Facilitative diffusion

Calcium is absorbed from the intestine by facilitated diffusion and active transport. In the former, Ca " moves from the mucosal to the serosal compartments along a concentration gradient. The active transport system requires a cation pump. In both processes, a calcium-binding protein (CaBP) is thought to be required for the transport. Synthesis of CaBP is activated by 1,25-DHCC. In the active transport, release of Ca " from the mucosal cell into... 

Table 1 fists many metal borides and their observed melting points. Most metals form mote than one boride phase and borides often form a continuous series of solid solutions with one another at elevated temperatures thus close composition control is necessary to achieve particular properties. The relatively small size of boron atoms facilitates diffusion. 

FIGURE 10.3 Passive diffusion and facilitated diffusion may be distinguished graphically. The plots for facilitated diffusion are similar to plots of enzyme-catalyzed processes (Chapter 14) and they display saturation behav-... 

Glucose Transport in Erythrocytes Occurs by Facilitated Diffusion... 

Does this transport operate by passive diffusion or by facilitated diffusion ... 

Phloretin is the aglycon of phlorizin and inhibits the facilitated diffusion of glucose catalyzed by GLUT1 or GLUT4. It has been used to terminate the uptake of glucose in timed assays with isolated membranes or reconstituted transporters. 

Physiologically muscle-derived NO regulates skeletal muscle contractility and exercise-induced glucose uptake. nNOS is located at the plasma membrane of skeletal muscle and facilitates diffusion of NO to the vasculature to regulate muscle perfusion. 

The passage of a small and/or highly lipophilic molecule through the membrane phospholipid bilayer according to the gradient of its concentrations across the plasma membrane. It is slower than facilitated diffusion, which, however, also follows the gradient of solute concentrations across the membrane. 

Molecules can passively traverse the bilayer down electrochemical gradients by simple diffusion ot by facilitated diffusion. This spontaneous movement toward equilibrium contrasts with active transport, which requires energy because it constitutes movement against an electrochemical gradient. Figure 41-8 provides a schematic representation of these mechanisms. 

PLASMA MEMBRANES ARE INVOLVED IN FACILITATED DIFFUSION, ACTIVE TRANSPORT, OTHER PROCESSES... 

Facilitated diffusion and active transport share many features. Both appear to involve carrier proteins, and both show specificity for ions, sugars, and amino acids. 

Mutations in bacteria and mammalian cells (including some that result in human disease) have supported these conclusions. Facilitated diffusion and active transport resemble a substrate-enzyme reaction except that no covalent interaction occurs. These points of resemblance are as follows (1) There is a specific binding site for the solute. (2) The carrier is saturable, so it has a maximum rate of transport (V Figure 41-11). (3) There is a binding constant (Al) ) for the solute, and... 

Figure 41-11. A comparison of the kinetics of carrier-mediated (facilitated) diffusion with passive diffusion. The rate of movement in the latter is directly proportionate to solute concentration, whereas the process is saturable when carriers are involved. The concentration at half-maximal velocity is equal to the binding constant (KJ of the carrier for the solute. maximal rate.)...

Some specific solutes diffuse down electrochemical gradients across membranes more rapidly than might be expected from their size, charge, or partition coefficients. This facilitated diffusion exhibits properties distinct from those of simple diffusion. The rate of facilitated diffusion, a uniport system, can be saturated ie, the number of sites involved in diffusion of the specific solutes appears finite. Many facihtated diffusion systems are stereospecific but, fike simple diffusion, require no metabolic energy. 

As described earlier, the inside-outside asymmetry of membrane proteins is stable, and mobifity of proteins across (rather than in) the membrane is rare therefore, transverse mobility of specific carrier proteins is not likely to account for facilitated diffusion processes except in a few unusual cases. 

Certain solutes, eg, glucose, enter cells by facilitated diffusion, along a downhill gradient from high to low concentration. Specific carrier molecules, or transporters, are involved in such processes. 

Unphosphorylated functioning according to Fig. 5 catalyzes facilitated diffusion of mannitol across the membrane. The same process has been reported for purified II reconstituted in proteoliposomes [70]. The relevance of this activity in terms of transport of mannitol into the bacterial cell is probably low, but it may have important implications for the mechanism by which E-IIs catalyze vectorial phosphorylation. It would indicate that the transmembrane C domain of Il is a mannitol translocating unit which is somehow coupled to the kinase activity of the cytoplasmic domains. We propose that the inwardly oriented binding site which is in contact with the internal water phase (Ecyt Mtl, see Fig. 5) is the site from where mannitol is phosphorylated when transport is coupled to phosphorylation. Meehan-... 

S. typhimurium normally transport trehalose via the galactose permease and are able to grow on this substrate in the complete absence of PTS phosphorylating activity. However, in S. typhimurium which lack a functional galactose permease, IfMan appears to be able to transport trehalose [78]. There is no evidence that trehalose is phosphorylated in this process, again pointing to Il "-dependent transport in the facilitated diffusion mode. 

Postma and Stock [81] showed that HPr or E-I mutants were unable to grow on PTS carbohydrates suggesting that transport without phosphorylation did not take place in apparent contradiction with the studies presented above. The explanation may be that facilitated diffusion via PTS carriers is observed only in abnormal situations, carbohydrate being transported by the incorrect PTS carrier (galactose via the mannose carrier) or transport via a mutated carrier. Efflux, which also reflects facilitated diffusion, is more common for PTS carriers. 

The history of observations of efflux associated with PTS carriers is nearly as old as PTS itself. Gachelin [82] reported that A -ethylmaleimide inactivation of a-methyl-glucoside transport and phosphorylation in E. coli was accompanied by the appearance of a facilitated diffusion movement of both a-methylglucoside and glucose in both directions, uptake and efflux. His results could not discriminate, however, between one carrier operating in two different modes, active transport for the native carrier and facilitated diffusion for the alkylated carrier, or two distinct carriers. Haguenauer and Kepes [83] went on to show that alkylation of the carrier was not even necessary to achieve efflux NaF treatment which inhibits P-enolpyruvate synthesis was sufficient but this study did not address the question of one carrier or two. 

The phosphorylation of cytoplasmic sugar and the facilitated diffusion from the cytoplasm to the periplasm are catalyzed by the E-IIs under conditions where they are also active in the vectorial phosphorylation reaction. Therefore, the former two activities should be integral parts of any kinetic scheme representing the mechanism of E-IIs. Such a scheme should explain how vectorial phosphorylation, transport coupled to phosphorylation, is still achieved while the uncoupled pathways are integral parts of the scheme. 

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