Injury, Recovery and Death in Relation to Conductivity and Permeability

It would seem, therefore, that deformation may rup- ture the surface lavers of certain structures and cause their contents to diffuse out. If the diffusing substances meet others, from which they were separated by the semi- permeable surface layer before it was ruptured, it is 65 In many cases rupture of the plasma membrane causes the proto- plasm to disintegrate and mix with the surrounding medium. In other cases the surface layer is at once reconstituted.

easy to see that reactions may occur which may produce the responses characteristic of mechanical stimulation. The occurrence of such reactions seems probable, since many cases are known where substances in close juxta- position are prevented from reacting by the presence of semipermeable layers: when these layers are destroyed (by crushing the cells) the reaction at once takes place. If these processes occur it is evident that purely physical alterations in the protoplasm can give rise to chemical changes. Responses to contact and mechanical stimuli may thus be explained; and since gravitational stimuli involve deformation of the protoplasm we may extend this conception to geotropism.66

In this conception of mechanical stimulation the essential things are (1) substances which are more or less completely prevented from reacting by semipermea- ble surfaces, (2) a deformation of the protoplasm suffi- cient to produce in some of these surfaces a rupture which is not at once repaired, (3) a resulting reaction which produces the characteristic response to the stimulus. Some authors (particularly Lillie and Hober)67 as- sume that stimulation is always associated with a change in permeability, whereby the cell surface (which is

60 Small (1918) has made experiments on geotropism by inserting electrodes (a short distance apart) near the tip of the root and measuring the electrical conductivity before and during stimulation. He states that when the root is placed at angle to the vertical the resistance of both the upper and lower sides decreases, but the decrease is less in the upper side. He interprets this to mean that the permeability of both sides increases, but the increase is less in the upper side: in consequence it is more turgid and thus causes the downward bending of the tip.

Owing to the source of error in this method it is difficult to judge of the value of the results. It seems highly probable that upon changing position of the root there would be a movement of liquids and gases in the intercellular spaces which would change the conductivity. assumed to be permeable in the resting state to kations, but impermeable to anions)68 suddenly becomes permea- ble to anions, and hence becomes electrically negative. This hypothesis has been favorably received in some quarters, but according to, Loeb and Beutner,69 such as- sumptions are unnecessary. The conclusions of Loeb and Beutner are based on accurate quantitative experiments and in addition they have succeeded in imitating artificially some of the most important phenomena, both qualitatively and quantitatively.

It may be added that the facts mentioned above70 show that the electric current passes readily through the proto- plasm. This could not be the case if it were not permeable to both anions and kations. For further information, the reader is referred to the literature quoted, as a detailed discussion lies outside the scope of the present work. The facts set forth in this chapter indicate that changes in permeability may be followed by determining electrical conductivity. These alterations are evidently important, since they may affect all the fundamental life- processes. It has been shown that a study of such changes by means of electrical measurements makes it possible to treat such conceptions as vitality, injury, recovery, and death in a quantitative manner. It also enables us to predict the behavior of tissues, especially in respect to injury and recovery, and leads directly to a quantitative theory of the mechanism of certain fundamental life processes.

68 The idea that the cell surface may be permeable to only one kind of ions was suggested by Ostwald (1890). ALCOCK, N. H. 1906. The action of anesthetics on living tissue. I. Action on the isolated nerve. II. The frog's skin. Froc. Roy. 8oc., BECKER, G. F. and VAN ORSTRAND, C. E. 1909. Smithsonian mathematical tables, hyperbolic functions. Washington. BERNARD, CLAUDE. 1879. Lecons sur les phenomenes de la vie communs aux animaux et aux vegetaux. 1:28. Paris. BEUTNER, R. 1912, A. Potential differences at the junction of immiscible

BEUTNER, R. 1913, B. Water-immiscible organic liquids as central con- ductors in galvanic cells. Trans. Am. Electrochem Soc. 23:401. BEUTNER, R. 1913, C. Neue Erscheinungen der Elektrizitiitserregung welsche einige bioelektrische Phanomene erklaren. Ztschr. f. Elek- possible applications of physico-chemical reactions. Jour. Am. Chem. BEUTNER, R. 1913, E. New electric properties of a semi-permeable mem- brane of copper ferrocyanide. Jour. Phys. Chem. 17:344. BEUTNER, R. 1920. Die Enstehung elektrischer Strome in lebenden

BOTAZZI, F. 1911. Das Cytoplasma und die Korpersiifte, in H. Winter- stein's Handbuch der vergl. Physiol. I. BUGARSKY, S. and TANGL, F. 1897. Eine Methode zur Bestimmung des relativen Volums der Blutkorperchen und des Plasmas. Zentralbl. f. of ether on the respiration and growth of Bacillus subtilis. Jour. *No attempt has been made to give a complete bibliography. In general only those papers are listed which are mentioned in the text. BROOKS, M. M. 1919. Comparative studies on respiration. VIII. The respiration of B. subtilis in relation to antagonism. Jour. Gen. Phys- iol. 2:5.

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ologischen Vorgange ini menschlichen Korper auf denen der psycho- galvanische Reflex beruht. Munch. Mediz. Wochenschr. 43:2389. GOLDMANN, E. E. 1912. Die aussere und innere Sekretionen des Organ- determination of the freezing point of vegetable saps. Plant World GRAY, J. 1913, A. The effects of hypertonic solutions on the eggs of GRAY, J. 1913, B. The electrical conductivity of fertilized and unfertil- ized eggs. Jour. Marine Biol. Assoc. 10:50. GRAY, J. 1915. Note on the relation of spermatozoa to electrolytes and its bearing on fertilization. Quart. Jour. Micros. Soc. 61:119.

GRAY, J. 1916. The electrical conductivity of echinoderm eggs and its bearing on the problems of fertilization and artificial parthenogenisis. cycle alternating current in the measurement of the conductivity of electrolytes. Am. Jour. Bot. 4:411. GREEN, N. B. 1918. The effect of ions of sodium chloride and calcium chloride upon the electrical conductivity of certain colloid mixtures. effect of anaesthetics and other substances on the respiration of Asper-

effects of antagonistic salts on the respiration of Aspergillus niger. upon the hydrogen ion concentration. Jour. Biol. Chem. 26:515. HAAS, A. R. C. 1916, B. The permeability of living cells to acids and HALL, R. E., arid ADAMS, L. H. 1919. Application of the thermionic am- plifier to conductivity measurements. Jour. Am. Chem. Soc. 41:1515. HANSTEEN-CRANNER, B. 1910. Ueber das Verhalten der Kulturpflanzen zu den Bodensalzen. I und II. Prings. Jahrb. f. w. Bot. 47:289. HANSTEEN-CRANNER, B. 1914. Ueber das Verhalten der Kulturpflanzen

zu den Bodensalzen III. Beitriige zur Biochemie und Physiologic der Zellwand lebender Zellen. Prings. Jahrb. f. w. Bot. 53:536. HANSTEIN, J. 1870. Ueber die Bewegungsercheinungen des Zellkerns in ihren Beziehungen zum Protoplasma. Sitzungsber d. niederrhein. resembling sea urchin eggs in certain of their physical properties. HARVEY, E. N. 1912, B. A new type of artificial cell suitable for per- meability and other biochemical studies. Biochem. Bull. 2:50. HARVEY, E. N. 1914. The permeability of cells for acids. Internat.

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