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'''Marginal zone B cells''' (MZ B cells) are noncirculating mature [[B cell]]s that in humans segregate anatomically into the [[marginal zone]] (MZ) of the [[spleen]]<ref>{{cite journal | author = Martin F, Kearney JF | year = 2002 | title = Marginal-zone B cells | journal = Nat Rev Immunol | volume = 2 | issue = 5| pages = 323–335 | doi=10.1038/nri799 | pmid = 12033738}}</ref> and certain other types of [[lymphoid tissue]].<ref name="pmid23348416">{{cite journal | vauthors = Cerutti A, Cols M, Puga I | title = Marginal zone B cells: virtues of innate-like antibody-producing lymphocytes | journal = Nature Reviews. Immunology | volume = 13 | issue = 2 | pages = 118–32 | date = February 2013 | pmid = 23348416 | pmc = 3652659 | doi = 10.1038/nri3383 }}</ref> The MZ B cells within this region typically express low-affinity polyreactive [[B-cell receptor|B-cell receptors]] (BCR), high levels of [[IgM]], [[Toll-like receptor|Toll-like receptors]] (TLRs), [[CD21]], [[CD1]], [[CD9]], [[CD27]] with low to negligible levels of secreted-[[IgD]], [[CD23]], [[CD5 (protein)|CD5]], and [[CD11b]] that help to distinguish them phenotypically from [[follicular B cell|follicular (FO) B cells]] and [[B-1 cell|B1 B cells]].<ref name="pmid23348416" /><ref name=":4">{{Cite journal| |
'''Marginal zone B cells''' (MZ B cells) are noncirculating mature [[B cell]]s that in humans segregate anatomically into the [[marginal zone]] (MZ) of the [[spleen]]<ref>{{cite journal | author = Martin F, Kearney JF | year = 2002 | title = Marginal-zone B cells | journal = Nat Rev Immunol | volume = 2 | issue = 5| pages = 323–335 | doi=10.1038/nri799 | pmid = 12033738| s2cid = 22573840 }}</ref> and certain other types of [[lymphoid tissue]].<ref name="pmid23348416">{{cite journal | vauthors = Cerutti A, Cols M, Puga I | title = Marginal zone B cells: virtues of innate-like antibody-producing lymphocytes | journal = Nature Reviews. Immunology | volume = 13 | issue = 2 | pages = 118–32 | date = February 2013 | pmid = 23348416 | pmc = 3652659 | doi = 10.1038/nri3383 }}</ref> The MZ B cells within this region typically express low-affinity polyreactive [[B-cell receptor|B-cell receptors]] (BCR), high levels of [[IgM]], [[Toll-like receptor|Toll-like receptors]] (TLRs), [[CD21]], [[CD1]], [[CD9]], [[CD27]] with low to negligible levels of secreted-[[IgD]], [[CD23]], [[CD5 (protein)|CD5]], and [[CD11b]] that help to distinguish them phenotypically from [[follicular B cell|follicular (FO) B cells]] and [[B-1 cell|B1 B cells]].<ref name="pmid23348416" /><ref name=":4">{{Cite journal|last1=Won|first1=Woong-Jai|last2=Kearney|first2=John F.|date=2002-06-01|title=CD9 is a unique marker for marginal zone B cells, B1 cells, and plasma cells in mice|url=https://pubmed.ncbi.nlm.nih.gov/12023357/|journal=Journal of Immunology (Baltimore, Md.: 1950)|volume=168|issue=11|pages=5605–5611|doi=10.4049/jimmunol.168.11.5605|issn=0022-1767|pmid=12023357|doi-access=free}}</ref> |
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MZ B cells are innate-like B cells specialized to mount rapid T-independent, but also T-dependent responses against blood-borne pathogens.<ref name=":0">{{Cite journal| |
MZ B cells are innate-like B cells specialized to mount rapid T-independent, but also T-dependent responses against blood-borne pathogens.<ref name=":0">{{Cite journal|last1=Turner|first1=Vivian M.|last2=Mabbott|first2=Neil A.|date=July 2017|title=Ageing adversely affects the migration and function of marginal zone B cells|url=https://pubmed.ncbi.nlm.nih.gov/28369800/|journal=Immunology|volume=151|issue=3|pages=349–362|doi=10.1111/imm.12737|issn=1365-2567|pmc=5461100|pmid=28369800}}</ref> They are also known to be the main producers of [[Immunoglobulin M|IgM]] antibodies in human.<ref name=":3" /> |
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== Development and Differentiation == |
== Development and Differentiation == |
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The spleen's [[marginal zone]] contains multiple subtypes of [[Macrophage|macrophages]] and [[Dendritic cell|dendritic cells]] interlaced with the MZ B cells; it is not fully formed until 2 to 3 weeks after birth in rodents and 1 to 2 years in humans.<ref>{{cite journal|author=MacLennan IC, Bazin H, Chassoux D|display-authors=etal|year=1985|title=Comparative analysis of the development of B cells in marginal zones and follicles|journal=Adv Exp Med Biol|volume=186|pages=139–144}}</ref> In humans, but not rodents, marginal zone B cells are also located in the inner wall of the subcapsular sinus of lymph nodes, the epithelium of tonsillar crypts, and the sub-epithelial area of mucosa-associated lymphoid tissues including the sub-epithelial dome of intestinal [[Peyer's patch|Peyer's patches]].<ref name="pmid23348416" /> Human MZ B cells are also present in peripheral blood, suggesting that they recirculate.<ref name=":1">{{Cite journal| |
The spleen's [[marginal zone]] contains multiple subtypes of [[Macrophage|macrophages]] and [[Dendritic cell|dendritic cells]] interlaced with the MZ B cells; it is not fully formed until 2 to 3 weeks after birth in rodents and 1 to 2 years in humans.<ref>{{cite journal|author=MacLennan IC, Bazin H, Chassoux D|display-authors=etal|year=1985|title=Comparative analysis of the development of B cells in marginal zones and follicles|journal=Adv Exp Med Biol|volume=186|pages=139–144|doi=10.1007/978-1-4613-2463-8_17|pmid=3876699|isbn=978-1-4612-9495-5}}</ref> In humans, but not rodents, marginal zone B cells are also located in the inner wall of the subcapsular sinus of lymph nodes, the epithelium of tonsillar crypts, and the sub-epithelial area of mucosa-associated lymphoid tissues including the sub-epithelial dome of intestinal [[Peyer's patch|Peyer's patches]].<ref name="pmid23348416" /> Human MZ B cells are also present in peripheral blood, suggesting that they recirculate.<ref name=":1">{{Cite journal|last1=Palm|first1=Anna-Karin E.|last2=Friedrich|first2=Heike C.|last3=Kleinau|first3=Sandra|date=2016-06-09|title=Nodal marginal zone B cells in mice: a novel subset with dormant self-reactivity|journal=Scientific Reports|language=en|volume=6|issue=1|pages=27687|doi=10.1038/srep27687|pmid=27277419|pmc=4899733|issn=2045-2322|doi-access=free}}</ref> However, in mice they seem to be noncirculating and only limited to follicular shuttling.<ref name="pmid23348416" /> |
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In rodents, MZ B cells are recognized as [[Immunoglobulin M|IgM]]<sup>high</sup>[[Immunoglobulin D|IgD]]<sup>low</sup>[[CD21]]<sup>high</sup>[[CD23]]<sup>low</sup> population of B cells. They are furthermore distiguished by expression of [[CD9]]<ref name=":4" /> and [[CD27]] (in human).<ref name="pmid23348416" /> In mice, MZ B cells characteristically express high levels of [[CD1D|CD1d]], which is a MHC class-I like molecule involved in presentation of lipid molecules to [[Natural killer T cell|NKT cells]].<ref name=":2" /> Unlike FO B cells, MZ B cells express polyreactive [[B-cell receptor|BCRs]] that bind to multiple [[Pathogen-associated molecular pattern|microbial molecular patterns]]. Additionally they express high levels of [[Toll-like receptor|TLRs]].<ref name="pmid23348416" /> |
In rodents, MZ B cells are recognized as [[Immunoglobulin M|IgM]]<sup>high</sup>[[Immunoglobulin D|IgD]]<sup>low</sup>[[CD21]]<sup>high</sup>[[CD23]]<sup>low</sup> population of B cells. They are furthermore distiguished by expression of [[CD9]]<ref name=":4" /> and [[CD27]] (in human).<ref name="pmid23348416" /> In mice, MZ B cells characteristically express high levels of [[CD1D|CD1d]], which is a MHC class-I like molecule involved in presentation of lipid molecules to [[Natural killer T cell|NKT cells]].<ref name=":2" /> Unlike FO B cells, MZ B cells express polyreactive [[B-cell receptor|BCRs]] that bind to multiple [[Pathogen-associated molecular pattern|microbial molecular patterns]]. Additionally they express high levels of [[Toll-like receptor|TLRs]].<ref name="pmid23348416" /> |
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In specimens where the tyrosine kinase for [[PTK2B|Pyk-2]] has been knocked-out, marginal zone B-cells will fail to develop while [[B-1 cell]]s will still be present. MZ B-cells are the only B-cells dependent on [[NOTCH2]] signaling for proliferation.<ref name="Paul0807">{{cite book|last1=Hardy|first1=Richard|title=Fundamental Immunology|publisher=Lippincott Williams & Wilkins|year=2008|isbn=0-7817-6519- |
In specimens where the tyrosine kinase for [[PTK2B|Pyk-2]] has been knocked-out, marginal zone B-cells will fail to develop while [[B-1 cell]]s will still be present. MZ B-cells are the only B-cells dependent on [[NOTCH2]] signaling for proliferation.<ref name="Paul0807">{{cite book|last1=Hardy|first1=Richard|title=Fundamental Immunology|publisher=Lippincott Williams & Wilkins|year=2008|isbn=978-0-7817-6519-0|editor1-last=Paul|editor1-first=William|edition=6th|location=Philadelphia|pages=237–269|chapter=Chapter 7: B Lymphocyte Development and Biology|type=Book}}</ref> |
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== Activation and Function == |
== Activation and Function == |
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Similar to [[B-1 cell|B1 B cells]], MZ B cells can be rapidly recruited into the early adaptive immune responses in a [[T cell]] independent manner.<ref name="Paul0807" /> The MZ B cells are especially well positioned as a first line of defense against systemic blood-borne antigens that enter the circulation and become trapped in the spleen.<ref>{{cite journal|author=Balazs M, Martin F, Zhou T|display-authors=etal|year=2002|title=Blood dendritic cells interact with splenic marginal zone B cells to initiate T-independent immune responses|journal=Immunity|volume=17|issue=3|pages=341–352|doi=10.1016/s1074-7613(02)00389-8}}</ref> While large blood-borne antigens are captured by [[Dendritic cell|dendritic cells]], circulating [[Granulocyte|granulocytes]] or MZ [[Macrophage|macrophages]], smaller blood-borne antigens may directly interact with MZ B cells situated on the exterior of the marginal sinus.<ref name="pmid23348416" /><ref name=":0" /> MZ B cells shuttle between the blood-filled marginal zone for antigen collection and the follicle for antigen delivery to [[follicular dendritic cells]]. In mice, it has been show that these cells shear flow via the [[LFA-1]] integrin ligand [[ICAM-1]] and adhere or migrate down the flow via the [[VLA-4]] integrin ligand [[VCAM-1]]. While [[CXCR5]]/[[CXCL13]] signaling is required for MZ B cells to enter the follicle, [[Sphingosine-1-phosphate|Sphingosine-1-phosphate signaling]] is required for MZ B cells exit from the follicle.<ref>{{Cite journal| |
Similar to [[B-1 cell|B1 B cells]], MZ B cells can be rapidly recruited into the early adaptive immune responses in a [[T cell]] independent manner.<ref name="Paul0807" /> The MZ B cells are especially well positioned as a first line of defense against systemic blood-borne antigens that enter the circulation and become trapped in the spleen.<ref>{{cite journal|author=Balazs M, Martin F, Zhou T|display-authors=etal|year=2002|title=Blood dendritic cells interact with splenic marginal zone B cells to initiate T-independent immune responses|journal=Immunity|volume=17|issue=3|pages=341–352|doi=10.1016/s1074-7613(02)00389-8|pmid=12354386}}</ref> While large blood-borne antigens are captured by [[Dendritic cell|dendritic cells]], circulating [[Granulocyte|granulocytes]] or MZ [[Macrophage|macrophages]], smaller blood-borne antigens may directly interact with MZ B cells situated on the exterior of the marginal sinus.<ref name="pmid23348416" /><ref name=":0" /> MZ B cells shuttle between the blood-filled marginal zone for antigen collection and the follicle for antigen delivery to [[follicular dendritic cells]]. In mice, it has been show that these cells shear flow via the [[LFA-1]] integrin ligand [[ICAM-1]] and adhere or migrate down the flow via the [[VLA-4]] integrin ligand [[VCAM-1]]. While [[CXCR5]]/[[CXCL13]] signaling is required for MZ B cells to enter the follicle, [[Sphingosine-1-phosphate|Sphingosine-1-phosphate signaling]] is required for MZ B cells exit from the follicle.<ref>{{Cite journal|last1=Tedford|first1=Kerry|last2=Steiner|first2=Michael|last3=Koshutin|first3=Stanislav|last4=Richter|first4=Karin|last5=Tech|first5=Laura|last6=Eggers|first6=Yannik|last7=Jansing|first7=Inga|last8=Schilling|first8=Kerstin|last9=Hauser|first9=Anja Erika|last10=Korthals|first10=Mark|last11=Fischer|first11=Klaus-Dieter|date=2017-12-22|title=The opposing forces of shear flow and sphingosine-1-phosphate control marginal zone B cell shuttling|journal=Nature Communications|language=en|volume=8|issue=1|pages=2261|doi=10.1038/s41467-017-02482-4|pmid=29273735|pmc=5741619|issn=2041-1723|doi-access=free}}</ref> |
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MZ B cells respond to a wide spectrum of T-independent, but also T-dependent antigens. It is believed that MZ B cells are especially reactive to microbial polysaccharide antigens of [[encapsulated bacteria]] such as ''[[Streptococcus pneumoniae]]'', ''[[Haemophilus influenzae]]'' and ''[[Neisseria meningitidis]]''. TLRs often activate MZ B cells after recognizing [[Pathogen-associated molecular pattern|microbial molecular structures]] in cooperation with the [[B-cell receptor|BCR]].<ref name=":1" /> These innate-like B cells provide rapid first line of defence against blood-borne pathogens and produce [[Antibody|antibodies]] of wide specificity before the induction of T-cell-dependent high-affinity antibody responses. Therefore MZ B cells may play an important role in the prevention of [[sepsis]].<ref name=":2">{{Cite journal| |
MZ B cells respond to a wide spectrum of T-independent, but also T-dependent antigens. It is believed that MZ B cells are especially reactive to microbial polysaccharide antigens of [[encapsulated bacteria]] such as ''[[Streptococcus pneumoniae]]'', ''[[Haemophilus influenzae]]'' and ''[[Neisseria meningitidis]]''. TLRs often activate MZ B cells after recognizing [[Pathogen-associated molecular pattern|microbial molecular structures]] in cooperation with the [[B-cell receptor|BCR]].<ref name=":1" /> These innate-like B cells provide rapid first line of defence against blood-borne pathogens and produce [[Antibody|antibodies]] of wide specificity before the induction of T-cell-dependent high-affinity antibody responses. Therefore MZ B cells may play an important role in the prevention of [[sepsis]].<ref name=":2">{{Cite journal|last1=Hendricks|first1=Jacobus|last2=Bos|first2=Nicolaas A.|last3=Kroese|first3=Frans G.M.|date=2018|title=Heterogeneity of Memory Marginal Zone B Cells|journal=Critical Reviews in Immunology|volume=38|issue=2|pages=145–158|doi=10.1615/CritRevImmunol.2018024985|issn=1040-8401|pmc=5989013|pmid=29887727}}</ref> MZ B cells also display a lower activation threshold than their FO B cell counterparts with heightened propensity for [[plasma cell]] differentiation that contributes further to the accelerated primary antibody response.<ref name="pmid23348416" /><ref>{{cite journal|author=Lopes-Carvalho T, Foote J, Kearney JF|year=2005|title=Marginal zone B cells in lymphocyte activation and regulation|journal=Curr Opin Immunol|volume=17|issue=3|pages=244–250|doi=10.1016/j.coi.2005.04.009|pmid=15886113}}</ref> They have been acknowledged as the main producers of [[Immunoglobulin M|IgM]] antibodies in human.<ref name=":3" /> |
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They are not only important for antibody-response toward invading pathogens, but also in maintaining homeostasis via opsonization of dead cells and cellular debris.<ref name=":3">{{Cite journal| |
They are not only important for antibody-response toward invading pathogens, but also in maintaining homeostasis via opsonization of dead cells and cellular debris.<ref name=":3">{{Cite journal|last1=Appelgren|first1=Daniel|last2=Eriksson|first2=Per|last3=Ernerudh|first3=Jan|last4=Segelmark|first4=Mårten|date=2018-10-02|title=Marginal-Zone B-Cells Are Main Producers of IgM in Humans, and Are Reduced in Patients With Autoimmune Vasculitis|journal=Frontiers in Immunology|volume=9|page=2242|doi=10.3389/fimmu.2018.02242|issn=1664-3224|pmc=6190848|pmid=30356862}}</ref> Moreover, MZ B cells are potent [[Antigen-presenting cell|antigen-presenting cells]], which are able to activate [[T helper cell|CD4+ T cells]] more effectively than FO B cells due to their elevated expression levels of [[MHC class II]], [[CD80]] and [[CD86]] molecules.<ref name="pmid23348416" /><ref name=":1" /> |
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Deficiencies of MZ B cells are associated with higher risk of [[pneumococcal infection]], [[meningitis]] and insufficient antibody response to capsular polysaccharides.<ref name="pmid23348416" /><ref name=":0" /> |
Deficiencies of MZ B cells are associated with higher risk of [[pneumococcal infection]], [[meningitis]] and insufficient antibody response to capsular polysaccharides.<ref name="pmid23348416" /><ref name=":0" /> |
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Revision as of 23:10, 25 January 2021
Marginal zone B cells (MZ B cells) are noncirculating mature B cells that in humans segregate anatomically into the marginal zone (MZ) of the spleen[1] and certain other types of lymphoid tissue.[2] The MZ B cells within this region typically express low-affinity polyreactive B-cell receptors (BCR), high levels of IgM, Toll-like receptors (TLRs), CD21, CD1, CD9, CD27 with low to negligible levels of secreted-IgD, CD23, CD5, and CD11b that help to distinguish them phenotypically from follicular (FO) B cells and B1 B cells.[2][3]
MZ B cells are innate-like B cells specialized to mount rapid T-independent, but also T-dependent responses against blood-borne pathogens.[4] They are also known to be the main producers of IgM antibodies in human.[5]
Development and Differentiation
The spleen's marginal zone contains multiple subtypes of macrophages and dendritic cells interlaced with the MZ B cells; it is not fully formed until 2 to 3 weeks after birth in rodents and 1 to 2 years in humans.[6] In humans, but not rodents, marginal zone B cells are also located in the inner wall of the subcapsular sinus of lymph nodes, the epithelium of tonsillar crypts, and the sub-epithelial area of mucosa-associated lymphoid tissues including the sub-epithelial dome of intestinal Peyer's patches.[2] Human MZ B cells are also present in peripheral blood, suggesting that they recirculate.[7] However, in mice they seem to be noncirculating and only limited to follicular shuttling.[2]
In rodents, MZ B cells are recognized as IgMhighIgDlowCD21highCD23low population of B cells. They are furthermore distiguished by expression of CD9[3] and CD27 (in human).[2] In mice, MZ B cells characteristically express high levels of CD1d, which is a MHC class-I like molecule involved in presentation of lipid molecules to NKT cells.[8] Unlike FO B cells, MZ B cells express polyreactive BCRs that bind to multiple microbial molecular patterns. Additionally they express high levels of TLRs.[2]
In specimens where the tyrosine kinase for Pyk-2 has been knocked-out, marginal zone B-cells will fail to develop while B-1 cells will still be present. MZ B-cells are the only B-cells dependent on NOTCH2 signaling for proliferation.[9]
Activation and Function
Similar to B1 B cells, MZ B cells can be rapidly recruited into the early adaptive immune responses in a T cell independent manner.[9] The MZ B cells are especially well positioned as a first line of defense against systemic blood-borne antigens that enter the circulation and become trapped in the spleen.[10] While large blood-borne antigens are captured by dendritic cells, circulating granulocytes or MZ macrophages, smaller blood-borne antigens may directly interact with MZ B cells situated on the exterior of the marginal sinus.[2][4] MZ B cells shuttle between the blood-filled marginal zone for antigen collection and the follicle for antigen delivery to follicular dendritic cells. In mice, it has been show that these cells shear flow via the LFA-1 integrin ligand ICAM-1 and adhere or migrate down the flow via the VLA-4 integrin ligand VCAM-1. While CXCR5/CXCL13 signaling is required for MZ B cells to enter the follicle, Sphingosine-1-phosphate signaling is required for MZ B cells exit from the follicle.[11]
MZ B cells respond to a wide spectrum of T-independent, but also T-dependent antigens. It is believed that MZ B cells are especially reactive to microbial polysaccharide antigens of encapsulated bacteria such as Streptococcus pneumoniae, Haemophilus influenzae and Neisseria meningitidis. TLRs often activate MZ B cells after recognizing microbial molecular structures in cooperation with the BCR.[7] These innate-like B cells provide rapid first line of defence against blood-borne pathogens and produce antibodies of wide specificity before the induction of T-cell-dependent high-affinity antibody responses. Therefore MZ B cells may play an important role in the prevention of sepsis.[8] MZ B cells also display a lower activation threshold than their FO B cell counterparts with heightened propensity for plasma cell differentiation that contributes further to the accelerated primary antibody response.[2][12] They have been acknowledged as the main producers of IgM antibodies in human.[5]
They are not only important for antibody-response toward invading pathogens, but also in maintaining homeostasis via opsonization of dead cells and cellular debris.[5] Moreover, MZ B cells are potent antigen-presenting cells, which are able to activate CD4+ T cells more effectively than FO B cells due to their elevated expression levels of MHC class II, CD80 and CD86 molecules.[2][7]
Deficiencies of MZ B cells are associated with higher risk of pneumococcal infection, meningitis and insufficient antibody response to capsular polysaccharides.[2][4]
Memory
In humans the splenic marginal zone B cells have evidence of somatic hypermutation in their immunoglobulin genes, indicating that they have been generated through a germinal centre reaction to become memory cells. While naive MZ B cells produce low-affinity IgM antibodies, memory MZ B cells express high-affinity Ig molecules. Besides unswitched cells (IgM+), class switched B cells can be found in the human and rodent marginal zone (IgG+ and IgA+). In humans, MZ B cells express CD27, which is a member of the TNF-receptor family expressed by human memory B cells.[8]
Role in Autoimmune diseases
Many of MZ B cell-receptors are self-reactive, which may be a factor that contributes to their expansion in some autoimmune diseases. On the other side, aiding in clearance of self-antigens is considered an important mechanism to prevent development of autoimmune diseases. The role of expanded self-reactive MZ B cells has been observed on mice models of lupus, diabetes and arthritis.[7] However, their levels in human vasculitis are reduced.[5]
Role in Tumors
Marginal zone B cells are the malignant cells in marginal zone lymphomas, a heterogeneous group of generally indolent lymphomas.[13]
References
- ^ Martin F, Kearney JF (2002). "Marginal-zone B cells". Nat Rev Immunol. 2 (5): 323–335. doi:10.1038/nri799. PMID 12033738. S2CID 22573840.
- ^ a b c d e f g h i j Cerutti A, Cols M, Puga I (February 2013). "Marginal zone B cells: virtues of innate-like antibody-producing lymphocytes". Nature Reviews. Immunology. 13 (2): 118–32. doi:10.1038/nri3383. PMC 3652659. PMID 23348416.
- ^ a b Won, Woong-Jai; Kearney, John F. (2002-06-01). "CD9 is a unique marker for marginal zone B cells, B1 cells, and plasma cells in mice". Journal of Immunology (Baltimore, Md.: 1950). 168 (11): 5605–5611. doi:10.4049/jimmunol.168.11.5605. ISSN 0022-1767. PMID 12023357.
- ^ a b c Turner, Vivian M.; Mabbott, Neil A. (July 2017). "Ageing adversely affects the migration and function of marginal zone B cells". Immunology. 151 (3): 349–362. doi:10.1111/imm.12737. ISSN 1365-2567. PMC 5461100. PMID 28369800.
- ^ a b c d Appelgren, Daniel; Eriksson, Per; Ernerudh, Jan; Segelmark, Mårten (2018-10-02). "Marginal-Zone B-Cells Are Main Producers of IgM in Humans, and Are Reduced in Patients With Autoimmune Vasculitis". Frontiers in Immunology. 9: 2242. doi:10.3389/fimmu.2018.02242. ISSN 1664-3224. PMC 6190848. PMID 30356862.
{{cite journal}}: CS1 maint: unflagged free DOI (link) - ^ MacLennan IC, Bazin H, Chassoux D; et al. (1985). "Comparative analysis of the development of B cells in marginal zones and follicles". Adv Exp Med Biol. 186: 139–144. doi:10.1007/978-1-4613-2463-8_17. ISBN 978-1-4612-9495-5. PMID 3876699.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ a b c d Palm, Anna-Karin E.; Friedrich, Heike C.; Kleinau, Sandra (2016-06-09). "Nodal marginal zone B cells in mice: a novel subset with dormant self-reactivity". Scientific Reports. 6 (1): 27687. doi:10.1038/srep27687. ISSN 2045-2322. PMC 4899733. PMID 27277419.
- ^ a b c Hendricks, Jacobus; Bos, Nicolaas A.; Kroese, Frans G.M. (2018). "Heterogeneity of Memory Marginal Zone B Cells". Critical Reviews in Immunology. 38 (2): 145–158. doi:10.1615/CritRevImmunol.2018024985. ISSN 1040-8401. PMC 5989013. PMID 29887727.
- ^ a b Hardy, Richard (2008). "Chapter 7: B Lymphocyte Development and Biology". In Paul, William (ed.). Fundamental Immunology (Book) (6th ed.). Philadelphia: Lippincott Williams & Wilkins. pp. 237–269. ISBN 978-0-7817-6519-0.
- ^ Balazs M, Martin F, Zhou T; et al. (2002). "Blood dendritic cells interact with splenic marginal zone B cells to initiate T-independent immune responses". Immunity. 17 (3): 341–352. doi:10.1016/s1074-7613(02)00389-8. PMID 12354386.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Tedford, Kerry; Steiner, Michael; Koshutin, Stanislav; Richter, Karin; Tech, Laura; Eggers, Yannik; Jansing, Inga; Schilling, Kerstin; Hauser, Anja Erika; Korthals, Mark; Fischer, Klaus-Dieter (2017-12-22). "The opposing forces of shear flow and sphingosine-1-phosphate control marginal zone B cell shuttling". Nature Communications. 8 (1): 2261. doi:10.1038/s41467-017-02482-4. ISSN 2041-1723. PMC 5741619. PMID 29273735.
- ^ Lopes-Carvalho T, Foote J, Kearney JF (2005). "Marginal zone B cells in lymphocyte activation and regulation". Curr Opin Immunol. 17 (3): 244–250. doi:10.1016/j.coi.2005.04.009. PMID 15886113.
{{cite journal}}: CS1 maint: multiple names: authors list (link) - ^ Bron D, Meuleman N (September 2019). "Marginal zone lymphomas: second most common lymphomas in older patients". Current Opinion in Oncology. 31 (5): 386–393. doi:10.1097/CCO.0000000000000554. PMID 31246587.