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Transport of H+, K+, Na+ and Ca++ in Streptococcus

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Summary

The streptococci differ from other bacteria in that cation translocations (with the possible exception of one of the K+ uptake systems) occur by primary transport systems, i.e., by cation pumps which use directly the free energy released during hydrolysis of chemical bonds to power transport. Transport systems in other bacteria, especially for Na+ and Ca++, are often secondary, using the free energy of another ion gradient to drive cation transport. In streptococci H+ efflux occurs via the F1F0-ATPase. This enzyme is composed of eight distinct subunits. Three of the subunits are embedded in the membrane and form a H+ channel; this is called the F0 portion of the enzyme. The other five subunits form the catalytic part of the enzyme, called F1, which faces the cytoplasm and can easily be stripped from the membrane. Physiologically, this enzyme functions as a H+-ATPase, pumping protons out of the cell to form an electrochemical proton gradient, \(\Delta \tilde \mu _H + \). The F1F0-ATPase, however, is fully reversible and if supplied with Pi, ADP and a \(\Delta \tilde \mu _H + \) + of sufficient magnitude (ca −200 mv) catalyzes the synthesis of ATP.

Streptococcus faecalis can accumulate K+ and establish a gradient of 50 000:1 (in>out) under some conditions. Uptake occurs by two transport systems. The dominant, constitutive system requires both an electrochemical proton gradient and ATP to operate. The minor, inducible K+ transport system, which has many similarities to the K+-ATPase of the Kdp transport system found in Escherichia coli, requires only ATP to power K+ uptake.

Sodium extrusion occurs by a Na+/H+-ATPase. Exchange is electroneutral and there is no requirement for a \(\Delta \tilde \mu _H + \). The possibility that the Na+/H+-ATPase may consist of two parts, a catalytic subunit and a Na+/H+ antiport subunit, is suggested by the finding that damage to the Na+ transport system either through mutation or protease action leads to the appearance of \(\Delta \tilde \mu _H + \)-requiring Na+/H+ antiporter activity.

Ca++ like Na+ is extruded from metabolizing, intact cells. Transport requires no \(\Delta \tilde \mu _H + \) but does require ATP. Reconstitution of Ca++ transport activity with accompanying Ca++-stimulated ATPase activity into proteoliposomes suggests that Ca++ is transported by a Ca++-translocating ATPase.

Where respiring organelles and bacteria use secondary \(\Delta \tilde \mu _H + \) transport systems the streptococci have developed cation pumps. The streptococci, which are predominantly glycolyzing bacteria, generate a \(\Delta \tilde \mu _H + \) much inferior to that of respiring organisms and organelles. The cation pumps may have developed simply in response to an inadequate \(\Delta \tilde \mu _H + \).

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Abbreviations

\(\Delta \tilde \mu _H + \) :

electrochemical potential of protons

Δψ:

membrane potential

δpH:

pH gradient

Δp:

proton-motive force

DCCD:

N,Na1-dicyclohexlcarbodiimide

TCS:

tetrachlorosalicylanilide

FCCP:

carbonylcyanide-p-trifluoromethylphenylhydrazone

CCCP:

carbonylcyanie-m-chlorophenylhydrazone

TPMP+ :

triphenylmethyl phosphonium ion

DDA+ :

dibenzyldimethylammonium ion

Hepes:

4-(2-hydroxyethyl)-1-piperazineethanesulfonic acid

EGTA:

ethyleneglycol-bis (amino-ethyl-ether)-N,N′-tetraacetic acid

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Heefner, D.L. Transport of H+, K+, Na+ and Ca++ in Streptococcus . Mol Cell Biochem 44, 81–106 (1982). https://doi.org/10.1007/BF00226893

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