Maus-IgM-Standardsatz, A-F

Natural immunoglobulin M (IgM) antibodies are pentameric or hexameric macroimmunoglobulins and have been highly conserved throughout evolution. IgMs are initially expressed during B cell ontogeny and are the first antibodies to be secreted upon exposure to foreign antigens. The IgM multimer has either 10 (pentamer) or 12 (hexamer) antigen binding domains consisting of paired μ heavy chains with four constant domains, each with a single variable domain paired with a corresponding light chain. Although the antigen-binding affinities of natural IgM antibodies are typically lower than that of IgG, their polyvalency allows for high avidity binding and efficient complement binding to induce complement-dependent cell lysis. 

The high avidity of IgM antibodies makes them particularly efficient at binding antigens that are present in low amounts. and non-protein antigens such as carbohydrates or lipids present on microbial surfaces. Pentameric IgM antibodies also contain a connecting (J) chain that stabilizes the pentameric structure and allows binding to multiple receptors. One of these receptors, the polymeric immunoglobulin receptor (pIgR), is responsible for transcytosis from the vasculature to the mucosal surfaces of the lungs and gastrointestinal tract. Several naturally occurring IgM antibodies have been investigated in clinical trials as therapeutics, and a new class of molecules, engineered IgM antibodies with enhanced binding and/or additional functional properties, are being evaluated in humans. Here we review the significant advances that have been made in understanding biology, structure, function, manufacture,

Introduction to Immunoglobulin M (IgM)

During humoral immune responses, immunoglobulins of the IgM, IgD, IgG, IgA, and IgE isotypes can be produced, each expressing a unique profile of effector functions capable of mediating host defenses against invading pathogens. Macro-immunoglobulin, IgM, is initially produced as a surface-bound molecule and expressed in early B-cell differentiation. Later in the immune response, IgM is produced by plasma cells and excreted as soluble pentamers with 10 antigen binding sites and the connecting chain (J) or as hexamers with 12 antigen binding sites and no connecting chain (J chain). IgM has a molecular weight of approximately 900 or 1050 kDa for the pentamer or hexamer, respectively

Because of the polyvalent nature of IgMs, they may have higher avidity for antigen than the bivalent IgG. In addition to neutralizing pathogens, IgM antibodies are highly effective at activating complement to selectively lyse cells and pathogens.

Our understanding of the biology, structure, and functional relationships of IgM antibodies has progressed so far that this antibody isotype can be used therapeutically; however, challenges associated with their manufacture remain. Here we review the progress and therapeutic potential for this class of antibodies, as well as the potential for new classes of genetically engineered IgM antibodies.

 History and Discovery of IgM

Humoral immunity has been studied since the late 19th century when George Nuttall  discovered that animal immune sera could kill bacteria. 

  • Subsequent analysis of the immune serum using technologies such as electrophoresis and ultracentrifugation enabled the biochemical characterization of the various proteins that might mediate immunity, leading to the discovery of immunoglobulins. 
  • Originally, these serum components were assigned as α-globulin, β-globulin, and γ-globulin fractions to name the proteins in order of their electrophoretic mobility
  • The first description of IgM antibodies was in 1939 by Kabat et al.  who evaluated the molecular weight of antibodies produced in horse, cow, pig, monkey and human serum after immunization with pneumococci. 
  • Because of the size (approximately 990 kDa), the new antibody was named γ-macroglobulin. In 1944, it was discovered by Waldenstrom and later independently by Kunkel that γ-macroglobulins are expressed at high levels in patients with multiple myeloma [.
  •  They identified that the γ-macroglobulin migrated close to β-globulin in patient sera using immunoelectrophoresis and ultracentrifugation techniques. In the 1960s, methods were developed to induce plasmacytomas in mice that produced uniform immunoglobulins, including γ-macroglobulin-producing plasmacytomas, recapitulating the data observed in patients with multiple myeloma
  • Because the immunoglobulins discovered during this period were given arbitrary names, the World Health Organization defined a nomenclature system for antibody isotypes in 1964. As a consequence, γ-macroglobulin was renamed IgM and the M referred to “macroglobulin”.

Evolution of IgM antibodies

Immunoglobulins, including IgM antibodies, are found in all jawed vertebrates (gnathosomas), which evolved from jawless fish (agnathans) about 550 million years ago . Similar to mammals, IgM expression precedes the expression of other antibody isotypes, although in teleosts IgD and IgT are the only other isotypes present. The phylogeny of the immunoglobulin heavy and light chain isotypes is shown in figure 2 . However, within certain species there are marked differences in the structure of the IgM antibodies produced. For example, the predominant form of IgM antibodies in mice and humans is pentameric in structure and contains a J chain that stabilizes the pentamer, but hexamers and monomers can also be detected. However , IgM antibodies in frogs, for example  Xenopus,  have a hexameric structure, although  Xenopus  IgM has been reported to contain a J chain [  14  ]. In contrast, IgM from bony fish predominantly forms a tetrameric structure, while the IgM produced by cartilaginous fish such as sharks has a pentameric structure.

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Mouse IgG1 Standards Set, A-F

F077 Cygnus Technologies 1 ml 493.2 EUR

Mouse IgG2a Standards Set, A-F

F078 Cygnus Technologies 1 ml 493.2 EUR

Mouse IgG2b Standards Set, A-F

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Human IgM Standards A-F

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HSA Standards Set, A-E

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A549 Standards Set, A-F

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Insulin Standards Set, A-G

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NS/0 Standards Set, A-G

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CHO HCP Standards Set, A-F

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SF9 HCP Standards Set, A-H

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E.coli HCP Standards Set, A-F

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Goat IgG Standards Set, A-F

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Protein A Standards Set, A-E

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Protein A Standards Set, A-G

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Insulin-US Standards Set, A-F

F282 Cygnus Technologies 1 ml 585.6 EUR

P.fluorescens HCP Standards Set, A-F

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SP2/0 HCP Standards Set, A-F

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Vero Cell HCP Standards Set, A-F

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Hellma Cuvette Calibration Standards Set - EACH

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Pichia pastoris HCP Standards Set, A-F

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Bovine Plasma Proteins Standards Set, A-F

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Mettler-Toledo density standards set of - PK10

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Pichia pastoris HCP 2G Standards Set, A-F

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Protein A Mix-N-Go Standards Set, A-H

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Tosoh R40/R28 Protein A Standards Set, A-H

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Set of 2 ICP Drinking Water Standards - EACH

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EXOCET Standards

EXOCET-SD-1 SBI each 149 EUR

Protein standards

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Mouse Chemokine Panel, Antigen Standards: Pre-Mixed

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Mouse Th1/Th2 Panel, Antigen Standards: Pre-Mixed

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Mouse Inflammation Panel, Antigen Standards: Pre-Mixed

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Mouse Inflammation Panel, Antigen Standards: Pre-Mixed

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Mouse Inflammation Panel, Antigen Standards: Pre-Mixed

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Mouse Inflammation Panel, Antigen Standards: Pre-Mixed

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Mouse TH1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Mouse TH1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Mouse TH1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Mouse TH1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Mouse TH1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Size Calibration Standards

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Size Calibration Standards

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MRC5 Standards A-F

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AccuBlue® High Sensitivity dsDNA Standards, set of eight, 0.5 mL each

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Lyophilized Exosome Standards

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Lyophilized Exosome Standards

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Lyophilized Exosome Standards

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Lyophilized Exosome Standards

ExoUrine ImmunoStep 100 μg 364.2 EUR

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Density Standards 6ml - PK10

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Set of 8 Sulfur Oil Standards in Base Oil 20 100 grams - EACH

DSS8-SET Scientific Laboratory Supplies EACH 851.85 EUR

CHO DNA Standards (A-H)

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E.coli DNA Standards A-H

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Chlorine meter standards - EACH

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AccuBlue® Broad Range dsDNA Quantitation Standards, set of nine, 0.5 mL each

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AccuBlue® Broad Range dsDNA Quantitation Standards, set of nine, 0.5 mL each

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Protein A Standards A-H

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NS/0 DNA Standards (A-H)

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Set of Bromide Standards for Calibration of Ion Selective Electrodes - EACH

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Set of Cyanide Standards for Calibration of Ion Selective Electrodes - EACH

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Set of 8 Sulfur in Oil Standards in no.2 DieselFuel 100 grams - EACH

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BHK Cell HCP Standards A-F

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Set of Fluoride Standards for Calibration of Ion Selective Electrodes - EACH

AS-F9-SET Scientific Laboratory Supplies EACH 367.2 EUR

Bovine Serum Albumin Standards Kit

20830003-1 Glycomatrix 1 Kit 75.66 EUR

Pre-diluted BSA (bovine serum albumin) standards (1 set) for BCA Optima Protein Assay kit

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Per. C6Ð’ Cell HCP Standards A-F

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boite de 100 filtres plissés standards, 600 mm

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boite de 100 filtres plissés standards, 650 mm

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EZ‐PRRSV MPX 4.0 Quantification Standards

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Human Chemokine Panel, Antigen Standards: Pre-Mixed

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Pre-diluted BGG (bovine gamma globulin) standards (1 set: 2000, 1000, 500, 125, 62.5, 31.25, and 0 ng/ml BGG) for BCA Optima Protein Assay kit

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Purified Human C1q protein for ELISA or Standards

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Purified Human C1q protein for ELISA or Standards

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Purified, Human S100 protein for ELISA or Standards

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Human Inflammation Panel, Antigen Standards: Pre-Mixed

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Human Inflammation Panel, Antigen Standards: Pre-Mixed

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Human Inflammation Panel, Antigen Standards: Pre-Mixed

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Human Inflammation Panel, Antigen Standards: Pre-Mixed

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Human Inflammation Panel, Antigen Standards: Pre-Mixed

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Purified, Bovine S100 protein for ELISA or Standards

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Lyophilized Exosome fromm A-549 cell line Standards

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LIMITED QTY-Nucleoside Standards 8-Oxo-dG (1 mg)

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LIMITED QTY-Nucleoside Standards 8-Oxo-dA (1 mg)

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Human Adipsin/Factor D protein for ELISA or Standards

ADN15-N Alpha Diagnostics 5 ug 315.6 EUR

Human Th1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Human Th1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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Human Th1/Th2/Th17 Panel, Antigen Standards: Pre-Mixed

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paquet de 1000 filtres à plat standards 64g/m², 100 mm

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Nitrated MW Standards, Western Blot (SOD, SOD Dimer, BSA, Myosin)

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Nitrated MW Standards, Western Blot (SOD, SOD Dimer, BSA, Myosin)

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Human CD8 + T-Cell Cytokines Panel, Antigen Standards: Pre-Mixed

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Purified Human beta-2 microglobulin (B2M) protein for ELISA/Standards

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AccuClear® Ultra High Sensitivity dsDNA Quantitation Kit with 8 DNA Standards (1000 assays)

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Enhanced Green Fluorescent Proteins (EGFP) protein for ELISA or Standards

EGFP16-R Alpha Diagnostics 10 ug 300 EUR

Enhanced Green Fluorescent Proteins (EGFP) protein for ELISA or Standards

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Human Paraoxonase 1 (PON) Standards QQ and RR Phenotypes (2 x 0.3 mL)

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Human Paraoxonase 1 (PON) Standards QQ and RR Phenotypes (2 x 0.3 mL)

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Mouse Immunoglobulin M (IgM) Antibody Pair Kit (with Standard)

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Mouse Immunoglobulin M (IgM) Antibody Pair Kit (with Standard)

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Mouse Immunoglobulin M (IgM) Antibody Pair Kit (with Standard)

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Mouse Immunoglobulin M (IgM) Antibody Pair Kit (with Standard)

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LIMITED QTY-Dinucleotide Standards Formamido Guanine Dimer d (PfpG) (50 µg)

0801210 Zeptometrix 50 µg 142.37 EUR

LIMITED QTY-Dinucleotide Standards Formamido Cytidine Dimer d (PfpC) (50 µg)

0801211 Zeptometrix 50 µg 142.37 EUR

LIMITED QTY-Dinucleotide Standards Formamido Thymine Dimer d (PfpT) (50 µg)

0801212 Zeptometrix 50 µg 142.37 EUR

LIMITED QTY-Dinucleotide Standards Formamido Adenine Dimer d (PfpA) (50 µg)

0801213 Zeptometrix 50 µg 142.37 EUR

Pre-diluted BSA (bovine serum albumin) standards (750 ug/ml) for BCA Optima Protein Assay kit

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LIMITED QTY-Dinucleotide Standards Thymine Glycol Cytidine Dimer d (TgpC) (50µg)

0801216 Zeptometrix 50µg 142.37 EUR

LIMITED QTY-Dinucleotide Standards Thymine Glycol Guanine Dimer d (TgpG) (50 µg)

0801214 Zeptometrix 50 µg 142.37 EUR

LIMITED QTY-Dinucleotide Standards Thymine Glycol Thymine Dimer d (TgpT) (50 µg)

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LIMITED QTY-Dinucleotide Standards Thymine Glycol Adenine Dimer d (TgpA) (50 µg)

0801217 Zeptometrix 50 µg 142.37 EUR

Pre-diluted BSA (bovine serum albumin) standards (250, 125, 62.5, 31.25, and 0.0 ug/ml) for BCA Optima Protein Assay kit

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Lyophilized Exosome Standards - BFP (blue) Fluorescent EVs from HEK293 cells (CD81-BFP)

HBM-HEK-BFP-81 HansaBioMed 1x10^9 vial 226.04 EUR

Lyophilized Exosome Standards - EGFP (green) Fluorescent EVs from HEK293 cells (CD63-EGFP)

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Lyophilized Exosome Standards - EGFP (green) Fluorescent EVs from HEK293 cells (CD81-EGFP)

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Lyophilized Exosome Standards - EGFP (green) Fluorescent EVs from HEK293 cells (CD9-EGFP)

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BSA and BGG Stock standards (1 vial each @ 2 mg/ml) for BCA Optima Protein Assay kit

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Recombinant (E. coli) Yellow Fluorescent Proteins (YFP) protein for ELISA or Standards (>98%)

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Lyophilized Exosome Standards - mCherry (red) Fluorescent EVs from HEK293 cells (CD63-mCherry)

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Lyophilized Exosome Standards - mCherry (red) Fluorescent EVs from HEK293 cells (CD81-mCherry)

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Recombinant (E. coli) Red Fluorescent Proteins (RFP/dsRed) protein for ELISA or Standards (>98%)

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Lyophilized Exosome Standards - double labeled Fluorescent EVs from HEK293 cells (EGFP and mCherry)

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BCA Optima Protein Assay kit with Pre-diluted BSA standards and BCA Strip plates (5), 500 tests

BCAO-521 Alpha Diagnostics 1 kit 270 EUR

 It is unclear why hexamer IgM was produced, but it was possible that J chain synthesis might be limiting [  17  ]. In addition, there are examples of IgM where the expressed µ chain lacks the cysteine ​​in the tail that is required for proper insertion into the IgM structure [  18  ,  19  ]. Interestingly, pentameric IgM that does not contain J chain can also be present in humans and mice . Indeed, we have observed hexamers, ie pentamer mixtures, produced from recombinant IgM derived from CHO cells in the absence of transfected J chain ( and unpublished observations).

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