Salt Tolerance of Archaeal Extremely Halophilic Lipid Membranes

The membranes of extremely halophilic Archaea are characterized by the abundance of a diacidic phospholipid, archaetidylglycerol methylphosphate (PGP-Me), which accounts for 50–80 mol% of the polar lipids, and by the absence of phospholipids with choline, ethanolamine, inositol, and serine head groups. These membranes are stable in concentrated 3–5 m NaCl solutions, whereas membranes of non-halophilic Archaea, which do not contain PGP-Me, are unstable and leaky under such conditions. By x-ray diffraction and vesicle permeability measurements, we demonstrate that PGP-Me contributes in an essential way to membrane in the polar of and and in the m to By of polar halophilic and non-halophilic Archaea are leaky and PGP-Me to in of with the PGP-Me The are by the x-ray which that and and NaCl an with of which stable to in m NaCl and to in m by the and the the membrane in halophilic Archaea to of the head of PGP-Me, to the PGP-Me head The membranes of extremely halophilic Archaea are characterized by the abundance of a diacidic phospholipid, archaetidylglycerol methylphosphate (PGP-Me), which accounts for 50–80 mol% of the polar lipids, and by the absence of phospholipids with choline, ethanolamine, inositol, and serine head groups. These membranes are stable in concentrated 3–5 m NaCl solutions, whereas membranes of non-halophilic Archaea, which do not contain PGP-Me, are unstable and leaky under such conditions. By x-ray diffraction and vesicle permeability measurements, we demonstrate that PGP-Me contributes in an essential way to membrane in the polar of and and in the m to By of polar halophilic and non-halophilic Archaea are leaky and PGP-Me to in of with the PGP-Me The are by the x-ray which that and and NaCl an with of which stable to in m NaCl and to in m by the and the the membrane in halophilic Archaea to of the head of PGP-Me, to the PGP-Me head a of the to NaCl of 3–5 m under of halophilic and and membrane and the to the halophilic Archaea by and membranes to and permeability The membrane by NaCl for by a in for in by a in and a in and in of Archaea, and the membrane of the in the in and are characterized by and and in to the and of and to an to the of the the to the membranes of extremely halophilic Archaea that with ethanolamine, inositol, and serine head are and contain a phospholipid, archaetidylglycerol with with methylphosphate (PGP-Me), PGP-Me, archaetidylglycerol archaetidylglycerol PGP-Me, archaetidylglycerol archaetidylglycerol which an of methylphosphate and PGP-Me accounts for of the polar membrane of and by in and in the which PGP-Me, the phospholipids are The contain archaetidylglycerol and of of the phospholipids PGP-Me and in Archaea in of polar and with of the and with of the and with of the and with of the of polar PGP-Me and with of the in a of abundance in the diacidic PGP-Me to a in the of that contributes to membrane in the of PGP-Me, we a the of of polar and which by PGP-Me of and that the with the PGP-Me of the These by x-ray diffraction measurements, which that the polar and and lipids, by a PGP-Me of an stable to m NaCl in the whereas lipids, which PGP-Me, by the and the NaCl the and the x-ray diffraction demonstrate a of PGP-Me for the in and Archaea in are in The of and by the and and polar by and The polar by and the of with that The to to that of in The of PGP-Me and in the are in PGP-Me and of in of polar PGP-Me the of of and The the of of of PGP-Me and a in and of PGP-Me and polar of by PGP-Me and and a of for PGP-Me and of by in under a of and under for of in and for and to the for in and a an by a by the of a of the in a an a of of of m with m m and m The the a of by the the the by of by of the of of NaCl for the and for the m The with of of the to in m of with NaCl by to with a and with of the NaCl m m m m of in m not of in m not in a and and in with a of and the under for with by and by and by the The m The for and for to the x-ray The and x-ray NaCl and m NaCl with and m NaCl to and the in the of a for of the of we to of the that not and of such a of the to in the the which not in way the the x-ray x-ray and a of the and and in the a of The a and by with to the to that the of the NaCl a to the of the membrane and to the of PGP-Me the membrane we the of of polar extremely and non-halophilic Archaea, which by PGP-Me an we the of for in of stable which not of of of with concentrated NaCl a a of the of concentrated NaCl and the of by the with These the polar of the polar of the stable and the m NaCl the polar of the are stable NaCl and of with in the NaCl of in m NaCl for are in by the and which the PGP-Me halophilic and non-halophilic which PGP-Me, The and are by with leaky PGP-Me that to the membrane to a of PGP-Me with in to the to to the of to in of Archaea in the polar of Archaea of in m NaCl The in the PGP-Me in for for the of with with the the to in the NaCl to m an in in by and the in with of not to of By halophilic and non-halophilic to NaCl of the of the with PGP-Me of to the of These that PGP-Me do not in concentrated NaCl and a of and to that of of of to in m NaCl that PGP-Me and with to to the membrane and in by of PGP-Me and polar of in m NaCl The PGP-Me in the are for in and of in m of and polar PGP-Me and polar for a of the of PGP-Me to to membranes in of polar are unstable in with that the of PGP-Me in to in to for the stable and These demonstrate the of PGP-Me to the membranes in with NaCl the NaCl with that of of in m of and These that NaCl and the PGP-Me whereas and not of the the with in to m NaCl of m in the and m NaCl in the we to with NaCl to stable to vesicle and m NaCl not and in of the of m NaCl These the that the of PGP-Me of for the membrane and in the of PGP-Me, we the of polar the and the a of NaCl and of to and to in the for the in of to to of the x-ray with the the of the the absence of by x-ray in the The of that with These of which are not and not such The and of the in absence of NaCl an for the of of of a the of with head in the and with of the NaCl the of not by the NaCl to with the of to the of NaCl and the of the x-ray of by and polar for in the in absence of m m in m NaCl a and in m of and in m NaCl a m The of with of characterized by the stable in the to to the of m NaCl and by with of not stable and a of an and a with The not and the of not by a with of the and with of the and The by a in the of that are are with a of the of are m m of the of are m m the of m the of with to the m and a of to and by the of for the These are and a a of The of with that extremely to which are under the and the not by the By with the of m NaCl not by a of a with a of with the of the a by a of and in and that the to in that PGP-Me, a phospholipid, which accounts for a of the polar in an essential in the membrane NaCl of The of of the with the membrane PGP-Me The for the polar of the and which PGP-Me for the for the polar of halophilic and non-halophilic such which contain the polar of in in to the of PGP-Me the of the and with PGP-Me and PGP-Me stable and to m the polar of halophilic and non-halophilic are unstable and NaCl polar to stable NaCl to m in with that stable in NaCl the polar of the Archaea and By to vesicle polar NaCl such and m NaCl not and in of the of m NaCl to the x-ray and polar and NaCl an with of which stable to in m NaCl and to in m NaCl with the of to under the conditions. by the and the NaCl that are to under in with the the that the membrane of extremely in by are that to the membrane for the and to that of of PGP-Me by the of lipids, which to leaky the of PGP-Me under the membrane in the to the of PGP-Me with x-ray that the of stable in to by a of m NaCl with to in of with in the for the a that PGP-Me a to an in of m NaCl with PGP-Me stable in m NaCl not in m and that and a the polar head to the head of with the a of to the and to of and the of the not that and the to PGP-Me membranes the PGP-Me in that the of halophilic Archaea by the of of the are of Archaea and are not to that the of such with that PGP-Me and which and do not to m and which in m NaCl and of and m the of that the of head to the that of the of of PGP-Me in to of that the of PGP-Me of the methylphosphate in the polar head of PGP-Me, the PGP-Me and The that the head of PGP-Me for the membrane by by x-ray which in the of and polar NaCl of m NaCl in of a a with of that the the in absence of NaCl and that the m the NaCl to m NaCl in of the in that a in that the and with a for of not by a of NaCl m to that of which by in under such in a of of the of by the of with head groups. in that PGP-Me, a with head of the polar and a for the of the to PGP-Me which to a with of m NaCl that the of with of and for which with of NaCl that polar to which stable and not in the m in with the of the of polar m NaCl by of the and and stable to NaCl in with the x-ray and the that the of by the of polar to in a of NaCl and The PGP-Me a in the to PGP-Me are in halophilic and non-halophilic Archaea are by a The of the PGP-Me NaCl that and of the the to the polar head of PGP-Me and to a of the by PGP-Me, of for x-ray that a to for of to to the of and a of the to NaCl of 3–5 m under of halophilic and and membrane and the to the halophilic Archaea by and membranes to and permeability The membrane by NaCl for by a in for in by a in and a in and in of Archaea, and the membrane of the in the in and are characterized by and and in to the and of and to an to the of the the to the membranes of extremely halophilic Archaea that with ethanolamine, inositol, and serine head are and contain a phospholipid, archaetidylglycerol with with methylphosphate (PGP-Me), PGP-Me, archaetidylglycerol archaetidylglycerol PGP-Me, archaetidylglycerol archaetidylglycerol which an of methylphosphate and PGP-Me accounts for of the polar membrane of and by in and in the which PGP-Me, the phospholipids are The contain archaetidylglycerol and of of abundance in the diacidic PGP-Me to a in the of that contributes to membrane in the of PGP-Me, we a the of of polar and which by PGP-Me of and that the with the PGP-Me of the These by x-ray diffraction measurements, which that the polar and and lipids, by a PGP-Me of an stable to m NaCl in the whereas lipids, which PGP-Me, by the and the NaCl the and the x-ray diffraction demonstrate a of PGP-Me for the in and Archaea in are in The of and by the and and polar by and The polar by and the of with that The to to that of in The of PGP-Me and in the are in PGP-Me and of in of polar PGP-Me the of of and The the of of of PGP-Me and a in and of PGP-Me and polar of by PGP-Me and and a of for PGP-Me and of by in under a of and under for of in and for and to the for in and a an by a by the of a of the in a an a of of of m with m m and m The the a of by the the the by of by of the in m of with NaCl by to with a and with of the NaCl m m m m of in m not of in m not in a and and in with a of and the under for with by and by and by the The m The for and for to the x-ray The and x-ray NaCl and m NaCl with and m NaCl to and the in the of a for of the of we to of the that not and of such a of the to in the the which not in way the the x-ray x-ray and a of the and and in the a of The a and by with to the to that the of the and Archaea in are in The of and by the and and polar by and The polar by and the of with that The to to that of in The of PGP-Me and in the are in PGP-Me and of in of polar PGP-Me the of of and The the of of of PGP-Me and a in and of PGP-Me and polar of by PGP-Me and and a of for PGP-Me and of by in under a of and under for of in and for and to the for in and a an by a by the of a of the in a an a of of of m with m m and m The the a of by the the the by of by of the in m of with NaCl by to with a and and in with a of and the under for with by and by and by the The m The for and for to the x-ray The and x-ray NaCl and m NaCl with and m NaCl to and the in the of a for of the of we to of the that not and of such a of the to in the the which not in way the the x-ray x-ray and a of the and and in the a of The a and by with to the to that the of the NaCl a to the of the membrane and to the of PGP-Me the membrane we the of of polar extremely and non-halophilic Archaea, which by PGP-Me an we the of for in of stable which not of of of with concentrated NaCl a a of the of concentrated NaCl and the of by the with These the polar of the polar of the stable and the m NaCl the polar of the are stable NaCl and of with in the NaCl of in m NaCl for are in by the and which the PGP-Me halophilic and non-halophilic which PGP-Me, The and are by with leaky PGP-Me that to the membrane to a of PGP-Me with in to the to to the of to in for the of with with the the to in the NaCl to m an in in by and the in with of not to of By halophilic and non-halophilic to NaCl of the of the with PGP-Me of to the of These that PGP-Me do not in concentrated NaCl and a of and to that of of of to in m NaCl that PGP-Me and with to to the membrane and in by of PGP-Me and polar of in m NaCl The PGP-Me in the are for in and of in m of and polar PGP-Me and polar for a of the of PGP-Me to to membranes in of polar are unstable in with that the of PGP-Me in to in to for the stable and These demonstrate the of PGP-Me to the membranes in with NaCl the NaCl with that of of in m of and These that NaCl and the PGP-Me whereas and not of the the with in to m NaCl of m in the and m NaCl in the we to with NaCl to stable to vesicle and m NaCl not and in of the of m NaCl These the that the of PGP-Me of for the membrane and in the of PGP-Me, we the of polar the and the a of NaCl and of to and to in the for the in of to to of the x-ray with the the of the the absence of by x-ray in the The of that with These of which are not and not such The and of the in absence of NaCl an for the of of of a the of with head in the and with of the NaCl the of not by the NaCl to with the of to the of NaCl and the of the x-ray of by and polar for in the in absence of m m in m NaCl a and in m of and in m NaCl a m The of with of characterized by the stable in the to to the of m NaCl and by with of not stable and a of an and a with The not and the of not by a with of the and with of the and The by a in the of that are are with a of the of are m m of the of are m m the of m the of with to the m and a of to and by the of for the These are and a a of The of with that extremely to which are under the and the not by the By with the of m NaCl not by a of a with a of with the of the a by a of and in and that the to NaCl a to the of the membrane and to the of PGP-Me the membrane we the of of polar extremely and non-halophilic Archaea, which by PGP-Me an we the of The for in of stable which not of of of with concentrated NaCl a a of the of concentrated NaCl and the of by the with These the polar of the polar of the stable and the m NaCl the polar of the are stable NaCl and of with in the NaCl of in m NaCl for are in by the and which the PGP-Me halophilic and non-halophilic which PGP-Me, The and are by with leaky PGP-Me that to the membrane to a of PGP-Me with in to the to to the of to in for the of with with the the to in the NaCl to m an in in by and the in with of not to of By halophilic and non-halophilic to NaCl of the of the with PGP-Me of to the of These that PGP-Me do not in concentrated NaCl and a of and to that of of of to in m NaCl that PGP-Me and with to to the membrane a of the of PGP-Me to to membranes in of polar are unstable in with that the of PGP-Me in to in to for the stable and These demonstrate the of PGP-Me to the membranes in with NaCl the NaCl with that of of in m of and These that NaCl and the PGP-Me whereas and not of the the with in to m NaCl of m in the and m NaCl in the we to with NaCl to stable to vesicle and m NaCl not and in of the of m NaCl These the that the of PGP-Me of for the membrane and in the of PGP-Me, we the of polar the and the a of NaCl and of to and to in the for the in of to to of the x-ray with the the of the the absence of by x-ray in the The of that with These of which are not and not such The and of the in absence of NaCl an for the of of of a the of with head in the and with of the NaCl the of not by the NaCl to with the of to the of NaCl and the of the m The of with of characterized by the stable in the to to the of m NaCl and by with of not stable and a of an and a with The not and the of not by a with of the and with of the and The by a in the of that are are with a the of m the of with to the m and a of to and by the of for the These are and a a of The of with that extremely to which are under the and the not by the By with the of m NaCl not by a of a with a of with the of the a by a of and in and that the to in that PGP-Me, a phospholipid, which accounts for a of the polar in an essential in the membrane NaCl of The of of the with the membrane PGP-Me The for the polar of the and which PGP-Me for the for the polar of halophilic and non-halophilic such which contain the polar of in in to the of PGP-Me the of the and with PGP-Me and PGP-Me stable and to m the polar of halophilic and non-halophilic are unstable and NaCl polar to stable NaCl to m in with that stable in NaCl the polar of the Archaea and By to vesicle polar NaCl such and m NaCl not and in of the of m NaCl to the x-ray and polar and NaCl an with of which stable to in m NaCl and to in m NaCl with the of to under the conditions. by the and the NaCl that are to under in with the the that the membrane of extremely in by are that to the membrane for the and to that of of PGP-Me by the of lipids, which to leaky the of PGP-Me under the membrane in the to the of PGP-Me with x-ray that the of stable in to by a of m NaCl with to in of with in the for the a that PGP-Me a to an in of m NaCl with PGP-Me stable in m NaCl not in m and that and a the polar head to the head of with the a of to the and to of and the of the not that and the to PGP-Me membranes the PGP-Me in that the of halophilic Archaea by the of of the are of Archaea and are not to that the of such with that PGP-Me and which and do not to m and which in m NaCl and of and m the of that the of head to the that of the of of PGP-Me in to of that the of PGP-Me of the methylphosphate in the polar head of PGP-Me, the PGP-Me and The that the head of PGP-Me for the membrane by by x-ray which in the of and polar NaCl of m NaCl in of a a with of that the the in absence of NaCl and that the m the NaCl to m NaCl in of the in that a in that the and with a for of not by a of NaCl m to that of which by in under such in a of of the of by the of with head groups. in that PGP-Me, a with head of the polar and a for the of the to PGP-Me which to a with of m NaCl that the of with of and for which with of NaCl that polar to which stable and not in the m in with the of the of polar m NaCl by of the and and stable to NaCl in with the x-ray and the that the of by the of polar to in a of NaCl and The PGP-Me a in the to PGP-Me are in halophilic and non-halophilic Archaea are by a The of the PGP-Me NaCl that and of the the to the polar head of PGP-Me and to a of the by PGP-Me, of for x-ray that a to for of to to the of and The in that PGP-Me, a phospholipid, which accounts for a of the polar in an essential in the membrane NaCl of The of of the with the membrane PGP-Me The for the polar of the and which PGP-Me for the for the polar of halophilic and non-halophilic such which contain the polar of in in to the of PGP-Me the of the and with PGP-Me and PGP-Me stable and to m the polar of halophilic and non-halophilic are unstable and NaCl polar to stable NaCl to m in with that stable in NaCl the polar of the Archaea and By to vesicle polar NaCl such and m NaCl not and in of the of m NaCl to the x-ray and polar and NaCl an with of which stable to in m NaCl and to in m NaCl with the of to under the conditions. by the and the NaCl that are to under in with the the that the membrane of extremely in by are that to the membrane for the and to that of of PGP-Me by the of lipids, which to leaky the of PGP-Me under the membrane in the to the of PGP-Me with x-ray that the of stable in to by a of m NaCl with to in of with in the for the a that PGP-Me a to an in of m NaCl with PGP-Me stable in m NaCl not in m and that and a the polar head to the head of with the a of to the and to of and the of the not that and the to PGP-Me membranes the PGP-Me in that the of halophilic Archaea by the of of the are of Archaea and are not to that the of such with that PGP-Me and which and do not to m and which in m NaCl and of and m the of that the of head to the that of the The of of PGP-Me in to of that the of PGP-Me of the methylphosphate in the polar head of PGP-Me, the PGP-Me and The that the head of PGP-Me for the membrane by by x-ray which in the of and polar NaCl of m NaCl in of a a with of that the the in absence of NaCl and that the m the NaCl to m NaCl in of the in that a in that the and with a for of not by a of NaCl m to that of which by in under such in a of of the of by the of with head groups. in that PGP-Me, a with head of the polar and a for the of the to PGP-Me which to a with of m NaCl that the of with of and for which with of NaCl that polar to which stable and not in the m in with the of the of polar m NaCl by of the and and stable to NaCl in with the x-ray and the that the of by the of polar to in a of NaCl and The PGP-Me a in the to PGP-Me are in halophilic and non-halophilic Archaea are by a The of the PGP-Me NaCl that and of the the to the polar head of PGP-Me and to a of the by PGP-Me, of for x-ray that a to for of to to the of and x-ray and with by of by The and the of the of and the of of by the of of for and and for x-ray

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