scholarly journals THE KINETICS OF PENETRATION

1934 ◽  
Vol 18 (2) ◽  
pp. 229-234 ◽  
Author(s):  
S. E. Kamerling ◽  
W. J. V. Osterhout

To imitate cells which have ceased to grow we have made models in which artificial sap is separated from the external solution by a non-aqueous layer (representing the protoplasm). A stream of CO2 is bubbled through the artificial sap to imitate its production by the living cell. Potassium passes from the external solution through the non-aqueous layer into the artificial sap and there reacts with CO2 to form KHCO3: its rate of entrance depends on the supply of CO2. Hence the increase of volume depends on the supply of CO2 (as is probably true of the living cell). By regulating the supply of CO2 and the osmotic pressure we are able to keep the volume and composition of the artificial sap approximately constant while maintaining a higher concentration of potassium than in the external solution. In these respects the model resembles certain mature cells which have ceased to grow.

1932 ◽  
Vol 16 (1) ◽  
pp. 157-163 ◽  
Author(s):  
W. J. V. Osterhout

In a model consisting of a non-aqueous layer (representing the protoplasm) placed between an inner, more acid, aqueous layer (representing the sap) and an outer, more alkaline, aqueous solution (representing the external solution bathing a living cell) the penetration of potassium creates an outwardly directed potential against which potassium continues to diffuse inward, thereby increasing the outward potential. This continues until the steady state is reached. The potassium sets up less potential in entering than in escaping and the net result is an outwardly directed potential. A similar process appears to take place in certain living cells.


1932 ◽  
Vol 15 (6) ◽  
pp. 667-689 ◽  
Author(s):  
W. J. V. Osterhout ◽  
W. M. Stanley

Inasmuch as attempts to explain accumulation by the Donnan principle have failed in the case of Valonia, a hypothesis of the steady state has been formulated to explain what occurs. In order to see whether this hypothesis is in harmony with physico-chemical laws attempts have been made to imitate its chief features by means of a model. The model consists of a non-aqueous layer (representing the protoplasmic surface) placed between an alkaline aqueous phase (representing the external solution) and a more acid aqueous phase (representing the cell sap). The model reproduces most of the features of the hypothesis. Attention may be called to the following points. 1. The semipermeable surface is a continuous non-aqueous phase. 2. Potassium penetrates by combining with an acid HX in the non-aqueous layer to form KX which in turn reacts with an acid HA in the sap to form KA. Since KX is little dissociated in the non-aqueous layer potassium appears to pass through it chiefly in molecular form. 3. The internal composition depends on permeability, e.g., sodium penetrates less rapidly than potassium and in consequence potassium predominates over sodium in the "artificial sap." The order of penetration in the model is the same as in Valonia, i.e., K > Na > Ca > Mg, and Cl > SO4, but the quantitative resemblance is not close, e.g., the difference between potassium and sodium, and chloride and sulfate is much less in the model. 4. The formation of KA and NaA in the sap raises its osmotic pressure and water enters. 5. The concentration of potassium and sodium and the osmotic pressure become much greater inside than outside. For example, potassium may become 200 times as concentrated inside as outside. 6. No equilibrium occurs but a steady state is reached in which water and salt enter at the same rate so that the composition of the sap remains constant as its volume increases. 7. Since no equilibrium occurs there is a difference of thermodynamic potential between inside and outside. At the start the thermodynamic potential of KOH is much greater outside than inside. This difference gradually diminishes and in the steady state has about the same value as in Valonia. The difference in pH value between the internal and external solutions is also similar in both cases (about 2 pH units). 8. Accumulation does not depend on the presence of molecules or ions inside which are unable to pass out. One important feature of the hypothesis is not seen in the model: this is the exchange of HCO3 for Cl-. Experiments on this point are in progress.


1934 ◽  
Vol 17 (3) ◽  
pp. 469-480 ◽  
Author(s):  
W. J. V. Osterhout ◽  
S. E. Kamerling ◽  
W. M. Stanley

In some living cells the order of penetration of certain cations corresponds to that of their mobilities in water. This has led to the idea that electrolytes pass chiefly as ions through the protoplasmic surface in which the order of ionic mobilities is supposed to correspond to that found in water. If this correspondence could be demonstrated it would not prove that electrolytes pass chiefly as ions through the protoplasmic surface for such a correspondence could exist if the movement were mostly in molecular form. This is clearly shown in the models here described. In these the protoplasmic surface is represented by a non-aqueous layer interposed between two aqueous phases, one representing the external solution, the other the cell sap. The order of penetration through the non-aqueous layer is Cs > Rb > K > Na > Li. This will be recognized as the order of ionic mobilities in water. Nevertheless the movement is mostly in molecular form in the nonaqueous layer (which is used in the model to represent the protoplasmic surface) since the salts are very weak electrolytes in this layer. The chief reason for this order of penetration lies in the fact that the partition coefficients exhibit the same order, that of cesium being greatest and that of lithium smallest. The partition coefficients largely control the rate of entrance since they determine the concentration gradient in the non-aqueous layer which in turn controls the process of penetration. The relative molecular mobilities (diffusion constants) in the non-aqueous layer do not differ greatly. The ionic mobilities are not known (except for K+ and Na+) but they are of negligible importance, since the movement in the non-aqueous layer is largely in molecular form. They may follow the same order as in water, in accordance with Walden's rule. Ammonium appears to enter faster than its partition coefficient would lead us to expect, which may be due to rapid penetration of NH3. This recalls the apparent rapid penetration of ammonium in living cells which has also been explained as due to the rapid penetration of NH3. Both observation and calculation indicate that the rate of penetration is not directly proportional to the partition coefficient but increases somewhat less rapidly. Many of these considerations doubtless apply to living cells.


1934 ◽  
Vol 17 (4) ◽  
pp. 507-516 ◽  
Author(s):  
W. J. V. Osterhout ◽  
S. E. Kamerling

A model is described which throws light on the mechanism of accumulation. In the model used an external aqueous phase A is separated by a non-aqueous phase B (representing the protoplasm) from the artificial sap in C. A contains KOH and C contains HCl: they tend to mix by passing through the non-aqueous layer but much more KOH moves so that most of the KCl is formed in C, where the concentration of potassium becomes much greater than in A. This accumulation is only temporary for as the system approaches equilibrium the composition of A approaches identity with that of C, since all the substances present can pass through the non-aqueous layer. Such an approach to equilibrium may be compared to the death of the cell as the result of which accumulation disappears. During the earlier stages of the experiment potassium tends to go in as KOH and at the same time to go out as KCl. These opposing tendencies do not balance until the concentration of potassium inside becomes much greater than outside (hence potassium accumulates). The reason is that KCl, although its driving force be great, moves very slowly in B because its partition coefficient is low and in consequence its concentration gradient in B is small. This illustrates the importance of partition coefficients for penetration in models and in living cells. It also indicates that accumulation depends on the fact that permeability is greater for the ingoing compound of the accumulating substance than for the outgoing compound. Other things being equal, accumulation is increased by maintaining a low pH in C. Hence we may infer that anything which checks the production of acid in the living cell may be expected to check accumulation and growth. This model recalls the situation in Valonia and in most living cells where potassium accumulates as KCl, perhaps because it enters as KOH and forms KA in the sap (where A is an organic anion). In some plants potassium accumulates as KA but when HCl exists in the external solution it will tend to enter and displace the weaker acid HA (if this be carbonic it can readily escape): hence potassium may accumulate to a greater or less extent as KCl. Injury of the cell may produce a twofold effect, (1) increase of permeability, (2) lessened accumulation. The total amount of electrolyte taken up in a given time will be influenced by these factors and may be greater than normal in the injured cell or less, depending somewhat on the length of the interval of time chosen.


1927 ◽  
Vol 10 (5) ◽  
pp. 659-664 ◽  
Author(s):  
Morton McCutcheon ◽  
Baldwin Lucke

1. The rate of exosmosis of water was studied in unfertilized Arbacia eggs, in order to bring out possible differences between the kinetics of exosmosis and endosmosis. 2. Exosmosis, like endosmosis, is found to follow the equation See PDF for Equation, in which a is the total volume of water that will leave the cell before osmotic equilibrium is attained, x is the volume that has already left the cell at time t, and k is the velocity constant. 3. The velocity constants of the two processes are equal, provided the salt concentration of the medium is the same. 4. The temperature characteristic of exosmosis, as of endomosis, is high. 5. It is concluded that the kinetics of exosmosis and endosmosis of water in these cells are identical, the only difference in the processes being in the direction of the driving force of osmotic pressure.


1927 ◽  
Vol 10 (6) ◽  
pp. 883-892 ◽  
Author(s):  
John H. Northrop

It is shown that by combining the osmotic pressure and rate of diffusion laws an equation can be derived for the kinetics of osmosis. The equation has been found to agree with experiments on the rate of osmosis for egg albumin and gelatin solutions with collodion membranes.


Recent work on the osmotic pressure of the hen’s egg has introduced a sense of uncertainty as to the value of the many comparisons which have been made between osmotic pressures of the blood, body fluids, and surrounding media. The uncertainty pertains not to theory but to a simple matter of fact and, as this involves that most fundamental datum for biological theory—viz., the state of the water in the living cell—there is urgent need to have it cleared up. The fact in dispute is the freezing point of the yolk and white of the bird’s egg. Atkins in 1909 by measurements, obviously made with the greatest care, found “no difference between the freezing point of white and yolk of the same egg and a mixture of white and yolk gave the same depression.” Atkins (1909) used the ordinary Beckmann technique and so, too, did Straub (1929) twenty years later, but with a surprisingly different result for he found a constant difference between white and yolk of the hen’s egg amounting on the average to —0·15° C. A. V. Hill (1930) confirmed Straub’s (1929) finding by a different method. He compared the fall in temperature caused by evaporation with that of water and from the difference calculated the osmotic pressure. Howard (1932) using the Beckmann method again found no difference in the freezing point of white and yolk. In these measurements the yolk was puddled by stirring so that at sometime or another the structure was broken down. Yolk is not only a chemical complex but it is alive, gross mechanical disturbance might, therefore, have the effect it usually has on living cells and cause chemical breakdown with consequent fall of the freezing point. Hale’s experiments were designed to explore this possibility by observing directly the freezing point of intact yolk and white.


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