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Big Mouth (TV series)
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Big Mouth
Created by
Nick Kroll
Andrew Goldberg
Mark Levin
Jennifer Flackett
Starring
Nick Kroll
John Mulaney
Jessi Klein
Jason Mantzoukas
Jenny Slate
Fred Armisen
Maya Rudolph
Jordan Peele
Music by
Mark Rivers
Opening theme
"Changes" by Charles Bradley
Country of origin
United States
Original language(s)
English
No. of seasons
2
No. of episodes
20 (list of episodes)
Production
Executive producer(s)
Nick Kroll
Andrew Goldberg
Mark Levin
Jennifer Flackett
Producer(s)
Nate Funaro
Running time
25–28 minutes
Production company(s)
Danger Goldberg Productions
Good at Bizness, Inc.
Fathouse Industries
Titmouse, Inc.
Distributor
Netflix
Release
Original network
Netflix
Original release
September 29, 2017 (2017-09-29) – present (present)
External links
Website
Big Mouth is an American adult animated sitcom created by Nick Kroll, Andrew Goldberg, Mark Levin, and Jennifer Flackett based on Kroll and Goldberg's tweenage years growing up in Westchester County, New York, with Kroll voicing his fictional self. The first season consisting of ten episodes premiered on Netflix on September 29, 2017. The second season also was released on October 5, 2018.
Contents
1 Summary
2 Cast
2.1 Recurring
3 Episodes
3.1 Season 1 (2017)
3.2 Season 2 (2018)
4 Production
5 Release
6 Reception
6.1 Critical reception
6.2 Accolades
7 References
8 External links
Summary[edit]
The series follows a group of 7th graders, including best friends Nick Birch and Andrew Glouberman, as they navigate their way through puberty, masturbation, and sexual arousal in the suburbs of New York City. Acting as sex-based shoulder angels are the hormone monsters: Maurice, who pesters Andrew and occasionally Nick and Jay; and Connie, who pesters Jessi and occasionally Missy. Throughout the series, the kids interact with people and objects who are personified in one way or another and offer helpful, albeit confusing, advice in their puberty-filled lives including the ghost of Duke Ellington, a French accented Statue of Liberty, a pillow capable of getting pregnant, and even Jessi's own genitals.
Cast[edit]
Nick Kroll[1] as:
Nick Birch, an almost-adolescent boy living with loving and overprotective parents. He also has an older brother named Judd and an older sister named Leah. He is arrogant and selfish, traits that are represented in the way he treated his girlfriend, Gina.
Maurice the Hormone Monster, a monster who follows Andrew around and is responsible for his brooding sexual desires, which frequently occur at inappropriate moments.
Coach Steve, the middle school gym class teacher, who is incompetent and overly talkative, often trying to get his colleagues—or even the students—to hang out (often to no avail). It is heavily implied that the coach may be mentally disabled; he cannot read. It is revealed in season 2 that he is a virgin, though he ultimately loses his virginity to Jay's mom, by whom he is sexually approached and with whom he has a one night stand.
Rick the Hormone Monster, an old, dysfunctional yet enthusiastic hormone monster who follows Coach Steve and later Nick.
Lola, an intense popular girl who is terrified of losing her social status and the approval of her best friend Devin.
The Jansen Twins, a pair of girls from the Netherlands who are new to the school.
Kroll also voices Lady Liberty, a ladybug, Joe Walsh, the ghosts of Picasso and Richard Burton, and Sylvester Stallone.
John Mulaney as Andrew Glouberman, a young boy who spends much of his time furtively masturbating.
Mulaney also voices Mint, Grandpa Andrew, Babe the Blue Ox, and Detective Florez.
Jessi Klein as Jessi Glaser, a sarcastic and smart girl. Her father, named Greg, is an unemployed stoner and her mother, named Shannon, is secretly dating another woman. She gets her period for the first time on a school trip to the Statue of Liberty. Though Jessi has kissed Nick before, specifically in the first episode "Ejaculation", she later has many French kissing make-out sessions with Jay. She, like Andrew, is Jewish, as she has a Bat Mitzvah in episode 9, "I Survived Jessi's Bat Mitzvah".
Jason Mantzoukas as Jay Bilzerian, an Armenian American boy who is obsessed with magic and sex. He has sex with a pillow that he made into a sex toy. His mom is mentally ill, his dad is a divorce lawyer, and his two older brothers are named Kurt and Val. He is obsessed with Nick's mother and father and wishes for a loving family.
Mantzoukas also voices Guy Bilzerian and the ghost of Socrates.
Jenny Slate as Missy Foreman-Greenwald, a girl who is nerdy and kind.
Fred Armisen as Elliot Birch, Nick's loving father, who frequently shares inappropriate stories.
Armisen also voices the ghost of Antonin Scalia, a bus driver, and Stavros.
Maya Rudolph as:
Connie the Hormone Monstress, who follows Jessi (and sometimes Missy and Nick) around and encourages her to embrace her wildest desires.
Diane Birch, Nick's loving mother, who babies him.
Rudolph also voices a bath mat and the ghosts of Elizabeth Taylor and Whitney Houston.
Jordan Peele as the Ghost of Duke Ellington, the late jazz musician whose spirit now lives in Nick's attic and sometimes offers him advice.
Peele also voices the ghost of Freddie Mercury, Atlanta Claus, a DJ, Missy's father Cyrus, Patrick Ewing and Jay's pitbull, Featuring Ludacris.
Recurring[edit]
Paula Pell as Barbara Glouberman, Andrew's mother.
Richard Kind as Marty Glouberman, Andrew's grumpy father and lover of scallops.
Andrew Rannells as Matthew, a flamboyantly gay student with a love of drama and spreading gossip.
June Diane Raphael as Devin, a popular student at the high school.
Gina Rodriguez as Gina Alvarez, a very physically developed girl on Missy and Jessi's soccer team whose sudden pubertal development causes a stir at school. (season 2)
John Gemberling as Security Guard/Tyler the Hormone Monster
David Thewlis as Shame Wizard, the Hormone Monster's mortal enemy who haunts the kids, stoking their deepest shame. (season 2)
Jak Knight as DeVon, the boyfriend of Devin and another popular student.
Neil Casey as Lars/Detective Dumont
Joe Wengert as Caleb, another 7th grader, possibly autistic.
Seth Morris as Greg Glaser, the perpetually stoned father of Jessi Glaser and the husband of Shannon Glaser.
Jessica Chaffin as Shannon Glaser, the mother of Jessi Glaser and the unfaithful wife of Greg Glaser.
Rob Huebel as Mr. Lizer
Kat Dennings as Leah Birch, the older sister of Nick Birch and younger sister of Judd Birch.
Kristen Bell as Pam the pillow/Uninterested Girl
Heather Lawless as Jenna Bilzerian
Chelsea Peretti as Monica Foreman-Greenwald
Mark Duplass as Val Bilzerian, an older, bullying brother to Jay Bilzerian. Duplass also voices a clerk.
Paul Scheer as Kurt Bilzerian, another older, bullying brother to Jay Bilzerian.
Nathan Fillion as himself
Kristen Wiig as Jessi's genitalia
Nasim Pedrad as Fatima, the female character in the book The Rock of Gibraltar.
Carlos Alazraqui as Gustavo, the male character in the book The Rock of Gibraltar.
Alia Shawkat as Roland, a friend of Nick Birch's who lives in Manhattan, New York.
Mae Whitman as Tallulah Levine, a friend of Leah's who takes a liking to Nick.
Zach Woods as Daniel, a boy Leah is interested in. Woods also voices a sock.
Jon Hamm as Scallops
Jack McBrayer as Nick's pubic hair #1
Craig Robinson as Nick's pubic hair #2
Jon Daly as Judd Birch, the dark yet caring older brother of Nick and Leah Birch, and the son of Diane and Elliot Birch.
Rosa Salazar as Miss Benitez, a 7th grade teacher.
Michaela Watkins as Cantor Dina Reznick, Shannon Glaser's love interest.
Jean Smart as Depression Kitty
Bobby Cannavale as Gavin the Hormone Monster
Episodes[edit]
Season 1 (2017)[edit]
No.
overall
No. in
season
Title
Directed by
Written by
Original release date
1
1 "Ejaculation" Joel Moser Nick Kroll & Andrew Goldberg & Jennifer Flackett & Mark Levin September 29, 2017 (2017-09-29)
As Andrew falls under the spell of the Hormone Monster, Maury, his buddy Nick becomes obsessed with the lack of changes in his own body. The gang go to the school dance.
2
2 "Everybody Bleeds" Bryan Francis Kelly Galuska September 29, 2017 (2017-09-29)
A field trip goes from awkward to mortifying as Jessi is pressured into being Nick's girlfriend, then gets her first period—inside the Statue of Liberty.
3
3 "Am I Gay?" Mike L. Mayfield Joe Wengert September 29, 2017 (2017-09-29)
A trailer for a new movie starring The Rock leaves Andrew questioning his sexuality. Jessi and Nick's budding romance turns ugly fast.
4
4 "Sleepover: A Harrowing Ordeal of Emotional Brutality" Joel Moser Jess Dweck & Victor Quinaz September 29, 2017 (2017-09-29)
While the boys head to Jay's for a testosterone-fueled sleepover, Jessi throws a party of her own and gets caught up in mean-girl drama.
5
5 "Girls Are Horny Too" Bryan Francis Emily Altman September 29, 2017 (2017-09-29)
A steamy novel making the rounds at school inspires Jessi to explore her own sexuality—and leaves the boys totally confused about what girls want.
6
6 "Pillow Talk" Mike L. Mayfield Peter A. Knight September 29, 2017 (2017-09-29)
Nick drags Andrew into the city to visit an old camp crush, but the day doesn't go as planned. Jay embarks on an emotional journey with his pillow.
7
7 "Requiem for a Wet Dream" Joel Moser Duffy Boudreau September 29, 2017 (2017-09-29)
Andrew struggles to keep the Hormone Monster in check as he works with Missy on a science fair project. Jessi and Matthew have fun at Jay's expense.
8
8 "The Head Push" Bryan Francis Emily Altman & Jennifer Flackett & Mark Levin September 29, 2017 (2017-09-29)
When Leah hosts a party for the high school drama crowd, Nick wrestles with a romantic dilemma, and Andrew and Missy witness a disturbing scene.
9
9 "I Survived Jessi's Bat Mitzvah" Mike L. Mayfield Kelly Galuska September 29, 2017 (2017-09-29)
Family blowups, secret liaisons and an unflattering dress make Jessi's bat mitzvah a night to remember.
10
10 "The Pornscape" Joel Moser Gil Ozeri September 29, 2017 (2017-09-29)
Andrew tries to drown his sorrows in a sea of porn, only to become hopelessly addicted. Coach Steve becomes a suspect in a murder spree. Jessi and Jay run away together. Meanwhile, Andrew is arrested and goes to prison.
Season 2 (2018)[edit]
No.
overall
No. in
season
Title
Directed by
Written by
Original release date
11
1 "Am I Normal?" Bob Suarez Andrew Goldberg October 5, 2018 (2018-10-05)
Andrew struggles to blend in after a growth spurt, Nick worries his hormones are out of whack, and Jessi and Jay get a taste of the teen runaway life.
12
2 "What Is It About Boobs?" Bryan Francis Kelly Galuska October 5, 2018 (2018-10-05)
A classmate's booming bosom sends the boys into a rapture -- and leaves the girls with a raging case of insecurity. Coach Steve drops a bombshell.
13
3 "The Shame Wizard" Joel Moser Victor Quinaz October 5, 2018 (2018-10-05)
As Nick tries to figure out whether he has a shot with Gina, a mortifying encounter with Leah sends Andrew into a spiral of shame.
14
4 "Steve the Virgin" Bob Suarez Joe Wengert October 5, 2018 (2018-10-05)
Coach Steve finally gets his chance to do sex on a lady, Nick and Andrew go on a double date, and Jessi's rebellious streak leads to trouble.
15
5 "The Planned Parenthood Show" Bryan Francis Emily Altman October 5, 2018 (2018-10-05)
An argument in sex ed class turns into a skit-tastic journey through the worlds of Planned Parenthood, contraceptives, and STDs.
16
6 "Drug Buddies" Joel Moser Gil Ozeri October 5, 2018 (2018-10-05)
Jessi and Nick dip into her dad's stash of edibles, with mind-bending results. Andrew searches for the right moment to break up with Lola.
17
7 "Guy Town" Bob Suarez Joe Wengert October 5, 2018 (2018-10-05)
While helping Greg move into a seedy bachelor complex, the boys and their dads debate what women want -- and what it means to be a man.
18
8 "Dark Side of the Boob" Bryan Francis Kelly Galuska October 5, 2018 (2018-10-05)
A school sleepover turns messy as Nick dishes on a makeout session, and the Shame Wizard works his miserable magic on the whole class.
19
9 "Smooch or Share" Joel Moser Alex Rubens October 5, 2018 (2018-10-05)
Nick tries to lighten the mood at the sleepover with a new twist on Truth or Dare. The Shame Wizard meets his match. Jessi vows to change her ways.
20
10 "The Department of Puberty" Bob Suarez Gil Ozeri October 5, 2018 (2018-10-05)
After traveling through a magic portal to puberty headquarters, Nick seeks out a new Hormone Monster, and Jessi bonds with the Depression Kitty.
Production[edit]
TV writer Andrew Goldberg and screenwriter-directors Jennifer Flackett and Mark Levin approached Nick Kroll, Goldberg's best friend since childhood, with the idea to develop a show about going through puberty.[2] Kroll and Goldberg used their divergent pubertal experiences as a centerpiece of the show, because Kroll was a late bloomer while Goldberg went through the physical changes of puberty very early.[3] Many of their lived experiences are featured in the show, such as Kroll's first kiss, and Goldberg's parents waxing his mustache.[2] The show also includes an experience of their childhood friend, Lizzie, who the character of Jessi is based on, getting her first period on a school trip to the Statue of Liberty. According to Kroll in an interview on NPR, "[Big Mouth] takes an equal look at what it's like for girls and women, the process of going through puberty, which I think has not been quite as explored in most popular culture."[2]
Netflix announced they had picked up Big Mouth in June 2016.[4] Flackett, Levin, Goldberg, and Kroll all serve as executive producers on the show.
Release[edit]
The first season consisting of ten episodes premiered on Netflix on September 29, 2017.[5][6] On October 24, 2017, it was confirmed that a second season had been ordered,[7] which was released on October 5, 2018.[8]
Reception[edit]
Critical reception[edit]
Big Mouth has received critical acclaim since its official release. On Rotten Tomatoes, the series has an approval rating of 100% based on 21 reviews, with the critical consensus reading, "Big Mouth's simplistic animation and scatological humor belie its finely sketched characters and smart, empathetic approach to the messiness of adolescence."[9] At Metacritic, the show currently holds a rating of 80 out of 100, based on 6 critics, indicating "generally favorable reviews".[10]
Vox described the show as sharp and jarring depicting the awkwardness of pubescence, postpubescence and prepubescence.[11]
Rotten Tomatoes gave Season 2 a rating of 100% based on 21 reviews, with the critical consensus reading "Poignantly repulsive, Big Mouth continues to confront the awkwardness of adolescence with foul-mouthed glee and an added layer of maturity."[12]
Erik Adams from The A.V. Club awarded the second season an 'A-'. Adams praised the cast, especially Thewlis, and the experimentation of this season, commenting that "it’s shown that it deserves to be included in any conversation about TV’s animated greats."[13]
Accolades[edit]
Year
Award
Category
Nominee(s)
Result
Ref.
2018 Annie Award Best General Audience Television/Broadcast Production Big Mouth Nominated
2018 Primetime Emmy Award Outstanding Original Music and Lyrics Mark Rivers (for "Totally Gay") Nominated
References[edit]
Jump up ^ Wagmeister, Elizabeth (June 13, 2016). "Netflix Greenlights Animated Series from Nick Kroll". Variety. Retrieved June 13, 2016.
^ Jump up to: a b c Gross, Terry. "Nick Kroll and John Mulaney Relive Raging Hormones and First Kisses in 'Big Mouth'". NPR.org. NPR. Retrieved 6 October 2018.
Jump up ^ Schneider, Michael. "'Big Mouth' Creator Nick Kroll on Convincing Netflix to Let Him Make a 'Perverted Wonder Years' — Turn It On Podcast". IndieWire.com. Indie Wire. Retrieved 6 October 2018.
Jump up ^ Andreeva, Nellie. "Netflix Orders 'Big Mouth' Animated Series From Nick Kroll & Andrew Goldberg". Deadline.com. Retrieved 6 October 2018.
Jump up ^ Andreeva, Nellie (June 13, 2016). "Netflix Orders 'Big Mouth' Animated Series From Nick Kroll & Andrew Goldberg". Deadline. Retrieved August 31, 2016.
Jump up ^ Farley, Rebecca (August 23, 2017). "Exclusive: Meet The Hormone Monster, The Personification Of Puberty In Big Mouth". Refinery29. Retrieved August 23, 2017.
Jump up ^ Petski, Denise (October 24, 2017). "'Big Mouth' Renewed For Season 2 At Netflix". Deadline. Retrieved December 2, 2017.
Jump up ^ "'Big Mouth' season 2 premiere date, more puberty horrors revealed in Netflix teaser". EW.com. Retrieved 2018-08-29.
Jump up ^ "Big Mouth: Season 1". Rotten Tomatoes. Retrieved February 21, 2018.
Jump up ^ "Big Mouth", Metacritic, retrieved October 14, 2017
Jump up ^ "Netflix's Big Mouth takes a sharp, surprisingly joyful look at the gross time that is puberty". Vox. Retrieved 2017-10-15.
Jump up ^ "Big Mouth: Season 2". Rotten Tomatoes. Retrieved October 15, 2018.
Jump up ^ "Big Mouth returns, faster, funnier, and filthier than anything else on TV". The AV Club. Retrieved October 20, 2018.
External links[edit]
Big Mouth on Netflix
Big Mouth on IMDb
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Extracellular matrix
From Wikipedia, the free encyclopedia
Jump to navigation Jump to search
Extracellular matrix
Illustration depicting extracellular matrix (basement membrane and interstitial matrix) in relation to epithelium, endothelium and connective tissue
Details
Identifiers
Latin
Matrix Extracellularis
Acronym(s)
ECM
MeSH
D005109
TH
H2.00.03.0.02001
Anatomical terms of microanatomy
[edit on Wikidata]
In biology, the extracellular matrix (ECM) is a collection of extracellular molecules secreted by support cells that provides structural and biochemical support to the surrounding cells.[1][2] Because multicellularity evolved independently in different multicellular lineages, the composition of ECM varies between multicellular structures; however, cell adhesion, cell-to-cell communication and differentiation are common functions of the ECM.[3]
The animal extracellular matrix includes the interstitial matrix and the basement membrane.[4] Interstitial matrix is present between various animal cells (i.e., in the intercellular spaces). Gels of polysaccharides and fibrous proteins fill the interstitial space and act as a compression buffer against the stress placed on the ECM.[5] Basement membranes are sheet-like depositions of ECM on which various epithelial cells rest. Each type of connective tissue in animals has a type of ECM: collagen fibers and bone mineral comprise the ECM of bone tissue; reticular fibers and ground substance comprise the ECM of loose connective tissue; and blood plasma is the ECM of blood.
The plant ECM includes cell wall components, like cellulose, in addition to more complex signaling molecules.[6] Some single-celled organisms adopt multicellular biofilms in which the cells are embedded in an ECM composed primarily of extracellular polymeric substances (EPS).[7]
Contents
[hide]
1 Structure
1.1 Proteoglycans
1.1.1 Heparan sulfate
1.1.2 Chondroitin sulfate
1.1.3 Keratan sulfate
1.2 Non-proteoglycan polysaccharide
1.2.1 Hyaluronic acid
1.3 Proteins
1.3.1 Collagen
1.3.2 Elastin
1.4 Extracellular vesicles
1.5 Other
1.5.1 Fibronectin
1.5.2 Laminin
2 Development
3 Physiology
3.1 Stiffness and elasticity
3.1.1 Effect on gene expression
3.1.2 Effect on differentiation
3.1.3 Durotaxis
4 Function
4.1 Cell adhesion
5 Clinical significance
6 In plants
7 In pluriformea + filozoa
8 History
9 See also
10 References
11 Further reading
Structure[edit]
Components of the ECM are produced intracellularly by resident cells and secreted into the ECM via exocytosis.[8] Once secreted, they then aggregate with the existing matrix. The ECM is composed of an interlocking mesh of fibrous proteins and glycosaminoglycans (GAGs).
Proteoglycans[edit]
Glycosaminoglycans (GAGs) are carbohydrate polymers and mostly attached to extracellular matrix proteins to form proteoglycans (hyaluronic acid is a notable exception, see below). Proteoglycans have a net negative charge that attracts positively charged sodium ions (Na+), which attracts water molecules via osmosis, keeping the ECM and resident cells hydrated. Proteoglycans may also help to trap and store growth factors within the ECM.
Described below are the different types of proteoglycan found within the extracellular matrix.
Heparan sulfate[edit]
Heparan sulfate (HS) is a linear polysaccharide found in all animal tissues. It occurs as a proteoglycan (PG) in which two or three HS chains are attached in close proximity to cell surface or ECM proteins.[9][10] It is in this form that HS binds to a variety of protein ligands and regulates a wide variety of biological activities, including developmental processes, angiogenesis, blood coagulation, and tumour metastasis.
In the extracellular matrix, especially basement membranes, the multi-domain proteins perlecan, agrin, and collagen XVIII are the main proteins to which heparan sulfate is attached.
Chondroitin sulfate[edit]
Chondroitin sulfates contribute to the tensile strength of cartilage, tendons, ligaments, and walls of the aorta. They have also been known to affect neuroplasticity.[11]
Keratan sulfate[edit]
Keratan sulfates have a variable sulfate content and, unlike many other GAGs, do not contain uronic acid. They are present in the cornea, cartilage, bones, and the horns of animals.
Non-proteoglycan polysaccharide[edit]
Hyaluronic acid[edit]
Hyaluronic acid (or "hyaluronan") is a polysaccharide consisting of alternating residues of D-glucuronic acid and N-acetylglucosamine, and unlike other GAGs, is not found as a proteoglycan. Hyaluronic acid in the extracellular space confers upon tissues the ability to resist compression by providing a counteracting turgor (swelling) force by absorbing significant amounts of water. Hyaluronic acid is thus found in abundance in the ECM of load-bearing joints. It is also a chief component of the interstitial gel. Hyaluronic acid is found on the inner surface of the cell membrane and is translocated out of the cell during biosynthesis.[12]
Hyaluronic acid acts as an environmental cue that regulates cell behavior during embryonic development, healing processes, inflammation, and tumor development. It interacts with a specific transmembrane receptor, CD44.[13]
Proteins[edit]
Collagen[edit]
Collagens are the most abundant protein in the ECM. In fact, collagen is the most abundant protein in the human body[14][15] and accounts for 90% of bone matrix protein content.[16] Collagens are present in the ECM as fibrillar proteins and give structural support to resident cells. Collagen is exocytosed in precursor form (procollagen), which is then cleaved by procollagen proteases to allow extracellular assembly. Disorders such as Ehlers Danlos Syndrome, osteogenesis imperfecta, and epidermolysis bullosa are linked with genetic defects in collagen-encoding genes.[8] The collagen can be divided into several families according to the types of structure they form:
Fibrillar (Type I, II, III, V, XI)
Facit (Type IX, XII, XIV)
Short chain (Type VIII, X)
Basement membrane (Type IV)
Other (Type VI, VII, XIII)
Elastin[edit]
Elastins, in contrast to collagens, give elasticity to tissues, allowing them to stretch when needed and then return to their original state. This is useful in blood vessels, the lungs, in skin, and the ligamentum nuchae, and these tissues contain high amounts of elastins. Elastins are synthesized by fibroblasts and smooth muscle cells. Elastins are highly insoluble, and tropoelastins are secreted inside a chaperone molecule, which releases the precursor molecule upon contact with a fiber of mature elastin. Tropoelastins are then deaminated to become incorporated into the elastin strand. Disorders such as cutis laxa and Williams syndrome are associated with deficient or absent elastin fibers in the ECM.[8]
Extracellular vesicles[edit]
In 2016, Huleihel et al., reported the presence of DNA, RNA, and Matrix-bound nanovesicles (MBVs) within ECM bioscaffolds.[17] MBVs shape and size were found to be consistent with previously described exosomes. MBVs cargo includes different protein molecules, lipids, DNA, fragments, and miRNAs. Similar to ECM bioscaffolds, MBVs can modify the activation state of macrophages and alter different cellular properties such as; proliferation, migration and cell cycle. MBVs are now believed to be an integral and functional key component of ECM bioscaffolds.
Other[edit]
Fibronectin[edit]
Fibronectins are glycoproteins that connect cells with collagen fibers in the ECM, allowing cells to move through the ECM. Fibronectins bind collagen and cell-surface integrins, causing a reorganization of the cell's cytoskeleton to facilitate cell movement. Fibronectins are secreted by cells in an unfolded, inactive form. Binding to integrins unfolds fibronectin molecules, allowing them to form dimers so that they can function properly. Fibronectins also help at the site of tissue injury by binding to platelets during blood clotting and facilitating cell movement to the affected area during wound healing.[8]
Laminin[edit]
Laminins are proteins found in the basal laminae of virtually all animals. Rather than forming collagen-like fibers, laminins form networks of web-like structures that resist tensile forces in the basal lamina. They also assist in cell adhesion. Laminins bind other ECM components such as collagens and nidogens.[8]
Development[edit]
There are many cell types that contribute to the development of the various types of extracellular matrix found in plethora of tissue types. The local components of ECM determine the properties of the connective tissue.
Fibroblasts are the most common cell type in connective tissue ECM, in which they synthesize, maintain, and provide a structural framework; fibroblasts secrete the precursor components of the ECM, including the ground substance. Chondrocytes are found in cartilage and produce the cartilagenous matrix. Osteoblasts are responsible for bone formation.
Physiology[edit]
Stiffness and elasticity[edit]
The ECM can exist in varying degrees of stiffness and elasticity, from soft brain tissues to hard bone tissues. The elasticity of the ECM can differ by several orders of magnitude. This property is primarily dependent on collagen and elastin concentration,[18] and it has recently been shown to play an influential role in regulating numerous cell functions.
Cells can sense the mechanical properties of their environment by applying forces and measuring the resulting backlash.[19] This plays an important role because it helps regulate many important cellular processes including cellular contraction,[20] cell migration,[21] cell proliferation,[22] differentiation[23] and cell death (apoptosis).[24] Inhibition of nonmuscle myosin II blocks most of these effects,[23][21][20] indicating that they are indeed tied to sensing the mechanical properties of the ECM, which has become a new focus in research during the past decade.
Effect on gene expression[edit]
Differing mechanical properties in ECM exert effects on both cell behaviour and gene expression. Although the mechanism by which this is done has not been thoroughly explained, adhesion complexes and the actin-myosin cytoskeleton, whose contractile forces are transmitted through transcellular structures are thought to play key roles in the yet to be discovered molecular pathways.[20]
Effect on differentiation[edit]
ECM elasticity can direct cellular differentiation, the process by which a cell changes from one cell type to another. In particular, naive mesenchymal stem cells (MSCs) have been shown to specify lineage and commit to phenotypes with extreme sensitivity to tissue-level elasticity. MSCs placed on soft matrices that mimic brain differentiate into neuron-like cells, showing similar shape, RNAi profiles, cytoskeletal markers, and transcription factor levels. Similarly stiffer matrices that mimic muscle are myogenic, and matrices with stiffnesses that mimic collagenous bone are osteogenic.[23]
Durotaxis[edit]
Main article: Durotaxis
Stiffness and elasticity also guide cell migration, this process is called durotaxis. The term was coined by Lo CM and colleagues when they discovered the tendency of single cells to migrate up rigidity gradients (towards more stiff substrates)[21] and has been extensively studied since. The molecular mechanisms behind durotaxis are thought to exist primarily in the focal adhesion, a large protein complex that acts as the primary site of contact between the cell and the ECM.[25] This complex contains many proteins that are essential to durotaxis including structural anchoring proteins (integrins) and signaling proteins (adhesion kinase (FAK), talin, vinculin, paxillin, α-actinin, GTPases etc.) which cause changes in cell shape and actomyosin contractility.[26] These changes are thought to cause cytoskeletal rearrangements in order to facilitate directional migration.

Function[edit]
Due to its diverse nature and composition, the ECM can serve many functions, such as providing support, segregating tissues from one another, and regulating intercellular communication. The extracellular matrix regulates a cell's dynamic behavior. In addition, it sequesters a wide range of cellular growth factors and acts as a local store for them.[4] Changes in physiological conditions can trigger protease activities that cause local release of such stores. This allows the rapid and local growth factor-mediated activation of cellular functions without de novo synthesis.
Formation of the extracellular matrix is essential for processes like growth, wound healing, and fibrosis. An understanding of ECM structure and composition also helps in comprehending the complex dynamics of tumor invasion and metastasis in cancer biology as metastasis often involves the destruction of extracellular matrix by enzymes such as serine proteases, threonine proteases, and matrix metalloproteinases.[4][27]
The stiffness and elasticity of the ECM has important implications in cell migration, gene expression,[28] and differentiation.[23] Cells actively sense ECM rigidity and migrate preferentially towards stiffer surfaces in a phenomenon called durotaxis.[21] They also detect elasticity and adjust their gene expression accordingly which has increasingly become a subject of research because of its impact on differentiation and cancer progression.[29]
Cell adhesion[edit]
Many cells bind to components of the extracellular matrix. Cell adhesion can occur in two ways; by focal adhesions, connecting the ECM to actin filaments of the cell, and hemidesmosomes, connecting the ECM to intermediate filaments such as keratin. This cell-to-ECM adhesion is regulated by specific cell-surface cellular adhesion molecules (CAM) known as integrins. Integrins are cell-surface proteins that bind cells to ECM structures, such as fibronectin and laminin, and also to integrin proteins on the surface of other cells.
Fibronectins bind to ECM macromolecules and facilitate their binding to transmembrane integrins. The attachment of fibronectin to the extracellular domain initiates intracellular signalling pathways as well as association with the cellular cytoskeleton via a set of adaptor molecules such as actin.[5]
Clinical significance[edit]
See also: Regenerative medicine
Extracellular matrix has been found to cause regrowth and healing of tissue. Although the mechanism of action by which extracellular matrix promotes constructive remodeling of tissue is still unknown, researchers now believe that Matrix-bound nanovesicles (MBVs) are a key player in the healing process.[17][30] In human fetuses, for example, the extracellular matrix works with stem cells to grow and regrow all parts of the human body, and fetuses can regrow anything that gets damaged in the womb. Scientists have long believed that the matrix stops functioning after full development. It has been used in the past to help horses heal torn ligaments, but it is being researched further as a device for tissue regeneration in humans.[31]
In terms of injury repair and tissue engineering, the extracellular matrix serves two main purposes. First, it prevents the immune system from triggering from the injury and responding with inflammation and scar tissue. Next, it facilitates the surrounding cells to repair the tissue instead of forming scar tissue.[31]
For medical applications, the ECM required is usually extracted from pig bladders, an easily accessible and relatively unused source. It is currently being used regularly to treat ulcers by closing the hole in the tissue that lines the stomach, but further research is currently being done by many universities as well as the U.S. Government for wounded soldier applications. As of early 2007, testing was being carried out on a military base in Texas. Scientists are using a powdered form on Iraq War veterans whose hands were damaged in the war.[32]
Not all ECM devices come from the bladder. Extracellular matrix coming from pig small intestine submucosa are being used to repair "atrial septal defects" (ASD), "patent foramen ovale" (PFO) and inguinal hernia. After one year 95% of the collagen ECM in these patches is replaced by the normal soft tissue of the heart.[33]
Extracellular matrix proteins are commonly used in cell culture systems to maintain stem and precursor cells in an undifferentiated state during cell culture and function to induce differentiation of epithelial, endothelial and smooth muscle cells in vitro. Extracellular matrix proteins can also be used to support 3D cell culture in vitro for modelling tumor development.[34]
A class of biomaterials derived from processing human or animal tissues to retain portions of the extracellular matrix are called ECM Biomaterial.
In plants[edit]
Plant cells are tessellated to form tissues. The cell wall is the relatively rigid structure surrounding the plant cell. The cell wall provides lateral strength to resist osmotic turgor pressure, but it is flexible enough to allow cell growth when needed; it also serves as a medium for intercellular communication. The cell wall comprises multiple laminate layers of cellulose microfibrils embedded in a matrix of glycoproteins, including hemicellulose, pectin, and extensin. The components of the glycoprotein matrix help cell walls of adjacent plant cells to bind to each other. The selective permeability of the cell wall is chiefly governed by pectins in the glycoprotein matrix. Plasmodesmata (singular: plasmodesma) are pores that traverse the cell walls of adjacent plant cells. These channels are tightly regulated and selectively allow molecules of specific sizes to pass between cells.[12]
In pluriformea + filozoa[edit]
The extracellular matrix functionality of animals/metazoa developed in the common ancestor of the pluriformea and filozoa, after the Ichthyosporea diverged.
History[edit]
The importance of the extracellular matrix has long been recognized (Lewis, 1922), but the usage of the term is more recent (Gospodarowicz et al., 1979).[35][36][37][38]
See also[edit]
Perineuronal net
Interstitium
References[edit]
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^ Jump up to: a b 'Pixie dust' helps man grow new finger
Jump up ^ HowStuffWorks, Humans Can Regrow Fingers? In 2009, the St. Francis Heart Center announced the use of the extracellular matrix technology in repair surgery. Archived March 10, 2007, at the Wayback Machine.
Jump up ^ "First Ever Implantation of Bioabsorbable Biostar Device at DHZB". DHZB NEWS. December 2007. Retrieved 2008-08-05. The almost transparent collagen matrix consists of medically purified pig intestine, which is broken down by the scavenger cells (macrophages) of the immune system. After about 1 year the collagen has been almost completely (90-95%) replaced by normal body tissue: only the tiny metal framework remains. An entirely absorbable implant is currently under development.
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Further reading[edit]
[ ANAT3231 Lecture 08 Extracellular Matrix] - Lecture about extracellular matrix from UNSW Cell Biology website.
Extracellular matrix: review of its roles in acute and chronic wounds
Usage of Extracellular Matrix from pigs to regrow human extremities
"The Extracellular Matrix of Animals", from Chapter 19 of The Molecular Biology of the Cell, 4th edition, Alberts et al.
Biology, John W. Kimball. An online Biology textbook.
[3], The man who grew a finger, By Matthew Price, BBC news
Sound Medicine - Heart Tissue Regeneration - July 19 interview discussing ECM and its uses in cardiac tissue repair (requires MP3 playback) [broken link].
Growing Body Parts - A December 2009 report by 60 Minutes
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matrix
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EMILIN1
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Protein: scleroproteins
Extracellular matrix
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type II (COL2A1)
type III
type V
COL5A1
COL5A2
COL5A3
COL24A1
COL26A1
Other
FACIT: type IX
COL9A1
COL9A2
COL9A3
type XII (COL12A1)
COL14A1
COL16A1
COL19A1
COL20A1
COL21A1
COL22A1
basement membrane: type IV
COL4A1
COL4A2
COL4A3
COL4A4
COL4A5
COL4A6
multiplexin: COL15A1
type XVIII
COL18A1
Endostatin
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COL17A1
COL23A1
COL25A1
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COL6A1
COL6A2
COL6A3
COL6A5
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type VIII
COL8A1
COL8A2
type X (COL10A1)
type XI
COL11A1
COL11A2
COL27A1
COL28A1
Enzymes
Prolyl hydroxylase/Lysyl hydroxylase
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ADAMTS2
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beta
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FAM20C
ECM1
Matrix gla protein
Tectorin
TECTA
TECTB
Other
Keratin/Cytokeratin
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Cartilage oligomeric matrix protein
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