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It is assumed that the proton gradient imposes an active conformation on TonB. Upon interaction with FhuA, conformational energy is transmitted to FhuA, and TonB is converted to the inactive ground state, ready to be activated again. The functionally important histidine residue (His20) [7] in the cytoplasmic membrane part of TonB may be the site where protons bind and induce an allosteric transition. If there is a proton- conducting channel in TonB, the channel should be open for activation, and it should be closed after activation. ExbD could be involved in closing the proton channel and in deprotonation of TonB in that Asp25 of ExbD accepts the proton of His20. Asp25 has been localized within or at the inner border of the cytoplasmic membrane, and it is essential for ExbD and TonB activity since conversion to Asn abolishes TonB-relatedactivities [7].
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