scholarly journals CHANGES OF APPARENT IONIC MOBILITIES IN PROTOPLASM

1939 ◽  
Vol 22 (3) ◽  
pp. 417-427 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Partition Coefficient ◽  
Action Current ◽  
Protoplasmic Surface ◽  
Outer Protoplasmic Surface ◽  
Ionic Mobilities

In Nitella, as in Halicystis, guaiacol increases the mobility of Na+ in the outer protoplasmic surface but leaves the mobility of K+ unaffected. This differs from the situation in Valonia where the mobility of Na+ is increased and that of K+ is decreased. The partition coefficient of Na+ in the outer protoplasmic surface is increased and that of K+ left unchanged. Recovery after the action current is delayed in the presence of guaiacol and the action curves are "square topped."

scholarly journals MECHANICAL RESTORATION OF IRRITABILITY AND OF THE POTASSIUM EFFECT

1935 ◽  
Vol 18 (5) ◽  
pp. 687-694 ◽  
Author(s):  
S. E. Hill ◽  
W. J. V. Osterhout
Keyword(s):  
Distilled Water ◽  
Action Current ◽  
Protoplasmic Surface ◽  
Normal Behavior ◽  
Inner Surface ◽  
Potassium Effect ◽  
Outer Protoplasmic Surface

Treatment of Nitella with distilled water apparently removes from the cell something which is responsible for the normal irritability and the potassium effect, (i.e. the large P.D. between a spot in contact with 0.01 M KCl and one in contact with 0.01 M NaCl). Presumably this substance (called R) is partially removed from the protoplasm by the distilled water. When this has happened a pinch which forces sap out into the protoplasm can restore its normal behavior. The treatment with distilled water which removes the potassium effect from the outer protoplasmic surface does not seem to affect the inner protoplasmic surface in the same way since the latter retains the outwardly directed potential which is apparently due to the potassium in the sap. But the inner surface appears to be affected in such fashion as to prevent the increase in its permeability which is necessary for the production of an action current. The pinch restores its normal behavior, presumably by forcing R from the sap into the protoplasm.

scholarly journals DIFFERING RATES OF DEATH AT INNER AND OUTER SURFACES OF THE PROTOPLASM

1944 ◽  
Vol 28 (1) ◽  
pp. 23-36 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Hydrostatic Pressure ◽  
Partition Coefficient ◽  
The Other ◽  
Protoplasmic Surface ◽  
Inner Surface ◽  
Potassium Effect ◽  
Outer Protoplasmic Surface ◽  
The Difference

When protoplasm dies it becomes completely and irreversibly permeable and this may be used as a criterion of death. On this basis we may say that when 0.2 M formaldehyde plus 0.001 M NaCl is applied to Nitella death arrives sooner at the inner protoplasmic surface than at the outer. If, however, we apply 0.17 M formaldehyde plus 0.01 M KCl death arrives sooner at the outer protoplasmic surface. The difference appears to be due largely to the conditions at the two surfaces. With 0.2 M formaldehyde plus 0.001 M NaCl the inner surface is subject to a greater electrical pressure than the outer and is in contact with a higher concentration of KCl. In the other case these conditions are more nearly equal so that the layer first reached by the reagent is the first to become permeable. The outer protoplasmic surface has the ability to distinguish electrically between K+ and Na+ (potassium effect). Under the influence of formaldehyde this ability is lost. This is chiefly due to a falling off in the partition coefficient of KCl in the outer protoplasmic surface. At about the same time the inner protoplasmic surface becomes completely permeable. But the outer protoplasmic surface retains its ability to distinguish electrically between different concentrations of the same salt, showing that it has not become completely permeable. After the potential has disappeared the turgidity (hydrostatic pressure inside the cell) persists for some time, probably because the outer protoplasmic surface has not become completely permeable.

scholarly journals NATURE OF THE ACTION CURRENT IN NITELLA

1943 ◽  
Vol 27 (1) ◽  
pp. 61-68 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Partial Response ◽  
Outer Surface ◽  
Base Line ◽  
Normal Amount ◽  
Action Current ◽  
Typical Response ◽  
Total Potential ◽  
Protoplasmic Surface ◽  
Outer Protoplasmic Surface

When a stimulus arrives before recovery is complete there may be no response or only a partial response. A typical response appears to involve an immediate loss of potential at the inner protoplasmic surface but not at the outer surface. As long as recovery is incomplete only a part of the total potential is located at the inner protoplasmic surface and the loss of this part of the total potential can cause only a partial response; i.e., one of smaller magnitude than the normal. Even after the action curve has returned to the base line recovery may be incomplete and the response only a partial one. The return of the action curve to the base line means a recovery of total potential but if part of this is located at the outer protoplasmic surface and if this part is not lost when stimulation occurs the response can be only a partial one. During recovery there is a shift of potential from the outer to the inner protoplasmic surface. Not until this shift is completed can recovery be called complete. The response to stimulation then becomes normal because the loss of potential reaches the normal amount. In many cases the partial responses appear to conform to the all-or-none law. In other cases this is doubtful.

scholarly journals NATURE OF THE ACTION CURRENT IN NITELLA

1935 ◽  
Vol 18 (4) ◽  
pp. 499-514 ◽  
Author(s):  
W. J. V. Osterhout ◽  
S. E. Hill
Keyword(s):  
Single Peak ◽  
Longitudinal Flow ◽  
Action Current ◽  
Protoplasmic Surface ◽  
Short Period ◽  
Outer Protoplasmic Surface

Several forms of the action curve are described which might be accounted for on the ground that the outer protoplasmic surface shows no rapid electrical change. This may be due to the fact that the longitudinal flow of the outgoing current of action is in the protoplasm instead of in the cellulose wall. Hence the action curve has a short period with a single peak which does not reach zero. On this basis we can estimate the P.D. across the inner and outer protoplasmic surfaces separately. These P.D.'s can vary independently. In many cases there are successive action currents with incomplete recovery (with an increase or decrease or no change of magnitude). Some of the records resemble those obtained with nerve (including bursts of action currents and after-positivity).

scholarly journals KINETICS OF PENETRATION

1934 ◽  
Vol 17 (3) ◽  
pp. 469-480 ◽  
Author(s):  
W. J. V. Osterhout ◽  
S. E. Kamerling ◽  
W. M. Stanley
Keyword(s):  
Partition Coefficient ◽  
Partition Coefficients ◽  
Molecular Form ◽  
External Solution ◽  
Living Cells ◽  
Protoplasmic Surface ◽  
Weak Electrolytes ◽  
Ionic Mobilities ◽  
Aqueous Layer ◽  
Kinetics Of

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.

scholarly journals ACTION CURVES WITH SINGLE PEAKS IN NITELLA IN RELATION TO THE MOVEMENT OF POTASSIUM

1940 ◽  
Vol 23 (6) ◽  
pp. 743-748 ◽  
Author(s):  
W. J. V. Osterhout ◽  
S. E. Hill
Keyword(s):  
Outer Surface ◽  
Distilled Water ◽  
Protoplasmic Surface ◽  
Outer Protoplasmic Surface ◽  
Two Peaks

In Nitella the action curve has two peaks, apparently because both protoplasmic surfaces (inner and outer) are sensitive to K+. Leaching in distilled water makes the outer surface insensitive to K+. We may therefore expect the action curve to have only one peak. This expectation is realized. The action curve thus obtained resembles that of Chara which has an outer protoplasmic surface that is normally insensitive to K+. The facts indicate that the movement of K+ plays an important part in determining the shape of the action curve.

scholarly journals THE ROLE OF WATER IN PROTOPLASMIC PERMEABILITY AND IN ANTAGONISM

1956 ◽  
Vol 39 (6) ◽  
pp. 963-976 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Surface Layer ◽  
Water Content ◽  
Active Substance ◽  
External Medium ◽  
Surface Active Substance ◽  
Potassium Ions ◽  
Protoplasmic Surface ◽  
Outer Protoplasmic Surface ◽  
Aqueous Surface ◽  
Surface Active

The behavior of the cell depends to a large extent on the permeability of the outer non-aqueous surface layer of the protoplasm. This layer is immiscible with water but may be quite permeable to it. It seems possible that a reversible increase or decrease in permeability may be due to a corresponding increase or decrease in the water content of the non-aqueous surface layer. Irreversible increase in permeability need not be due primarily to increase in the water content of the surface layer but may be caused chiefly by changes in the protoplasm on which the surface layer rests. It may include desiccation, precipitation, and other alterations. An artificial cell is described in which the outer protoplasmic surface layer is represented by a layer of guaiacol on one side of which is a solution of KOH + KCl representing the external medium and on the other side is a solution of CO2 representing the protoplasm. The K+ unites with guaiacol and diffuses across to the artificial protoplasm where its concentration becomes higher than in the external solution. The guaiacol molecule thus acts as a carrier molecule which transports K+ from the external medium across the protoplasmic surface. The outer part of the protoplasm may contain relatively few potassium ions so that the outwardly directed potential at the outer protoplasmic surface may be small but the inner part of the protoplasm may contain more potassium ions. This may happen when potassium enters in combination with carrier molecules which do not completely dissociate until they reach the vacuole. Injury and recovery from injury may be studied by measuring the movements of water into and out of the cell. Metabolism by producing CO2 and other acids may lower the pH and cause local shrinkage of the protoplasm which may lead to protoplasmic motion. Antagonism between Na+ and Ca++ appears to be due to the fact that in solutions of NaCl the surface layer takes up an excessive amount of water and this may be prevented by the addition of suitable amounts of CaCl2. In Nitella the outer non-aqueous surface layer may be rendered irreversibly permeable by sharply bending the cell without permanent damage to the inner non-aqueous surface layer surrounding the vacuole. The formation of contractile vacuoles may be imitated in non-living systems. An extract of the sperm of the marine worm Nereis which contains a highly surface-active substance can cause the egg to divide. It seems possible that this substance may affect the surface layer of the egg and cause it to take up water. A surface-active substance has been found in all the seminal fluids examined including those of trout, rooster, bull, and man. Duponol which is highly surface-active causes the protoplasm of Spirogyra to take up water and finally dissolve but it can be restored to the gel state by treatment with Lugol solution (KI + I). The transition from gel to sol and back again can be repeated many times in succession. The behavior of water in the surface layer of the protoplasm presents important problems which deserve careful examination.

scholarly journals CHANGES OF APPARENT IONIC MOBILITIES IN PROTOPLASM

1936 ◽  
Vol 20 (1) ◽  
pp. 13-43 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Sea Water ◽  
Chemical Compounds ◽  
Monomolecular Layer ◽  
Normal Cells ◽  
Protein Film ◽  
Organic Ions ◽  
Protoplasmic Surface ◽  
Short Latent Period ◽  
Ionic Mobilities ◽  
Cell Changes

In normal cells of Valonia the order of the apparent mobilities of the ions in the non-aqueous protoplasmic surface is K > Cl > Na. After treatment with 0.01 M guaiacol (which does not injure the cell) the order becomes Na > Cl > K. As it does not seem probable that such a reversal could occur with simple ions we may assume provisionally that in the protoplasmic surface we have to do with charged complexes of the type (KXI)+, (KXII)+, where XI and XII are elements or radicals, or with chemical compounds formed in the protoplasm. When 0.01 M guaiacol is added to sea water or to 0.6 M NaCl (both at pH 6.4, where the concentration of the guaiacol ion is negligible) the P.D. of the cell changes (after a short latent period) from about 10 mv. negative to about 28 mv. positive and then slowly returns approximately to its original value (Fig. 1, p. 14). This appears to depend chiefly on changes in the apparent mobilities of organic ions in the protoplasm. The protoplasmic surface is capable of so much change that it does not seem probable that it is a monomolecular layer. It does not behave like a collodion nor a protein film since the apparent mobility of Na+ can increase while that of K+ is decreasing under the influence of guaiacol.

scholarly journals THE MECHANISM OF ACCUMULATION IN LIVING CELLS

1952 ◽  
Vol 35 (4) ◽  
pp. 579-594 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Osmotic Pressure ◽  
Outer Surface ◽  
Organic Molecules ◽  
Sea Water ◽  
Relative Rate ◽  
Point Of View ◽  
External Concentration ◽  
Protoplasmic Surface ◽  
A Cell ◽  
Outer Protoplasmic Surface

When a compound enters a living cell until its activity becomes greater inside than outside, it may be said to accumulate. Since it moves from a region where its activity is relatively low to a region where its activity is relatively high, it is evident that work must be done to bring this about. The following explanation is suggested to account for accumulation. The protoplasmic surface is covered with a non-aqueous layer which is permeable to molecules but almost impermeable to ions. Hence free ions cannot enter except in very small numbers. The experiments indicate that ions combine at the outer surface with organic molecules (carrier molecules) and are thus able to enter freely. If upon reaching the aqueous protoplasm these molecules are decomposed or altered so as to set the ions free, the ions must be trapped since they cannot pass out except in very small numbers. If we adopt this point of view we can suggest answers to some important questions. Among these are the following: 1. Why accumulation is confined to electrolytes. This is evident since only ions will be trapped. 2. Why ions appear to penetrate against a gradient. Actually there is no such penetration since the ions enter in combination with molecules. The energy needed to raise the activity of entering compounds is furnished by the reactions involved in the process of accumulation. 3. Why, in absence of injury, ions do not come out when the cell is placed in distilled water. Presumably the outgoing ions will combine at the outer surface with carrier molecules and then move inward in the same way as ions coming from without. 4. Why the relative rate of penetration falls off as the external concentration increases. This is because the entrance of ions is limited by the number of carrier molecules but no such limitation exists when ions move outward since they can do so without combining with carrier molecules. 5. Why accumulation is promoted by constructive metabolism which is needed to build up the organic molecules and by destructive metabolism which brings about their decomposition. 6. Why measuring the mobilities of ions in the outer protoplasmic surface does not enable us to predict the relative rate of entrance of ions. We find for example in Nitella that K+ has a much higher mobility than Na+ but the accumulation of these ions does not differ greatly. This is to be expected if they enter by combining with molecules at the surface. Only if K+ is able to combine preferentially will it accumulate preferentially. 7. Why ions may come out in anoxia and at low temperatures. If these conditions depress the formation of carrier molecules and their decomposition in the protoplasm, the balance between intake and outgo of ions will be disturbed and relatively more may come out. 8. Why the excess of internal over external osmotic pressure is less in sea water than in fresh water. As the external concentration of ions increases the rate of intake does not increase in direct proportion since the number of carrier molecules does not increase and this slows down the relative rate of intake of ions. But it does not slow down the rate of exit of ions since they need not combine with carrier molecules in order to pass out. Hence the excess of ions inside will be relatively less as the concentration of external ions increases. 9. How water is pumped from solutions of higher to solutions of lower osmotic pressure. If metabolism and consequently accumulation is higher at one end of a cell than at the other, the internal osmotic pressure will be higher at the more active end and this makes it possible for the cell to pump water from solutions of higher osmotic pressure at the more active end to solutions of lower osmotic pressure at the less active, as shown experimentally for Nitella. This might help to explain the action of kidney cells and the production of root pressure in plants.

scholarly journals EFFECTS OF HYDROXYL ON NEGATIVE AND POSITIVE CELLS OF NITELLA

1945 ◽  
Vol 29 (1) ◽  
pp. 43-56 ◽  
Author(s):  
W. J. V. Osterhout
Keyword(s):  
Fatty Acids ◽  
Fatty Acid ◽  
Dilute Solution ◽  
Pore System ◽  
Positive Direction ◽  
Negative Cell ◽  
Protoplasmic Surface ◽  
Potassium Effect ◽  
Ionic Mobilities

Remarkable changes are brought about by KOH in transforming negative cells of Nitella (showing dilute solution negative with KOH) to positive cells (showing dilute solution positive with KOH). NaOH is less effective as a transforming agent. This might be explained on the ground that the protoplasm contains an acid (possibly a fatty acid) which makes the cell negative and which is dissolved out more rapidly by KOH than by NaOH, as happens with the fatty acids in ordinary soaps. Part of a negative cell can be changed to positive by exposure to KOH while the untreated portion remains negative. After exposure to KOH the potential the protoplasm has when in contact with NaCl may increase. At the same time there may be an increase in the potassium effect; i.e., in the change of P.D. in a positive direction observed when 0.01 M KCl is replaced by 0.01 M NaCl. In some cases the order of ionic mobilities is uK > vOH > uNa. This shows that the protoplasmic surface cannot be a pore system: for in such a system all cations must have greater mobilities than all anions or vice versa.

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