterson, who claims that Coulson's experiences are just a dream, and that he is dying on an operating table after the Battle of New York. Coulson overcomes his fear, and is saved by the real Peterson, who responds to Deke's call and arrives with several other S.H.I.E.L.D. agents. Coulson and Peterson fight off manifestations of Vrellnexians, Lash, and Hive, before managing to contain the rift. Later, the team organize a wedding ceremony for Fitz and Simmons. General Hale, who is investigating sightings of Johnson, finds evidence that Deke is related to Fitz and Simmons.

101
13
"Principia"
Brad Turner
Craig Titley
March 16, 2018 (2018-03-16)
2.07[12]

When the rift begins to reopen, Fitz sends the team to search for more gravitonium. The team hunt down a lead in Baton Rouge; Mack reunites with his school mate, Tony Caine, who has been helping to redeem scientists coerced into working for Hydra through Cybertek Industries. Caine states that the gravitonium was in transport aboard the Principia, a Cybertek ship that went missing. Deke realizes the gravitonium may have been activated by the lightning storm, sending the Principia into the sky. Following Deke's theory, the team find the Principia suspended in the atmosphere and discover most of the gravitonium is gone, leaving only a small portion to sustain the ship. As Mack collects the remaining gravitonium, a group of Hale's robotic soldiers ambush them. The team successfully escapes the ship before it plummets back to Earth. Upon hearing his mother's saying in an exchange between Simmons and Yo-Yo, Deke realizes Fitz and Simmons are his grandparents. Meanwhile, General Hale continues to form her team by attempting to recruit Werner von Strucker, whose memory had been enhanced after S.H.I.E.L.D. revived him from a vegetative state. While Strucker initially rejects Hale's offer, Ruby convinces him to stay.

102
14
"The Devil Complex"
Nina Lopez-Corrado
Matt Owens
March 23, 2018 (2018-03-23)
2.07[13]

When Simmons is attacked by a manifestation of an astronaut, Fitz accelerates work on sealing the rift but struggles to figure out how to compress the gravitonium. Under stress, Fitz encounters his Framework alter ego, The Doctor, who taunts him. On the Zephyr, Coulson, May, and Agent Piper capture General Hale; however, she reveals that her capture was intentional, and threatens to detonate explosives strapped to Creel if S.H.I.E.L.D. fails to comply. Anton Ivanov arrives, and Coulson agrees to accompany them despite May's unease. Simmons, Mack, and Yo-Yo are attacked by one of the robots that Mack brought back from the Principia. While dealing with technical malfunctions, Johnson is rendered unconcious by another robot and awakens restrained by The Doctor. Simmons finds Fitz trying to remove Johnson's inhibitor. She deduces that The Doctor was not a result of the fear dimension, but a projection of Fitz's internal turmoil. He reprogrammed the robots to isolate Johnson, whose powers he uses to compress the gravitonium and seal the rift completely. Deke attempts to comfort Simmons, while also revealing himself as their grandson. Later, Hale speaks to her superior, who hands her a vial of odium and reminds her of her Hydra allegiance.

103
15
"Rise and Shine"
Jesse Bochco
Iden Baghdadchi
March 30, 2018 (2018-03-30)
1.88[14]

28 years prior, a young Hale was assigned by Daniel Whitehall to infiltrate the U.S. Air Force, and also to be impregnated via in vitro fertilization with Ruby to create Hydra's next leader. In the present, Hale fails to convince a recently-awakened Talbot to release Hydra contraband. She also informs Coulson that an alien alliance known as the Confederacy had made contact with Hydra and offered to assist Earth in an impending invasion; she intends to betray the alliance by using Whitehall's particle-infusion chamber and gravitonium to create a Destroyer of Worlds. Though she intends for it to be Ruby, she suggests Johnson may be a better fit. Upon hearing this, Coulson denies her offer and reveals his trip to the future, but she doesn't believe him. Back at the Lighthouse, May and Johnson struggle to deduce Hale's motivations and locate Coulson. Johnson, who has been left as the de facto head of S.H.I.E.L.D., refuses to trust Fitz and goes in search of Robin. Simmons tells Fitz about their relation to Deke, suggesting they will survive despite the odds.

104
16
"Inside Voices"[15]
Salli Richardson-Whitfield
Mark Leitner
April 6, 2018 (2018-04-06)
TBD

105
17
"The Honeymoon"[16]
Garry A. Brown
James C. Oliver & Sharla Oliver
April 13, 2018 (2018-04-13)
TBD

106
18
"All Roads Lead..."[17]
Jennifer Lynch
George Kitson
April 20, 2018 (2018-04-20)
TBD

tag: table

text: Cast and characters[edit]

text: Main[edit]

text:
Clark Gregg as Phil Coulson[18]
Ming-Na Wen as Melinda May[18]
Chloe Bennet as Daisy Johnson / Quake[19]
Iain De Caestecker as Leo Fitz[18]
Elizabeth Henstridge as Jemma Simmons[18]
Henry Simmons as Alphonso "Mack" MacKenzie[18]
Natalia Cordova-Buckley as Elena "Yo-Yo" Rodriguez[20]

tag: ul

text: Recurring[edit]

text:
Jeff Ward as Deke Shaw[21][22]
Joel Stoffer as Enoch[23]
Eve Harlow as Tess[22]
Dominic Rains as Kasius[23]
Florence Faivre as Sinara[23]
Pruitt Taylor Vince as Grill[22]
Coy Stewart as Flint[22]
Catherine Dent as Hale[24]
Dove Cameron as Ruby[24]
Brian Patrick Wade as Carl Creel[25]
Spencer Treat Clark as Werner von Strucker[26]
Adrian Pasdar as Glenn Talbot[27]

text: Notable guests[edit]

text:
Lola Glaudini as Polly Hinton[28]
Nick Blood as Lance Hunter[29]
Briana Venskus as Piper[25]
J. August Richards as Mike Peterson / Deathlok[30]
Maximilian Osinski as Davis[31]
Zach McGowan as Anton Ivanov / The Superior[32]
Reed Diamond as Daniel Whitehall[27]
Adam Faison as Jasper Sitwell[27]
Joey Defore as Wolfgang von Strucker[27]

text: Production[edit]

text: Development[edit]

text: In January 2017, ahead of the mid-season premiere of the fourth season, ABC's Channing Dungey said that she was "very bullish" about S.H.I.E.L.D.'s future, feeling that "the episodes just keep getting better and stronger".[33] The series was renewed for a fifth season of 22 episodes on May 11, with ABC looking to lower the cost of the series moving forward,[34][35] by reducing its budget and licensing fee.[36] Asked whether she had considered giving the season a shorter episode order, Dungey noted that every season of the series had been 22 episodes long, and she felt its prior success in delayed viewership and overseas justified continuing that. She added that the "show has continued to grow creatively every season. I feel like last season [was] its strongest creatively yet. I'm very excited for what we have planned for Season 5."[37] It was reported that Disney, the parent company of Marvel Television, ABC Studios, and ABC, had given a mandate to ABC to renew Agents of S.H.I.E.L.D. "despite the desire by some at the network to end the series".[38]

tag: p

text: Writing[edit]

text: In May 2017, ahead of the fifth season renewal, showrunner and executive producer Jed Whedon said the writers were not sure what would happen in the season, and that it would be "by the seat of our pants."[39] The season explores the ramifications of Phil Coulson making a deal with Ghost Rider at the end of the previous season, as well as the groundwork laid by Aida for humans to fear S.H.I.E.L.D. and Inhumans. Whedon stated, "The public perception of S.H.I.E.L.D. is at an all-time low, so we have not resolved that, and there will be still more fallout from it."[40] Moving beyond the Framework reality established at the end of season four, Tancharoen said the emotional impact from the characters' experiences would "be something that resonates throughout the season", especially for Fitz and Mack.[41] When asked if the season would be broken into pods as with season four, executive producer Jeffrey Bell said, "A 22-episode arc is a lot for people to hold onto. By breaking it up into either smaller arcs or different pods, by introducing a set of antagonists and putting them down, or moving from space to space, our experience has been that it's something the viewers enjoy, and it makes it a little easier to digest when you're telling some of these stories." However, Whedon noted that it would depend on how the season would be aired as to where the story is broken up.[40] The season ultimately was broken into two pod story arcs, with each pod having a different emphases, but the whole season having an arc "that will pay off".[42] The writers for the season began work at the end of May 2017.[19]

text: First pod[edit]

text: The first pod of the season was informally referred to by the production team as S.H.I.E.L.D. in Space.[42] Ahead of the season premiere, executive producer Maurissa Tancharoen said that "every year we reset the series, and this year we definitely knew that it would be the most giant reset to date [with the characters going to space]. Just creatively across the board for everyone, art direction, all of it, our sets, you'll see an overhaul."[43] On the move to space, Whedon commented, "Last year was about tearing everybody apart. We spent a lot of time doing that ... So our goal this year was... putting [this] family together in an intense situation [that] will end up causing drama internally, inevitably.... we've spent this many years with them, let's throw them on the craziest roller coaster adventure we could think of."[44] The main characters eventually return home to their time period.[45]

text: Second pod[edit]

text: The second pod of the season includes the series' 100th episode, which Whedon and Tancharoen described as a "game changer" that would "shake up" the rest of the pod emotionally, to the point that "nothing will be the same after" it.[46] Bell added the episode featured "a device that grows out of our current storyline and plot that allows us to look back and reflect on where we've come from, turn over a couple cards that people will be excited about and then also celebrate the show and people on it."[47] Loeb also felt the episode "finishes up some stories that perhaps [viewers] didn't even know we haven't revealed".[48] The episode featured Fitz and Simmons getting married, which Tancharoen said was included because "it was about damn time. After 100 episodes, they needed to get married." Coulson's deal with Ghost Rider was also revealed, which was to have the Kree serum that revived him after his death be burned off, resulting in his chest wound slowly killing him. Whedon said that Coulson has "come to terms with it. It's something he actually had to come to terms with a long time ago when he was discovering the T.A.H.I.T.I. Project and everything that had been done to him. I don't think he wants to go through any of that again. He's ready for nature to take its course... he seems as much at peace with it as you can be considering there's so much he's done in a world that thinks he's dead."[30]

text: At the end of February 2018, the writers were planning the end of the pod, and were planning for the final episode to be able to serve as both a season and series finale, with some elements that could be adjusted based on whether the series was renewed for a sixth season or not. Whedon added, "we're ready for if this is the end. We're definitely going to make it rewarding either way."[46]

text: Casting[edit]

text: Main cast members Clark Gregg, Ming-Na Wen, Chloe Bennet, Iain De Caestecker, Elizabeth Henstridge, and Henry Simmons return from previous seasons as Phil Coulson, Melinda May, Daisy Johnson / Quake, Leo Fitz, Jemma Simmons, and Alphonso "Mack" MacKenzie, respectively.[18] Before the 2017 New York Comic-Con, it was revealed that Natalia Cordova-Buckley had been promoted to series regular for the season, after recurring in the past two seasons as Elena "Yo-Yo" Rodriguez.[20] De Caestecker also portrays "The Doctor", the version of Fitz from the Framework reality.[49]

text: In September 2017, former series regular Nick Blood was announced as returning to reprise his role of Lance Hunter. Blood left the series during the third season, to star in the spin-off series Marvel's Most Wanted, which never came to fruition.[29] Adrianne Palicki, who portrayed Bobbi Morse and also left in the third season for Most Wanted with Blood, expressed interest in October 2017 in returning to guest star in the season, saying she "would absolutely come back" if asked.[50] Also returning from earlier in the series are Joel Stoffer as Enoch,[23] who was simply credited as "silhouetted man" in his previous appearance,[51] and Lola Glaudini as Polly Hinton.[28] Polly's daughter Robin also appears, portrayed by multiple actresses,[52] after the character made an uncredited appearance in "Ascension":[53] Lexy Kolker appears as 7-years-old Robin, Ava Kolker appears as 12-year-old Robin,[52] and Willow Hale appears as an older Robin, when she is also known as The Seer.[52][54]

text: Additional returning actors include Brian Patrick Wade as Carl Creel,[25] J. August Richards as Mike Peterson / Deathlok,[30] Spencer Treat Clark as Werner von Strucker,[26] Zach McGowan as Anton Ivanov / The Superior,[32] Adrian Pasdar as Glenn Talbot, Reed Diamond as Daniel Whitehall, and Raquel Gardner as Carla Talbot,[27] along with Briana Venskus and Maximilian Osinski as S.H.I.E.L.D. agents Piper and Davis.[25][31] The characters Lash and Hive return from earlier seasons for the series' 100th episode.[55][30] Younger versions of Jasper Sitwell and Wolfgang von Strucker also appear, portrayed by Adam Faison and Joey Defore, respectively.[27] Older versions of the characters had been respectively portrayed by Maximiliano Hernández (in the MCU films and first season of the series) and Thomas Kretschmann (in the MCU films).[56][57]

text: Jeff Ward was cast in a recurring role in August 2017,[21] and was revealed in October to be portraying Deke Shaw, alongside other newcomers Eve Harlow as Tess, Coy Stewart as Flint, and Pruitt Taylor Vince as Grill.[22] The next month, Marvel revealed that Dove Cameron had joined the season in an unspecified role,[58] which was revealed in January 2018 to be the character Ruby, the daughter of Catherine Dent's General Hale.[24] Other recurring guests for the season include Dominic Rains as Kasius, Florence Faivre as Sinara, Jay Hunter as a Kree watch commander, Tunisha Hubbard as Ava,[23] and Shontae Saldana as Candice Lee.[25]

text: Design[edit]

text: After leaving the series during the fourth season, costume designer Ann Foley returned for the first two episodes of the fifth season, before handing over to Whitney Galitz, who had assisted her on the previous few seasons, and Christann Chanell.[59] The opening for "Orientation" is reminiscent of the sequence for "4,722 Hours", forgoing the title card and having the typeface silently fading onto the screen.[60] The subsequent episodes of the season feature a title card with the series name against a backdrop of a destroyed Earth.[61]

text: Filming[edit]

text: Filming for the season began on July 20, 2017.[62][63] In May 2017, Gregg expressed interest in directing an episode in the season,[64] and confirmed that September he would be directing an episode,[65] which was the sixth of the season, "Fun & Games". Gregg felt the idea of directing was "daunting" as he would need to be doing "five weeks of double duty", needing to act in addition to the various aspects of directing the episode.[66] He reached out to fellow S.H.I.E.L.D. directors Kevin Tancharoen, Billy Gierhart, and Jesse Bochco to get tips on filming for the series.[67] Bennet felt Gregg was able to bring out performances from the cast "that another director probably wouldn't have been able to because he knows the characters so well."[68] Gregg added that he had "shorthand with [the other actors] about what we want to try to do, ways the script could give us a chance to push us into new territory. They all also have deep and interesting ideas that helped me." Gregg also received a deeper appreciation for some of the other departments on the series he normally does not encounter on a regular basis and the work they contribute to each episode.[66] The scene for Fitz and Simmons' wedding in "The Real Deal" was filmed in "a very remote location" in Placerita Canyon State Park.[69]

text: Marvel Cinematic Universe tie-ins[edit]

text: In terms of having connections to Marvel's Inhumans, Whedon felt it was doubtful, but noted "we also want there to be a bigger reward for people who watch all of [the MCU series], so they'll start to see that there's lines connecting it. It's more fun for people who are playing the whole game, but if you don't, you'll still have a great rollercoaster ride."[70] In November 2017, Whedon said that the season would not have moved the characters to space if the recent MCU films like Guardians of the Galaxy Vol. 2 and Thor: Ragnarok had not been exploring that part of the MCU, but that this would remain a "thematic tie". He explained that they had moved on from the overt tie-ins with the MCU films that the series utilized in earlier seasons because "we have our own mythology. That started to be much more interesting to us, and hopefully to the audience, just digging that stuff up. No one wants to come to see our show to see another show."[43] Whedon added in March 2018 that the then-upcoming release of Avengers: Infinity War would similarly "open a new playground" for the series to explore.[71]

text: The season features the Kree, an alien race that previously appeared in the first, second and third seasons, and in the film Guardians of the Galaxy.[72] The fictional element Gravitonium also returns, having first appeared in the first season.[73]

text: Release[edit]

text: Broadcast[edit]

text: The season began airing in the United States on ABC on December 1, 2017,[68] beginning once Inhumans finished airing its episodes.[74] It is set to run for 22 episodes,[35] with a short hiatus for the airing of the 2018 Winter Olympics between "Past Life" and "All the Comforts of Home".[45] The season is intended to conclude on May 18, 2018.[75]

text: The fifth season moved to a Friday timeslot, alongside the seventh season of ABC's Once Upon a Time, with ABC chief Channing Dungey explaining, "We've turned Friday into more of a destination for our fantasy and science fiction fans. Once Upon a Time and S.H.I.E.L.D. are airing on the same night for the first time, which is giving many fans of both shows what they've been asking for for a long time."[76] Andy Kubitz, executive vice president for program planning and scheduling at ABC, added that ABC had "confidence that [the S.H.I.E.L.D.] core audience... [would] travel with it" to its new Friday time slot. He continued, "The great thing about Friday night for these shows is it gives three days of downtime for a lot of these younger viewers to be able to catch up on it. You've got Saturday and Sunday viewing that will be able to be counted into our C3 to help us monetize it."[36]

text: Marketing[edit]

text: The main cast of the season appeared at New York Comic Con on October 7, 2017, where they promoted the season and debuted the first 20 minutes of the first episode of the season.[77][68] A shortened version of the footage was released by ABC and Marvel on November 26, 2017, as a sneak peek before the season premiere later that week.[78] The cast and executive producers will appear at WonderCon 2018 on March 24, to promote the rest of the season, and show a sneak peek of an upcoming episode.[79] Marvel released three roundtable discussions in support of the series reaching 100 episodes. Hosted by Patton Oswalt, who has portrayed The Koenigs in the series, the first was released on March 8, 2018 with the women of the series. The second, which release on March 9, featured the full cast, while the third debuted on March 12 with the executive producers. All premiered on Marvel.com while also being available on Marvel and the series' social media platforms and Marvel's YouTube channel.[80] The executive producers, main cast members except Gregg, and Ward appeared at WonderCon 2018 to promote the remainder of the season and answer fan questions. A clip from the end of "Rise and Shine" was also shown.[71]

text: "The Road to 100"[edit]

text: In December 2017, Marvel announced the "Marvel's Agents of S.H.I.E.L.D.: The Road to 100" art program to commemorate the series reaching 100 episodes. The program features five posters, one for each season of the series, representing "key pivotal moments of each season". The art also appeared as variant covers to select titles published by Marvel Comics in March 2018. Megan Thomas Bradner, Vice President of Development and Production, Live Action at Marvel Television, called reaching 100 episodes "quite a feat... and it felt special enough to share with the fans that got us here. By working with some of our favorite comic artists, we felt we could show a large variety of meaningful moments and the characters that helped get us to a 100." She added that the program "commemorates these creators, the actors, the characters and a hundred amazing stories".[81]

text: The first poster, highlighting season one, was created by Dale Keown. It features four panels, showcasing the moment Coulson and Grant Ward approached Skye in her van from the pilot, Coulson learning he survived the Battle of New York from the T.A.H.I.T.I. program and a drug from a half-dissected Kree alien corpse, the moment Grant Ward revealed he was a Hydra agent by killing Victoria Hand, and Fitz giving Simmons the last remaining oxygen tank and declaring his love for her in the season one finale.[81] The second poster, highlighting season two, was created by Daniel Acuña. The scenes depicted are Coulson writing the Inhuman map on a wall; Daisy coming out of her Terrigenesis; the S.H.I.E.L.D. team, Bobbi Morse, Lance Hunter, and the "real S.H.I.E.L.D." logo; and May cradling the deceased body of Katya Belyakov.[82] The third poster, highlighting season three, was created by Nick Bradshaw. The poster shows the S.H.I.E.L.D. team and Hive, along with the Hydra logo breaking the S.H.I.E.L.D. logo and its tentacles covering much of the poster. Specific moments highlighted include Fitz and Simmons after she has been pulled from the portal to Maveth, Coulson killing Ward on Maveth, and the formation of the Secret Warriors.[83] The fourth poster, highlighting season four, was created by Rahzzah. It highlighted the introduction of Ghost Rider, Life Model Decoys (LMD) and the Framework reality, Jeffrey Mace sacrificing himself in the Framework, and Coulson's kiss with the May LMD.[84] The fifth and final poster, highlighting this season, was created by Stonehouse. Flanked on either side are Kasius and Sinara, with the S.H.I.E.L.D. team and Deke in the center in front of the destroyed Earth of the future as well as Fitz reuniting with Simmons.[85]

text: Reception[edit]

text: Ratings[edit]

text:
No.
Title
Air date
Rating/share
(18–49)
Viewers
(millions)
DVR
(18–49)
DVR viewers
(millions)
Total
(18–49)
Total viewers
(millions)

1/2
"Orientation"
December 1, 2017
0.7/3
2.54[1]
0.6
1.72
1.3
4.27[86]

3
"A Life Spent"
December 8, 2017
0.5/2
1.93[2]
0.7
1.68
1.2
3.61[87]

4
"A Life Earned"
December 15, 2017
0.5/2
1.84[3]
N/A
N/A
N/A
N/A

5
"Rewind"
December 22, 2017
0.6/3
2.40[4]
N/A
N/A
N/A
N/A

6
"Fun & Games"
January 5, 2018
0.7/3
2.49[5]
0.7
N/A
1.4[88]
N/A

7
"Together or Not at All"
January 12, 2018
0.6/3
2.31[6]
0.7
1.75
1.3
4.07[89]

8
"The Last Day"
January 19, 2018
0.6/3
2.41[7]
0.8
1.82
1.4
4.23[90]

9
"Best Laid Plans"
January 26, 2018
0.6/3
2.27[8]
0.6
1.72
1.2
4.00[91]

10
"Past Life"
February 2, 2018
0.6/3
2.22[9]
0.6
1.74
1.2
3.96[92]

11
"All the Comforts of Home"
March 2, 2018
0.5/2
1.90[10]
0.7
1.68
1.2
3.58[93]

12
"The Real Deal"
March 9, 2018
0.5/2
2.04[11]
0.7
1.72
1.2
3.78[94]

13
"Principia"
March 16, 2018
0.5/2
2.07[12]
0.6
1.58
1.1
3.65[95]

14
"The Devil Complex"
March 23, 2018
0.5/2
2.07[13]
TBD
TBD
TBD
TBD

15
"Rise and Shine"
March 30, 2018
0.5/2
1.88[14]
TBD
TBD
TBD
TBD

text: Critical response[edit]

text: The review aggregator website Rotten Tomatoes reports a 100% approval rating, with an average rating of 7.71 out of 10, based on 11 reviews.[96]

text: Reviewing the premiere episodes, Merrill Barr of Forbes felt moving the series to space was not a reboot of the series, rather "a continuation, in some form, of the story set-up at the end of last season. The show is not different. The team is still trying to save the world. They're just doing it in a new place and new way. Overall, there's a lot to love about what S.H.I.E.L.D. is doing in the new season. It's a crazy ride fans are going to be glad they've stayed aboard for. The "Agents of Hydra" run [at the end of season four] was a bit weak. The space run is so far proving to be anything but."[97] Jon Negroni of We Got This Covered praised the show for its delightful twists and turns and for not relying on what's already worked, adding that "it's a testament to the show's ongoing experimentation that we can watch these characters get to space and believe that this is far from their final frontier." He felt, however, that there was "little room for pathos and seeds for character development" and that "we know almost nothing about where these individuals are at emotionally because of how fast we blazed through the finale."[98] Alex McLevy of The A.V. Club praised the move to space, saying that "if season four was Agents of S.H.I.E.L.D. firing on all cylinders, especially during its stellar 'Agents of Hydra' arc in the Framework, then this is the look of a show that knows it has mastered its storytelling, and is confidently expanding the scope of its ambition."[99]

text: Accolades[edit]

text:
Year
Award
Category
Recipient
Result
Ref.

2018
TVLine's Performer of the Week
Performance in "The Last Day"
Ming-Na Wen
Nominated
[100]

Visual Effects Society Awards
Outstanding Visual Effects in a Photoreal Episode
Mark Kolpack, Sabrina Arnold, David Rey, Kevin Yuille, Gary D'Amico for "Orientation, Part 1"
Nominated
[101]

Saturn Awards
Best Superhero Television Series
Agents of S.H.I.E.L.D.
Pending
[102]


document_title: Chicago Crossover

question_text: who plays erin lindsay's brother on chicago pd

question_tokens: who plays erin lindsay 's brother on chicago pd

text: Chicago Crossover

text: Plot[edit]

text: The case opens up where the Chicago Fire episode "Nobody Touches Anything", left off; where the CFD firefighters rescued a man from a burning building and in his possession, he held a box full of child pornography, a box which was dropped during Lieutenant Kelly Severide's (Taylor Kinney) attempt to save the man's life. The CFD then turned this evidence over to the Chicago police. Detective Erin Lindsay has a personal stake in the case: her half-brother is involved somehow and in New York City. Benson gets a call from Voight that his team is on a plane to Manhattan to help the SVU solve the case.

text: Lindsay explains to the SVU about how her half-brother Teddy Courtney (Lou Taylor Pucci) went missing when they were younger and how her mother thought that somehow he ran away to New York. Benson tells Lindsay that NCMEC ran Teddy’s face through their system and they found more images, and concluded that he was connected to a pedophile ring. While searching for Teddy they learn from the National Center for Missing and Exploited Children that another young boy, Henry Thorne (Christian Godwin), is being victimized on a live stream, although at the time no one could track down where the feed originated from. They learn from the photos found in the Chicago fire that the ring is called the "Chess and Checkers Club" and it originated out of Chicago back in 2004; NCMEC pushing the federal government to shut down the website, but to no avail.

text: Later the detectives discover through advanced facial recognition that Teddy has a juvenile record and went under the name Teddy Voight, who five years prior was arrested for solicitation. Lindsay explains to Detective Jay Halstead (Jesse Lee Soffer) that they knew Sgt. Voight before. SVU detectives Nick Amaro (Danny Pino) and Amanda Rollins (Kelli Giddish) go to a youth center where Teddy was remanded and learn that Teddy frequently ran away from the center. Security guard George Turner (Danny Mastrogiorgio) doesn't remember Teddy, but Miss Bagley (Donnetta Lavinia Grays) recommends that the detectives talk to Jocelyn Cerpaski (Isabel Shill), who left the center to start a better life. Jocelyn initially doesn't want to talk to the detectives, but she directs them to the piers where Teddy solicits. Rollins and Halstead find him and bring him in.

text: Lindsay tries to talk to Teddy, but he refuses. Benson tells Lindsay that they have him on narcotics possession. Benson tries to get Teddy to open up about what happened to him by letting him know that another boy is in danger, but he remains silent. Amaro and Rollins go back to Jocelyn, and learn that Teddy assaulted her. Later, Benson and Lindsay push Teddy, reassuring that he raped Jocelyn because his abusers forced him to. Teddy breaks down and admits what he did, reasoning that if he hadn't, the pedophile ring would have had someone else hurt her worse. Teddy explains to Benson and Lindsay there was someone they were all afraid of and that at the youth center, one of the security guards helped him escape—George Turner.

text: Benson, Halstead and Lindsay go back to the center to pick up Turner for questioning. He tries to run from them, and in the ensuing struggle Lindsay is injured by a passing bicyclist and hospitalized. When she awakes, Voight is at her bedside, telling her to go back to Chicago and vowing to help Teddy. Voight later shows up at SVU and immediately bursts into the interrogation room, where Amaro and Rollins are trying to question Turner. Voight threatens Turner, causing Benson to intervene; she threatens to have Voight arrested if he touches Turner again. However, Benson lets Turner believe that she is going to let Voight hurt him in order to scare him into revealing what he knows. Turner tells them some details involving toll fees that their prime suspect had to pay in order to get back into the city. Voight and Benson go to Teddy, who still refuses to remember that part of his life, so they go back to Jocelyn, undercover as Henry's parents in order to get her to talk. They take her back to SVU, where the detectives put her in an interrogation room with Teddy and a live feed of Henry. She remembers that they drove her across a long bridge, that she was blindfolded, and that she heard the sound of tires on metal. She and Teddy also remember a name: Bob Clinton (Mark H. Dold).

text: The detectives soon learn that Clinton is somewhere on Staten Island, and Benson and Voight rescue Henry and arrest Clinton. Meanwhile, Detectives Amaro and Fin Tutuola (Ice-T)  arrest two others that seem to be involved. At the squad room, Rollins comes up with a laptop, showing video of another child who had been abducted by the ring, this time a young girl. Benson and Voight try to lean on Clinton, who asks for full immunity before he divulges anything. Benson decides to lock him up instead.

text: Hours after Benson sends Clinton to The Tombs, Clinton is shived in a shower while in protective custody by a Lester Davis, who was awaiting trial on armed robbery. SVU learns that Davis got a phone call from a no-name cell phone purchased in Chicago. Voight was told by Linsday that in Chicago, Andrew Llewellyn (the fire victim who had the child porn photos), and the officer guarding him were shot and killed. Benson comments that their suspect fled back to Chicago.

text: Production[edit]

text: "Chicago Crossover" was written by long-time Law & Order franchise writer Ed Zuckerman (teleplay), SVU executive producer/show runner Warren Leight and Law & Order: SVU creator Dick Wolf, and was directed by Steve Shill. This episode marks the first episode to credit Dick Wolf as a writer since the 100th episode of SVU, "Control", back in the fifth season. In this episode, Raúl Esparza (Assistant District Attorney Rafael Barba) and newcomer Peter Scanavino (Detective Sonny Carisi) were absent.

text: Concept and writing[edit]

text: In May 2014, NBC President, Robert Greenblatt announced that Chicago PD, Chicago Fire, and also SVU, could have another crossover episode, after SVU and CPD briefly crossed-over with SVU's fifteenth season's episode, "Comic Perversion" which guest starred Sophia Bush, who portrayed her CPD character, Detective Erin Lindsay; while Ice-T & Kelli Giddish briefly portrayed their characters respectively in the episode "Conventions" of Chicago PD, two episodes which Leight and Olmstead wouldn't call a true 'crossover'.[1]

text: On September 29, creator Dick Wolf said of the crossover, "Warren Leight and Matt Olmstead are two of the best writers in the business. We wanted to figure out a way to create a crime that connects all three shows. Warren, Matt and I came up with a story that was big and unique enough to transit to all three."[2] SVU show runner/EP Warren Leight noted the three-show crossover was Wolf's idea, although Leight said that both he and Chicago Fire/PD show runner/EP Matt Olmstead had their uncertainties about the crossover. Leight admitted, "I think my first reaction was anxiety, I know the fans will like it, but I know how much extra work is involved and also, legitimately these are two different kinds of police shows and I wasn’t sure how the cross-pollination would work."[3][4]

text: Olmstead said of crafting the crossover, "What Warren and I did was jump in, I knew what I was going to hand off to him at the end of [Chicago] Fire and he told me what he was going to hand back over to me at PD and then we fine-tuned and figured out the logistics.” Leight explained that each show's writers wrote for their own show and said that he and Olmstead were on the phone a lot for the plotting stages. "We sent the [Chicago PD] script to Warren to vet since he knows the [SVU] characters better than we do," said Olmstead, Leight jumped in adding, “If there was something I felt was wrong for one of our characters in the PD part, I made my opinions known.” Olmstead assured that it was the same with Leight in writing for the CPD characters, "There was a respectful vetting of both scripts. When I read the SVU script of our characters, it sounded very true, so there weren't any doubts."[4]

text: Leight and Olmstead however worried about having the audience have to tune in to three shows, "That’s a big fear," admits Leight, "But this is still in the Dick Wolf universe where each episode has to stand alone and also link. You don’t want to have a three minute ‘previously on Chicago Fire’ to open up ‘SVU.’ You want that to be as short and clear as possible.  Our episode is stand alone yet the baton passes and there are enough unresolved parts for PD to get another hour out of it. That’s an interesting writing challenge – how much to reveal when and how to stage the investigation and the information over 80 minutes. One good thing what we have is that our episode goes right into PD. Our hope that the audience comes and stays for two hours and that we've made it compelling enough for them to do that."[4]

text:
"You want to see Voight and Benson, you want to see Batman and Superman together for an extended period, working through things. So that’s what we’re doing here."
—SVU showrunner Warren Leight to TV Guide.[3]

tag: div

text: Olmstead and Leight noted about tension and 'fireworks' between Sgts. Benson and Voight, both showrunners were also anxious to pair the two commanding officers on-screen, “There are two very different types of fireworks that we could be talking about." SVU's Leight started, "They have two very different approaches to interrogations and procedure. I think we all know Voight can be more physical and Olivia is a more empathetic detective. Those kind of fireworks take place in both episodes and they are among the most fun scenes in SVU and CPD.” Olmstead added with Leight, “When the Benson character shows up the first time during a pretty grim investigation there’s a smile on [Voight’s] face, he’s happy to see her. There’s this immediate chemistry, immediate shared affection between two very similar characters.[4] There’s a real bond right away between those two characters." Olmstead said in a separate interview with TV Guide, "More than anybody he's ever encountered, I think he respects this person because it's almost like a female version of Voight," Olmstead says. "When the case is starting to go south on the Chicago side and she calls to check in and says, 'What do you need?' he says, 'I need your help.' And he would never admit that to anybody. He's being vulnerable. And her response is, 'I'll get on the next flight out.' So, they just have an affinity towards each other."[3]

text: Continuing storylines[edit]

text: The case is very personal for Detective Lindsay (Bush) because her missing half-brother Teddy (Pucci) might be somehow involved. "It really knocks her sideways," Olmstead says of Lindsay's personal connection to the case. "She admits that she's just trying to keep her head on straight because she has so many competing emotions while trying to be objective and do her job. ... There's some hard-earned advice that Benson is able to impart to Lindsay."[3]

text: There is a slow-burning relationship between Detectives Lindsay and Halstead, Olmstead teased, "[There will be] further slow burning, because, as has been made clear to them by Voight, he doesn't tolerate in house romances." And showrunner Leight teased about sexual tension in TV Guide on November 11, "Lindsay and Halstead have this unrequited crush on each other, and you have Rollins and Amaro with a down-low affair going on. There's a moment or two when it seems like one group notices something about the other. We tease a little bit with that." Leight adds that going to Chicago will bring out a new side to the characters, particularly Amaro. "[He is] much more physical than he gets to be on our show," he adds.[5]

text: The cast members of all three shows spoke highly of the event. Pino called the crossover "a Dick Wolf universe, colliding."[6] Mariska Hargitay was excited when she got the script, "For me reading the Chicago PD script, I was very, very excited about it. Sort of not knowing what it was going to be. And he [Jason Beghe] kept saying 'have you read it, it's so great', and then I read it, and I was absolutely thrilled."[7] Kelli Giddish noted how unprecedented a move the three-show crossover was, "People are really going to enjoy seeing us all kind of mix-and-match. And it's really lucky that we've got three shows on the air right now that we can all do that with, I mean that's really cool."[8]

text: Filming[edit]

text: The shooting of the SVU episode "Chicago Crossover" began on October 9, 2014 in New York City.[9] SVU cast members Hargitay, Pino, and Giddish shot the CPD conclusion episode starting October 22 in Chicago.[10]

text: Leight said of the tight filming schedule in Observer, "We had Kelly and Danny in Chicago for four days and Mariska there for two days. Then we had Halstead (Soffer) and Voight (Beghe) here for four days and Lindsay (Bush) here for two days. It made production very difficult and it meant our next [SVU] episode had to be a legal one to allow for scheduling."[4]

text: Casting[edit]

text: It was announced on September 29, 2014 that three members of Chicago P.D.'s cast would be guest starring in the SVU hour; Sophia Bush (portraying Detective Erin Lindsay), Jesse Lee Soffer (portraying Detective Jay Halstead), and Jason Beghe (portraying Sergeant Henry "Hank" Voight).[2] On November 10, 2014, TV Guide noted that Lou Taylor Pucci would be portraying Lindsay's estranged half-brother, Teddy Courtney. Pucci guest starred in a Warren Leight written episode of Law & Order: Criminal Intent, "Cruise to Nowhere" as Joey Frost.[3]

text: The showrunners mention in their interviews that the same togetherness the cast showed on-screen, they had it behind the scenes as well. "The most unexpected and rewarding development in filming the crossover episodes was seeing how close the actors from the different shows became," says CPD's Olmstead, "Between set-ups they stuck together as a group and they socialized after wrap. It brought the shows closer."[4] In TV Guide Olmstead continued, "One thing that I didn't really expect was the camaraderie with the actors, you never saw actors standing by themselves, waiting for their line. They were always in groups. They hung out as a group after they filmed. There was a lot of travel and working weekends, but the feedback I got from everybody was what a special event it was for them just as actors. I know that when we do it again — and I'm sure that we'll do it at some point down the road — it will not be a hard sell."[3]

text: Danny Mastrogiorgio who portrayed George Turner in "Chicago Crossover"; the actor has guest starred on SVU three times before, in the season thirteen episode, "Home Invasions" as a young boy's father, tenth season episode, "Lunacy" as 'the Cabbie Rapist', Orlando McTeer, and in the second season episode "Noncompliance" as Earl Miller/Gilmore as well. Prior to that, Mastrogiorgio has worked with SVU show runner Warren Leight, guest starring on Law & Order: CI as a murderous drug lord named Testarossa in the episode "Purgatory".

text: Reception[edit]

text: Ratings[edit]

text: In its original American broadcast on November 12, 2014, "Chicago Crossover" was viewed by 10.01 million viewers and acquired a 2.4 rating/7% share in the age 18–49 demographic. "Chicago Crossover" was the second watched program on NBC that night, retaining more than 95% viewership of its lead-in for the night, The Voice. "Chicago Crossover" was the second ranked program in that timeslot, under ABC's Modern Family, but above Criminal Minds on CBS by 0.1 in the 18-49 age demo.[11]

text: Critical response[edit]

text: The crossover in its entirety got extremely positive reviews from both critics and fans alike, praising how the show were meshed together to make the three-hour crossover along with praising the cast and crew members of the shows as well. Narsimha Chintaluri of TV Fanatic said in a positive review, "It was a bold, and probably complicated, move to tie these three shows together, but the premise definitely delivered a strong episode of SVU." Chintaluri also noted how the characters interacted in scenes, "The interpersonal dynamics were entertaining to be privy too. There were subtle moments, such Rollins and Amaro saying "we get it," after Halstead denied any sort of relationship with Lindsay, and then there were the not so subtle moments where Voight almost simulated raping a suspect to get his point across."[12]

text: Matt Carter of cartermatt.com gave the episode a "B+" rating, commenting "This story was about chasing a criminal ring doing unspeakable things to New York City, and there were a number of memorable moments that we saw throughout this. Obviously, seeing Benson completely take some of these people (including Voight) to task was a highlight, but there were some very cool moments that came out of seeing most of these people together." Carter compared this season's crossover to last with, "some of the crossovers between “SVU” and “Chicago PD” just didn't feel right, and mostly because we did not know many of the people well enough yet. This hour really changed that, and we felt the weight of the investigation. Plus, it is so much easier to root for many of the people when you know who they are."[13] The showrunners of both shows promised to deliver true crossover, "They were true to their word here as the episode zipped along and every character was successfully integrated into the narrative in a satisfying and in a manner that kept each character completely true to his/her nature. Well done, gentlemen." said Anne Easton of Observer.[14]

text: Easton continued, "While the intermingling of established SVU and CPD characters was great to watch, it would be a huge oversight not to give a commendation of some sort to actor Lou Taylor Pucci whose performance as Lindsay’s extremely damaged brother, Teddy, really tied the two shows together. His portrayal of the fragile victim, and victimizer, was equal parts cringe worthy and heat wrenching, a combination that is incredibly difficult to pull off. Watching him struggle to come to terms with the direction his life had taken gave the narrative a deeper meaning than had this just been a case of the week." Easton commented on the chemistry between Sgts. Benson (Hargitay) and Voight (Beghe), "They butted heads, they worked things out, and they made things happen, all in the name of protecting their respective turfs and solving the case. It was fun to see these two go toe-to-toe, each pushing the other to explore new tactics in interrogation. Their chemistry, while not really ignited on the personal side, was clearly the type of which ‘work husband and wife’ partnerships are made."[14]


document_title: Octopus

question_text: how many different kinds of octopuses are there

question_tokens: how many different kinds of octopuses are there

text: Octopus

text: Etymology and pluralisation

text: The scientific Latin term octopus was derived from Ancient Greek ὀκτώπους or ὀκτά-, a compound form of ὀκτώ (oktō, "eight") and πούς (pous, "foot"), used for example by Alexander of Tralles (c. 525-c. 605) for the common octopus.[3][4][5][6] The standard pluralised form of "octopus" in English is "octopuses" /ˈɒktəpəsɪz/;[7] the Ancient Greek plural ὀκτώποδες, "octopodes" /ɒkˈtɒpədiːz/, has also been used historically.[8] The alternative plural "octopi" – which misguidedly assumes it is a Latin "-us"-word – is considered grammatically incorrect,[9][10][11][12] but is used frequently enough to be acknowledged by the descriptivist Merriam-Webster 11th Collegiate Dictionary and Webster's New World College Dictionary. The Oxford English Dictionary lists "octopuses", "octopi", and "octopodes", in that order, calling "octopodes" rare and noting that "octopi" is based on a misunderstanding.[13] New Oxford American Dictionary (3rd Edition 2010) lists "octopuses" as the only acceptable pluralisation, and indicates that "octopodes" is still occasionally used, but that "octopi" is incorrect.[14]

text: Anatomy and physiology

text: Size

text: The giant Pacific octopus (Enteroctopus dofleini) is often cited as the largest known octopus species. Adults usually weigh around 15 kg (33 lb), with an arm span of up to 4.3 m (14 ft).[15] The largest specimen of this species to be scientifically documented was an animal with a live mass of 71 kg (156.5 lb).[16] Much larger sizes have been claimed for the giant Pacific octopus:[17] one specimen was recorded as 272 kg (600 lb) with an arm span of 9 m (30 ft).[18] A carcass of the seven-arm octopus, Haliphron atlanticus, weighed 61 kg (134 lb) and was estimated to have had a live mass of 75 kg (165 lb).[19][20] The smallest species is Octopus wolfi, which is around 2.5 cm (1 in) and weighs less than 1 g (0.035 oz).[21]

text: External characteristics

text: The octopus is bilaterally symmetrical along its dorso-ventral axis; the head and foot are at one end of an elongated body and function as the anterior (front) of the animal. The head includes the mouth and brain. The foot has evolved into a set of flexible, prehensile appendages, known as arms, that surround the mouth and are attached to each other near their base by a webbed structure.[22] The arms can be described based on side and sequence position (such as L1, R1, L2, R2) and divided into four pairs.[23][22] The bulbous and hollow mantle is fused to the back of the head and is known as the visceral hump; it contains most of the vital organs.[24][25] The mantle cavity has muscular walls and contains the gills; it is connected to the exterior by a funnel or siphon.[22][26] The mouth of an octopus, located underneath the arms, has a sharp hard beak.[25]

text: The skin consists of a thin outer epidermis with mucous cells and sensory cells, and a connective tissue dermis consisting largely of collagen fibres and various cells allowing colour change.[22] Most of the body is made of soft tissue allowing it to lengthen, contract, and contort itself. The octopus can squeeze through tiny gaps; even the larger species can pass through an opening close to 2.5 cm (1 in) in diameter.[25] Lacking skeletal support, the arms work as muscular hydrostats and contain longitudinal, transverse and circular muscles around a central axial nerve. They can extend and contract, twist to left or right, bend at any place in any direction or be held rigid.[27][28]

text: The interior surfaces of the arms are covered with circular, adhesive suckers. The suckers allow the octopus to anchor itself or to manipulate objects. Each sucker is usually circular and bowl-like and has two distinct parts: an outer shallow cavity called an infundibulum and a central hollow cavity called an acetabulum, both of which are thick muscles covered in a protective chitinous cuticle. When a sucker attaches to a surface, the orifice between the two structures is sealed. The infundibulum provides adhesion while the acetabulum remains free, and muscle contractions allow for attachment and detachment.[29][30]

text: The eyes of the octopus are large and are at the top of the head. They are similar in structure to those of a fish and are enclosed in a cartilaginous capsule fused to the cranium. The cornea is formed from a translucent epidermal layer and the slit-shaped pupil forms a hole in the iris and lies just behind. The lens is suspended behind the pupil and photoreceptive retinal cells cover the back of the eye. The pupil can be adjusted in size and a retinal pigment screens incident light in bright conditions.[22]

text: Some species differ in form from the typical octopus body shape. Members of the suborder Cirrina have stout gelatinous bodies with webbing that reaches near the tip of their arms, and two large fins above the eyes, supported by an internal shell. Fleshy papillae or cirri are found along the bottom of the arms, and the eyes are more developed.[31][32]

text: Circulatory system

text: Octopuses have a closed circulatory system, where the blood remains inside blood vessels. Octopuses have three hearts; a systemic heart that circulates blood round the body and two branchial hearts that pump it through each of the two gills. The systemic heart is inactive when the animal is swimming and thus it tires quickly and prefers to crawl.[33][34] Octopus blood contains the copper-rich protein haemocyanin to transport oxygen. This makes the blood very viscous and it requires considerable pressure to pump it round the body; octopuses' blood pressures can exceed 75 mmHg.[33][34][35] In cold conditions with low oxygen levels, haemocyanin transports oxygen more efficiently than haemoglobin. The haemocyanin is dissolved in the plasma instead of being carried within blood cells, and gives the blood a bluish colour.[33][34]

text: The systemic heart has muscular contractile walls and consists of a single ventricle and two atria, one for each side of the body. The blood vessels consist of arteries, capillaries and veins and are lined with a cellular endothelium which is quite unlike that of most other invertebrates. The blood circulates through the aorta and capillary system, to the vena cavae, after which the blood is pumped through the gills by the auxiliary hearts and back to the main heart. Much of the venous system is contractile, which helps circulate the blood.[22]

text: Respiration

text: Respiration involves drawing water into the mantle cavity through an aperture, passing it through the gills, and expelling it through the siphon. The ingress of water is achieved by contraction of radial muscles in the mantle wall, and flapper valves shut when strong circular muscles force the water out through the siphon.[36] Extensive connective tissue lattices support the respiratory muscles and allow them to expand the respiratory chamber.[37] The lamella structure of the gills allows for a high oxygen uptake; up to 65% in water at 20⁰C.[38] Water flow over the gills correlates with locomotion, and an octopus can propel its body when it expels water out of its siphon.[37][35]

text: The thin skin of the octopus absorbs additional oxygen.[39] When resting, around 41% of an octopus's oxygen absorption is through the skin. This decreases to 33% when it swims, as more water flows over the gills; skin oxygen uptake also increases. When it is resting after a meal, absorption through the skin can drop to 3% of its total oxygen uptake.[39]

text: Digestion and excretion

text: The digestive system of the octopus begins with the buccal mass which consists of the mouth, pharynx, radula and salivary glands.[40] The radula is a spiked, tongue-like organ made of chitin.[25] Food is broken down and is forced into the oesophagus by two lateral extensions of the esophageal side walls in addition to the radula. From there it is transferred to the gastrointestinal tract, which is mostly suspended from the roof of the mantle cavity by numerous membranes. The tract consists of a crop, where the food is stored; a stomach, where food is ground down; a caecum where the now sludgy food is sorted into fluids and particles and which plays an important role in absorption; the digestive gland, where liver cells break down and absorb the fluid and become "brown bodies"; and the intestine, where the accumulated waste is turned into faecal ropes by secretions and blown out of the funnel via the rectum.[40][41]

text: During osmoregulation, fluid is added to the pericardia of the branchial hearts. The octopus has two nephridia (equivalent to vertebrate kidneys) which are associated with the branchial hearts; these and their associated ducts connect the pericardial cavities with the mantle cavity. Before reaching the branchial heart, each branch of the vena cava expands to form renal appendages which are in direct contact with the thin-walled nephridium. The urine is first formed in the pericardial cavity, and is modified by excretion, chiefly of ammonia, and selective absorption from the renal appendages, as it is passed along the associated duct and through the nephridiopore into the mantle cavity.[22][42]

text: Nervous system and senses

text: The octopus (along with cuttlefish) has the highest brain-to-body mass ratios of all invertebrates, and is also greater than many vertebrates.[43] It has a highly complex nervous system, only part of which is localised in its brain, which is contained in a cartilaginous capsule.[44] Two-thirds of an octopus's neurons are found in the nerve cords of its arms, which show a variety of complex reflex actions that persist even when they have no input from the brain.[45] Unlike vertebrates, the complex motor skills of octopuses are not organised in their brain via an internal somatotopic map of its body, instead using a nonsomatotopic system unique to large-brained invertebrates.[46]

text: Like other cephalopods, octopuses can distinguish the polarisation of light. Colour vision appears to vary from species to species, for example being present in O. aegina but absent in O. vulgaris.[47] Researchers believe that opsins in the skin can sense different wavelengths of light and help the creatures choose a coloration that camouflages them, in addition to light input from the eyes.[48] Other researchers hypothesize that cephalopod eyes in species which only have a single photoreceptor protein may use chromatic aberration to turn monochromatic vision into color vision, though this sacrifices image quality.[49] This would explain pupils shaped like the letter U, the letter W, or a dumbbell, as well as explaining the need for colorful mating displays.[50]

text: Attached to the brain are two special organs called statocysts (sac-like structures containing a mineralised mass and sensitive hairs), that allow the octopus to sense the orientation of its body. They provide information on the position of the body relative to gravity and can detect angular acceleration. An autonomic response keeps the octopus's eyes oriented so that the pupil is always horizontal.[22] Octopuses may also use the statocyst to hear sound. The common octopus can hear sounds between 400 Hz and 1000 Hz, and hears best at 600 Hz.[51]

text: Octopuses also have an excellent sense of touch. The octopus's suction cups are equipped with chemoreceptors so the octopus can taste what it touches. Octopus arms do not become tangled or stuck to each other because the sensors recognise octopus skin and prevent self-attachment.[52]

text: The arms contain tension sensors so the octopus knows whether its arms are stretched out, but this is not sufficient for the brain to determine the position of the octopus's body or arms. As a result, the octopus does not possess stereognosis; that is, it does not form a mental image of the overall shape of the object it is handling. It can detect local texture variations, but cannot integrate the information into a larger picture. The neurological autonomy of the arms means the octopus has great difficulty learning about the detailed effects of its motions. It has a poor proprioceptive sense, and it knows what exact motions were made only by observing the arms visually.[53]

text: Ink sac

text: The ink sac of an octopus is located under the digestive gland. A gland attached to the sac produces the ink, and the sac stores it. The sac is close enough to the funnel for the octopus to shoot out the ink with a water jet. Before it leaves the funnel, the ink passes through glands which mix it with mucus, creating a thick, dark blob which allows the animal to escape from a predator.[54] The main pigment in the ink is melanin, which gives it its black colour.[55] Cirrate octopuses lack the ink sac.[31]

text: Lifecycle

text: Reproduction

text: Octopuses are gonochoric and have a single, posteriorly-located gonad which is associated with the coelom. The testis in males and the ovary in females bulges into the gonocoel and the gametes are released here. The gonocoel is connected by the gonoduct to the mantle cavity, which it enters at the gonopore.[22] An optic gland creates hormones that cause the octopus to mature and age and stimulate gamete production. The gland may be triggered by environmental conditions such as temperature, light and nutrition, which thus control the timing of reproduction and lifespan.[56][57]

text: When octopuses reproduce, the male uses a specialised arm called a hectocotylus to transfer spermatophores (packets of sperm) from the terminal organ of the reproductive tract (the cephalopod "penis") into the female's mantle cavity.[58] The hectocotylus in benthic octopuses is usually the third right arm, which has a spoon-shaped depression and modified suckers near the tip. In most species, fertilisation occurs in the mantle cavity.[22]

text: The reproduction of octopuses has been studied in only a few species. One such species is the giant Pacific octopus, in which courtship is accompanied, especially in the male, by changes in skin texture and colour. The male may cling to the top or side of the female or position himself beside her. There is some speculation that he may first use his hectocotylus to remove any spermatophore or sperm already present in the female. He picks up a spermatophore from his spermatophoric sac with the hectocotylus, inserts it into the female's mantle cavity, and deposits it in the correct location for the species, which in the giant Pacific octopus is the opening of the oviduct. Two spermatophores are transferred in this way; these are about one metre (yard) long, and the empty ends may protrude from the female's mantle.[59] A complex hydraulic mechanism releases the sperm from the spermatophore, and it is stored internally by the female.[22]

text: About forty days after mating, the female giant Pacific octopus attaches strings of small fertilised eggs (10,000 to 70,000 in total) to rocks in a crevice or under an overhang. Here she guards and cares for them for about five months (160 days) until they hatch.[59] In colder waters, such as those off of Alaska, it may take as much as 10 months for the eggs to completely develop.[60]:74 The female aerates the eggs and keeps them clean; if left untended, many eggs will not hatch.[61] She does not feed during this time and dies soon afterwards. Males become senescent and die a few weeks after mating.[62]

text: The eggs have large yolks; cleavage (division) is superficial and a germinal disc develops at the pole. During gastrulation, the margins of this grow down and surround the yolk, forming a yolk sac, which eventually forms part of the gut. The dorsal side of the disc grows upwards and forms the embryo, with a shell gland on its dorsal surface, gills, mantle and eyes. The arms and funnel develop as part of the foot on the ventral side of the disc. The arms later migrate upwards, coming to form a ring around the funnel and mouth. The yolk is gradually absorbed as the embryo develops.[22]

text: Most young octopuses hatch as paralarvae and are planktonic for weeks to months, depending on the species and water temperature. They feed on copepods, arthropod larvae and other zooplankton, eventually settling on the ocean floor and developing directly into adults with no distinct metamorphoses that are present in other groups of mollusc larvae.[22] Octopus species that produce larger eggs – including the southern blue-ringed, Caribbean reef, California two-spot, Eledone moschata[63] and deep sea octopuses – do not have a paralarval stage, but hatch as benthic animals similar to the adults.[60]:74–75[64]

text: In the argonaut (paper nautilus), the female secretes a fine, fluted, papery shell in which the eggs are deposited and in which she also resides while floating in mid-ocean. In this she broods the young, and it also serves as a buoyancy aid allowing her to adjust her depth. The male argonaut is minute by comparison and has no shell.[65]

text: Lifespan

text: Octopuses have a relatively short life expectancy; some species live for as little as six months. The giant Pacific octopus, one of the two largest species of octopus, may live for as much as five years. Octopus lifespan is limited by reproduction: males can live for only a few months after mating, and females die shortly after their eggs hatch. Octopus reproductive organs mature due to the hormonal influence of the optic gland but result in the inactivation of their digestive glands, typically causing the octopus to eventually die from starvation.[66]:276–277 Experimental removal of both optic glands after spawning was found to result in the cessation of broodiness, the resumption of feeding, increased growth, and greatly extended lifespans.[67]

text: Distribution and habitat

text: Octopuses live in every ocean, and different species have adapted to different marine habitats. As juveniles, common octopuses inhabit shallow tide pools. The Hawaiian day octopus (Octopus cyanea) lives on coral reefs; argonauts drift in pelagic waters. Abdopus aculeatus mostly lives in near-shore seagrass beds. Some species are adapted to the cold, ocean depths. The spoon-armed octopus (Bathypolypus arcticus) is found in abyssal plains at depths of 1,000 m (3,300 ft), and Vulcanoctopus hydrothermalis lives near hydrothermal vents at 2,000 m (6,600 ft).[24] The cirrate species are often free-swimming and live in deep-water habitats.[32] No species are known to live in fresh water.[68]

text: Behaviour and ecology

text: Most species are solitary when not mating,[69] though a few are known to occur in high densities and with frequent interactions, signaling, mate defending and eviction of individuals from dens. This is likely the result of abundant food supplies combined with limited den sites.[70] Octopuses hide in dens, which are typically crevices in rocky outcrops or other hard structures, though some species burrow into sand or mud. Octopuses are not territorial but generally remain in a home range; they may leave the area in search of food. They can use navigation skills to return to a den without having to retrace their outward route.[71] They are not known to be migratory.[72]

text: Octopuses bring captured prey back to the den where they can eat it safely. Sometimes the octopus catches more prey than it can eat, and the den is often surrounded by a midden of dead and uneaten food items. Other creatures, such as fish, crabs, molluscs and echinoderms, often share the den with the octopus, either because they have arrived as scavengers, or because they have survived capture.[73]

text: Feeding

text: Nearly all octopuses are predatory; bottom-dwelling octopuses eat mainly crustaceans, polychaete worms, and other molluscs such as whelks and clams; open-ocean octopuses eat mainly prawns, fish and other cephalopods.[74] Major items in the diet of the giant Pacific octopus include bivalve molluscs such as the cockle Clinocardium nuttallii, clams and scallops, and crustaceans such as crabs and spider crabs. Prey that it is likely to reject include moon snails, because they are too large, and limpets, rock scallops, chitons and abalone, because they are too securely fixed to the rock.[73]

text: A bottom-dwelling octopus typically moves among the rocks and feels through the crevices. The creature may make a jet-propelled pounce on prey and pull it towards the mouth with its arms, the suckers restraining it. Small prey may be completely trapped by the webbed structure. Octopuses usually inject crustaceans like crabs with a paralysing saliva then dismember them with their beaks.[74][75] Octopuses feed on shelled molluscs either by forcing the valves apart, or by drilling a hole in the shell to inject a nerve toxin.[76][75] It used to be thought that the hole was drilled by the radula, but it has now been shown that minute teeth at the tip of the salivary papilla are involved, and an enzyme in the toxic saliva is used to dissolve the calcium carbonate of the shell. It takes about three hours for O. vulgaris to create a 0.6 mm (0.024 in) hole. Once the shell is penetrated, the prey dies almost instantaneously, its muscles relax, and the soft tissues are easy for the octopus to remove. Crabs may also be treated in this way; tough-shelled species are more likely to be drilled, and soft-shelled crabs are torn apart.[77]

text: Some species have other modes of feeding. Grimpoteuthis has a reduced or non-existent radula and swallows prey whole.[31] In the deep-sea genus Stauroteuthis, some of the muscle cells that control the suckers in most species have been replaced with photophores which are believed to fool prey by directing them towards the mouth, making them one of the few bioluminescent octopuses.[78]

text: Locomotion

text: Octopuses mainly move about by relatively slow crawling, with some swimming in a head-first position. Jet propulsion, or backwards swimming, is their fastest means of locomotion, followed by swimming and crawling.[79] When in no hurry, they usually crawl on either solid or soft surfaces. Several arms are extended forwards, some of the suckers adhere to the substrate and the animal hauls itself forwards with its powerful arm muscles, while other arms may push rather than pull. As progress is made, other arms move ahead to repeat these actions and the original suckers detach. During crawling, the heart rate nearly doubles, and the animal requires ten or fifteen minutes to recover from relatively minor exercise.[27]

text: Most octopuses swim by expelling a jet of water from the mantle through the siphon into the sea. The physical principle behind this is that the force required to accelerate the water through the orifice produces a reaction that propels the octopus in the opposite direction.[80] The direction of travel depends on the orientation of the siphon. When swimming, the head is at the front and the siphon is pointed backwards, but when jetting, the visceral hump leads, the siphon points towards the head and the arms trail behind, with the animal presenting a fusiform appearance. In an alternative method of swimming, some species flatten themselves dorso-ventrally, and swim with the arms held out sideways, and this may provide lift and be faster than normal swimming. Jetting is used to escape from danger, but is physiologically inefficient, requiring a mantle pressure so high as to stop the heart from beating, resulting in a progressive oxygen deficit.[79]

text: Cirrate octopuses cannot produce jet propulsion and rely on their fins for swimming. They have neutral buoyancy and drift through the water with the fins extended. They can also contract their arms and surrounding web to make sudden moves known as "take-offs". Another form of locomotion is "pumping", which involves symmetrical contractions of muscles in their webs producing peristaltic waves. This moves the body slowly.[31]

text: In 2005, Adopus aculeatus and veined octopus (Amphioctopus marginatus) were found to walk on two arms, while at the same time mimicking plant matter.[81] This form of locomotion allows these octopuses to move quickly away from a potential predator without being recognised.[79] A study of this behaviour led to the suggestion that the two rearmost appendages may be more accurately termed "legs" rather than "arms".[82] Some species of octopus can crawl out of the water briefly, which they may do between tide pools while hunting crustaceans or gastropods or to escape predators.[83][84] "Stilt walking" is used by the veined octopus when carrying stacked coconut shells. The octopus carries the shells underneath it with two arms, and progresses with an ungainly gait supported by its remaining arms held rigid.[85]

text: Intelligence

text: Octopuses are highly intelligent, but the extent of their intelligence and learning capability are not well defined.[86][87][88][89] Maze and problem-solving experiments have shown evidence of a memory system that can store both short- and long-term memory. It is not known precisely what contribution learning makes to adult octopus behaviour. Young octopuses learn nothing from their parents, as adults provide no parental care beyond tending to their eggs until the young octopuses hatch.[60]:75

text: In laboratory experiments, octopuses can be readily trained to distinguish between different shapes and patterns. They have been reported to practise observational learning,[90] although the validity of these findings is contested.[86][87] Octopuses have also been observed in what has been described as play: repeatedly releasing bottles or toys into a circular current in their aquariums and then catching them.[91] Octopuses often break out of their aquariums and sometimes into others in search of food.[83][92][93] They have even boarded fishing boats and opened holds to eat crabs.[88] The veined octopus collects discarded coconut shells, then uses them to build a shelter, an example of tool use.[85][94][95]

text: Camouflage and colour change

text: Octopuses use camouflage when hunting, and to avoid predators. To do this they use specialised skin cells which change the appearance of the skin by adjusting its colour, opacity, or reflectivity. Chromatophores contain yellow, orange, red, brown, or black pigments; most species have three of these colours, while some have two or four. Other colour-changing cells are reflective iridophores and white leucophores.[96] This colour-changing ability is also used to communicate with or warn other octopuses.[97]

text: Octopuses can create distracting patterns with waves of dark coloration across the body, a display known as the "passing cloud". Muscles in the skin change the texture of the mantle to achieve greater camouflage. In some species, the mantle can take on the spiky appearance of algae; in others, skin anatomy is limited to relatively uniform shades of one colour with limited skin texture. Octopuses that are diurnal and live in shallow water have evolved more complex skin than their nocturnal and deep-sea counterparts.[97]

text: Defence

text: Aside from humans, octopuses may be preyed on by fishes, seabirds, sea otters, pinnipeds, cetaceans, and other cephalopods.[98] Octopuses typically hide or disguise themselves by camouflage and mimicry; some have conspicuous warning coloration (aposematism) or deimatic behaviour.[97] An octopus may spend 40% of its time hidden away in its den. When the octopus is approached, it may extend an arm to investigate. 66% of Enteroctopus dolfleini in one study had scars, with 50% having amputated arms.[98] The blue rings of the highly venomous blue-ringed octopus are hidden in muscular skin folds which contract when the animal is threatened, exposing the iridescent warning.[99] The Atlantic white-spotted octopus (Callistoctopus macropus) turns bright brownish red with oval white spots all over in a high contrast display.[100] Displays are often reinforced by stretching out the animal's arms, fins or web to make it look as big and threatening as possible.[101]

text: Once they have been seen by a predator, they commonly try to escape but can also use distraction with an ink cloud ejected from the ink sac. The ink is thought to reduce the efficiency of olfactory organs, which would aid evasion from predators that employ smell for hunting, such as sharks. Ink clouds of some species might act as pseudomorphs, or decoys that the predator attacks instead.[102]

text: When under attack, some octopuses can perform arm autotomy, in a manner similar to the way skinks and other lizards detach their tails. The crawling arm may distract would-be predators. Such severed arms remain sensitive to stimuli and move away from unpleasant sensations.[103] Octopuses can replace lost limbs.[104]

text: Some octopuses, such as the mimic octopus, can combine their highly flexible bodies with their colour-changing ability to mimic other, more dangerous animals, such as lionfish, sea snakes, and eels.[105][106]

text: Pathogens and parasites

text: The diseases and parasites that affect octopuses have been little studied, but cephalopods are known to be the intermediate or final hosts of various parasitic cestodes, nematodes and copepods; 150 species of protistan and metazoan parasites have been recognised.[107] The Dicyemidae are a family of tiny worms that are found in the renal appendages of many species;[108] it is unclear whether they are parasitic or are endosymbionts. Coccidians in the genus Aggregata living in the gut cause severe disease to the host. Octopuses have an innate immune system, and the haemocytes respond to infection by phagocytosis, encapsulation, infiltration or cytotoxic activities to destroy or isolate the pathogens. The haemocytes play an important role in the recognition and elimination of foreign bodies and wound repair. Captive animals have been found to be more susceptible to pathogens than wild ones.[109] A gram-negative bacterium, Vibrio lentus, has been found to cause skin lesions, exposure of muscle and death of octopuses in extreme cases.[110]

text: Evolution

text: Fossil history and phylogeny

text: Cephalopods have existed for 500 million years and octopus ancestors were in the Carboniferous seas 300 million years ago. The oldest known octopus fossil is Pohlsepia, which lived 296 million years ago. Researchers have identified impressions of eight arms, two eyes, and possibly an ink sac.[111] Octopuses are mostly soft tissue, and so fossils are relatively rare. Octopuses, squids and cuttlefish belong to the clade Coleoidea. They are known as "soft-bodied" cephalopods, lacking the external shell of most molluscs and other cephalopods like the Nautiloids and the extinct Ammonoidea.[112] Octopuses have eight arms like other coleoids but lack the extra specialised feeding appendages known as tentacles which are longer and thinner with suckers only at their club-like ends.[113][114][115] The vampire squid (Vampyroteuthis) also lacks tentacles but has sensory filaments.[116]

text: Two possible extant cephalopod phylogenies, based on genetics studies by Strugnell et al. 2007, are shown in the possible cladograms.[117]

text:
Cephalopods

Nautiloids

Nautilus

Coleoids

Octopods

Vampyroteuthis

Squids and cuttlefish

text:
Cephalopods

Nautiloids

Nautilus

Coleoids

Squids and cuttlefish

Vampyroteuthis

Octopods

text: Taxonomy

text: The scientific name Octopoda was first coined and given as the order of octopuses in 1818 by English biologist William Elford Leach,[118] who classified them as Octopoida the previous year.[2] The Octopoda consists of around 300 known species[119] and can be divided into two suborders, the Incirrina and the Cirrina. The incirrate octopuses (the majority of species) lack the cirri and paired swimming fins of the cirrates.[32] In addition, the internal shell of incirrates is either present as a pair of stylets or absent altogether.[120]

text:
Order Octopoda

Life restoration of Keuppia levante, an extinct species from the Cretaceous

Genus †Keuppia (incertae sedis)
Genus †Palaeoctopus (incertae sedis)
Genus †Paleocirroteuthis (incertae sedis)
Genus †Pohlsepia (incertae sedis)
Genus †Proteroctopus (incertae sedis)
Genus †Styletoctopus (incertae sedis)
Suborder Cirrina: finned deep-sea octopus

Family Opisthoteuthidae: umbrella octopus

The flapjack octopus (Opisthoteuthis californiana)

Family Cirroctopodidae
Family Cirroteuthidae
Family Stauroteuthidae

Suborder Incirrina

Superfamily Octopodoidea[121]

Family Amphitretidae
Family Bathypolypodidae
Family Eledonidae
Family Enteroctopodidae
Family Megaleledonidae Taki, 1961
Family Octopodidae

Superfamily Argonautoidea

Shelled argonaut, Argonauta argo

Family Alloposidae: seven-arm octopus
Family Argonautidae: argonauts
Family Ocythoidae: tuberculate pelagic octopus
Family Tremoctopodidae: blanket octopus

text: RNA editing

text: Octopuses and other coleoid cephalopods are capable of greater RNA editing (which involves changes to the nucleic acid sequence of the primary transcript of RNA molecules) than any other organisms. Editing is concentrated in the nervous system and affects proteins involved in neural excitability and neuronal morphology. More than 60% of RNA transcripts for coleoid brains are recoded by editing, compared to less than 1% for a human or fruit fly. Coleoids rely mostly on ADAR enzymes for RNA editing, which requires large double-stranded RNA structures to flank to the editing sites. Both the structures and editing sites are conserved in the coleoid genome and the mutation rates for the sites are severely hampered. Hence, greater transcriptome plasticity has come as the cost of slower genome evolution. High levels of RNA editing do not appear to be present in more basal cephalopods or other molluscs.[122][123]

text: Relationship to humans

text: Cultural references

text: Ancient seafaring people were aware of the octopus, as evidenced by certain artworks and designs. For example, a stone carving found in the archaeological recovery from Bronze Age Minoan Crete at Knossos (1900 – 1100 BC) has a depiction of a fisherman carrying an octopus.[124] The terrifyingly powerful Gorgon of Greek mythology has been thought to have been inspired by the octopus or squid, the octopus itself representing the severed head of Medusa, the beak as the protruding tongue and fangs, and its tentacles as the snakes.[125] The Kraken are legendary sea monsters of giant proportions said to dwell off the coasts of Norway and Greenland, usually portrayed in art as a giant octopus attacking ships. Linnaeus included it in the first edition of his 1735 Systema Naturae.[126][127] A Hawaiian creation myth says that the present cosmos is the last of a series which arose in stages from the ruins of the previous universe. In this account, the octopus is the lone survivor of the previous, alien universe.[128] The Akkorokamui is a gigantic octopus-like monster from Ainu folklore.[129]

text: A battle with an octopus plays a significant role in Victor Hugo's book Travailleurs de la mer (Toilers of the Sea), relating to his time in exile on Guernsey.[130] Ian Fleming's 1966 short story collection Octopussy and The Living Daylights, and the 1983 James Bond film were partly inspired by Hugo's book.[131]

text: Japanese erotic art, shunga, includes ukiyo-e woodblock prints such as Katsushika Hokusai's 1814 print Tako to ama (The Dream of the Fisherman's Wife), in which an ama diver is sexually intertwined with a large and a small octopus.[132][133] The print is a forerunner of tentacle erotica.[134] The biologist P. Z. Myers noted in his science blog, Pharyngula, that octopuses appear in "extraordinary" graphic illustrations involving women, tentacles, and bare breasts.[135][136]

text: Since it has numerous arms emanating from a common centre, the octopus is often used as a symbol for a powerful and manipulative organisation, usually negatively.[137]

text: Danger

text: Octopuses generally avoid humans, but incidents have been verified. For example, a 2.4-metre (8 ft) Pacific octopus, said to be nearly perfectly camouflaged, "lunged" at a diver and "wrangled" over his camera before it let go. Another diver recorded the encounter on video.[138]

text: All species are venomous, but only blue-ringed octopuses have venom that is lethal to humans.[139] Bites are reported each year across the animals' range from Australia to the eastern Indo-Pacific Ocean. They bite only when provoked or accidentally stepped upon; bites are small and usually painless. The venom appears to be able to penetrate the skin without a puncture, given prolonged contact. It contains tetrodotoxin, which causes paralysis by blocking the transmission of nerve impulses to the muscles. This causes death by respiratory failure leading to cerebral anoxia. No antidote is known, but if breathing can be kept going artificially, patients recover within 24 hours.[140][141] Bites have been recorded from captive octopuses of other species; they leave swellings which disappear in a day or two.[142]

text: Fisheries and cuisine

text: Octopus fisheries exist around the world with total catches varying between 245,320 and 322,999 metric tons from 1986 to 1995.[143] The world catch peaked in 2007 at 380,000 tons, and fell by a tenth by 2012.[144] Methods to capture octopuses include pots, traps, trawls, snares, drift fishing, spearing, hooking and hand collection.[143] Octopus is eaten in many cultures and is a common food on the Mediterranean and Asian coasts.[145][146] The arms and sometimes other body parts are prepared in various ways, often varying by species or geography. Live octopuses are eaten in several countries around the world, including the US.[147][148] Animal welfare groups have objected to this practice on the basis that octopuses can experience pain.[149] Octopuses have a food conversion efficiency greater than that of chickens, making octopus aquaculture a possibility.[150]

text: In science and technology

text: In classical Greece, Aristotle (384 – 322 BC) commented on the colour-changing abilities of the octopus, both for camouflage and for signalling, in his Historia animalium: "The octopus ... seeks its prey by so changing its colour as to render it like the colour of the stones adjacent to it; it does so also when alarmed."[151] Aristotle noted that the octopus had a hectocotyl arm and suggested it might be used in sexual reproduction. This claim was widely disbelieved until the 19th century. It was described in 1829 by the French zoologist Georges Cuvier, who supposed it to be a parasitic worm, naming it as a new species, Hectocotylus octopodis.[152][153] Other zoologists thought it a spermatophore; the German zoologist Heinrich Müller believed it was "designed" to detach during copulation. In 1856 the Danish zoologist Japetus Steenstrup demonstrated that it is used to transfer sperm, and only rarely detaches.[154]

text: Octopuses offer many possibilities in biological research, including their ability to regenerate limbs, change the colour of their skin, behave intelligently with a distributed nervous system, and make use of 168 kinds of protocadherins (humans have 58), the proteins that guide the connections neurons make with each other. The California two-spot octopus has had its genome sequenced, allowing exploration of its molecular adaptations.[155] Having independently evolved mammal-like intelligence, octopuses have been compared to hypothetical intelligent extraterrestrials.[156] Their problem-solving skills, along with their mobility and lack of rigid structure enable them to escape from supposedly secure tanks in laboratories and public aquariums.[157]

text: Due to their intelligence, octopuses are listed in some countries as experimental animals on which surgery may not be performed without anesthesia, a protection usually extended only to vertebrates. In the UK from 1993 to 2012, the common octopus (Octopus vulgaris) was the only invertebrate protected under the Animals (Scientific Procedures) Act 1986.[158] In 2012, this legislation was extended to include all cephalopods[159] in accordance with a general EU directive.[160]

text: Some robotics research is exploring biomimicry of octopus features. Octopus arms can move and sense largely autonomously without intervention from the animal's central nervous system. In 2015 a team in Italy built soft-bodied robots able to crawl and swim, requiring only minimal computation.[161] In 2017 a German company made an arm with a soft pneumatically controlled silicone gripper fitted with two rows of suckers. It is able to grasp objects such as a metal tube, a magazine, or a ball, and to fill a glass by pouring water from a bottle.[162]