The CD20 Calcium Channel Is Localized to Microvilli and Constitutively Associated with Membrane Rafts

CD20 is a B cell-specific membrane protein that functions in store-operated calcium entry and serves as a useful target for antibody-mediated therapeutic depletion of B cells. Antibody binding to CD20 induces a diversity of biological effects, some of which are dependent on lipid rafts. Rafts are isolated as low density detergent-resistant membranes, initially characterized using Triton X-100. We have previously reported that CD20 is soluble in 1% Triton but that antibodies induce the association of CD20 with Triton-resistant rafts. However, by using several other detergents to isolate rafts and by microscopic co-localization with a glycosylphosphatidylinositol-linked protein, we show in this report that CD20 is constitutively raft-associated. CD20 was distributed in a punctate pattern on the cell surface as visualized by fluorescence imaging and was also localized to microvilli by electron microscopy. association of CD20 with Triton-resistant rafts was and to but was dependent on the in 1% Triton a to association that as a of a in the CD20 CD20 is a B cell-specific membrane protein that functions in store-operated calcium entry and serves as a useful target for antibody-mediated therapeutic depletion of B cells. Antibody binding to CD20 induces a diversity of biological effects, some of which are dependent on lipid rafts. Rafts are isolated as low density detergent-resistant membranes, initially characterized using Triton X-100. We have previously reported that CD20 is soluble in 1% Triton but that antibodies induce the association of CD20 with Triton-resistant rafts. However, by using several other detergents to isolate rafts and by microscopic co-localization with a glycosylphosphatidylinositol-linked protein, we show in this report that CD20 is constitutively raft-associated. CD20 was distributed in a punctate pattern on the cell surface as visualized by fluorescence imaging and was also localized to microvilli by electron microscopy. association of CD20 with Triton-resistant rafts was and to but was dependent on the in 1% Triton a to association that as a of a in the CD20 CD20 is a B cell-specific protein that and a store-operated calcium entry that is to B cell B cell electron fluorescence B cell electron fluorescence CD20 is also target for in depletion of B using antibodies which and and membrane as lipid rafts Rafts are to in as for with that to the lipid which membrane for association of a protein are in detergents and on density characterized for is Triton and we previously that antibodies induce of CD20 the soluble of Triton cell the with association with rafts However, association of some using low of Triton other detergents and we that soluble in 1% Triton was in 1% CD20 localized to on density of a previously to for Triton-resistant rafts also the association of CD20 with CD20 to of a protein that is is the of Triton-resistant we on the of association and the of on cell surface and of of show that binding induce a in cell surface of which in the of binding is distributed on the B cell electron that CD20 was localized to membrane in to induce Triton of and we show that to the to the of cell but are Triton-resistant rafts is to of the of and the of are with the that CD20 is constitutively with lipid rafts on microvilli and that the of the association as a in the CD20 and B in in isolated using of fluorescence protein by and with of was to the of CD20 by CD20 the of the using by and with and antibodies and by and the of by was was and to using the protein to was B and was and a CD20 was as of and and of using the was by of and by using with of of was was with of of with protein was in of and and and and and in detergents Triton which was also and density with of the previously and for in of in with of in with of and of in and for the of the on for CD20 was as with the in 1% Triton and soluble and by for with cell cell with to for by and to with was to as using and visualized using the imaging was to the with by using a and using the in 1% for and with as with to and with of in of for B with for with the antibodies for and in 1% with and with imaging was with a to a was using the some imaging of was with with in of in and in the in in with for with the the and in was and in 1% in for in in and in with a on and with a electron in 1% with for in and in cell was in of and to a for with and on the of a electron with the of detergents other Triton a of CD20 in rafts in the of B in the of detergents in density by for the of the also for and and CD20 was by Triton as However, in Triton and in other detergents of CD20 to the localized to the in protein was in the in detergents of was in protein, was in the in of the and the CD20 the of in B with and the by with are in with in to a and of as by and with of in and the of of CD20 and on was to with and was by with the of was dependent on that the of in rafts was by with and for CD20 and of with and in the of for in by density and by with using cells. the of in was using are the in the of by with for as in and on density for and as are of of CD20 and with association of CD20 was in the we cell surface and with which induces of CD20 protein to in 1% Triton we B B and and with a and and with B cell pattern of CD20 on that to was to that with the was in the of CD20 CD20 was distributed the membrane but was with of in the of and also CD20 was with of to surface of CD20 was CD20 was and with which induces Triton of CD20 with We a in in to the of CD20 in the of distributed to CD20 on density and the pattern of the cell surface was the with the of CD20 in lipid we fluorescence to the of the glycosylphosphatidylinositol-linked protein, and with and the was with the of with was surface of CD20 and with with 1% with with for and are the and of the and of the as and with of as with 1% with with the antibodies and with in 1% and by density by with as with and antibodies visualized with using and and of to of as in that CD20 was on membrane this B with CD20 and and for was of in and with membrane with the of the membrane a of with membrane is in low of and of B in the of the membrane of which is to that reported previously for B in is localized to with and and for with and in the low is in the and a is in the in 1% CD20 is soluble in 1% binding induce of CD20 in this CD20 was in the low density of and in the density of CD20 was to depletion and by with of the on is for of the association that Triton of the association with rafts. We the this using a and previously in which the of CD20 the soluble of Triton by binding is as as the of CD20 in the the of CD20 using this B with the of soluble and and for the of CD20 of CD20 the soluble was by of the with the CD20 was by of CD20 in 1% with in 1% Triton on density and for CD20 with for the and was for in 1% Triton X-100. soluble and and by for are of in and in and in was by but is for of association of CD20 with 1% Triton-resistant rafts by the by by on a that is the of by a as a in are the in membrane rafts membrane also to the of lipid rafts in with for to CD20 Triton by as with cells. with and with the as by of and which CD20 Triton is of and with for the of for and with antibodies for in 1% Triton X-100. and and for CD20 as in are of in and and in Triton of association is that we and to in CD20 Triton the as by the of the which the of CD20 and by of and CD20 is by and we of and have on Triton of in and the of CD20 Triton of the CD20 to that the was of the was by to calcium with the in that of CD20 in 1% Triton of Triton of and show in the of CD20 Triton induce of the in CD20 However, and of a of CD20 Triton as of the CD20 Triton is but and with of with that of with of for and in 1% Triton X-100. and by and for the of CD20 by are of and with of by was on a fluorescence is the of we that and on CD20 and that the the as and are of several CD20 with we show that CD20 Triton and have of the CD20 in was in and to B as by using and by was in this is a with and with the other that the of and to induce Triton of association is of and to induce Triton of association that was and the of was of to cells. We that CD20 to the to the as of CD20 Triton by is with binding to to as by is and are to induce Triton of that CD20 is constitutively localized to lipid rafts is by and CD20 localized to low density detergent-resistant isolated using a low of Triton X-100. rafts using detergents that 1% a is that membrane are We for this using which is on B as is on and is in lipid rafts. low density isolated in of CD20 to rafts was as as by of in a membrane of CD20 the cell CD20 with as by of with a glycosylphosphatidylinositol-linked protein, was that CD20 and also rafts. rafts in B previously co-localization CD20 and was the of the detergents in this with to and in low density the Triton a for the of of Triton to some to is a in in other of a to the of with other to Triton and a the of to Triton and to some also to that CD20 is to with rafts with are for and of CD20 association is a in of CD20 in rafts. is a to in by the of CD20 to that is by with with this the of CD20 was to to the of membrane a of detergents of detergents of CD20 on microvilli is with co-localization with the which is also on to with also and with rafts the protein is also with lipid rafts that are soluble in Triton but in other detergents the several and to rafts on microvilli rafts a with that that with on association are in the of the therapeutic of antibodies of antibodies to with to induce Triton-resistant association and the of by CD20 which to cell is also a Antibody of rafts. the of was to to induce a to a to Triton was and by with of to a to of CD20 the of that a of of the store-operated calcium that is by in is by depletion as as by of that the of CD20 for of the in CD20 that association also calcium entry of the association with by a by in calcium have to calcium with in this was in the but was previously to induce the association of CD20 with Triton-resistant rafts a calcium entry and CD20 is a B cell-specific protein that and a store-operated calcium entry that is to B cell B cell electron fluorescence B cell electron fluorescence CD20 is also target for in depletion of B using antibodies which and and membrane as lipid rafts Rafts are to in as for with that to the lipid which membrane for association of a protein are in detergents and on density characterized for is Triton and we previously that antibodies induce of CD20 the soluble of Triton cell the with association with rafts However, association of some using low of Triton other detergents and we that soluble in 1% Triton was in 1% CD20 localized to on density of a previously to for Triton-resistant rafts also the association of CD20 with CD20 to of a protein that is is the of Triton-resistant we on the of association and the of on cell surface and of of show that binding induce a in cell surface of which in the of binding is distributed on the B cell electron that CD20 was localized to membrane in to induce Triton of and we show that to the to the of cell but are Triton-resistant rafts is to of the of and the of are with the that CD20 is constitutively with lipid rafts on microvilli and that the of the association as a in the CD20 and B in in isolated using of fluorescence protein by and with of was to the of CD20 by CD20 the of the using by and with and antibodies and by and the of by was was and to using the protein to was B and was and a CD20 was as of and and of using the was by of and by using with of of was was with of of with protein was in of and and and and and in detergents Triton which was also and density with of the previously and for in of in with of in with of and of in and for the of the on for CD20 was as with the in 1% Triton and soluble and by for with cell cell with to for by and to with was to as using and visualized using the imaging was to the with by using a and using the in 1% for and with as with to and with of in of for B with for with the antibodies for and in 1% with and with imaging was with a to a was using the some imaging of was with with in of in and in the in in with for with the the and in was and in 1% in for in in and in with a on and with a electron in 1% with for in and in cell was in of and to a for with and on the of a electron and B in in isolated using of fluorescence protein by and with of was to the of CD20 by CD20 the of the using by and with and antibodies and by and the of by was was and to using the protein to was B and was and a CD20 was as of and and of using the was by of and by using with of of was was with of of with protein was in of Triton and and and and Rafts and in detergents Triton which was also and density with of the previously and for in of in with of in with of and of in and for the of the on for CD20 was as with the in 1% Triton and soluble and by for with cell cell with to for by and to with was to as using and visualized using the imaging was to the with by using a and using the in 1% for and with as with to and with of in of for B with for with the antibodies for and in 1% with and with imaging was with a to a was using the some imaging of was with with in of in and in the in in with for with the the and in was and in 1% in for in in and in with a on and with a electron in 1% with for in and in cell was in of and to a for with and on the of a electron with the of detergents other Triton a of CD20 in rafts in the of B in the of detergents in density by for the of the also for and and CD20 was by Triton as However, in Triton and in other detergents of CD20 to the localized to the in protein was in the in detergents of was in protein, was in the in of the and the CD20 the of in B with and the by with are in with in to a and of as by and with of in and the of of CD20 and on was to with and was by with the of was dependent on that the of in rafts was by with and for CD20 and of CD20 and with association of CD20 was in the we cell surface and with which induces of CD20 protein to in 1% Triton we B B and and with a and and with B cell pattern of CD20 on that to was to that with the was in the of CD20 CD20 was distributed the membrane but was with of in the of and also CD20 was with of to surface of CD20 was CD20 was and with which induces Triton of CD20 with We a in in to the of CD20 in the of distributed to CD20 on density and the pattern of the cell surface was the with the of CD20 in lipid we fluorescence to the of the glycosylphosphatidylinositol-linked protein, and with and the was with the of with was surface of CD20 and with with 1% with with for and are the and of the and of the as and with of as with 1% with with the antibodies and with in 1% and by density by with as with and antibodies visualized with using and and of to of as in that CD20 was on membrane this B with CD20 and and for was of in and with membrane with the of the membrane a of with membrane is in low of and of B in the of the membrane of which is to that reported previously for B in is localized to with and and for with and in the low is in the and a is in the in 1% CD20 is soluble in 1% binding induce of CD20 in this CD20 was in the low density of and in the density of CD20 was to depletion and by with of the on is for of the association that Triton of the association with rafts. We the this using a and previously in which the of CD20 the soluble of Triton by binding is as as the of CD20 in the the of CD20 using this B with the of soluble and and for the of CD20 of CD20 the soluble was by of the with the CD20 was by of CD20 in 1% with in 1% Triton on density and for CD20 with for the and was for in 1% Triton X-100. soluble and and by for are of in and in and in was by but is for of association of CD20 with 1% Triton-resistant rafts by the by by on a that is the of by a as a in are the in membrane rafts membrane also to the of lipid rafts in with for to CD20 Triton by as with cells. with and with the as by of and which CD20 Triton is of and with for the of for and with antibodies for in 1% Triton X-100. and and for CD20 as in are of in and and in Triton of association is that we and to in CD20 Triton the as by the of the which the of CD20 and by of and CD20 is by and we of and have on Triton of in and the of CD20 Triton of the CD20 to that the was of the was by to calcium with the in that of CD20 in 1% Triton of Triton of and show in the of CD20 Triton induce of the in CD20 However, and of a of CD20 Triton as of the CD20 Triton is but and with of with that of with of for and in 1% Triton X-100. and by and for the of CD20 by are of and with of by was on a fluorescence is the of we that and on CD20 and that the the as and are of several CD20 with we show that CD20 Triton and have of the CD20 in was in and to B as by using and by was in this is a with and with the other that the of and to induce Triton of association is of and to induce Triton of association that was and the of was of to cells. We that CD20 to the to the as of CD20 Triton by is with binding to to as by is and are to induce Triton of CD20 with the of detergents other Triton a of CD20 in rafts in the of B in the of detergents in density by for the of the also for and and CD20 was by Triton as However, in Triton and in other detergents of CD20 to the localized to the in protein was in the in detergents of was in protein, was in the in of the and the CD20 the of in B with and the by with are in with in to a and of as by and with of in and the of of CD20 and on was to with and was by with the of was dependent on that the of in rafts was by with and for CD20 and of CD20 and with association of CD20 was in the we cell surface and with which induces of CD20 protein to in 1% Triton we B B and and with a and and with B cell pattern of CD20 on that to was to that with the was in the of CD20 CD20 was distributed the membrane but was with of in the of and also CD20 was with of to surface of CD20 was CD20 was and with which induces Triton of CD20 with We a in in to the of CD20 in the of distributed to CD20 on density and the pattern of the cell surface was the with the of CD20 in lipid we fluorescence to the of the glycosylphosphatidylinositol-linked protein, and with and the was with the of with was CD20 to of as in that CD20 was on membrane this B with CD20 and and for was of in and with membrane with the of the membrane a of with membrane is in low of and of B in the of the membrane of which is to that reported previously for B in in 1% CD20 is soluble in 1% binding induce of CD20 in this CD20 was in the low density of and in the density of CD20 was to depletion and by with of the on is for of the association that Triton of the association with rafts. We the this using a and previously in which the of CD20 the soluble of Triton by binding is as as the of CD20 in the the of CD20 using this B with the of soluble and and for the of CD20 of CD20 the soluble was by of the with the CD20 was by association of CD20 with 1% Triton-resistant rafts by the by by on a that is the of by a as a in are the in membrane rafts membrane also to the of lipid rafts in with for to CD20 Triton by as with cells. with and with the as by of and which Triton of association is that we and to in CD20 Triton the as by the of the which the of CD20 and by of and CD20 is by and we of and have on Triton of in and the of CD20 Triton of the CD20 to that the was of the was by to calcium with the in that of CD20 in 1% Triton of Triton of and show in the of CD20 Triton induce of the in CD20 However, and of a of CD20 Triton as of the we that and on CD20 and that the the as and are of several CD20 with we show that CD20 Triton and have of the CD20 in was in and to B as by using and by was in this is a with and with the other that the of and to induce Triton of association is of and to induce Triton of association that was and the of was of to cells. We that CD20 to the to the as of CD20 Triton by is with binding to to as by is and are to induce Triton of that CD20 is constitutively localized to lipid rafts is by and CD20 localized to low density detergent-resistant isolated using a low of Triton X-100. rafts using detergents that 1% a is that membrane are We for this using which is on B as is on and is in lipid rafts. low density isolated in of CD20 to rafts was as as by of in a membrane of CD20 the cell CD20 with as by of with a glycosylphosphatidylinositol-linked protein, was that CD20 and also rafts. rafts in B previously co-localization CD20 and was the of the detergents in this with to and in low density the Triton a for the of of Triton to some to is a in in other of a to the of with other to Triton and a the of to Triton and to some also to that CD20 is to with rafts with are for and of CD20 association is a in of CD20 in rafts. is a to in by the of CD20 to that is by with with this the of CD20 was to to the of membrane a of detergents of CD20 on microvilli is with co-localization with the which is also on to with also and with rafts the protein is also with lipid rafts that are soluble in Triton but in other detergents the several and to rafts on microvilli rafts a with that that with on association are in the of the therapeutic of antibodies of antibodies to with to induce Triton-resistant association and the of by CD20 which to cell is also a Antibody of rafts. the of was to to induce a to a to Triton was and by with of to a to of CD20 the of that a of of the store-operated calcium that is by in is by depletion as as by of that the of CD20 for of the in CD20 that association also calcium entry of the association with by a by in calcium have to calcium with in this was in the but was previously to induce the association of CD20 with Triton-resistant rafts a calcium entry and that CD20 is constitutively localized to lipid rafts is by and CD20 localized to low density detergent-resistant isolated using a low of Triton X-100. rafts using detergents that 1% a is that membrane are We for this using which is on B as is on and is in lipid rafts. low density isolated in of CD20 to rafts was as as by of in a membrane of CD20 the cell CD20 with as by of with a glycosylphosphatidylinositol-linked protein, was that CD20 and also rafts. rafts in B previously co-localization CD20 and was We the of the detergents in this with to and in low density the Triton a for the of of Triton to some to is a in in other of a to the of with other to Triton and a the of to Triton and to some also to that CD20 is to with rafts with are for and of CD20 association is a in of CD20 in rafts. is a to in by the of CD20 to that is by with with this the of CD20 was to to the of membrane a of CD20 on microvilli is with co-localization with the which is also on to with also and with rafts the protein is also with lipid rafts that are soluble in Triton but in other detergents the several and to rafts on microvilli rafts a with that that with on association are in the of the therapeutic of antibodies of antibodies to with to induce Triton-resistant association and the of by CD20 which to cell is also a Antibody of rafts. the of was to to induce a to a to Triton was and by with of to a to of CD20 the of that a of of the store-operated calcium that is by in is by depletion as as by of that the of CD20 for of the in CD20 that association also calcium entry of the association with by a by in calcium have to calcium with in this was in the but was previously to induce the association of CD20 with Triton-resistant rafts a calcium entry and We and for and for with

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