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dC
                                                       –1
       Passive Transport by Means of    J diff ! A ! D ! ! " [mol ! s ]  [1.2]
                                                dx
       Diffusion
                                       where C is the molar concentration and x is the
       Diffusion is movement of a substance owing to  distance traveled during diffusion. Since the
       the random thermal motion (brownian move-  driving “force”—i.e., the concentration gradient
       ment) of its molecules or ions (! A1) in all  (dC/dx)—decreases with distance, as was ex-
       directions throughout a solvent. Net diffusion  plained above, the time required for diffusion
       or selective transport can occur only when the  increases exponentially with the distance
    Fundamentals and Cell Physiology  concentration gradient—i.e., at equilibrium—  if the above-water partial pressure of free O 2
                                       traveled (t # x ). If, for example, a molecule
       solute concentration at the starting point is
                                                2
                                       travels the first µm in 0.5 ms, it will require 5 s
       higher than at the target site. (Note: uni-
       directional fluxes also occur in absence of a
                                       to travel 100 µm and a whopping 14 h for 1 cm.
                                        Returning to the previous example (! A2),
       but net diffusion is zero because there is equal
                                       diffusion (! A2) is kept constant, the Po 2 in the
       flux in both directions.) The driving force of
                                       water and overlying gas layer will eventually
       diffusion is, therefore, a concentration gra-
       dient. Hence, diffusion equalizes concentra-
                                       equalize and net diffusion will cease (diffusion
       tion differences and requires a driving force:
                                       equilibrium). This process takes place within
                                       the body, for example, when O 2 diffuses from
       passive transport (= downhill transport).
         Example: When a layer of O 2 gas is placed
                                       and when CO 2 diffuses in the opposite direc-
       on water, the O 2 quickly diffuses into the water
       along the initially high gas pressure gradient
                                       tion (! p. 120).
                                        Let us imagine two spaces, a and b (! B1)
       (! A2). As a result, the partial pressure of O 2  the alveoli of the lungs into the bloodstream
    1
       (Po 2) rises, and O 2 can diffuse further  containing different concentrations (C a " C b)
       downward into the next O 2-poor layer of water  of an uncharged solute. The membrane sepa-
       (! A1). (Note: with gases, partial pressure is  rating the solutions has pores ∆x in length and
       used in lieu of concentration.) However, the  with total cross-sectional area of A. Since the
       steepness of the Po 2 profile or gradient (dPo 2/  pores are permeable to the molecules of the
       dx) decreases (exponentially) in each sub-  dissolved substance, the molecules will diffuse
       sequent layer situated at distance x from the  from a to b, with C –C = ∆C representing the
                                                  a
                                                     b
       O 2 source (! A3). Therefore, diffusion is only  concentration gradient. If we consider only the
       feasible for transport across short distances  spaces a and b (while ignoring the gradients
       within the body. Diffusion in liquids is slower  dC/dx in the pore, as shown in B2, for the sake
       than in gases.                  of simplicity), Fick’s first law of diffusion
                             –1
         The diffusion rate, J diff (mol·s ), is the  (Eq. 1.2) can be modified as follows:
       amount of substance that diffuses per unit of  ∆C
                                                       –1
       time. It is proportional to the area available for  J diff ! A ! D !  ∆x  [mol ! s ].  [1.3]
       diffusion (A) and the absolute temperature (T)
       and is inversely proportional to the viscosity  In other words, the rate of diffusion increases
       (η) of the solvent and the radius (r) of the dif-  as A, D, and ∆C increase, and decreases as the
       fused particles.                thickness of the membrane (∆x) decreases.
         According to the Stokes–Einstein equation,  When diffusion occurs through the lipid
       the coefficient of diffusion (D) is derived from T,  membrane of a cell, one must consider that hy-
       η, and r as                     drophilic substances in the membrane are
                                       sparingly soluble (compare intramembrane
              R ! T
                     2
                       –1
         D !       [m ! s ],     [1.1]  gradient in C1 to C2) and, accordingly, have a
           N A · 6π ! r ! η
                                       hard time penetrating the membrane by
       where  R  is  the  general  gas  constant  means of “simple” diffusion. The oil-and-water
                   –1
       (8.3144 J·K –1  · mol ) and N A Avogadro’s con-  partition coefficient (k) is a measure of the lipid
                 23
                     –1
       stant (6.022 · 10 mol ). In Fick’s first law of  solubility of a substance (! C).
       diffusion (Adolf Fick, 1855), the diffusion rate
   20
       is expressed as
                                                                   !
       Despopoulos, Color Atlas of Physiology © 2003 Thieme
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