text:
White chocolate
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Chocolate derivative that does not contain cocoa solids
This article is about the confection. For other uses, see White chocolate (disambiguation).
White chocolate
Pieces of white chocolate
Type
Chocolate derivative
Main ingredients
cocoa butter, sugar, milk solids
Ingredients generally used
salt
Cookbook: White chocolate
Media: White chocolate
White chocolate is a chocolate derivative. It commonly consists of cocoa butter, sugar and milk solids and is characterized by a pale yellow or ivory appearance. The melting point of cocoa butter, white chocolate's only cocoa bean component, is high enough to keep white chocolate solid at room temperature, just like milk chocolate or dark chocolate.
Contents
1 Composition
2 Regulations
3 History
4 See also
5 References
Composition[edit]
This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (August 2018) (Learn how and when to remove this template message)
White chocolate does not contain non-fat cocoa solids, the primary constituent of chocolate liquor — chocolate in its raw, unsweetened form. During manufacturing, the dark-colored solids of the cocoa bean are separated from its fatty content, as with milk chocolate and dark chocolate. As a result, this cocoa butter is the only cacao ingredient in white chocolate. Because it contains no cocoa solids, white chocolate contains only trace amounts of the stimulants theobromine and caffeine. White chocolate may include additional flavorings, such as vanilla.[1]
Regulations[edit]
Regulations govern what may be marketed as "white chocolate": In the United States, since 2004, white chocolate must be (by weight) at least 20% cocoa butter, 14% total milk solids, and 3.5% milk fat, and no more than 55% sugar or other sweeteners.[2] Before this date, American firms required temporary marketing permits to sell white chocolate. The European Union has adopted the same standards, except that there is no limit on sugar or sweeteners.[3]
History[edit]
In the 1930s, the white chocolate Milkybar was launched in Europe by Swiss company Nestlé.[4]
In North America, Kuno Baedeker developed a white chocolate while working for the Merckens Chocolate Company in 1945.[5]
See also[edit]
Food portal
Types of chocolate
References[edit]
Wikimedia Commons has media related to White chocolate.
Jump up ^ Blumberg, Naomi. "Chocolate". Encyclopædia Britannica Online. Retrieved 28 July 2015.
Jump up ^ "Title 21 Chapter I Subchapter B Part 163 of the Code of Federal Regulations". United States Government Publishing Office. 24 February 2017. Retrieved 27 February 2017.
Jump up ^ "Directive 2000/36/EC of the European Parliament and of the Council of 23 June 2000 relating to cocoa and chocolate products intended for human consumption". Retrieved 27 October 2010.
Jump up ^ "The History Of White Chocolate". The Nibble. The World’s Best White Chocolate. 1 April 2008. Retrieved 2 August 2013.
Jump up ^ "Chocolate-Loving Couple Settled Here" (PDF). Lake Placid News (PDF). Lake Placid, New York. 19 March 1987. p. 8. Retrieved 2 August 2013.
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Category: Chocolate
Retrieved from "https://en.wikipedia.org/w/index.php?title=White_chocolate&oldid=863429672"
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text:
Blue Eyes Crying in the Rain
From Wikipedia, the free encyclopedia
Jump to: navigation, search
"Blue Eyes Crying in the Rain"
Single by Willie Nelson
from the album Red Headed Stranger
Released
July 1975
Format
7"
Recorded
February 9–12, 1975
Genre
Country
Length
2:18
Label
Columbia
Songwriter(s)
Fred Rose
Producer(s)
Willie Nelson
Willie Nelson singles chronology
"Sister's Coming Home"
(1974) "Blue Eyes Crying in the Rain"
(1975) "Fire and Rain"
(1976)
"Sister's Coming Home"
(1974) "Blue Eyes Crying in the Rain"
(1975) "Fire and Rain"
(1976)
Music video
"Blue Eyes Crying In The Rain" on YouTube
"Blue Eyes Crying in the Rain" is a song written by songwriter Fred Rose. Originally performed by Roy Acuff, the song has been covered by many artist; such as Hank Williams Sr. and Charlie Pride. Also the song was later recorded by Willie Nelson as part of his 1975 album Red Headed Stranger. Both the song and album would become iconic in country music history, and jump start Nelson's success as a singer and recording artist.
Contents
[hide]
1 Song history
2 Chart success and legacy
3 Chart performance
4 Later versions
5 Sources
5.1 References
6 External links
Song history[edit]
Originally recorded in 1947 by Acuff, "Blue Eyes Crying in the Rain" was recorded by Hank Williams in 1951 for the Mother's Best Flour Hour. Other early remakes of the song were made by Donn Reynolds (MGM single - June 1957), Ferlin Husky (album Ferlin's Favorites - November 1959), Slim Whitman (album Country Favorites - 1959), Gene Vincent (recorded October 15, 1958/ album Crazy Times! -1960), Bill Anderson (album...Sings Country Heart Songs - January 15, 1962), John D. Loudermilk (album Country Love Songs Plain and Simply Sung Country Love Songs Plain and Simply Sung - August 1968), (Hank Snow)] (album, 'Greatest Hits" ) and Conway Twitty (album Hello Darlin' - June 1970).
The version by Willie Nelson, recorded for his 1975 concept album, Red Headed Stranger, which was about a fugitive preacher on the run from the law after killing his wife, was lauded by country music historian Bill Malone as "a fine example of clean, uncluttered country music, [with] a spare arrangement that could have come straight out of the 1940s."[1] Rolling Stone noted the song was delivered with his "jazz-style phrasing" and was "the beating heart of Red Headed Stranger."[2]
A music video for the song was created in 1986 to promote the motion picture adaptation of the album.
Chart success and legacy[edit]
Prior to the success of "Blue Eyes Crying in the Rain," Nelson had enjoyed widespread success primarily as a songwriter, with such songs as "Crazy" (Patsy Cline) and "Hello Walls" (Faron Young). As a performer, meanwhile, Nelson had hit the Top 10 of the Billboard magazine Hot Country Singles chart just twice; it had happened in 1962, once as a solo artist ("Touch Me") and again as part of a duet with Shirley Collie ("Willingly"). Thereafter, Nelson had approached the Top 20 on occasion, but went 13 years without a Top 10 hit.
In October 1975, the song became Nelson's first No. 1 hit as a singer, and at year's end was the third-biggest song of 1975 on the Billboard Hot Country Singles chart.[3] In addition, the song gained modest airplay on Top 40 radio, reaching number 21 on the Billboard Hot 100.[4]
In 2004, Rolling Stone ranked "Blue Eyes Crying in the Rain" #302 on its list of the 500 Greatest Songs of All Time.[5]
Chart performance[edit]
Chart (1975)
Peak
position
US Hot Country Songs (Billboard)[6] 1
US Billboard Hot 100[7] 21
Canadian RPM Country Tracks 2
Canadian RPM Top Singles 40
Canadian RPM Adult Contemporary Tracks 9
Australian Kent Music Report 57
Preceded by
"Daydreams About Night Things" by Ronnie Milsap Billboard Hot Country Singles
number-one single
October 4, 1975 – October 11, 1975 Succeeded by
"Hope You're Feelin' Me (Like I'm Feelin' You)" by Charley Pride
Later versions[edit]
Elvis Presley finally recorded "Blue Eyes Crying in the Rain" (having sung it privately with friends and family for years while accompanying himself on piano) in the Jungle Room at Graceland on 7 February 1976. This was the last known song that Elvis Presley sang (at the piano in the rest area of his Racquetball Court located to the rear of Graceland) before his death on August 16, 1977.[8]
Ken Kincaid, a New Zealand singer recorded a version which appears on the B side of his 1982 single "Cindy", CBS BA 222 941.[9][10]
Linda Good's version of the song was featured on the compilation album Sony/ATV Nashville Classic Covers, Vol.1 (2008).[11]
A live version by Willie Nelson and Shania Twain was included on Nelson's 2003 live album Live & Kickin': Willie Nelson and Friends.[12]
UB40 covered the song in their 2013 album Getting Over the Storm.[13]
Sources[edit]
References[edit]
Jump up ^ Malone, Bill, "The Smithsonian Collection of Classic Country Music" ((booklet included with The Smithsonian Collection of Classic Country Music 8-volume set). Smithsonian Institution, 1981).
Jump up ^ ain "Blue Eyes Crying in the Rain," from "The RS 500 Greatest Songs of All Time," Rolling Stone, November 2004.
Jump up ^ [1] Billboard magazine Hot Country Singles year-end chart - 1975.
Jump up ^ Whitburn, Joel (2004). The Billboard Book of Top 40 Hits: Eighth Edition. Record Research. p. 450.
Jump up ^ [2] "The RS 500 Greatest Songs of All Time," Rolling Stone.
Jump up ^ "Willie Nelson – Chart history" Billboard Hot Country Songs for Willie Nelson.
Jump up ^ "Willie Nelson – Chart history" Billboard Hot 100 for Willie Nelson.
Jump up ^ "Rescued Chile miner tours Graceland as Elvis fan". Foxnews. 7 January 2011. Retrieved 13 July 2013.
Jump up ^ 45cat Ken Kincaid - Discography
Jump up ^ The New Zealand Archive of Film, Television and Sound Ngā Taonga Whitiāhua Me Ngā Taonga Kōrero Catalogue → 18497, Cindy
Jump up ^ iTunes Sony/ATV Nashville Classic Covers: Volume One, Various Artists
Jump up ^ "Flashback: See Shania Twain, Willie Nelson Sing 'Blue Eyes Crying'". Rolling Stone. August 25, 2016.
Jump up ^ Discogs UB40 – Getting Over The Storm
External links[edit]
Lyrics of this song at MetroLyrics
[hide]
v
t
e
Willie Nelson singles discography
1950s
and
1960s
1957
"No Place For Me" / "Lumberjack"
1959
"Man With the Blues" / "The Storm Has Just Begun"
1960
"What a Way to Live" / "Misery Mansion"
"Nite Life" / "Rainy Day Blues"
1961
"The Part Where I Cry" / "Mr. Record Man"
1962
"Willingly" / "Chain of Love"
"Touch Me" / "Where My House Lives"
"Wake Me When It's Over" / "There's Gonna Be Love In My House"
"You Dream About Me" / "Is This My Destiny"
1963
"Half a Man" / "The Last Letter"
"Take My Word" / "Feed it a Memory"
1964
"How Long is Forever" / "You Took My Happy Away"
"Am I Blue" / "There'll Be No Teardrops Tonight"
"River Boy" / "Opportunity to Cry"
"I Never Cared For You" / "You Left Me (A Long Time Ago)"
"Pretty Paper" / "What A Merry Christmas This Could Be"
1965
"She's Not For You" / "Permanently Lonely"
"Healing Hands of Time" / "One Day at the Time"
"I Just Can't Let You Say Goodbye" / "And So Will You, My Love"
1966
"One In A Row" / "San Antonio Rose"
"Colombus Stockade Blues" / "He Sits at my Table"
"I'm Still Not Over You"/ "I Love You Because"
1967
"The Party's Over" / "Make Way for a Better Man"
"Blackjack County Chain" / "Some Other World"
"San Antonio" / "To Make a Long Story Short"
1968
"Little Things" / " Sweet Memories"
"Good Times" / "Where Do You Stand"
"Johnny One Time" / "She's Still Gone"
1969
"Bring Me Sunshine" / "Don't Say Love or Nothing"
"I Hope So" / "Right or Wrong"
1970s
1970
"Once More With Feeling" / "Who Do I Know in Dallas"
"Laying My Burdens Down" / "Truth Number One"
1971
"I'm a Memory" / "Fire and Rain"
"Yesterday's Wine" / "Me and Paul"
1972
"Words Don't Fit the Picture" / "A Moment"
1973
"Shotgun Willie" (mono) / "Sad Songs and Waltzes"
"Stay All Night (Stay a Little Longer)" / "Devil in a Sleepin' Bag"
1974
"I Still Can't Believe You're Gone" / "Heaven and Hell"
"Bloody Mary Morning" / "After the Fire is Gone"(with Tracy Nelson)
"Sister's Comin' Home" (mono) / "Pick Up The Tempo"
1975
"Blue Eyes Crying in the Rain" / "Bandera"
1976
"Remember Me (When the Candle Lights Are Gleaming)" / "Time of the Preacher"
"I'd Have to Be Crazy" (mono) / "Amazing Grace"
"If You've Got the Money I've Got the Time" / "The Sound in Your Mind"
1977
"Uncloudy Day" / "Precious Memories"
"I Love You A Thousand Ways" / "Mom and Dad's Waltz"
1978
"Georgia on My Mind" / "On The Sunny Side Of The Street"
"Blue Skies" / "Moonlight in Vermont"
"All of Me" / "Unchained Melody"
1979
"Whiskey River" / "Under the Double Eagle"
"September Song" / "Don't Get Around Much Anymore"
"White Christmas" / "Blue Christmas"
1980s
1980
"Help Me Make It Through the Night" / "The Pilgrim, Chapter 33"
"My Heroes Have Always Been Cowboys" / "Rising Stars"
"Midnight Rider" / "So You Think You're A Cowboy"
"On the Road Again"/ "Jumpin' Cotton Eyed Joe"(performed by Johnny Gimble)
1981
"Angel Flying Too Close to the Ground" / "I Guess I've Come to Live Here in Your Eyes"
"Mona Lisa" / "Twinkle, Twinkle Little Star"
"I'm Gonna Sit Right Down and Write Myself a Letter" / "Over the Rainbow"
"Heartaches of a Fool" / "Uncloudy Day"
1982
"Always on My Mind" / "The Party's Over"
"Let It Be Me" / "Permanently Lonely"
"Last Thing I Needed First Thing This Morning" / "Old Fords and a Natural Stone"
1983
"Little Old Fashioned Karma" / "Beer Barrel Polka"
"Why Do I Have to Choose" / "Would You Lay with Me (In a Field of Stone)"
"Take It to the Limit" / "Till I Gain Control Again"
"Without a Song"
1984
"City of New Orleans (Unedited version)" / "Why Are You Pickin' On Me"
1985
"Forgiving You Was Easy" / "You Wouldn't Cross the Street"
"Me and Paul" / "I Let My Mind Wander"
1986
"Living in the Promiseland" / "Bach Minuet in G"
"I'm Not Trying to Forget You" / "I've Got the Craziest Feeling"
1987
"Partners After All" / "Home Away From Home"
"Heart of Gold" / "So Much Like My Dad"
"Island in the Sea" / "There Is No Easy Way"
1988
"Nobody There But Me" / "Wake Me When It's Over"
"Spanish Eyes" (with Julio Iglesias) / "Ole Buttermilk Sky"
"Twilight Time" / "Ac-Cent-Tchu-Ate the Positive"
1989
"Nothing I Can Do About It Now" / "If I Were a Painting"
"There You Are" / "Spirit"
"Is The Better Part Over" / "Mr. Record Man"
1990s
1990
"The Highway" / "Spirit"
"Ain't Necessarily So" / "I Never Cared For You" ("Ain't Necessarily So" also on CD Single)
1991
"The Piper Came Today" / "(I Don't Have a Reason) To Go To California Anymore" ("The Piper Came Today" also on CD Single)
"Ten With a Two" / "You Decide"
1993
"Graceland"
"Still is Still Moving to Me" / "Valentine"
1995
"Turn Me Loose and Let Me Swing"
1998
"I Never Cared for You"
2000s
and
2010s
2002
"Mendocino County Line" (with Lee Ann Womack) / "Maria (Shut Up and Kiss Me)" (Both single tracks released on CD and 7" record)
2003
"Wurlitzer Prize" (with Norah Jones)
"Beer for My Horses" (with Toby Keith)
2005
"I'm a Worried Man" (with Toots Hibbert)
"The Harder They Come"
2006
"You Don't Know Me"
"Cowboys Are Frequently, Secretly Fond of Each Other"
2008
"Gravedigger"
"You Don't Think I'm Funny Anymore"
2011
"The Scientist"
2012
"Roll Me Up and Smoke Me When I Die"
"Just Breathe"
"Come On Back Jesus"
2013
"From Here to the Moon and Back" (with Dolly Parton)
"Grandma's Hands" (with Mavis Staples)
"It Won't Be Long" (with The Secret Sisters)
"Somewhere Between" (with Loretta Lynn)
2014
"The Wall"
"Bring It On"
"Laws of Nature"
"Who'll Buy My Memories"
2015
"It's All Going to Pot"
Book:Willie Nelson
Category:Willie Nelson
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B cell
From Wikipedia, the free encyclopedia
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This article is about the immune system cell. For the electrical cell, see Battery (vacuum tube). For β-cell of pancreas, see beta cell.
Basic B cell function: bind to an antigen, receive help from a cognate helper T cell, and differentiate into a plasma cell that secretes large amounts of antibodies
3D rendering of a B cell
B cells, also known as B lymphocytes, are a type of white blood cell of the lymphocyte subtype.[1] They function in the humoral immunity component of the adaptive immune system by secreting antibodies.[1] Additionally, B cells present antigen (they are also classified as professional antigen-presenting cells (APCs)) and secrete cytokines.[1] In mammals, B cells mature in the bone marrow, which is at the core of most bones.[2] In birds, B cells mature in the bursa of Fabricius, a lymphoid organ. (The "B" from B cells comes from the name of this organ, where it was first discovered by Chang and Glick,[2] and not from bone marrow as commonly believed).
B cells, unlike the other two classes of lymphocytes, T cells and natural killer cells, express B cell receptors (BCRs) on their cell membrane.[1] BCRs allow the B cell to bind to a specific antigen, against which it will initiate an antibody response.[1]
Contents
[hide]
1 Development
2 Activation
2.1 T cell-dependent activation
2.2 T cell-independent activation
2.3 Memory B cell activation
3 B cell types
4 B cell-related pathology
5 Epigenetic
6 References
Development[edit]
B cells develop from hematopoietic stem cells (HSCs) that originate from bone marrow.[3] HSCs first differentiate into multipotent progenitor (MPP) cells, then common lymphoid progenitor (CLP) cells.[3] From here, their development into B cells occurs in several stages (shown in image to the right), each marked by various gene expression patterns and immunoglobulin H chain and L chain gene loci arrangements, the latter due to B cells undergoing V(D)J recombination as they develop.[4]
Early B cell development: from stem cell to immature B cell
B cells undergo two types of selection while developing in the bone marrow to ensure proper development. Positive selection occurs through antigen-independent signaling involving both the pre-BCR and the BCR.[5][6] If these receptors do not bind to their ligand, B cells do not receive the proper signals and cease to develop.[5][6] Negative selection occurs through the binding of self-antigen with the BCR; If the BCR can bind strongly to self-antigen, then the B cell undergoes one of four fates: clonal deletion, receptor editing, anergy, or ignorance (B cell ignores signal and continues development).[6] This negative selection process leads to a state of central tolerance, in which the mature B cells don't bind with self antigens present in the bone marrow.[4]
To complete development, immature B cells migrate from the bone marrow into the spleen as transitional B cells, passing through two transitional stages: T1 and T2.[7] Throughout their migration to the spleen and after spleen entry, they are considered T1 B cells.[8] Within the spleen, T1 B cells transition to T2 B cells.[8] T2 B cells differentiate into either follicular (FO) B cells or marginal zone (MZ) B cells depending on signals received through the BCR and other receptors.[9] Once differentiated, they are now considered mature B cells, or naive B cells.[8]
Transitional B cell development: from immature B cell to MZ B cell or mature (FO) B cell
While immature and during the T1 phase, B cells express BCR of class IgG, but BCR expression changes to the classes IgM and IgD after transition into the T2 phase and while mature up to activation.[8]
Activation[edit]
B cell activation occurs in the secondary lymphoid organs (SLOs), such as the spleen and lymph nodes.[1] After B cells mature in the bone marrow, they migrate through the blood to SLOs, which receive a constant supply of antigen through circulating lymph.[10] At the SLO, B cell activation begins when the B cell binds to an antigen via its BCR.[11] Of the three B cell subsets, FO B cells preferentially undergo T cell-dependent activation while MZ B cells and B1 B cells preferentially undergo T cell-independent activation.[12]
B cell activation: from immature B cell to plasma cell or memory B cell
B cell activation is enhanced through the activity of CD21, a surface receptor in complex with surface proteins CD19 and CD81 (all three are collectively known as the B cell coreceptor complex).[13] When a BCR binds an antigen tagged with a fragment of the C3 complement protein, CD21 binds the C3 fragment, co-ligates with the bound BCR, and signals are transduced through CD19 and CD81 to lower the activation threshold of the cell.[14]
T cell-dependent activation[edit]
Antigens that activate B cells with the help of T-cell are known as T cell-dependent (TD) antigens and include foreign proteins.[1] They are named as such because they are unable to induce a humoral response in organisms that lack T cells.[1] B cell response to these antigens takes multiple days, though antibodies generated have a higher affinity and are more functionally versatile than those generated from T cell-independent activation.[1]
Once a BCR binds a TD antigen, the antigen is taken up into the B cell through receptor-mediated endocytosis, degraded, and presented to T cells as peptide pieces in complex with MHC-II molecules on the cell membrane.[15] T helper (TH) cells, typically follicular T helper (TFH) cells, that were activated with the same antigen recognize and bind these MHC-II-peptide complexes through their T cell receptor (TCR).[16] Following TCR-MHC-II-peptide binding, T cells express the surface protein CD40L as well as cytokines such as IL-4 and IL-21.[16] CD40L serves as a necessary co-stimulatory factor for B cell activation by binding the B cell surface receptor CD40, which promotes B cell proliferation, immunoglobulin class switching, and somatic hypermutation as well as sustains T cell growth and differentiation.[1] T cell-derived cytokines bound by B cell cytokine receptors also promote B cell proliferation, immunoglobulin class switching, and somatic hypermutation as well as guide differentiation.[16] After B cells receive these signals, they are considered activated.[16]
Now activated, B cells participate in a two-step differentiation process that yields both short-lived plasmablasts for immediate protection and long-lived plasma cells and memory B cells for persistent protection.[12] The first step, known as the extrafollicular response, occurs outside lymphoid follicles but still in the SLO.[12] During this step activated B cells proliferate, may undergo immunoglobulin class switching, and differentiate into plasmablasts that produce early, weak antibodies mostly of class IgM.[17] The second step consists of activated B cells entering a lymphoid follicle and forming a germinal center (GC), which is a specialized microenvironment where B cells undergo extensive proliferation, immunoglobulin class switching, and affinity maturation directed by somatic hypermutation.[18] These processes are facilitated by TFH cells within the GC and generate both high-affinity memory B cells and long-lived plasma cells.[12] Resultant plasma cells secrete large amounts of antibody and either stay within the SLO or, more preferentially, migrate to bone marrow.[18]
T cell-independent activation[edit]
Main article: T independent antigen (TI)
Antigens that activate B cells without T cell help are known as T cell-independent (TI) antigens[1] and include foreign polysaccharides and unmethylated CpG DNA.[12] They are named as such because they are able to induce a humoral response in organisms that lack T cells.[1] B cell response to these antigens is rapid, though antibodies generated tend to have lower affinity and are less functionally versatile than those generated from T cell-dependent activation.[1]
As with TD antigens, B cells activated by TI antigens need additional signals to complete activation, but instead of receiving them from T cells, they are provided either by recognition and binding of a common microbial constituent to toll-like receptors (TLRs) or by extensive crosslinking of BCRs to repeated epitopes on a bacterial cell.[1] B cells activated by TI antigens go on to proliferate outside lymphoid follicles but still in SLOs (GCs do not form), possibly undergo immunoglobulin class switching, and differentiate into short-lived plasmablasts that produce early, weak antibodies mostly of class IgM, but also some populations of long-lived plasma cells.[19]
Memory B cell activation[edit]
Memory B cell activation begins with the detection and binding of their target antigen, which is shared by their parent B cell.[20] Some memory B cells can be activated without T cell help, such as certain virus-specific memory B cells, but others need T cell help.[21] Upon antigen binding, the memory B cell takes up the antigen through receptor-mediated endocytosis, degrades it, and presents it to T cells as peptide pieces in complex with MHC-II molecules on the cell membrane.[20] Memory T helper (TH) cells, typically memory follicular T helper (TFH) cells, that were derived from T cells activated with the same antigen recognize and bind these MHC-II-peptide complexes through their TCR.[20] Following TCR-MHC-II-peptide binding and the relay of other signals from the memory TFH cell, the memory B cell is activated and differentiates either into plasmablasts and plasma cells via an extrafollicular response or enter a germinal center reaction where they generate plasma cells and more memory B cells.[20][21] It is unclear whether the memory B cells undergo further affinity maturation within these secondary GCs.[20]
B cell types[edit]
Plasmablast - A short-lived, proliferating antibody-secreting cell arising from B cell differentiation.[1] Plasmablasts are generated early in an infection and their antibodies tend to have a weaker affinity towards their target antigen compared to plasma cell.[12] Plasmablasts can result from T cell-independent activation of B cells or the extrafollicular response from T cell-dependent activation of B cells.[1]
Plasma cell - A long-lived, non-proliferating antibody-secreting cell arising from B cell differentiation.[1] There is evidence that B cells first differentiate into a plasmablast-like cell, then differentiate into a plasma cell.[12] Plasma cells are generated later in an infection and, compared to plasmablasts, have antibodies with a higher affinity towards their target antigen due to affinity maturation in the germinal center (GC) and produce more antibodies.[12] Plasma cells typically result from the germinal center reaction from T cell-dependent activation of B cells, however they can also result from T cell-independent activation of B cells.[19]
Lymphoplasmacytoid cell - A cell with a mixture of B lymphocyte and plasma cell morphological features that is thought to be closely related to or a subtype of plasma cells. This cell type is found in pre-malignant and malignant plasma cell dyscrasias that are associated with the secretion of IgM monoclonal proteins; these dyscrasias include IgM monoclonal gammopathy of undetermined significance and Waldenström's macroglobulinemia.[22]
Memory B cell - Dormant B cell arising from B cell differentiation.[1] Their function is to circulate through the body and initiate a stronger, more rapid antibody response (known as the anamnestic secondary antibody response) if they detect the antigen that had activated their parent B cell (memory B cells and their parent B cells share the same BCR, thus they detect the same antigen).[21] Memory B cells can be generated from T cell-dependent activation through both the extrafollicular response and the germinal center reaction as well as from T cell-independent activation of B1 cells.[21]
Follicular (FO) B Cell (also known as a B-2 cell) - Most common type of B cell and, when not circulating through the blood, is found mainly in the lymphoid follicles of secondary lymphoid organs (SLOs).[12] They are responsible for generating the majority of high-affinity antibodies during an infection.[1]
Marginal zone (MZ) B cell - Found mainly in the marginal zone of the spleen and serves as a first line of defense against blood-borne pathogens, as the marginal zone receives large amounts of blood from the general circulation.[23] They can undergo both T cell-independent and T cell-dependent activation, but preferentially undergo T cell-independent activation.[12]
B-1 cell - Arises from a developmental pathway different from FO B cells and MZ B cells.[24] In mice, they predominantly populate the peritoneal cavity and pleural cavity, generate natural antibodies (antibodies produced without infection), defend against mucosal pathogens, and primarily exhibit T cell-independent activation.[24] A true homologue of mouse B-1 cells has not been discovered in humans, though various cell populations similar to B-1 cells have been described.[24]
B-2 cell - FO B cells and MZ B cells.[24]
Regulatory B (Breg) cell - An immunosuppressive B cell type that stops the expansion of pathogenic, pro-inflammatory lymphocytes through the secretion of IL-10, IL-35, and TGF-β.[25] Also, it promotes the generation of regulatory T (Treg) cells by directly interacting with T cells to skew their differentiation towards Tregs.[25] No common Breg cell identity has been described and many Breg cell subsets sharing regulatory functions have been found in both mice and humans.[25] It is currently unknown if Breg cell subsets are developmentally linked and how exactly differentiation into a Breg cell occurs.[25] There is evidence showing that nearly all B cell types can differentiate into a Breg cell through mechanisms involving inflammatory signals and BCR recognition.[25]
B cell-related pathology[edit]
Autoimmune disease can result from abnormal B cell recognition of self-antigens followed by the production of autoantibodies.[26] Autoimmune diseases where disease activity is correlated with B cell activity include scleroderma, multiple sclerosis, systemic lupus erythematosus, type 1 diabetes, and rheumatoid arthritis.[26]
Malignant transformation of B cells and their precursors can cause a host of cancers, including chronic lymphocytic leukemia (CLL), acute lymphoblastic leukemia (ALL), hairy cell leukemia, follicular lymphoma, non-Hodgkin's lymphoma, Hodgkin's lymphoma, and plasma cell malignancies such as multiple myeloma, Waldenström's macroglobulinemia, and certain forms of amyloidosis.[27][28]
Epigenetic[edit]
A study that investigated the methylome of B cells along their differentiation cycle, using whole-genome bisulfite sequencing (WGBS), showed that there is a hypomethylation from the earliest stages to the most differentiated stages. The largest methylation difference is between the stages of germinal center B cells and memory B cells. Furthermore, this study showed that there is a similarity between B cell tumors and long-lived B cells in their DNA methylation signatures.[29]
References[edit]
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