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PROTOPLASMA-MONOGRAPHIEN
VOUJMK
i>

OTTO HAHN, INVISIBLE RADIATIONS OF ORGANISMS

Protoplasma-Monographien
R. Chambers

Herausgogebeu von (New York), E.Faure'-Fremiet (Paris), H.Freundlich (London), E.Kiister
Lloyd (Montreal),
J.

(GieBen),

F. E.

H. Schade

(Kiel) f,

W.

Seifriz

(Philadelphia),

Spek (Heidelberg), W.
Redigiert von
F.

Stiles

(Birmingham)

Weber (Graz)

VOLUME

9

INVISIBLE

RADIATIONS

OF ORGANISMS
OTTO RAHN
PIIOJ'LSSOR OF JUfilKRlOLOfrr, COIIXKLL

FXIVLKStTY

WITH

.V2

ILI.rSTKATIONS

Berlin

Verlag von Gebruder Borntraeger
W35
Koester-Ufer 17

1936

INVISIBLE RADIATIONS

OF ORGANISMS

OTTO KAHN
PROFESSOR OF BACTERIOLOGY, CORNELL UNIVERSITY

WITH AN INTRODUCTION TO TUK PHYSICS OF RADIATION
BY

S1DNKY

\V.

IJAHNES

RESEAltCH ASSOCIVTE IN PHYSICS, UNIVERSITY OF RUCUKSTRR

WITH

52

II.IATSTRATIONS

Berlin
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insbesondero das Rroht dm- Ultt'rsetzung in fromde Sprachen,

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Copyright 193li

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Borntivipger in Berlin

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or of luring the male to the female. some disturbance of its mechanism. is They is are evident the healing of wounds. perhaps this Beta-mdi'cit'um controls the heart beat. Old age accompanied by complete the cause of old The. the invisible radiations of living organisms are of considerable physiological significance. The phosphorescent bacteria usually lose the ability to produce light when cultivated for some time on artificial media. without any apparent decrease in vitality. cessation of ultraviolet emission. Mutual influences of one species upon another Its role in the by radiation have been observed. The emission it of visible light is probably of no greater cell physiology of the organisms. is radiation tion. plays no essential role in the a distinct part in ill cell division and in growth. or used in the diagnosis of cancer. other investigators have contested these theories. While it is assumed by some biologists that it has the purpose of attracting the prey. perhaps only because the average man (not to mention woman) likes to believe in human radiations. of frightening enemies. loss of blood age. fireflies and glow worms.FOREWORD Visible biological radiations have always greatly attracted man's curiosity. and the illuminating organs of deep sea fish are among the well-known wonders of nature. the principal reason for the rejection of the discovery . Quite to the contrary. However. Biologists and physicists have always been suspicious of radiations from living organisms. They play importance than color. The biological I importance of this luminescence seems to be in no proportion to the impression it makes upon the human mind. phosphorescent meat. uniiiit went wood. it may is be that radiathe. metamorphosis amphibia has been demonstrated. Jinked with of cause of cancer.

a more conciliatory attitude has become noticeable since it has been shown that mitogonetic radiation not a mysterious force. in Chapter IV which discusses the various An approach be predicted from the results of physico-chemical to historical presentation is found methods used. The book deals almost exclusively with mitogenetic rays w hich r exist in the ultraviolet range of the spectrum. but the result of biochemical processes. GOKWITSCH had not discovered these emanations 10 years ago. Many simple chemical reactions have been found to emit weak is ultraviolet rays. though several factors responsible for negative results have been discovered. . it is tant. This very fact has been one of the authors' reasons for presenting the more important facts in this book. This had led to the fallacy that negative results disprove positive is quite evident that if two experimenters obtain different they cannot possibly iiave made the same experiment. but not really characteristic of the living is proportional to the potassium content. and the important task is to find out in what points the investigations differed. With a phenomenon so little understood as these biological radiations. it is not surprising that these apparent contradictions have not as yet been explained in every ease. #eta-ray emission from potassium it is biologically imporcell. and death as during life. ones. almost all happen to contain negative results. No definite proof for the emission of infrared rays found (if by organisms could be we limit the infrared to radiations near that of the is visible). The objection this country. of the subject matter is not historical. just as strong after The arrangement logical. of this new Another factor is responsible for the slow adoption influence in biology: practically all papers on this subject are published in foreign languages. to biological radiations has been strongest in but even here. Both results are correct. but attempt has been made to show that ultraviolet If radiation from living organisms is nothing at all strange. It results. and of the very few in English.VI of ultraviolet emission FOREWORD from living cells was the inability of some to repeat the positive experiments of others with the same results. they would An now investigations.

up to the beginning the book by HTKMPKLL. and to Miss A. BARNES for her ceaseless assistance in editing he!]) in tin's book. Europe. J. during a recent journey to A very brief summary is given at the end. The authors an? further under great obligations to Mrs. FFJKJUSON for her proof-reading. Fthaca. and to Professor MACJROCT for the kindness of sending original photo- graphs of his experiments for the reproduction in this book. A fairly complete may be found in papers which have been quoted in the text: we suppose that this includes the more important publications. One of the authors had occasion. The literature compiled at the end of this book refers only to those greatly. August OTTO RAHX SIDNEY W. BAKNKS . to see this field. MARGARET N. of 1932. This might be more useful in some respects than the customary Table of Contents. The authors are further under great. not meant to represent a compilation of all This would have increased its size list of references. obligation 011 to Professor ALEXANDER CUIIWITHOH for sound advice various points. (1032). many of the biologists and physicists working in and he wishes to acknowledge the many valuable suggestions he received from those convinced of mitogeiietic radiation as well as from those who are convinced of its non-existance.FOREWORD The book literature is VJf on this subject. chapter by chapter. of the entire book.

.

:H HI ... . . . Beta -radiation I). cellb . General statements B.. Analysis of radiation by dispeision into a spectrum F. . 9 IS E. .. morphological changes. -physical methods) .H A.. . II .. Phenomena observed upon the interaction The wave theory of radiant energ\ The <|uantuin theory of radiant energx . 54 95 99 Infra-red rays. J3 CHAPTER III . Injurious human radiations C. J). Ncerobiotic rays . EFFECT OF ULTRAVIOLET RADIATIONS UPON CELLS A. .. .. . yeawt bud method. Intensity measurements of visible and ultraviolet radiations . . Effect of radiations upon chemical reactions B. 10J . changes in yeast metabolism. . V CHAPTER I . . Effect of radiation from chemical reactions upon . .. . . rays (onion root method.. increase in cell numbers. . Mitogeiictie. . Physical sources. . . 101 . 23 CHAPTER SOURCES OF RADIANT ENERGY. I I C.. ."50 CHAPTKR IV METHODS OF OBSERVING BIOLOGICAL RADIATIONS ... . . . A. . B. .'33 Secondary radiation . Ohemieal sources 0.TABLE OF CONTENTS FOREWORD PHYSICS OK KADIATION A. . .. physico-chemical methods. . . .. . . B.. E. . . of radiation and mutter . . . . . . cell division in larger organisms. . 40 46 48 . Effect of monochromatic ultraviolet upon cells C.. 2 *J . . .

Adaptation to gradual increases in intensity CHAPTER A. 142 HO 148 Blood radiation Nerve radiation and conduction of stimuli 154 G.... OS Intensity of the mitogenetic effect E. . K. parthenogenesis) Symbiosis and parasitism . metamorphosis of amphibia and insects. 138 . Influence of diffuse daylight C. Unicellular organisms (emission at different physiological stages. Higher plants C.. I). Retardation through radiation I). 1. change of chromosome number. E. Relation hetween the intensities of radiation and of effect C. Secondary radiation 103 103 10S I .X TABLE OF CONTENTS Pigo CHAPTER V SPECIAL CHARACTER FSTICS OF BIOLOGICAL RADIATIONS A. The . Tissues of adult animals E. If. 120 127 I2S Mitogenetic effects in multicellular organisms CHAPTER Vll SIGNIFICANCE OF BIOLOGICAL RADIATIONS IN BIOLOGY. ... 122 123 The minimal The harmful intensity required for an effect effect of over-exposure 125 .. .. bacteria. VJ 119 120 ANALYSIS OF THE MITOGENETIC EFFECT necessity of a particular physiological stage B.. Intermittent radiation B. I U 115 117 F. The cancer problem..... Radiations other than mitogenetic 184 188 OUTLOOK SUMMARY OF THE BOOK AUTHOR INDEX SUBJECT INOEX 193 196 209 . Morphological effects produced by mitogenetic rays (yoasts. Storage of mitogenetic charges F. F. sea urchin larvae. . 101 170 173 177 Healing of wounds J.KON yeast detector V*. 131 B.. reaction 131 upon irradiation) . MEDICINE AND AGRICULTURE A.. Eggs and embryonic stages of higher animata D. Mechanism of the BA.

definite velocity. radio waves. Radiant energy might be given the alias of traveling energy. (2) occurs over an extended energy range. For one thing. radio waves are completely absorbed by a coarsely-woven copper wire screen. traveling through space at its particular speed of 3xl0 per second (speaking here of space in which the matter density is zero). To the best of our knowledge. Any form of wave motion can be made to exhibit the Protoplasms. . it is known that such energy is propagated through empty spaee. Radiation energy spectrum. visible light and gamma rays all travel with precisely this velocity. from one part of the radiant energy spectrum to another.CHAPTER 1 PHYSICS OF RADIATION A. The direction of travel (4) is rectilinear. Radiation (3) Radiant energy travels through space with a fixed. ^he studies of radiant energy have come several ideas about its nature. and. (1) GENERAL STATEMENTS From Radiation travels through empty space. The character of radiation varies greatly. Radiation exhibits the phenomena of interference. the lower-energy jr-ray* pass through perhaps an eighth of an inch of lead visible light is absorbed by a metal layer only a few atoms thick. The extremely high-energy gamma rays penetrate several inches of lead. known which can flow through it is the only form of energy matter-free spaee. finally. Moreover. come millions of miles of interstellar space. -MonograpMen IX: Kahn ] . is . as one can see. for it spends each instant of its existence 10 cm. The vast amounts of solar radiation life temperature that possible. which maintain the earth at such a from the sun through which contains but an infinitesmal density of matter. This speed is entirely independent of the character of the radiation.

This transformation obeys the law of conservation of energy. also see p. 16). PHENOMENA OBSERVED UPON THE INTERACTION OF RADIATION AND MATTER (1) Kef Joe ti on. are an example.2 CHAPTER of interference. 128). The beats heard when two tuning same frequency are struck. . In this case the angle equal to the angle of incidence. it is necessary that it be transferred into *the familiar- potential or kinetic energy of matter. (3) The mechanism of ab- Refraction. the number of ergs of radiant energy disappearing potential or kinetic energy is equal to the number of ergs of which appear. (4) called the Dispersion. The ratio of the is velocity of radiation in matter to the velocity in space index of refraction of the refracting substance. (2) Absorption. fails to take its toll from the is radiation incident No material substance known totally transparent to radiant energy. This statement is a all reminder that recognized measurements of energy are limited to energy associated with matter. (5) it is Radiation may be observed when. 2 a arid b. I phenomena Later will forks of nearly the (see p. sorption will be treated later (see p. the* velocity of energy which is passing through change of direction of the radiation (see fig. The phenomenon of dispersion occurs because the index of refraction of any transparent material depends also upon the wave length of the radiant energy which is passing through it. i. B. Matter never upon it. and the reflected in the plane of the incident energy. always some of the radiation is transmitted or absorbed by the reflector. and only when. allowed to interact with matter. Matter has the property of changing it. 18). To detect or measure radiant energy. This results in a 1). Radiant energy may be regularly reflected from a plane surface whose granular structure is small in compari- son with the of reflection energy is of the radiation. an example of interference exhibited by light be given. e. A beam of white light is dispersed into a colored band or spectrum suffers when passed through a prism since each wave length a different refraction (see figs. No perfect reflectors is wave length of radiation are known. thus.

refraction and dispersion. This sort of disturbance is called a wave train. it is a true radiation. (2) by wave motions in elastic media. a seen to speed along it. Figure 2. Refraction and Dispersion. for a short period of time this end of the rope is given a regular to-and-fro motion. C. it seems not impossible to effect a harmonious combination of these two into one which adequately covers all uho observed phenomena of radiant energy. Refraction. Air Figure 1. theory. or the .3) by material projectiles. based upon the well understood principles of wave motions in elastic solids.PHYSICS OF RADIATION 3 rize radiant These are only a few of the many phenomena which characteenergy in its passage through space and matter these must be explained by any theory of radiation. the wave theory and the quantum . If so. The question: WJiat is the nature of radiant energy ? has been nearly answered bv each of two different theories. namely (1) by the How or movement of definite masses of matter. absorption. or the drive rod on a locomotive. If. as will be demon- The strated in Chapter TV. These? may be illustrated by the as sound in air. The length of the individual waves. and to show reflection. hump in the rope is a disturbance as pictured in fig. As it will be pointed out later. a rather surprising thing happens (at least so to the uninitiated) . THE WAVE THEORY OF RADIANT ENERGY There are three ways in which energy may be transferred with the aid of matter. If a long stretched rope is given a blow at one of its supports. so-called "mitogenetic radiation" which is the principal subject of this book is said to proceed rectiliiiearly. 3 will travel along it with the same velocity as in the former case. is and of radiation following simple experiments. such as tides in the seas. such The wave theory (.

divided by tho wave length: Reflection. wave c. train will set it disappear. the train of waves will be reflected and will travel back along the rope with the same velocity and amplitude it had before reflection. not of little or no motion called nodes. Absorption. The rope appears to be divided When Figure 4. the energy carried by the wave train in has been transferred to the support. true waves at all. train in a rope. the wave motion. are the result of the interference of the original . spend its energy upon it and completely In this event. equal to the velocity. Standing Waves. Above: Waves in a rope. A. into vibrating segments called loops which are separated by points These standing waves. The frequency. or the number of waves passing a x ^ x is -^ . below: standing waves in a rope. ' ^_^^"- the distance from crest to crest or trough to v. fixed point per second.4 CHAPTER length. is I wave trough. If the support there is ideally rigid. 4). it is fastened to the rigid support. If the support is ideally non-rigid. There is still wave motion which may be illustrated another phenomenon of by waves in the rope. if the free end is kept moving with a uniform to-and-fro motion. so-called standing waves will be formed (see fig. A Figure short 3. Let this wave train be o bserved when it reaches the end of the rope.

The mechanical rope apparatus is frequently used as an analogy to the electric field of a point into which a stone has been dropped. 5). The wave trains traveling outward along each rope with the same velocity give the appearance of regularly growing or spreading concentric rings. Definition of a Charge of Electricity. . waves will travel out along each rope and the system will present somewhat the appearance of still water ' A left: Figure 5. and this excess is called an electric charge. If an object has equal amounts of the two. If more ropes are added to this system so that they are stretched equally m every direction in various planes and the central point is given a regular to-and-fro motion the system will give the appearance of expanding or growing spherical shells. Here the distance between the shells is again equal to the length of the waves in the individual ropes. If it has an excess of either kind it is said to be charged. embodies two kinds called positive and negative. Radiation from a central point. right: waves in a rope system. Tf the central point is given a regular up-anil-down motion. The rings are separated by a distance equal to the length of the waves. according to present ideas.PHYSICS OF RADIATION 5 and the reflected waves. " S uPP 08e a system of ropes is strung from a central point so they lie in a plane (see fig. Electricity. nodes. a rope system. Usually this charge is distributed over the surface of the object. (Charge. Their importance lies in the fact that they offer a very simple way of determining the wave length of the true waves which is equal to twice the distance between . it is said to be electrically neutral.

6 CHAPTER I In discussions of the effects of one charge upon another. We may then think. it is convenient to ignore the object and to think of the charge as being concentrated at one point. . Now. The train of waves which force it consists in variations in the direction of the line of is traveling along this line with the velocity of light. In fig. Suppose a charge of positive electricity is fixed in space at some pornt A. Now is known wave that the motion of an electric field. if the charge at A were given a rapid to-and-fro motion. We will now see why the three-dimensional system of ropes forms a rough mechanical analogy to the electric field of a point charge. These two wave trains lie in pianos perpendicular to each other. The waves in the electric field about the charge which has been set in oscillation arc known to be not the only waves present. and in related problems. field through space produces an associated magnetic this It theref on* follows that it train must have associated with a train of waves of magnetic. This is called a point charge. Moreover. The point ft may be anywhere in space in the vicinity of ^1 and still it is drawn directly toward /I. If a charge of negative electricity is brought to some point B. intensity. The wave theory predicts that the velocity of propagation of these waves should be independent of their wave length. surround electric charges. Radiation is associated with waves traveling through these fields. 6. it experience a force which tends to draw it straight toward A though the two were connected by an invisible stretched elastic cord. The Electric Field of a Point Charge. it would seem likely that waves should be formed and sped along will as the lines of force through space. if we like. Radiation is then nothing other than these electromagnetic waves. The classical electromagnetic wave along one line of force is represented in fig. of the space about A as filled or made up of these stretched elastic cords (called lines of force) extending outward in every direction from the charge at point A. such waves would be expected to exhibit all the pheno- mena of interference just as radiation does. 3 is represented the shape of one of the lines of force shortly after the charge has been given a few oscillations. This is the explanation offered by the wave theory of light concerning the manner in which radiant energy electric fields We know that is propagated through space.

7). for when a spark occurs the charge flows across the as to discharge the condenser. not only line of force but in all directions. 1 In fig. This process repeated many times a second amounts to a charge moving rapidly back and forth spark gap in such a way flow until the condenser is between the spheres of the gap or a charge closely-placed points. and S a spark gap. Experiments showing the wave nature of Hertzian waves. though the amount of radiant energy sent out in different directions varies. Plane ofElectrostatic Field Zero Figure is fi. distance from the oscillating charge Mxtor Figure 7. The charge then begins to reverse its direction of flow.PHYSICS OF RADIATION An along one oscillating charge radiates energy. radiated along the line of motion of the charge and the is radiated in a plane normal to the direction of motion. c represents a condenser. HEKTZ (1866) caused a charge to oscillate rapidly between ei. oscillating rapidly between two . 7. The con) denser \\as charged to such a difference of potential that a spark occurred between the balls of the spark gap.ergy maximum amount two closely-placed points 1 ) and found that energy was being ra-dia-tcd as He placed a metal plane some the result of the accelerations of the charge (sets tig. of course. Diagram ot an electromagnetic wave. Such an arrangement is known as an oscillatory circuit. and it continues to recharged with the difference of potential reversed.

Jight determined velocity of light. the silver compound was reduced. offered a beautifully direct experiment not only showed that light was a wave motion but way of measuring the wave length. at planes in between. Experiment showing the wave nature of light. the direct and the reflected beam interfering to cause standing waves. Definition of Intensity. 8) alternate exposed and unexposed layers At were found to exist throughout the depth of the emulsion planes where the direct and reflected beams interfered to form the node's of the standing waves. The intensity of radiation received from a source at a fixed point in space is defined as being the number of energy units (ergs) received per second by a square cm. These loops and nodes showed that the energy was being radiated in the form of transverse waves. When the.8 CHAPTER I and found regularly-spaced points between the radiating charge and the reflector at which the energy was alternately of a maximum and of a minimum value. it was immediately suggested that was nothing other than such waves. cut and the edge viewed under a microscope (see fig. When the velocity of these waves. which arc of the wave length of short radio waves. simple experiment performed by LLPPMA^N showed thatbe made to form standing waves and thus must have a wavelike nature. only of shorter wave length. Light of one wave length was allowed to visible light could A upon a fine-grained photographic emulsion which was backed by a reflecting layer of metallic silver. emulsion was developed. the silver compound was shine perpendicularly unaffected.chromatic light G -* standing Surface Figure 8. of surface placed normal to the radiation at that point (see . was found to equal the experimentallyMono . corresponding to the loops of This the standing waves.

a point. A K rr twice as far from the source. dispersion. reflection. faces or volumes. For these cases. cm 2 will be the same at any distance from the source With a point source.) 9. 9b. Experiment is the best means of determining the variation of intensity with distance in the region of space closely surrounding a source of finite size. 9 a).PHYSICS OF RADIATION fig. for points still closer to the source. receives only -. Illustration of the definition of intensity. been successful in explaining bo transferred through space it has predicted accurately the velocity of radiant energy the phenomof radiation has The wave theory radiant energy how may . ena of interference. A given atom. 9 a). for example. will . the classical theory fails and gives place to the quantum theory. will receive ergs per second while B. THE QUANTUM THEORY OP RADIATION It is Definitions. radiation generally being emitted by sur(This is commonly the rule* in biological radiat- Figure ions. however. 1. D. polarization and double refraction offer no difficulties. it will decrease as the second power of the distance between the radiating source and the point of measurement. the inverse square law holds only for distances so great that the source may be considered to be. For shorter distances the intensity may be roughly proportional to the reciprocal of the distance. In practice. When the radiation is strictly parallel. ergs. refraction. point surfaces are rare. . with respect to the emission and absorption of radiant energy. Thus. in fig. known that matter exhibits the curious behavior of discontinuity in processes in which it emits or absorbs radiant energy. the intensity may be independent of the distance. 9 the intensity per (fig.

these differences can occur only by differences in the size of each proIt has been shown that the energy of a jectile. column I are given the various wave lengths in h. known 10~ 10 m). Let us suppose (see fig. of its store of energy. it is of emission and absorption of quanta necessary to discuss briefly the atomic theory. Since they consist simply we have radio waves to gamma seen that the different kinds of radiation. that radiation is corpuscular in nature. e. This electron is Atomic Theory. For the understanding by matter. i. quantum can be given as the product of a universal constant. q. all travel with the same speed. as PLANCK'S constant and equal to 6. Likewise. Let us lay aside for the moment then this conception of the nature of radiation and consider the only other possible one. From our knowledge of electrostatics we know that the electron experiences a force of attraction toward q. 8 Thus. This phenomenon is one with which the wave theory of light is unable to cope. correspond to each frequency (E -hv). from a small positively charged particle. quantum energies.55 x!0~ 27 erg and the frequency of the equivalent electromagnetic seconds. only certain multiples of the unit energy into radiation. 2. Energy of quantum E hv. it will absorb radiant energy only when the energy comes in precisely the proper-sized amounts. of small units of energy. per second. Thus we think now of radiation as consisting of small energy projectiles which travel 10 through space with the familiar velocity of 3 X 1C cm. which E. In Table 1. an X-ray quantum is a 10" erg projectile.10 CHAPTER 1 of cgnvert. F. and those of the ultraviolet being about 2 to 10 times as large as those of visible light. These projectiles are of course non-material. is proportional to the number of units of charge possessed . wave. force. or quantum. Column Ji gives the corresfrom the equation obtained ponding frequencies ANGSTROM units (1 A where In the third column arc the the velocity of light. from rays. while a quantum c is of visible light has an energy value of but 1C" 12 ergs. 10) that an r. held at some distance.

/ CC rH X V N X (M CO I. CO OC r CO oooooooooooooocoooooooooo : / - -^ A x x y x x .PHYSICS OF RADIATION 11 AXXAXXXXX J <q co CO W t (^ CO l^ 1^ O5O5OOOXl-<NOOCOOOiO'NO '-O '-O "> I IQ |- rH <?-! ^' 3D < 1O H^ .< v > % / >.30 Oi OilOOirHOOH^OJOJ ' -' 1-- >O O HH -f H4 CO* ^1 rH 8rH q rH d d d co 10 o* o* PH o q ^ q q o d g o FH < - <N M * CO l^ 'JP O O rH *M I .

\ q \ x *> Figure 11. centripetal force upon it by q. For any value of r.10. However.charged body. fall toward and the electron into the positively. ' ^\ N Figure. 11).. Positive arid negative ative charges at the distance r . An example becomes the nucleus of the atom and the electron represents the hydrogen atom's one orbital electron. a corresponding orbital velocity can be calculated which will bring about the balance of the electrostatic attraction and the centripetal force of the electron. inversely proportional to the square of the distance. . of course. or f ~ ? r * Let us assume that q is heavy compared to the electron so that the electron will be the motile one of the two charges. Ef its orbital velocity is adjusted until its just equal to the force of attraction. F. and the charge q represents an electron. of such a system is furnished by the sun and the Here it is the balance between the earth's centripetal force due to its orbital velocity and the gravitational atractiori of the sun which keeps the* earth in its orbit. Suppose. which is thus the unit of positive earth. that is set travelling in a circular orbit about q as the *~ ~~~ 0-t <T T>_ ^Q electron *' . If the electron is released from its position. q. Electron in a circular orbit traveling about the positive \ * charge q. 11 represents a hydrogen atom. electricity. the distance r js 0. however.5xlO~ 8 cm. If the mass of q is 1. then fig. exerted then the electron will be in stable equilibrium remain- is ing indefinitely in this orbit.65 X 10~ 24 grams. q. From elementary considerations it would seem that the nucleus with would form a stable system also for values of r other than 0. from each other. ^ center (see fig. it is found experimentits orbital electron .12 CHAPTER is I by q and r. it will.5 X 10~ 8 cm.

out of the infinite number possible. From the radius. The atomic number is equal either to the number of nuclear or orbital electrons. . which the orbital electron frequents. Table 2 shows in what way the outer electrons group themselves in the orbital shells for the first eleven elements of the periodic table. and 2 electrons. this leaves the nucleus with a positive charge 4 and the whole atom electrically neutral. It can be seen that the atomic. i. see how Two orbital nuclei. For example. 4 Tlie orbital velocity of the electron decreases as ti increases. to the number of protons in the nucleus of the atom.5xlO~ 8 cm. The atomic weight of carbon is approximately 12 and that of oxygen is 10. weight is equal. In all atoms. can be easily calculated and this energy characterizes the cntrgy state of the atom. The usually unoccupied orbits are called virtual orbit*. and we will very shortly apply this idea in a discussion of the absorption and emission of radiant energy by atoms (see pp. the carbon atom is composed of a nucleus of 12 protons and electrons about which revolve 6 electrons in the various consists of 8 outer electrons shells. 14 and 1C). The oxygen atom and a nucleus which contains 1(> protons and 8 electrons. the number of protons in the nucleus exceeds the number of electrons by just the number hydrogen of orbital electrons. We have seen that the electron hydrogen atom can exist in different energy states. The atom lias the least energy when the is in the innermost orbit (state of least energy) and its energy increases as the electron exists in orbits of greater radius The innermost orbit is the preferred (states of higher energy). r lt of the innermost orbit of which we have just seen to be equal H to 0. it will be advantageous to other atoms beside hydrogen are constituted. In this case. The energy possessed by the atom or the* nucleus-electron s\ stem. e. one. 3. 2. the atom is said to be in its normal state. the electron inhabits this one the greater part of the time. for the case when the electron is in any one particular orbit. electrons complete the atom. The one Its nucleus consists of 4 simple next in simplicity is helium.PHYSICS OF RADIATION ally that there are only a 13 few orbits. and tlie limiting case n-> oo corresponds to an atom with its electron at rest at an infinite distance from the nucleus. we may the relation calculate the other possible orbits by r = n 2r l where n has any value 1. called protons. for all practical purposes. Before considering these topics.

the first circle is called the K shell and contains two electrons.CHAPTER Table 2. The central dot marks the nucleus. of the A't/ Jb'ig. The binding energy these is of negative charges about 1000 electron volts or this 1.6xlO~ 9 ergs. The next circle represeiitsthe L orbit. I Electron Configuration for the Elements from H to Na. about The outer dotted in the circles represent energy levels virtual orbits. i. solid circle there is but its showing the various one electron and is binding energy 5 electron volts. which are unoccupied 3. amount of energy z would be required to remove either one of the K electrons from the atom. normal Na atom. The Emission and Absorption of Radiation by its Atoms. which in Na of contains 8 electrons. In the last Diagram of the sodium atom shells. 12 is a wholly diagrammatic representation atom. energy states associated with the orbit occupied by . e. Figure 12. the binding energies which arc approximately 35 elec1 tron volts. We have seen that the hydrogen atom can exist in various electron.

Em En- In this equation h the universal constant known as PLANCK'S constant equalling 6. Table 3. it does so with the emission of a quantum (1) of radiant energy liv such that .. 4). It follows that the various quanta hv may be emitted uiid absorbed by the hydrogen atom.PHYSICS OF RADIATION 15 By and emission state this necessar}' mechanism can be explained the absorption of radiation. 4. 2. - 4 . -- En+. e is 10 =- where a constant which sec.99790v 10 cm. hr Km K where m hr . per the wave length tion resulting from or producing the energy change the atom will be given by the equation Em Kn in Table 3 gives the values of the first eight of the forty or so known energy states of the hydrogen atom. Knorgy obtained by electron shifts from normal to higher orbits in the Hydrogen atom .. 3 2. hv hr Since A light or = Em E2 -En. 3.. If the atom changes from an energy Em to a lower one En hi> . it (1) Inis .547 .Km E! m in 1. sees.< 10~ 27 erg. is equal to the speed of of the radia- 2. 3. = E m En E n to a higher state of can do so only by the absorption of a quantum of radiation hv of such energy value that If the atom changes from an energy state energy Em . 5 . and v is the frequency (sec p.

99796 x 10 10 X 6. Tf its energy is not one of these discrete values. g. the atom is said to be in an excited state. e. for the shift from the 6th to the 7th orbit. is used in shooting the =E X E n + ^mv 2 is its where m is the mass of the electron and r velocity (the term mv 2 2 represents the kinetic energy of the ejected electron). i. . is called a photoelectron. it is 178 500 A. it will not be absorbed -?. In this event. in the far ultraviolet. It will be.4 x 10. then it may be absorbed. of course.. the electron is in the orbit corresponding to the lower of these states. After this length of time the atom reverts to its normal state with the resulting emission of radiant energy. or hv Ku). at this moment.e. sufficient to shift the energy is greater than (Eoo electron beyond the outermost orbit. it may be absorbed and the surplus. An electron so ejected from an atom ionized.547 x 1C" 27 __ ~ ~~~ 161. states provided that. an electron spends most . if its energy happens to equal the energy difference between any two quantum . and the of its atom itself is said to be As has been stated before. as _ ~~ This wave length 2. The equation (1) states that an atom in the state E n will absorb a quantum hv and be raised to the energy state E m if the is precisely equal to the difference in the energy of the two states.unless its Kn). The life of an atom in an excited state is of the order of 10~ 7 to 10~ 9 seconds. is readily calculated from the data this table.16 CHAPTER I The wave length emitted by the hydrogen atom for the energy change in Ex E for example. If the quantum is larger than Km En. 1216 A. Supposing the energy of the quantum is slightly less than this difference will it be absorbed ? The answer is 110 there is no possibility that it will be. existence in the normal state. or is . E When it has been raised to a state of higher energy E n by virtue of the absorption of energy. hr (K<x> electron away from the atom. Changes between other light states result in the radiation of visible and still others produce radiation in the far infra red.ia 1216 xK)" 8 cm.

to changes in the rotational energy of the dipole molecule.PHYSICS OF RADIATION 17 These icmarks on emission and absorption of radiation apply not only to hydrogen but to the other atoms as well. (2) 13 represent a simple diatomic-.Mo nographien IX: Rah n 2 . while in the solid state it will give continous absorption. The absorption spectrum of a molecule is. These last remarks apply equally well to absorption. a contimioux spectrum is emitted which does not contain lines characteristic of the atom. and the third. vibrational and electronic? energy of the molecule. In this way originate the atomic spectra. The atoms are so mingle and close together that the outer virtual orbits interare distorted. in the simple theory. Pro to plasma. The radiation in the tirst group is ascribed. red. This is called thermal radiation since it is the result of the temperature of the source. Let (1) fig. GHZ) the ultraviolet regions. light bands replaced by dark.molecule such as NO. If the pressure is raised (also the case for solids). Molecular Spectra results from electrons jumping from one energy level to another. to simultaneous changes in rotational. and (3) the visible or A Figure 13. simple diatomic molecule. molecular spectra an* assumed to nrise from the motions of atoms or. An element ill the gaseous state will give a line absorption spectrum. Only the outer electrons of the more complicated atoms behave in a manner similar to the one electron of hydrogen. The great number of linos fall into groups which under low dispersion give the appearance* of bands. Such a molecule emits radiation the far infra red. of course. in three wave length regions: the near infra. They are emitted by sources (such as the mercury arc or a glow discharge tube) in which the gas is at sufficiently low pressure that the atoms arc not in contact with each other but for a small fraction of the time. ions which form the While atomic radiation molecule. The second group is ascribed to simultaneous changes of rotational and vibrational energy. better. Molecular spectra are in general exceedingly more complex than atomic spectra. its emission spectrum in reverse.

and this may. depending upon the temperature of the solid (see lower spectrum of fig. A given atom will emit only certain lines (its lme spectrum) and each one will have associated with it Atomic IliJ Molecular Jherma! 6000 1 . moo visible- A range Figure 14. 14). and thermal radiation. is still the reduction of the salt with the deposition of Without presenting the theory of the process. a certain energy give rise to (see upper spectrum of fig. together with the distribution of energy among these wave lengths. become a chemically just this. and an incandescent solid emits all wave lengths with varying intensity beyond a certain wave length. The most prominent effect is that of light on silver salts as in all photograph ic emulsions. it easy to see that the absorption of radiant energy might do For an absorption of energy means that^tho molecule must go into a state of higher energy-a less stable state. ANALYSIS OF RADIATION BY DISPERSION INTO A SPECTRUM The radiation emitted by a source is characterized by the wave lengths present. it is only necessary to remember the number of brown bottles that are used for storing chemicals. Kepresentation of atomic. molecular. a molecule may some such spectrum as that of the center strip. 14). The result is metallic silver.18 CHAPTER T In this connection. To realize that such an effect does exist. on the absorption of sufficient energy. the effect of radiation upon chemical reactions should be mentioned. . unstable state. E.

The simple optical spectrometer monochromatic illuminator.PHYSICS OF RADIATION 19 The various wave lengths present in the radiation emitted by a source are determined by dispersing the radiation into a spectrum. tho prism alone gives rise to a spectrum in which there is some overlapping of the (sec fig. below: a quartz double moiiochromator.lf prism J Figure 15. When a narrow beam of parallel light falls upon a prism. the prisms and lens must be of quartz.spectrometer. 15 B represents the optical system of a typical quartz double moiiochromator. This subchapter deals with the instruments and methods used to produce a radiation spectrum both in the visible and the ultraviolet. The next deals with the subject of measuring the intensity associated with the various wave lengths. lengths present in the beam suffer different deviations in passing through the prism. beams of light are generally divergent rather than parallel. in practice. Fig. The two instruments most frequently used to disperse light into a spectrum are the prism and the grating. A \ S. As a result. For work in the ultraviolet region. Tn most instruments the prisms may be rotated by some device which is connected to a drum or . each wave length emerges with a slight angular separation from its neighbors. the different wave. Since. By replacing the eye piece by a slit. 15A) prevents this in a lens which forms the by employing eyepiece a narrow image of the slit through which the light enters. Spectrometers above: a simple . the instrument may be used as a moiiochromator wave lengths.

20 calibrated in CHAPTER wave lengths. the intensity of the unwanted be reduced to practically ZATO. relative intensities of the lines in the ultraviolet spectrum of the quartz mercury arc as transmitted by such a monochromator. . This defect may be greatly reduced by using with the moiioehromator a filter having a ''cut off' on the short wave length side of the desired radiation. I Turning this drum to a certain wave length reading sets the prisms so that only this wave length is Table 4 gives the permitted to pass through the second slit. a discussion of their characteristics be omitted. An excellent given in a table (12 5) in Photoelectric Pheno- mena by HUGHES and Du BKTDCJE and the ultraviolet portion is reproduced here with their kind permission (see Table 5). Relative Intensities of II g Spectral Lines Uratings are also used to produce spectra of violet light but since their application is will visible 1 and ultra- more strictly confined to special work in spectroseopy. a Hilger quartz double moiioehromator. some degree of impurity seems unavoidable. In general. 1 ) Table 4. The wave lengths given in the table are those at which the filter ceases 1 ) In particular. some intensity of radiation of shorter 1 wave length than that desired is transmitted by the instrument. While monochromators are designed to give high spectral purity of the isolated light. \vave lengths list may is of such filters By so doing.

kShort Wave Cut-off Kilters *) standard filters of the Corning Glass Works. .PHYSICS OF RADIATION Table 5.

some data are given on the short wave length limit of the solar spectrum. Such a combination may give much greater intensity than could be obtained by the use of the monochromator. By using a suitable filter with such a source. Another experiment carried out is at 9000 meters showed energy present at 2897 A. The conclusion given that the intensity. we must keep in mind the possibility that extremely small intensities of the lower wavelengths exist in day light. Because of the great. Nevertheless. far beyond detection by photographic *) plates. Jii kw some cases it is possible to rind a source giving wideh separated hues in a desired wave length region. at the surface of the earth. This sharp cut-off is ascribed to the ozone present in the upper layers of the atmosphere. of the solar radiation of wave length 2900 A is not more than one -millionth of the intensity at 3150 A. This limit of the solar spectrum as recorded by^a photographic plate at various altitudes of from 50 to 4560 meters was found by one observer to be 2910 A. we are dealing with very low intensities. intensity of solar radiation and the possible effects of daylight on biological reactions. This is important since in biological radiations. it frequently occurs that but one line is transmitted. standard filters of the Corning Glass Works. though the absorption approaches asymptotically 100%.22 CHAPTER I Them* authors point out that a filter is seldom found in which the transmission changes from 50 per cent to 1 percent or less in 200 A". to transmit appreciably. .

Thermocouples left: a two-metal thermoelement. together with the absorption spectrum of oxygen. The Absorption of Ultraviolet (16. the photocell graphic plate. 10 b in .97 of This. counter.97 in Distilled Water mm) Absorption Wave Length l\ THE INTENSITY MEASUREMENT OF VrSFRLK ULTRA V OLET It A 1)1 ATION I AJSI) Arranged in order visible and of increasing sensitivity. which is proportional to the temperature difference of the two junctions. no experiments mm are carried below 1900 A.PHYSICS OF RADIATION 23 distilled water. the detectors of ultraviolet light are: the thermocouple. shows plainly why in biological radiations. right: a modern highsensitivity thermocouple. a current will flow in the circuit. Table 6. The Thermocouple: The composed of thermocouple consists of a fig. Table 6 gives the absorption of ultraviolet by 10. Figure 16. the photoelectric (1) and the photoelectric. circuit two dissimilar metals or alloys (sec U>a). Modern high-sensitivity If thermopiles are frequently built as diagrammed in fig. (J the junctions are maintained at different temperatures.

17 taken from C. vantage possessed by the thermopile* over the other means of intensity measurement is that its response is quite independent of the wave length of the radiation. the sensitivity is is low. 1931). perhaps 1 to 20 (see fig. 1 or '2 mm 2 in area. the other of bismuth-tin. K.E. Spectral sensitivity of Eastman plates cmvcs ( .MEES. emitted by a standard tungsten lam]). HALL (1926). paper rich in information on the characteristics of photographic A emulsions is that of L. The ad- 2. By using pieces of small dimensions. Characteristic curve of .24 CHAPTER I which A is a very light bit of thin gold loaf. Furthermore. it js often used to calibrate a source. of ultraviolet for instance. /*: Eastman 4O Eastman 35. Figure IS. the blackening of the emulsion for one wave length is proportional to the intensity only over a (2) The Photographic fest the defect just relatively short intensity range. (1. as is the heat loss by conduction : sensitive galvanometer. photographic pla tc. .soldered to the heavy leads.0 Log "'ijiiiiv Exposure 17. in terms of the energy in the form of visible light. D: Eastman Process. C.1 i: /: Eastman Speeds ay. the heat capacity of the instrument thus. high. IS). The gold leaf is supported by two tine wires. Photographic plates manimentioned. which are. namely that of giving a response which is not independent of wave length (see fig. A. or 1 - 10 2 ergs/cm 2/ . The disadvantage of the thermopile is its low sensitivity as compared to the other instruments for this type of measurement. JONES and V. Plate. For this reason.sec. such a thermopile will give a detectable deflection when the radiation falling upon the gold leaf has an intensity of approxi- Used with a mately IJyJO 10 cal/em 2 /sec. our of a bismuthantimony alloy.

the photoelectrons (see p. the SCHTMVNN plate. Bjffef>y lowest sensithity to which the photographic emulsion will respond is 12000 quanta /cm at a 2 /'see . cells is Photoelectric cells are of two kinds. lengths which easily About the /. Figure 19. different The sensitivity of the SCHUMANN plate for the ultraviolet depends upon the absence of gelatin. for example. They an covered with a very thin film of 1 a substance (a car boxy lie ester of dihydro-colloidin which lluoresees of furniture polish) strongly under the action of ultraviolet with the emission longer wave the penetrate* gelatin. This is many times greater than the linear portion the photographic emulsion curve which extends over an intensity range from I to 20. 2 .~>00 The Photoelectric ('ell. radiation falls upon a metal surface. roughly that of the photographic plate. 1 to ot however. even better plates have been obtained by sensitizing ordinary plates. current and intensity has been tested in properlyphotoelectric designed cells and found to hold over a range. 6alvanomefcr\ wave (3) length of 2. This coating is of the photoelectric clement. or 10 7 ergs/cm A.'sec. 10) form a photoelectric current which is proportional In ejected to the linear relationship between the intensity of the radiation. it is quite unsafe to assume a linear relation between photoelectric current and light intensity for any but The sensitivity of photoelectric specially-made cells. Gelatin has a strong absorption for wave lengths below 2800 A and becomes practically opaque even in very thin layers for wave lengths in in the ultraviolet. the neighborhood of 2000 A. vacuum and gas-filled. Na. Another wire is sealed in a side arm. With . A glass bulb (see inner surface a coating fig. 19) has deposited over most of its or Mg. Photoelectric < photoelectric cells. of intensities of from A 50 million. A . Recently. a battery main- tains the coating negative with respect to the wire at A. e. in contact with a wire which is sealed through tho glass wall of K the bulb. In principle they operate in the same way. Not all commercial cells give this behavior.PHYSICS OP RADIATION Plates have been evolved which are particularly sensitive to wave length regions. g. In fact.

such as oxygen. since these data Du BKIDUE). the "work function". Table 7 gives the photoelectric work functions. . The energy required to remove a photoelectron from a photoelectric surface is called it. In a photoelectric cell. ti is situation in the sensitive surface of a photoelectric cell further complicated because of the* impossibility of having this The surface consist of one kind of atom. W . If light is allowed to fall upon the metal surface. which is a measure of the photoelectric current. the galvanometer reads zero indicating that no current is flowing in the circuit. in such cells. hv. It is rare in this work to find the results of two The investigators coming within more than approximate agreement. The purest surfaces by distilling metals in a high vacuum. given. With the best high vacuum technique known today it is impossible to prevent contamination with various atoms. chiefly those of the gases prevalent in the air. and this may be transformed into a value in electron volts then called the photoelectric work function by the equation may may be regarded as representative. These electrons flowing through the wire cause the galvanometer to deflect. to Necessarily. (Further be found in Photoelectric Phenomena. The result is tluit 1F for a given metal depends considerably upon its history and the has been freed from gases.20 the CHAPTER I window covered so that no light can enter the cell. ir o for various metals as obtained by different investigators. photoelectrons are ejected from the atoms of the metal film and are attracted to the central wire. only a few metals being care with which it are prepared . of each quantum be measured must be greater than the energy required to surface. HUGHES and W= vv 12330 -^ i A in A ' first three columns show how the value of the long wave length limit or the work function depends upon the treatment given the surface. This equation applies to isolated atoms. that is remove an electron from the hv>Eoo E . the energy. hydrogen and nitrogen. The threshold wave length in A refers to the longest wave length which will eject photoelectrons from a given surface. the photoelcctrons must not only be removed from the atoms but must be shot away from the metal surface and eventually even through This requires a little more energy.

27 of the Metals Photoelectric Work Functions .PHYSICS OF RADIATION' Table 7.

The metal tube is connected to the negative terminal of a battery of perhaps 1000 or 1500 volts. such as is found in a radio receiver. the photoelectric-ally active element is deposited inside walls of a cylinder. . and the collector is a fine wire 4 Figure 20. on the Jn a counter. This momentary movement of charge is equivalent to a small current which when amplified 11 produces a "plunk in the loud speaker. In such an instrument. it will be accelerated toward the wire (which is positive by 1000 or 1500 volts) and in its course through the gas will ionize some of the atoms with which it collides. the counter gives a few counts per minute when no radiation from the source under experiment is falling upon it. and stretched along its axi. The number of "plunks" per second indicates the number of photoelectrons ejected per .s (sec with some gas to a pressure of about 10 cm of mercury. The electrons thus freed are also attracted toward the wire and in turn form more ions. Each time a photoeleetron is ejected from the walls of the tube. The lowest intensity which is measurable with a counter is about 10~ 9 ergs/cm 2 sec. These are due to cosmic radiation. ft and y rays from radiocative impurities in the metal of the wire and tube they are second. It is filled in the cut in the cylinder to let in the light.28 CHAPTER I The fourth column gives the best estimate that could be made for the various metals. (4) of the value of IF photoelectric tube merely a photoelectric cell of a special geometrical shape. of quanta While in use. making it much more sensitive than the photo cell in which photoelectron currents arc measured. the individual photoelectrons are recorded. The Photoelectric Counter. and the collecting wire to an amplifier. 20). Photoelectric tubi counter. . Slits may be insulated from the cylinder fig. In a very small fraction of a second all these negative* ions will reach the wire. or it may be allowed to shine ends of the cylinder. this number is proportional to the number striking the inner wall of the tube each second. local gamma radiation and a. The is counter or about 500 quanta /cm -/see.

es and shielded from the source of radiation. surface. Howewer. (). etc. "strays" or background radiation. is called the photoelectric t/icltl of the The photoelectric yield is the reciprocal of the effiof a surface. It for the photoelectric cell) only has been found that for the ordinary counter (as well as about 1 quantum in 10000 striking the walls ejects a photoelectron into the gas of the tube The number of quanta incident upon a surface* divided by the number ol photoelectrons ejected. At lion the counter is exposed alternately tor 5 ininiit. Fig. with which these counters are tilled A\J 11 reduce the sensitivity of the surface. o minute interval. 21 illustrates data taken with an aluminum counter by FRANK and RODIONOW (1931) to pro\e the radiation from chemical reactions and from tetani/ed muscle difference The dotted line line indicates the the total number of counts number of "strays" and the solid when the counter is exposed to The ordinal os are the number of impacts obtained per Finnic 21. .PHYSICS OF KAD1ATION called L>9 "dark counts". 91. some inert gas which a counter having a highly will not reduce the WERNER (1935) recommends 75%Ne and 25/ He. at. p. The source of radiation \\as O 7 -|. It should be possible to Jill sensitive surface with sensitivity. Yields of surfaces in photoelectric counters have never been as high as those in photoelectric cells because the active gases H.. A perfectly efficient surface would yield ciency one photoelectron for every incident quantum. More literature is quoted in Chapter IV. The between tlie number of counts when the counter is e\po<ed to and shielded from radiation is proportional to the intensity of the incident radiation. O the radiation. or the average number of quanta required to eject one photoelectron. a 5 minute exposure being alternated Avith 5 minutes of shielding. the experience of one of the authors shows that it is difficult to obtain sharply defined counts with this mixture.FeS<). the riuht a tctam/cd at the left the chemical reaction K 3 2 sartorius muscle of the frou.

yields of 100 or even 1000 timen these values Table arc 1 - considered to be good. S. Generally. Highest yields obtained with various surfaces at the maxima of thoir spectral distribution curve .30 CHAPTER I. PHYSICS OF RADIATION The values of Table H represent maximum yields obtained by experienced workers.

though the temperature remains within a few degrees of the. when an electric current is set up in a tube containing gas at low pressure. as in the slow oxidation of phosphorus or in the light of the firefly. However. is. the fact that all bodies emit at all times radiant energy they also absorb at all times radiant energy. 'Chough the chemical sources of radiation are more important because they show us that we may also be in expect radiations arc in much better known. and afterwards with those emitted by chemical reactions. from the oil lamp to the incandescent light arc based on this principle. PHYSICAL SOURCES Returning for the moment to the wave Thermal Radiation: theory of light. In this chapter. a division is made between physical and chemical sources of radiation. originate under widely varying condiWhen the temperature rises visible. may Warm bodies radiate heat. Too. biochemical processes. first. room. very high. the physical sources The? discussion begins therefore with the physical sources of rays. we remember that the oscillations of an electric charge result in the production of radiant energy. the tube will emit light. A. At ordinary . without great increases temperature. This radiation is due to the vibrations of the atoms (built of electric charges) of which the body is composed.CHAPTKR II SOURCES OF RADIANT ENERGY Radiant energy tions. There . in Chapter HI. the radiation becomes Practically all our sources of illumination. Lot us wee how this idea may be applied to the various sources of radiation with whieh we are familiar. the biological effects will also be first demonstrated with rays of physical origin. visible radiation may produced from chemical processes. followed by the chemical sources of such rays.

but solely upon its temperature. . e. It can be seen from these data that thermal radiators are poor sources of ultraviolet. Table 9. in a direction perpendicular to the surface. wave lengths Table 9 gives the energy radiated by a tungsten filament at various for the two temperatures 2500 and 3000 0. However. known as black body or thermal is The peculiarity of a thermal radiator that the distri- bution of energy among the wavelengths depends not at all upon the atoms or molecules of the body. and in some of the hotter stars to a blue-white eolor. from 1 om 2 of surface. that of the tungsten filament in an electric bulb. for 1 cm range of \vave lengths. Spectral Distribution of Thermal Radiation from Tungsten Wave Length in 1<\V (A) (watt/cm ! 3 ) 1 ) A 2500 (' 3000" (' 2000 2500 3000 3500 4000 4500 5000 5500 oOOO (5500 7000 Atomic Radiation: Everyone is familiar with the neon signs so frequently used at present for advertising purposes. per unit solid angle. g. whieh is being heated. These are nothing more than discharge tubes filled with neon or a mixture of neon and other gases.32 CHAPTER II temperatures. Radiation which arises as a result of of iron is the temperature of a body radiation. They are fitted with metal 1 ) Kw(A) is the rate of omission of energy in \\atts. At still higher temperatures. the color of the radiation is changed to nearly white. this radiation is of very long wave length. the radiation becomes visible when the temperature of the body reaeJies the witness the dull red color neighborhood of 500 C.

at the hydrogen-oxygen combination. shown that these are not By far more common is the emanation of ultraviolet light Recent investigations make it appear very probable that all chemical reactions emit part of their energy in the form of short ultraviolet rays. glow and discharge tubes depending upon the pressure of gas within them and the voltage necessary to make them function. c.SOURCES OF RADIANT ENERGY 33 terminals to which electrical potentials are applied. Haber has cases of light produced by heat." implies. or at the oxidation of pyrogallic acid. potentials high enough to cause ioiiization of the gas atoms in the tube. energy being liberated as visible light. as the name Occasionally. as will be shown later. Ordinarily. CHEMICAL SOURCES Most is of the chemical reactions i. namely the mercury arc and the hydrogen arc are of this type*. however. the reaction causes "exothermic. As examples may serve the light produced during the slow oxidation of phosphorus. 21). the energy emitted in form of heat. in water. are exothermic. This has been proven for such simple processes as JNaOH-l HC1 = NaCl+H 2 O. The latter two are strong sources of ultraviolet but suffer from the disadvantage that they are not sources of constant intensity. luminescence the 4 . However.e. The atoms lose and regain electrons many times a second with Such the emission of light each time an electron is regained. Other sources may depend simply upon the ioni/ation of air between two naked terminals. NaCl the radiations an very weak. but that part of the original energy of reaction is liberated in the form of visible light. 3 Protoplasma-Monographien IX: . such as the carbon arc or the iron or tungsten spark. The most practical sources of ultraviolet. and even for the solution of NaCl Na + +Cl~. which proceed spontaneously they liberate energy. at the reaction of potassium with water.it is the necessarily much different from room temperature electrical energy which causes the ionization of the atoms and the resultant emission of light. tubes are known variously as arc. i. altogether too weak Ordinarily. 1 to be registered by the photographic plate. B. In this type of source the temperature is not --. it has been possible to prove their existence by the GETGER-MULLKR counter which {see is essentially an extremely Rah n sensitive photoelectric cell fig.

vessels. Ultra -violet radiation from chemical reactions 1 a nionochromator. the number of photo-electrons was when exposed to radiation from proteolysis than without and in 7 experiments it was more than 3 times as large as the error. therefore it must be of a wave length shorter than 3500 A. Table 10. not from gla?ss containers. It was also observed at this time that in some of the reactions. This physical proof of light from chemical reactions has been ultra-violet radiation ficiently sensitive instrument. only the rays between 2000 and ) Radiation passed 2700 A were measured.34 Jn this way. trically also measured photo-electhe emission of ultra-violet light by several inorganic oxidations. and by pyrogallic acid in air. verified by GERLACH (1933) who showed that this radiation appears only from quart/. CHAPTER II number of FKAKK and RODIONOW (1932) proved that a common chemical oxidative reactions produced an 1 which could be demonstrated with a sufTable 10 gives their results. . In is all larger this. BABTH (1934) could also prove existence* of ultra-violet emission AUDTJBEKT and VAN DooKMAL (1933) tile known of from proteolysis which to give an immeasurably small heat of reaction. the emanation is greatly increased in the presence of diffuse day light. by the oxidation of alcohol with chromic acid. Recently. 1 . 25 experiments but one.

studied three in alkaline types of oxidation. C: agar blocks with yeast on the exposed side. The biological aspects of this acceleration will be dis- cussed in Chapter TV. The organisms most used in these experiments are yeasts the growth rate of which is accelerated by short ultraviolet rays under certain conditions. 22 shows the first attempt.SOURCES OF RADIANT ENERGY 35 tive counters will detect them. namely pyrogallic acid . solution It was by air. glucose -{. The original method consisted simply in placing before the eollimator slit of a quartz spectro- graph quartz tube with the reagents to bo & a tested.. milogonctic spectrum " 1 excitod muscle: B: tho quartz prism. and to substitute m . by KFUNK (1929) to obtain tho spectrum of a frog muscle. 3* . as far as can bo ascertained with this rather crude method. : T d^ "^ the yeast was permitted to grow for a short time order to bring out the growth rate differences. Each yeast block was thus exposed to a definite" range of the spectrum which could be determined fairly accurately.. and was then compared with the controls. photographic plate by a succession of tiuy blocks of nutrient agar on which yeast in the proper physiological condition was growing. each block receiving rays C knou-n wave length. this method. By blocks. in First attempt to obtain a Figure 22.KMnO4 and blood serum -| H C) a found that the growth was stimulated only on the two blocks None* receiving radiation from 22202280 and 2280 -2340 A of the other detector blocks differed appreciably from the controls. In fact. After the irradiation. These radiations are so very weak that only the most sensiHowever. Kig. living cells under cer- tain physiological conditions react very promptly upon irradiation in the wave length range 1800 2600 A. it would appear that all three oxidations gave the same spectrum.. KANNEC HESSE it (1931) working with yeast each representing approximately 50 A. From these results. they are so sensitive that it has been possible to use them in place of photographic plates for determining the spectra of such radiations.

The limit of error is + 15. Device for exposing yeast to successive ranges of the spectrum of f>0 A each. BBAUNSTETN and POTOZKY (1932) showed that the spectra of different cellophane. by means prepared a chamber of which corresponded exactly to the 50 A (fig. reproduced in Table 1 J The various spectra are not quite identical. By this simple instrument.36 CHAPTER The next advancement was the II division of the spectrum into separate strips of exactly 50 of glass needles and heavy A each. oxidations possessed certain specific regions besides the general The data of 7 separate experiments are oxidation spectrum. POTOZKY (1932). Induction Efiects obtained from 50 ANGSTKOM stups of the Spectra of various Oxidations Detector: Cell Numbers in liquid Yeast Cultures . 23) the sections divisions of her spectrograph. Table 11. and even the rather crude determination by 50 A strips shows . Figure 23. differences which remained typically constant when the experi- ments were repeated.

1()3). The position of this slit could be changed over the entire spectral range By this method. 110). There may be strip of 10 A. to divide the agar surface into such narrow strips and there was PONOMAREWA tilways the possibility of confusion p. 37 of diffuse daylight increases the intensity Cr 2 7 FeSO4 but does not affect 2 K j . the negative regions need not be examined. of course. .SOURCES OF RADIANT ENEKtiY It was also found that some reactions."> A 24). e. or. h'g. Thr spoctia of SOTTIC common biological i ('actions. g. Recently. PONOMAREWA could show that the glycolytie spectrum of blood consists of only 5 regions. and the amount of work is thus greatly reduced. only one small part could be studied at one time. On account of the very low intensity. Therefore. more precisely. In this way. and the progress of such analysis is slow. that only of the 60 spaces of 10 each manifested mitogcnctic effects (see . one narrow slit of 10 A. if the general spectrum has been investigated by the above-mentioned eoarser methods. the spectrum A still more detailed analysis was finally accomplished by It was impossible (1931) who used 10 A sections. PONOMAREWA by the spreading effect (see screened off all radiation except oxidation Hiiuar ( J'yrogallol) fermentation nucloase (phosphatasc) phosphate Heavauo protrolysis sum of all above rear! ions aniylnst 1 mallasc su erase re 24. DIOCKKR (1934) succeeded to to split the lines of 5 in a first double line of this spectrum into two different A each. However. of itself. this procedure is usually combined with intermittent radiation (see p. more than one spectral line.

and also with those of the lactic fermentation by Streptococci. They prepared phosphate from muscle. according to LUNDSGAARD. The spectral analysis was carried out by counting the total number of T 3 east cells (method. and another from the splitting of glycy Ugly cine by erepsin. GUKWITSCH (1932 a) and lysis. Since both enzymes split the phos- now phoric acid radical from the organic remainder. GURWITSCH concludes that to all there must be some process common giving off the same radiation. and its chemical decomof by means H 2 8O4 was the source of radiation. 74). but has . however. Aft<T some preliminary analysis by LYDIA GUKWITSCH (1931). The same lines have been found by GTTRWITSCH in the decomposition of lecithin by "lecithase" (unpublished. acid in phosphagen. (1932) attempted to analyze the a different type. acid by the pulp of adeno.carcinoma of a mouse has a spectrum decidely different from that of pro too it was determined by A. The final determination. The "nucleasc/' gives a very long wave length. in 10 The line 2000 2100 is strips. The two spectra proved to be exactly alike. BILLIU. This seems probable since it is As a consequence of these splendid findings.se phosphate through glyeeric aldehyde to methyl glyoxal is common to all three types of sugar decomposition. 1932). the method was number of frequently-occurring biological reactions. '24 together with that of glyeolysis and of an oxidation. phosphate cleavage. is also shown in fig. showed 8 with definite radiation. of organic. 72). A doubtful. generally assumed that the cleavage of the hexo. Two sets of data were obtained. and L. quoted from BRAUNSTETN and SEVKRTN. The authors assume that the source of radiation is the deamiiiisation of the amino-acid group (we also Table 23 p. The splitting of nucleic. This. and was considered negative by these authors. of these sugar decompositions.38 CHAPTER 11 The most important result. plays an important BRAUNSTKIN and SEVERJN of spectrum role in the ('a-creatm position energy for the working muscle. one with the digestion of serum albumin by the gastric juice of a dog. was the observation that these bands coincided exactly with those of the alcoholic fermentation by yeast. namely that of amino-groups coupled with phosphoric. the Russian school calls this the "phosphatase spectrum". K ANNECLIESSEK and SOLOWJ u\v (1932) produced the detailed used for a proteolytic spectrum. sec p.

SOURCES OP RADIANT ENERGY
subsequently been assumed as positive
in all

Russian publications.

The mitogenetic spectrum of the working muscle (FRANK, 1929) contained some linos which at the time could not be accounted for. and which now can bo explained by this phosphate cleavage.
These are the detailed spectra of simple chemical reactions published at present, as far as we have been able to ascertain. As oxidation spectra, the two double, lines of pyrogallol oxidation
1

have been inserted; these
Figure 25.
block tor ylasrt exposing bacterial cultures to the various wave lengths of the spectrum, to be nsod
quartz plate.* in front, taking the place of the photo^rapliv plate in the
\\ith
rt{)t>ctro,ur;i]>li.

arc 1

commonly used by

the Russian

Grooved

.Kig. 24 shows that only rarely is the Kven then, it produced by two different processes. should be realized that we are not dealing with true spectral lines, but with relatively broad regions, and that two identical strips do not necessarily indicate two identical lines, but rather two

workers for this purpose.
line

same

(or

more) proximate lines. There is also a spectrum shown which

is

the

sum

of all these

processes.
of

We

shall see later (p. 156) that the spectra, of nerves
all

frequently combine

of these lines, in addition to

some others

unknown

origin.

of the action of amylase and maltase, and of suerase (invertase) have been obtained by KLENITZKV and PKO KOFI EWA (1934). The lines of these two enzymes agree to a. much

The spectra

This

larger degree than any of the previously-mentioned processes. is to be expected from their chemical parallelism.

Another method of obtaining spectra, is that of WOLFF and (19,32) who used bacteria as detectors. They made a number of vertical grooves in a glass block which fitted into the camera of the spectrograph (see fig. 25) the grooves were covered by a quartz plate so that they became tiny pockets into which the

RAS

;

detector culture was placed for exposure. each one could be determined accurately.

The wave length

for

40

CHAPTER
By
this procedure,

Jl

they found the spectrum of neutralization

of acid

and

alkali to consist of three lines

a strong line between 1960 and 1090 a strong line between 2260 and 2300
Fig. 26 shows

A A

a weaker line between 2070 and 2090 A.
these* regions, and also the spectra obtained by RAS (quoted from RT'VSSEN, 1933) for the Bunseiiburner flame. This latter spectrum lias also been photographed, and had many more lines of longer wave length, some of which are shown

m^ISlMllBEii
I.

en 1*11 11SC

LI; spectrum

Fiyurc 26. Photographic and mito^ciietie spectra: burner flame, milo^enetic spectrum; //. same, photographic 111. Reaction tT 2 -|-(M 2 mito^enetic spectrum; IV, Reaction Na()H + H(H, mitogenetic Kpectrum.
,

here, but

The

figure also

photography failed entirely at the shorter ultraviolet. shows the biologically obtained spectrum of

the reaction ot hydrogen with chlorine. ft should not be gathered from this discussion that the
indicated lines represent the entire spectrum of these reactions. On the contrary, it is most probable that it extends to both sides
of the

narrow range shown here.
Radiation below 1900

We

arc limited, however, by our

indicators.

by

air (sec p. 23),

and that
;

A will be readily above 2600 A does not

absorbed even

produce mito-

therefore they can not be observed by the methods employed here. They may be capable of producing other biological effects hitherto unaccounted for, and may play an important part in chemical reactions.

genetic effects (see fig. 28)

A

good summary

of

the Russian studies of mitogenetic

spectra, including those of inorganic reactions, and with sufficient data to obtain a conception of the error of the method, has been

given by A. and L. GI T RWTTSCH (1934).

The
is

intensity of ultraviolet radiation

not proportional to the total energy liberated.

from a chemical reaction This was to be

SOURCES OF RADIANT KNE1KJY
1

41

expected because the same holds true for the emission of visible Strong mitogenetie effects can be light from chemical reactions. obtained from protein digestion by pepsin or from milk coagulation by rennet while RITBNEK found the heat of reaction of proteolysis to be immeasurably small. On the other hand, the heat of neutralization of aeid by alkali is so large that it can be observed even without a thermometer, yet
its

radiation
is

is

relatively weak.

by no means clear. LOKUNK'S criticism (1934) can be explained in a concrete example as follows: If a spectrum represents the radiant energy emitted by the reaction as such, then each reacting molecule must emit at least one quantum of the shortest wave length observed in the spectrum. The longer wave lengths might originate from the shorter ones by loss of definite amounts of energy. Thus, in the case of sucrose hydrolysis by invertasc (figure 24) each sucrose molecule must emit at least one quantum of the wave length 2020 A, i.e. of the
origin of the spectra

The

energy content 9.S

10~ 12 ergs (see Table 1). Then* are (>.l 6 1 X 10 23 in Ig of sucrose molecules in a grammoleeule, or --:<
t

'

10 2:i

The

k

minimal amount
then be
:

of radiant energy

from

1

g of sucrose would

H.I > 10 23
-

-

X

9.8 < 10~ 12
-

ergs

0.175

>

10 11 ergs
-

-- 0.042 , 10* cal

420

calories.

total energy liberated by this hydrolysis has been measured by RuiiNUJi (1913) by means of a BJUCKMANX thermometer in silverlined Dewar bottles, and was found to be 9.7 calories per

The

gram. This experimental value is only one-fiftieth of the minimum amount calculated. It seems impossible that appreciable amounts of energy could have been lost by the method used. We are compelled to the following alternative: Either, the spectra do not
originate from the reactions for which they are considered specific, but from some quantitatively unimportant side reaction; or, the

calculated
again.

amounts are actually

liberated, but arc at once absorbed

This latter explanation does not appear entirely impossible. biologist is familiar with "false equilibria", such as the stability of sugar in the presence of air, though it could be oxidized

The

with liberation of

much

be expected to take place spontaneously.

energy, and the process might, therefore, Modern chemistry
1

42

CHAPTER

II

explains this by the necessity of ''activation" of the molecule to make it react chemically. This activation requires energy. FRICKE (1934) in an introductory summary, makes the following general
estimates:

"Generally, energies of activation are of the order of 100,000

gram

gram molecule which equals 1.5x10 gram At ordinary temperature, the average per molecule. kinetic energy of a molecule is of the order of 10~ 21 gram calorics.
calories per calories
of 10~ 43 of Ihe molecules

19

Only the inconceivably small fraction have energies in excess of 10~ 10 gram

calories per molecule.

"The quantum theory gives as the reason that activation may be produced by radiation, the fact that the energy of the radiation is carried in a concentrated form, as quanta. The energy
of a
is

the

quantum of radiation is 5x 10~ 16 /A gram calories where "k wave length in the ANUSTKOM unit. The wave length has

to be reduced to the order of 3 ,000 A. before the quantum has the value L.5>, K)~ 19 gram calories which, as we saw, represents the usual value for the energy of activation per molecule."

The energy of activation necessary for the hydrolysis of sucrose has not been determined. If wo assume it to be of the
"usual value" as computed by FRICKTC it would bo equivalent to quantum of about 3,000 A. wave length per molecule. When this is absorbed, the snciose molecule hydrolyscs, and the amount
a
7 calories per gram. The energy will be absorbed at once by a neighboring sucrose molecule which becomes activated, and hydroly/ed, and thus the
I).

of energy thus released must be larger of the additional heat of reaction of

than that absorbed, because

free

In this way, all the liberated energy is again absorbed, except for the difference between heat of hydrolysis and heat of activation which we measure as heat of reaction.
reaction goes on.

However, then? is another small "leak". Of those molecules adjacent to the walls of the vessel, the energy may radiate into the wall rather than to another sucrose molecule, and these few quanta would leave the system, and produce a radiation if the
vessel is transparent. The amount of energy thus leaving the vessel would be extremely small, and would be of a wave length

equal or shorter than that required for activation.
If this

explanation of the mitogenetic spectra

is

correct,

they

would be an excellent means to measure the energy

of activation.

the suspension reacted normally again. it irradiated with the lines 3220. 47) which is then radiating with its own This would account for the difference in wave lengths as well as for the increase in intensity. Other examples have been given by WOLFF and KAS (193.SOURCES OF RADIANT ENERCV C. A very short action of living bacteriaThese suffices to change the broth to a "secondary sender". observed further that after long exposure to primary radiation. or systems. ease. but showed it when bacteria had grown in it. . At one end. however.of the r through a monochromator. A of the trough. 24). but is a property of certain chemical solutions. and L. the induced light may secondary radiation has a And most remarkable of be stronger in intensity than the primary 4 HOU1VC. but the wave length of this "secondary" radiation w as not the same as that. nor is it a case of fluorescence. These authors observed the same effect with sterile blood serum. which is the spectrum of nucleic acid hydrolysis (see fig. spectrum. This proves that it can not be merely a reflection of light. these liquids ceased to produce secondary radiation. The OU'it \MTscjr best explanation is most probably the one given by that the primary radiation induces some kind of chain reaction (see p.* SECONDARY RADIATION be recorded here which was first A phenomenon must believed to be typioal of living organisms. namely the emission of rays from 11 a solution as a response to "primary rays directed upon it. The first purely chemical effect of this kind was observed by A. on the In another next day.3240 from a copper arc.'U>). 45 minutes ex4 posure of a staphylococcus supscnsion to a strong primary sender had made it unfit to produce secondary rays. radiation of the nucleic acid could be observed. Very intense primary light caused a more rapid exhaustion. even after the bacteria themselves had been removed by filtration through a porcelain filter. WOLFF and HAS filtrates as such emit no primary radiation. A 3% solution of nucleic acid was was gelatinized in a glass trough. even 30 minutes exposure was sufficient to destroy the power of secondary radiation. because the wave length changed. 4. GfKwiTscit (1932 b) with nucleic acid. They found also that sterile nutrient broth did not produce secondary radiation. wave length than the primary. At the other end primary: it was between 2450 and 2500 A. for the shorter all.

. Intensity of Secondary Radiation of Nucleic Acid. to accelerate the The numbers in the table represent the growth of bacteria. i. which produced a mitogenetic effect at least 5 times as strong as the 1% solution. The lowest efficient concentration was 0. Bacteiial suspensions and their filtrates lose the power of secondary radiation upon heating. The more dilute they are. the secondary radiation becomes weaker.e. they do not transmit initogenetic rays. nucleic acid solutions do not. required to produce a "mitogenetie effect".44 CHAPTER II Nucleic acid solutions also cease to function when overexposed.solutions. c. measured by the time required to produce a "' " in 1 ogc ne e e f f e c t 1 1 1 The results are somewhat surprising. Perhaps this is brought about by the absorjtfi-ion overexposed solutions (see above). Table 12 represents an experiment with nucleic acid. by WOLFF and RAS. An important observation is that with increasing concentration of the acting substances. A similar relation was observed with bacterial suspensions. the stronger is the secondary radiation they emit upon excitation by some primary source. percentage increase in cells over the control. source of primary radiation was the reaction The intensity of secondary radiation from the nucleic acid dilutions was measured by the length of exposure. . and during this stage.02%. The increase in intensity by secondary radiation enabled of rays in WOLFF and RAS to construct an "amplifier" for mitogenetic rays. Strong solutions recover again after 1 or 2 days of "rest". it produced the same effect in one-fifth the time. Table 12. i. The of milk with rennet.

WOI/FF and KAS radiation is (li)34a) could show that proved polarized. and (2). statements really indicate an increase in intensity of radiation of the word. does not give a greatly increased intensity. obtained 1 a good mitogeiietic effect from the opposite side in 10 seconds. This means an amplification of 27 times. i. and they also usually. e. proteins." (1). The observation is added that one continuous column of the same length as all <> cuvettes together. 24) that the reciprocity law (requiring double exposure time for half the intensity) does not even hold for such simple reactions as those in the photographic plate when the intensity becomes very low. In his in the physical sense most recent summary. by the time required to produce a mitogeiietic? effect. secondary that polarized mito- geiietic rays exert a very much stronger effect Tt is not at all certain. . the same primary source used for direct irradiation of the same detector required 4. that the above upon organisms. the radiation is resonant. Its application to biological reactions is quite doubtful. urea.sorncKs OF RADIANT KNKRCY 4r> sion side They placed six quartz cuvettes filled with staphylococcus suspenby side. it travels from the irradiated part through the liquid The only published experimental proof for this is that by BE KOROSI (11)34) as far as the authors have been able to ascertain. such as glucose. medium to distances of several centimeters with the measurable speed of a few meters per second. the substrate reacts mostly upon those wave lengths which it emits when decomposed enzymatically. fats etc.5 minutes. therefore. : <*rK\\iTscii (1934) make>* the following statement "It has been found that all substrates capable of enzymatic cleavage. It must be kept in mind that the intensity of radiation lias been ascertained only biologically. and by irradiating one side of the series. react also upon mitogenetic radiation and become radiant. It has already been pointed out (p. This "secondary radiation" has certain properties of great interest . nucleic acid. Recently.

which is called a chain 1 reaction. by means of radiant energy from the sun. The ion pair tlms produced initiate being absorbed by a molecule. . however. TAYLOR. S. Just as a chemical synthesis may be brought about by heat. will produce chemical changes. If only one molecule becomes activated by an energy quantum of the right size. p. however. 1931. that radiant energy be. which term photochemical reactions. ionizes it. several hundred. with chlorophyl as the necessary ''transformer" of the energy. This type of reaction. These reactions are exothermic. or even several million molecules are changed for each energy quantum which is absorbed. All organic matter be induced lay adding is thus produced from CO 2 1J 2 O and nitrates.UHAFTKR III EFFECT OF ULTRAVIOLET RADIATIONS UPON CELLS A. the energy liberated by its reaction activate s another molecule. is usually explained in the following 101 8): in way (_H. The classical example by BOBENSTETN and NJJJKNST is the photochemical reaction between the gases H 2 and C1 2 The chain reaction can be written . 1009 The quantum. reactions which again produce ions. EFFECT OF RADIATIONS UPON CHEMICAL REACTIONS known that energy in the form of visible Tt has long been light. we or near this range. two independent. number of chemical reactions are known which require some radiant A energy to be initiated. Thus. absorbed by more than one molecule. many molecules may be changed though it is quite impossible' that ono quantum can b. . present to increase the energy level of each reacting molecule. so may it radiant enei'gy in the form of light. It is not necessary.

If there Here 100 times as much of the foreign substance. the chain length would be reduced to ten thousand molecules. 1 1 The same reasoning holds true for ehain reactions in solutions. In were not for those terminations.HOI + H H + 01 01 -I 2 -. Whenever a chain . The chain is terminated by the reactive molecules or atoms combining with each oilier. or stable compounds.EFFECT OF ULTRAVIOLET RADIATIONS I'l'OX CULLS 47 01 +H 2 2 HOI -j- H (11 Cl -}- H2 -- HOI | H Cl H + C1 01 HC1 f _= H 01 i OL Ho - HOI | +H 2 KOI |- H 01 + . as indicated in the above model. In the photochemical oxidation of Na 2 SO 3 to Na 2 S04 the quantum yield was about 100000. that would account for an average? chain length of one million molecules. this is practically the result. HOI HOI ! H h 01 3 = HOI + 01 H 2 -f H H ! 01 HOI. Under "chain" is understood the number of consecutive molecules entering into reaction. where the reaction liberates a large amount of energy.. but not by tertiary alcohols. sufficient to cause all molecules to fact. in the case of explosions. If it by reacting with other molecules io form one. The presence of foreign substances reacting with the components of the system is a common cause of cessation. quantum would be read with one another. . This reaction was inhibited by primary and secondary. If there were only one such molecule present for every million molecules of hydrogen and chlorine. The chain may be as "long" as one million molecules The "length" can be ascertained by determining the number of molecules changed for each quantum of light entering the system.

STAIR and H QUITE (1932) on erythema. This is the ease in normally nourished cells of animals. might produce a very noticeable effect in a living cell. which liberate energy constantly by meta- bolizing carbohydrates. 3000 A. fats. The known chain reactions are exothermic. just as the Cl 2 -molecule needed only one quan- tum of the right size to common. they synthetize new body substance endothermically. two molecules of the alcohols were oxidized to the corresponding aldehydes and ketones. prodigiosum . bacteria. by RIVERS and GATES (1928) on vaccine virus and fltaphylococcus aureus and by DUGGAR and HOLLAENBEK (1934) on Bact. It is not impossible. EFFECT OF MONOCHROMATIC ULTRAVIOLET UPON LIVING CELLS any of in the Ultraviolet light of mentioned effect previous chapters the different physical sources may have* a very distinct upon living cells. significant because the Ultraviolet light also kills bacteria and other microorganisms. that they be so if another source of energy available. This fact is major part of this book is concerned with mitogeiietic rays which are shorter than 2(500 A. By means of this energy. however.48 CHAPTER III was terminated. The most the only reaction in the coll which can be . etc. itlooks almost like* the reverse. Jt does not seem is imperative. i. however. Figure 27 shows the results by COBLENTZ. B. The best-known is the reddening of the by ultraviolet light. cell division this results in a more rapid multipli- cation. or an increased growth rate. Between these two maxima is a zone of very weak effects.. they grow. the cells of the skin will also react upon those below 2600 A which are not found in sunlight. and perhaps thus released is start the reaction with hydrogen. Strangely. oven a single quantum. that very small amounts of energy. A quantitative study of the relation between the intensity of the effect and the wave length has revealed that aside from the very marked erythema produced by waveskin lengths around. fungi. the intensity curve for the different wavelengths is quite different from that of the erythema effect. proteins or other organic. This could be accomplished by releasing a complex mechanism which needs only one or several quanta of a given size to be initiated. e. compounds.

The Comparative intensities of the killing effect ol different wave intensities are uniform for each individual organism. but vary greatly for the different curves. The result of the most extensive of the fig. and especially if the intensity is very greatly decreased. it is possible Figure 27. Protoplasma-Monographien IX: Rahn 4 . many 28. lengths. to obtain growtli stimulation under certain conditions which will be specified in Chapter IV. wave lengths and + indicates accelerated growth. If shorter wave lengths are taken into consideration. o means no effect. The effect of ultraviolet light of different different intensities upon yeast.EFFECT OF I'LTRA VIOLET RADIATIONS UPON CELLS 49 and the mosaic virus of tobacco. experiments of this nature is shown graphically in 10-* 10- Figure 28. Most of these investigations have been carried out no further than to a wave length of about 2500 A.

No effect is noticeable in . it could be shown that only radiation of less than 2700 A produced positive effects. wave C. The same authors found that even the dissolution of NaCl water produces a growth-stimulating radiation (Table 14). i. is dissolved in water. the number of cells has been increased approximately terial One 40-50%. or when palmitic acid is dissolved WOLFF and HAS of dissociation of salt concluded therefore that the process into ions is the source of ultraviolet. Each culture thus obtained light of one definite wave length. underneath which was the bacterial culture. A large number of similar experiments have been carried out by GITBTWTTSCH and his associates. to irradiate yeast cultures. increased the growth rate of yeast. employing a monoehromator. when sugar in alcohol. from the sparks of aluminum. absorption by quartz. zinc or cadmium. By varying the intensity as well as the wave length. usually as controls for They will be mentioned in the succeeding organic radiations. There seems to be very sities of different little difference in the limiting inten- lengths. . After exposure. Table 13 shows very distinctly in both experiments that an exposure of approximately 5 minutes to the radiation of the neutralization process has stimulated the growth. FRANK and KANNECUESSER (1930) used individual spectral lines. flowing from two tubes. microorganisms to the of the simplest examples is the stimulation of the bacgrowth rate by the emanations from the neutralization of NaOH with HOI. EFFECT OF RADIATION FROM CHEMICAL REACTIONS UPON LIVING CELLS effect The same result of irradiation with ultraviolet of which has been demonstrated above as the known wave lengths. Below 1900 A. chapters. these cultures were incubated for 2 hours. can of be produced also by exposing the cells emanation from chemical reactions. good effects can be obtained.50 CHAPTER III CHARITON. to unite on a quartz plate. e. This energy emission occurs only during the act of dissolving it ceases completely when all the salt is in solution. Other experiments have shown that even at 1900 A. water and air interferes with the experiment. WOLFF and HAS (1933 b) allowed these two chemicals.

having been exposed to ultraviolet light from the oxidation process. FERGUSON. one of quartz and one of glass. coll. the other received no stimulus since glass Table 14. Staphylocoeci exposed through quartz to the energy emanation from dissolving substances 4* . Simple oxidation processes also emit energy which can cause an increase in the growth rate of bacteria or yeasts. This is already cited in the method of obtaining oxidation spectra.Na('l f-H 2 O and counted 2 hours later In the same way. of the same culture of Bdtfcrrntu. 51 Staphylococci exposed through quartz to the energy emanations of the reaction NaOH-(-HCl -. bacterial growth was accelerated by exposure to metallic zinc in a solution of lead acetate or copper sulphate. and maj be further illustrated by an unpublished experiment of Miss A. each containing a sample. acid was oxidized with permanganate in a glass vessel. Above this were fastened two small covered dishes. J. Oxalic. The sample in the quartz vessel grew more rapidly.EFFECT OF ULTRAVIOLET RADIATIONS UPON CELLS Table 13.

NSCHINA (1029). Since all organisms display processes liberating energy. . exposed through quartz 149 Immediately after exposure 1 hour later 2 149 154 140 253 216 1735 3 940 3335 4 9085 What lysis correct for the holds true for the simpler chemical reactions.52 OH AFTER III absorbs the radiation (Table 15). ex- Development of a culture of Bacterium eoli after posure to emanations from the reaction C 2 4 H 2 + KMnO 4 =. Jnerease in the development of yeast cultures after exposure to the emanation from proteolytic processes Proteolytic profess Pereentual increase of exposed culture over control Egg white with pepsin. 31. .5% 20. - lfi. containing the data obtained by KARPASS and LA.5 All other enzymic processes which have been tested so far have yielded positive growth stimulation. Of 12 experiments. 20.1%. but after the bacteria once start to grow. it is only logical to assume that all living somewhat by the consideration that these organisms radiate.4% . There is the usual lag period of 2 hours. This statement must be modified ultraviolet rays are .1 %.6 H 2 O ('ells per ec. is also more complicated biochemical processes. of culture exposed through glass . 25. 10. the irradiated culture grows Table 15.8%. 36.4 %. only one was negative. 15.0 egg yolk with pancrcatin fibrin with gastric juiee 30. egg white with pancreatm egg yolk with pepsin . Protco- by enzymes yields an ultraviolet emanation which greatly stimulates the growth of yeast as seen in Table 16.K 2 ('O 3 + 2 MnO + 9 (<O 2 H.6 % %. 37. more rapidly. Table 16.7%.

. will not pass the skiu man or animals. Since we must expect ultraviolet radiations from very many biochemical processes. will be discussed in Chapters IV and VII. Which parts of the various animals and plants Attention radiate. for example. and.EFFECT OF ULTRAVIOLET RADIATIONS UPON CELLS 53 very readily absorbed. and since they may stimulate cell division they may play an extremely important role in the development of all living beings. of should be called here only to the fact that there may be radiations and growth stimulation inside of an organ or tissue without becoming noticeable outside this focus.

is organisms. 1 a) The onion root method his associates The root of an onion. the methods radiations will be discussed. and the division of certain cells in the animal body. to verify its existence by physical measurements. in 1923. and in the larvae of sea urchins. The detector root was placed in a narrow glass tube to permit . th* development of eggs. however. it increases the number of mitoses. In used in detecting and proving such By far the most extensive treatment and the given to mitogeiietic radiation because it is the most studied best understood. Its effect upon living organisms is very conspicuous. The Beta-radiation of living as well as dead organisms is only mentioned in passing. It accelerates the growth of yeasts and bacteria. A. as long as they have any noticeable metabolism. this chapter. The iiecrobiotic rays and' the injurious human radiations are. or mitosis. In onion roots. MITOGENETIC RADIATION This type of ultraviolet rays was discovered by GUKWITSCH He called them mitogeiietic because he observed that they stimulated cell division. This radiation is so weak that it was not possible for a long time. perhaps. It may cause morphological changes in yeasts and bacteria. was the first detector of the rays. The presentation in this chapter is largely historical. used by GTJBWITSCH and extensively from 1923 to 1928.CHAPTER IV METHODS OF OBSERVING BIOLOGICAL RADIATIONS The preceding chapters have shown that for physico-chemical we should expect ultraviolet radiations from ah living 1 reasons. Occasional exceptions to this arrangement could not be avoided. only special manifestations of mitogenetic radiation.

a few the tip. the number roots were of dividing nuclei was ascertained. tho detector root \vas fixed and stained. In the first experiments (1923). also placed in a glass tube so that it could be directed to point exactly at the growing tissue of the first mm root (fig. 29). J: fresh Totals Ji. GuRWiTsm found that the exposed to the biological radiation showed regularly more dividing nuclei than the opposite side. the . The Figure 29. Two to three hours after the beginning of irradiation. Tablu 17.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 55 from easy handling. in microtome sections.source of radiation was another onion root. 1925). The onion arrangement of roots in the first the experiments on milogenetie rays. Gi RWITSCII coined the word "nritogenelie" rays. same h ratal to (J0(l. and . For this reason. left in this position for one or two hours. The radiation of onion base pulp The numbers given are those of dividing nuclei in corresponding microtome sections of the exposed and the uncxposed side* of the deteetor root. and. was uncovered allowing it to be irradiated. Table 17 gives side of tho root the results of a test with the crushed base of an onion (A. The zone of meristematic growth. Totals ! Difference jUl|-3l l|-lL_l3 ll ()! 2 -l| 5 ! *j 4 4 .

MAUROF) of annelids Eggs Eggs of sea urchins before the 1st division (FRANK and SALKI\I>) be -fore the 2nd and 3rd divisions (SALKINP) Egg yolk (SoiiiN). 24 hours old (ANNA Giwwrrscif) Young tadpole heads. The following list contains compiled by GUBWITSCH (1929). GUBWITSCH. SIKBHRT.oxygen Corneal epithelium of starving rats. of chicken. SORIN) Blood of man (SIEBEBT.: tails. ( J i u- Neoplasms: carcinoma. pulp of tad})ole heads (ANJKTN. young plumulae of Hihnn- (FRANK and SALKIND) Potato tubers: leptom fascicles only (KISLIAK-STATKEWITSCH) Onion roots connected with the bulb (GuRwrrscn) Onion base pulp (A. Staphyloeoeci (MAGBOU) JJaciUvti antHiracoidcif. gills. sarcoma GUBWITSCH. KEITEB and GABOB) Spleen of young frogs ( GUBWITSCH) Bone marrow (SIEBERT) Bone marrow and lymph glands of young rats (SUSSMANOWITSCH) ( Resorbed tissue.56 CHAPTER JV This technique was used by Gi'Bwrrscu and a number of associates to search for mitogenetic rays in the entire organic world. Radiating organisms and tissues Bacteria Hdclcrium tuwcfacicns. and GABOK) Blood of frog and rat (GuitwiTscn.lactic acid -\. and L. radiation of hi the ceases Embryos tkiis amphibia morn la stage (ANIKIN) Plant seedlings: root tips. intestine of amphibian larvae during metamorphosis (BLACKER. cotyledons. but not of normal rats \VITSCH) (L. FKANK) Pulp from resting muscle |. BBOMLEY) . POTOZKV and Ctontracting muscle (SiEBERT. GESENIUS. the more important earlier observations. only during the first two days of incubation After establishment of circulation system. RIOITER and OABOII) Turnip pulp. SIEBERT. Bacterium murimorx (Acz) (SEWKRTZOWA*) Sarcina flara (BARON) Streptococcus fastis (MAGBOU) Yeast (BARON. It would scarcely be worthwhile to compile a complete list of organisms found to radiate.

METHODS OF OBSERVIXC lilOMUIICAL RADIATIONS 57 Regenerating tissue of salamander and angleworm (BL \CUKK. Considering the great claims which Gurwitseh and his associates made for their discovery. and many Russian workers. and considered the results of the Russian workers Among and of REITER and (TABOR to later. TAYLOR and HARVEY (1932). WARMER (1927). In order to decide whether the positive results could be considered experimental errors. in this be experimental errors. long Chicken embryo . POTO/KY and ZOGLIXA) Blood of asphyxiated frogs Blood of cancer patients Blood of starving rats (becomes radiant with glucose) (AM KIN.TEFF) Hydra: hypostom and budding /one. by MAcuioir (1927). SCHWEMMLE (1929) undertook statistical investigation. it was surprising that comparatively few biologists were sufficiently interested to repeat the experiments. not other parts. The onion root method. POTOZKY and ZOCJTJNA) Active tissues with chloral hydrate Active tissues with KCN Further details will be given in Chapter VIF. the radiant nature of this effect was thus fortified beyond doubt (see p. SAMARA. these. The most (192S). in several different laboratories. physical nature of the phenomenon. BORODIN (1930). greatest early support to the establishment of mitogcnetic rays was given through the extensive and thorough work These two authors tested the of RETTER and GABOR (192S). Non-radiating organisms and tissues Tissues of adult animals except brain.after '2 days (blood radiates) Blood serum (becomes radiant with oxyhemoglobin) (SniiiN) (or with traces of H 2 O 2 ) (ANIKIN. some obtained negative results so consistently that they denied the existence of mitogcnetic rays altogether. and verified especially the. a He concentrated graphically all results published by GimwiTsoH and the Russian school (about 200) . and recently by PAUL (1933). ROSSMAN* country. Positive results were obtained Loos (1930). 59). blood and acting muscle (most tissues have later been found to radiate slightly) Tadpoles over 2 cm. frequently quoted of these are and much SCHWA KZ (1928).

30). From the "not induced" roots. The error of the experiments by WACJNER (1927). as the total number increased (fig. the mitoses of two different of sides of the roots varied greatly. number of mitoses counted. and a few experiments supposed by these authors to prove induced mitogenetic effect are really within the limits ^ error. All these data gave doubtful or negative results. The error was Jj 10% when 500 mitoses were counted. REITKR and GABON'S experiments have a much greater error. and decreased. Figure 30. Abscissa: total Circles indicate a positive induction. "induced" and "not induced" roots. SciiWEMMUt. . The lino gives the limits of error. 20%. All results with onion root as detector by UuitwrrscH and Ordinate: percentage of increase over control.CHAPTER IV by plotting the percentage increase or decrease of mitoses of the exposed over the unexposed side of the root. of course. did not consider it proved that the effect is caused by mitogenetic rays. All results associates. and TAYLOR and HARVEY'S few experiments (1931) indicate a similar large error. he could compute the probable error of the method. 8rHWAiiz (J928). and ROSSMANN (1928 29) is even larger than that of KEITEK and GABOH'S. because of the possibility of other physiological factors affecting mitosis during the? experiments. against the total He distinguished only between number of mitoses counted. which had been claimed to prove mitogenetic radiation were found outside the limits of error. black dots indicate no mitogenetic effect.

by irradiating roots with known wave lengths of the spectrum. In fact. in pretation. MAUUOU). and partly through very thin glass. and it is very interesting that two distinctly different wave lengths have been claimed. SIEBERT. Besides a sharp maximum at 3400 A. and in trying to verify his prediction. of common glass. nor through gelatin even in effect in The mitogenetic very this thin. Then. and the only question was its inter- That under different physiological conditions. but not through thick layers of glass. no mitogenetic effect was observed. thin cellophane (STEMPELL). and concluded that he was dealing with an ultra-violet radiation of about 2200 A. (it'KAMTSric could obtain the effect through quartz. but not through very thin gelatin. PKANK and Gi/Jtwrrsen exposed onion roots to different lengths from physical sources. such as glass slides. It seems hardly possible to account for all of ultra-violet rays. Below this. and of water (GuKVvrrsc'ii. they found the range to Inbetween 3200 and 3500 A. they determined the mitogenetic efficiency of this part of the spectrum. of and Joiia glass. through thin animal or vegetable membranes. found onion roots as the h'rst reliable indicators. layers. both based upon apparently reliable by any agent other than A t data. thin plates of mica (UNITE it and GABOK). and obtained effects only from the spectrum between 1990 and mitogenetic wave 2370 A. a straight line and is proceeds reflected from glass and from a mercury surface (GrKWiTscii. negative results have been obtained by some investigators is not really surprising GTJRWITSCH had always claimed that the effect of one root upon the other was caused by rays. another smaller maximum was discovered near 2800 A. ft will pass through thin layers of quart/. however. and still noticeably through 5mm. of the wave length of these vital is that very question ra \s. must be considered established by a very large number of data. he predicted in 1922 radiation as a factor in mitosis. not even in the neighborhood of 2000 A which was considered by . They found this radiation to be transmitted through 8mm. different countries with different onions. By means of special filters. REITEJI and GXBOK). also through gelatin which indicates a wave length above 4 3000 A. Quite different were the results of RUITEK and GAIJOR (192H).METHODS OF OBSERVING BIOLOGICAL RADIATIONS The 59 effect as such.

of the older parts of a root will produce. 31).GO CHAPTER TV and FKANK as the only efficient region. All three experiments gave an increase in. periments (fig. 35). The results have already been shown in fig. letting the spectrum from roots or sarcoma tissue fall upon the length of an onion root. The first experiment was FRANK'S spectral analysis (1929) way (see The publication of with a spectrograph Tn three well-agreeing exusing yeast as detector (see p. experiments can now be explained by an error was not known at that time that irradiation technique. The very interesting observation was made that the apparently inert spectrum between the two maxima will prevent mitogenetic effects by the active wave lengths. no variation of intensity produced any effect. 27 p. 2400 A was emitted. REITKR and GABOR'S experiments caused the . mitoses at. FKANK and KANNEOTESSER (1930) of the effect of monochromatic light from physical sources upon yeast. namely the meristem near the root tip. be discarded. Then followed the detailed study by CUARI- TON. This last argument in favor of It is a wavelength near 3400 A must therefore. The rays outside of the maxima were entirely neutral. it could be shown that no radiation above of the radiation of the tetanizcd muscle. but at the only reactive part. REITER and GABOR determined further the wave length of mitogenetic rays by means of a spectrograph. Beyond 2600 A.the place where the wave lengths between 3200 and 3500 A had fallen on the root. even when the intensity of the "antagonistic 11 rays is only one-tenth of that of the wave lengths between 2900 and 3200 A show this inhibition.a mitogenetic effect not These last in It at the place of irradiation. Direct sunlight and ultra-violet arc light also inhibited mitogenetic rays. and none of the German authors' results could be verified. as has already been shown in Chapters TT and satisfactory 111. the deviating experiences of REITER and GABOR have never been accounted for in a really GURWITSOH. considered definitely esta Wished now that mitogenetic rays range between 1SOO and 2600 A. Special efforts were made to . However. All mitogenetic effects. 28. 49). 1929). because they contradicted all their own statements about the wavelength. The curve resembles somewhat that for erythema (fig.Russian workers to repeat them at once.

Figure 31. tinwave lengths of various radiations have been shown to be quite different. Irradiation of Onion Roots \\ith Monochromatic Spectral Light . Since the width of the agar blocks was not uniform. By these and other methods with a variety of indicators. Results of 3 experiments on tJu* spectrum of muscle radiation. but all of them wen* below 2ft(M) A No records are Table 18. witli the technique shown in figure 22. but show the general spectrum. 4 the shorter wave lengths were found to be efficient. and those in the region of 3400 A gave 110 ell'ect (see Table IS). detectors. it yielded only consistently negative Considering that RJCITEK and UABOK used onion roots as Again. but results. the results overlap partly. the experiments were repeated with onion roots.METHODS OF OBSERVING BIOLOGICAL RADIATIONS investigate the range around 3-400 01 A claimed to be so efficient by REITER and OABOR.

but rather. in a dark . the onion root as detector has been substituted by the time-saving yeast methods or by bacterial detectors. (4) Omission by GURWITSCH of the microtome sections showing a decreased number of mitoses when they happen to come between. sections showing an increased number. in a completely covered inoist chamber. but all after repeated removal. this author obtained also negative results. absorption by quartz and air in the spectrograph makes aecurate determinations impossible. 1932) observed that roots when removed from the water show symmetrical distribution of mitoses. pressing the opposite effect occurs. This is no proof that they do not exist. and after long continued and pressure were carefully avoided. With yeast and blood as senders. friction When MOTSSEJKWA denies the existence of mitogenetic effects in onion roots and explains GUIIWITSCH'S consistent results by several assumptions: (1) One-sided pressure of curved roots in the glass tube. Most of them do not mention the still more careful work by MAKUARETE PAUL (1933). and confirmed by WOLFF and KAK (p.62 CHAPTER IV given of wave lengths below 1900 A. It can hardly be doubted from the Lirge amount of speetra analyzed by the Russian school. Since 1928. and they arrive at quite different conclusions. PAUL fastened the small onions (hazelnut size) by means of gauze to perforated cork stoppers. no increase* of mitoses was observed upon exposure to another onion root. that mitogeiictio radiation consists essentially of the wave lengths between 1900 and 2500 A. and of less uniform roots when no effect is expected. 40). MOTSHEJEWA (1931. This careful study has been considered by many critics to be the final proof against mitogcnetic radiation. which then leads to negative results. Both papers describe the technique employed very carefully. Pressing or rubbing of the roots will increase the number of mitoses. that below this point. However. these were held above the ground by simple stands. (3) Selection of good roots for important experiments which results hi increased mitosis through pressure. they do not. two extensive recent investigations must be mentioned which concern the question whether onion roots can be used at as detectors. of roots (2) Light applied repeatedly in centralization which causes phototropic curving of the root and results in increased mitosis. Realizing the prompt reaction of roots to touching.

his first experiments. and theii exposed it to the radiating source for definite short periods. the yeast was spread on glass slides. the starting point for all future work with this type of detector. When steel. Almost always. the sender root was substituted by a needle of stainless the exposed root also turned downwards. dried and stained. The yeast was then incubated for from 1 to 2 hours to permit the radiation effect to develop. this percentage was higher in the irradiated culture than in unexposed controls. and it will be.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 63 room at 20 C.5 cm. but the microscopic analysis showed no increase in mitosis at the exposed side. and a very careful investigation showed that the number of mitoses on the exposed part of the root was distinctly larger than on the opposite half. usually not over 30 minutes. When the roots were 1 2. one could be exposed to a root from another onion without being cut or even touched and without the need of glass tube holders. When. . sender roots -were taken as controls. The original method has since been changed in some details by BAIION (1930) and GUBWITSCH (1932) (see p. and they showed a uniform and symmetrical distribution of mitoses. it was considered a proof of a raitogenetic effect. The measure was the percentage of yeast cells showing buds. the roots developed in the* air through the muslin in two to five days. b) The yeast bud method BAIION (1920) suggested that the rate of bud formation of yeast could be used as indicator of mitogenetie radiation. he spread yeast over the surface of solidified nutrient agar containing glucose. 66). After this. while the leaves grew through the hole in the cork. The paper verifies GURWITHOII'S principal experiment. The onions were never placed in water. the exposed root grew more rapidly than the other roots of the same onion. The object of PAIR'S investigation was the establishment of a good method for studying mitogenetie rays with onion roots. However. whether all mitotic The stages were included or only the more conspicuous ones. The number of examples given is not large enough to draw many more conclusions. the symmetrical distribution was disturbed. allowed it to grow for from 9 to 15 hours at room temperature. The exposed roots turn downwards. however. In. long and absolutely straight.

is 9. in an oxygen Finally. or excited muscle to radiate strongly while the resting. sender and detector were J separated by a quartz plate. 24%. results as "induction' effect. IV Table Effect of Yeast. in air When .li buds. expressed in pereeiits of the control value. of Sarcoma. In all his experiments. chemically the pulp of resting muscle to that of working muscle.5%. __ 100 (exposed control control) SIEUEKT (1928b) used this method for a number of interesting studies in physiology. quiet muscle did not do He attempted to produce radiation by changing this (Table 20). Table 19 gives some of his experiments. i. the increase when the exposed yeast shows 33% buds. he obtained positive results by using a very dilute CuSO 4 solution as oxygen catalyst. He observed the working.5% over the control. Tt is now customary to record. Moreover.. he succeeded by placing the acidified pulp atmosphere. and this is an increase of 37. since the addition of lactic acid alone would not produce radiation. and of Hone upon the Budding Intensity of Yeast Percentage of Buds in Yeast Marrow | The most extensive early data with this method have been by SIEBBRT (1928a).<>4 CHAPTER 19.e. as increase in buds of the exposed yeast over the control. The induction effect is 37. Thus. and the control. The numbers indicate the percentages of yeast cells published wit.

173). but blood and urine wont parallel. Anemia. and that there was a good parallelism between blood and urine radiation. ftiEBERT later (1930) concentrated his attention upon blood radiation. The experiments on the metamorphosis of amphibia and and on the healing of wounds in animals (p. With syphilis.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 20. The exceptions were distinctly. his associates. radiation varied. the majority showed no radiation of blood or urine. Thus started the first experiments about chemical relictions as the source of radiant energy (p. He verified the statement of LYDFA (lirjtwiTSOH and &ALKIND (1929) that blood of normal. patients after recent treatment with X-rays and isarnine blue. further. 65 Effect of Electrically Excited and of Hosting Frog Muscle upon the Budding Intensity of Yeast he finally found that JQQQTT KCN solution would prevent radiation. healthy people radiated He found. were all carried out by the yeast BLACKER and by insects (p. radiated. luucemia. Of 35 patients with cancer. 153). while that of cancer patients did not. 167). scarlatina) prevented radiation of blood. high fever (sepsis. pneumonia. Protoplasma-Monoffraphien IX: Rahn 5 . ho concluded that the source of radiation must be chemical. that urine. as well as of urine. 33). None of the other diseases tested caused loss of blood radiation (details see p.

gives the following directions for the yeast bud (1932. and are no more capable of developing spontaneously the maximal energy for development. The temperature is probably room temperature. drying and staining them. the liquid is distributed evenly over the agar Method: GntwiTscH method surface by careful tilting." The method of making smears to count the buds is not given in GTKWITSCII'S book in any detail. This method is very commonly used in mitogenetic investigations at the present time. This method has been varied by other authors. CUTCWITSCH recommends that the person counting the buds should not know which slide or experiment he has under the microscope. dition of the detector plate. it (j and should bo kept in mind here that on p. TUTHJLI^ and RAHX (1933) studied the mode of bud formation of Burgundy .. though beer and wine yeasts are occasionally menas room temperature. All conditions are quite precisely standardJng. tioned. delicate film of yeast. Milano.. It consists simply in smearing the yeast cells on a glass slide. manufacturers of the "hemoradiometer" of Protti's. nor 12 Neither the temperature nor the concentration oi the yeast suspension its ago is mentioned. The importance of the yeast agar blocks in the establishment of biological spectra lias already been mentioned on p.. and this may account for the good results of Italian in- vestigators. which was found in the lowest layers . 317: "The most appropriate stage of the detector plate corresponds to a thickness of the yeast growth of about 25 to 30 layers of cells (p. in order to give a better conception of the proper physiological conwe quote from the same hook of GimwiTsr. 35. even smaller buds. ized. After about 5 to 6 hours. . Some authors limit far layers. and remains so until about the twelfth hour. . \Vocan be certain that the lowest layers of cells which arc in immediate contact . Only those buds are counted which arc smaller than half of the full-grown cell. TKRZANO & C. It can hardly be with the nutrient . the surface is covered with a fine. and the surplus liquid is drawn off with a pipette. their counts to 1 Attention should be called here to a leaflet published by G. and is now sensitive. medium consist essentially of young cells in rapid multipliThe cells of the middle layers arc not in optimal condition. this prevents subconscious arbitrary decisions.66 CHAPTER IV bud method.n\y 1 p. . Gunwrrsoii mentions for the yeast The directions are probably meant primarily Xadsonia fulvesecus which has been used most commonly by the Russian workers. 7): Beer wort agar plates are flooded with a very fine suspension of yeast in beer wort. p. 31 H). cation . they arc not real resting forms as yet. 14. from the truth to deny any appreciable multiplication in the topmost However.

32. This detector is really quite different from BARON'S or GUKWITSCH'S described above. depends upon the With the Burgundy age of the culture used for the seeding. and new buds are not formed at once. The typical result including all sizes culture immediately after being transferred contains but very few buds. i.e. at this latter stage. 69). become quite ready for budding. could not increase the perp. or to incubate for several hours after exposure if exposure had taken place immediately after seeding. and then incubated for one hour (so that the mitogenetie itself such a detector elleet might manifest by an increase in buds) we should have bud formation beginning on the exposed plate Figure 32. All yeast cells are at the same stage readily. it was advisable to incubate the plate for about 2 hours before exposure (woe Table 21). inoculation with yeast from agar surface cultures is recommended. It is also seen that they soon reach a maximum percentage of buds. . In liquid cultures. ]f. of 67 A buds is shown in fig. the rate of cell division were accelerated it by mitogenetie rays. veast employed by TimnLL and liAHN. a plate seeded with a 24 hours old culture was a good detector immediately after seeding When a (> days old culture was used. The old yeast ceils which had ceased to multiply require some lime before their reproductive mechanism is working normally. the time interval between the seeding of the plate and the first active bud formation. commonly called the lag phase.METHODS OF OBSERVING BIOLOGICAL RADIATIONS yeast on raisin agar at 30 C. The "lag period". the old yeast cells retained their buds for many days while old cultures on agar surface lost them Since a low initial percentage of buds is very desirable. buds in while the control has not yet The development of an agar surface culture of wine yeast. During this rejuvenation process. exposed for 30 minutes. centage of buds (see also The most opportune time for using such a plate as detector is evidently about an hour or two before If bud formation is begins. cells" respond most promptly to mitogenetie rays.

It is quite permissable to count all buds because the percentage at the beginning is very low.5 hrs Percentage of Buds 24 hours .5 hrs: 2 hrs [2. This should make the counting easier. inactive cells to "dilute'' the counts. 1 Culture ! hr 1. The "mitogeiictic effect" is much greater than in the other method because there are no old. 6 days . . and the cells are far enough apart not to influence each other. In fact. Method by The yeast is TUTIIILL and KAHN (designed for Burgundy yeast): kept throughout the experiment at 30 C. KH PO4 and added. 2 or seeded raisins is is heated off. this type of detector is so different from the BAKON type that it may react differently in certain experiments. A 24 hours old culture in raisin extract 1 ) is flooded over a solidified..Age of Parent ( 1 I Age of culture when exposed hrs 10. both types are good. with very weak radiations. with 1 liter of water in steam for 45 minutes. As long as they are used merely as detectors to prove the existence of radiations.5.N type is more sensitive because the old cells act as "amplifiers" (see p. the resulting medium is Raisin agar: Melted 6% water agar is mixed with an equal volume of the above raisin extract and sterilized by heating for 20 minutes at yeast extract (or meat extract) are sterilized at 100 (\ The pH is about 4 to t. and a limitation in size is not necessary. On the other hand. 5 g. sterile raisin agar l ) Raisin extract: 1 pound of chopped.68 Table 21. CHAPTER IV Kffeut of Irradiating Yeast Surface Cultures after Incubation for Different Times : Length of Exposure 30 minutes Incubation after Exposure: 30 minutes . 127). of earliest rejuvenation when exposed. pressed made up to 5g. these are probably the largest mitogeiietie effects ever recorded. the extract 1 liter..5 hrs. the BARO. Probably.

In a growing culture. Let us. Here.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 69 plate. One to two hours incubation. The organisms art* killed by placing a cotton wad with tincture of iodine in the Petri dish. e. Soon after that. radiation from these growing cells may be reflected by the glass or quartz walls. i. to produce two cells of the same developmental stage. counted from the beginning of the exposure. by RICHARDS and TAYLOK (1932). the surplus liquid is drained off at once. was the most suitable time to bring out the differences. the agar becomes hydrolyzed under foils to solidify. On pressure. it may be advisable to explain this important point in more detail. and the culture permitted to develop lor 24 hours. but mostly to an error in tho cross section. the same is true with prolonged heating . The percentage of buds is without. the percentage of buds cannot be changed by a change in the growth rate. 4 with buds and one first approximation of a ''cross section" through a yeast population growing at a constant rate. This fluctuation is due partly to the arbitrary selection of 5 stages. LKJ uid cultures have also been used successfully. This surface growth is washed off with 5 cc. too. Protection against reflection is advisable in all such cases (see p. but fluctuates between 67 and 80%. water.e. and the slightly-stained is observed in situ. Half of the plate should be shaded to servo as control. for this discussion. When all cells are out of tho lag period. SO). the suspension dilution. there must 100 C. a coverglass can be placed on the agar surface. the surplus liquid is poured off. eliminating all possibility of breaking oil buds glass. Fig. and may thus produce radiation effects in controls as w*! as 1 m the exposed cultures. and produce buds at a constant rate. that increases in the bud percentage can be expected only during the lag phase (see fig. and with this agar are Hooded. and account of the high acidity. is then diluted 1 : 100 with sterile some sterile solidified plates of raisin The length of exposure will depend upon the intensity of the sender: 30 minutes proved a good time with young yeast cultures. the rule applies. yeast by smearing on Tn all methods where growing cells in glass or quartz containers are used as detectors. at 100. Since this has been overlooked by some experimenters. and the plates should be exposed within half an hour. the yeast cells are so far apart that the buds can be counted directly on the agar surface. 32).g. distinguish five equal periods in the complete coll division of the yeast. It requires 5 time units for each cell to complete the cycle. In these plates. of sterile water. 33 shows a not constant.

they have kept their de12 hours at 25 C while BARON used room tector cultures for 9 . and our present conceptions of the mechanism of cell division do not contradict it. However. p.4 to 66. If niitogenotic radiation should speed up the very first stage so much that the "tiny" buds never appeared. Whether it grows rapidly or slowly. and this lower level would continue as long as radiation accelerated the one particular stage. or from 74. whether the time unit is 30 to 00 minutes (at cellar temperatures) or 10 minutes (under optimal conditions). This would offer a good explanation for the "false mitogenetic depression" of (rUBWiTScn (1932. A change of the growth rate would not affect the bud percentage at all.70 necessarily be CHAPTER TV more cells of the young stages than of the old. all (jells of the first 5 units as a more appropriate cross we obtain the following picture: at a Yeast Culture Time units I Constant Growth Rate There is a fluctuation of only about 1/ . This percentage depends only upon the variety of yeast used. 210) and especially of SALKTND (1933) where the percentage of buds decreases while the total cell count increases under the stimulation of mitogenetic radiation. the percentage of buds remains 74 75%. the number of buds at the period would drop from 26" out of 35 to 18 out of 27. The reliability of the BAKIXN method has been doubted by NAKAI DZUMI and HCHKEIBKK (1931) who claimed to have followed BARON'S method explicitly.8%. It is rather probable that only a certain stage of the cell cycle is affected by mite-genetic rays. a change would become noticeable if only one certain stage of the cycle should be accelerated. Many observations suggest this. However. If we take flection.

and the differences between control and poisoned catalyst give the error which is computed here in two different ways: and The increases resulting from irradiation are very much larger than. The following computation is made from a study of the effect of upon organic catalysts (19281) KCN 1929). from the data on experiments without mitogenctic effect. Tin's can be done very easily. The claim of these authors that the error of the method has never been considered by the workers is entirely wrong.Russia. knows the error of his methods. A schematic representation of the bud formation of n yeast culture growing at a constant growth rate. . Every biologist .METHODS OF OBSERVING BTOUMJICAL RADIATIONS 71 temperature which may be as low as 13 C in . at least approx- The Russian investigators have repeatedly stated the imately error of their method. can be seen from the fact that in most of their experiments. however. the error. though they have not given all the data necessary for others to check their computation. That their cultures were far too old. Time Units 12 J V 5 6 /Uumber of full-grown cells Number of Percentage cells wfti buds 5 V # cf cells wrth buds 80% 67% Figure 33. regardless by what method it is computed. SiiflBERT e.5 to <S hours. g. the percentage of buds decreased distinctly 1 in 2. rarely publishes less than 10 experiments to prove an yone point.

34 p. It is the same as that used for counting bacteria except that more preferable media are beer wort agar or raisin agar. they shall bo treated separately in this chapter on Methods. and again after three hours' incubation. For this reason. to the radiation of blood diluted \\itli MgS()4 solution to prevent coagulation. An counting by the addition of H a SO4 example of the results and of the method of calculation is given in Table 22. by measuring by comparing the turbidity by means of a nephelometer. The plate count method has rarely been used. or by hemacytometer count. because it onion roots. the respiration. 1 . The hemacytometer has been used by the Russian workers as well as by HEINKMANN (1932) in his studies on blood radiation. better than any other of the* methods described in this book. they differ in the method of measuring since yeasts arc so much larger than bacteria. 116). The exposure lasts 5 minutes. There is a difference so fundamental that in a number of experiments. g. 0. All cultures are preserved for The control is counted twice. and it is made discontinuous by moving some object between blood and yeast at intervals of 2 seconds (see p. and may be influenced by secondary effects. the preceding pages were written. the buds of yeast ascertains cell divisions directly while the mitoses in cells. an equal amount of a 4% . (1) Yeasts: The number of yeast cells in a liquid may be ascertained by plate all the volume of count. Table. etc. HEINEMYNN'S method for testing the radiation of blood consists essentially in thr exposure of a 12-houi culture of beer yeast (temperature not mentioned) in liquid beer wort. This method is used with yeasts as well as with bacteria. The total number of yeast cells (counting each smallest bud as an individual) is determined with a hemacytometer at the start. it may seem that there should be no essential difference between the total To the number of cells or the percentage of buds as a measure of t hegrowth rate.5 ee. are indirect measurements of growth. of the exposed yeast suspension is then mixed with an equal amount of beer wort and incubated for three hours at 24 C. The number of cells is the best measure of the growth rate. While they are identical in principle. the yeast bud method indicates a decrease in the growth rate while the total cell count shows an increase (e. cells. both in the control and in the exposed culture.72 c) Detection CHAPTER IV cell by increase in number investigator not familiar with yeasts. 103).

0.2 ec. also giving quicker results than the plating method.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 22. and incubated for 4 hours at 2s C. an will be shown in Chapter VI 1. To avoid lag phase. BUAINKSS (quoted from Ontwrrscu. 73 Yeast Cells Counted in Homaeytometer after Irradiation by Blood be/ore and HEIJSJKMANJN[ emphasizes t-liat this method depends upon the physiological condition of the yeast. he tested each blood sample with two different yeast strains The results by this method verified all former experiences with blood radiation. are suitable as detectors. measured by means of a micropipette. period. and a definite} amount of the exposed culture is is added to 1 ee. i.control grows too rapidly. only cultures ]5 (1932) used beer yeast in a 20 hours old (temperature not given) which arc actively fermenting. and the method requires less time and less eye strain than the hernacytometer method. Volumetric Method: KALUNDAROFF strong wort (18 22 Balling). 1932. of fresh wort. in othor words. and the curves and data published by this author show so little deviation only because they are However. when the control increases to more than double* during the incubation. e. that the yeast must he in . He also added somo important new facts regarding radiation of the blood of old people and of patients with chronic tonsilitis.*J hours. This means. After exposure. g. the volume is sufficiently accurate to prove mitogenetic radiation. 17) has adapted the method for the small vnhimina available in 1 mitogenetic work. The accuracy is not greater than with plate counts. e. the yeast is distributed evenly. else it would double in less than errors from this source. This The . especially in regard to cancer patients. The measurement of the growth rate of yeast by cell volume has been studied in detail by LITAS (1924). and that no effect can be expected when the. p. presented in logarithms instead of actual numbers.

2 for cc. The yeast column of the exposed sample is compared uith that of the control. More complicated is the differential photoelectric. . nephelometer described by (U'RWTTsriT (1932. 17). This method lias been used occasionbacteriologists for several decades. arc centrifuged in pij>ettes measuring the volume of blood corpuscles (the illustrations of the Russian workers appear to be VAN ALLEN hematocrit commonly used tubes). in A growth still is more rapid method for estimating the amount the measurement of the turbidity of the culture of by means ally by of a nephelometer. Height of yeast column of centrifuged yeast cultures. and of their controls yeast cells are killed l>y adding O. KANJVEUIESSEK and HOLOWJEFF (1032) who used the determination of the spectrum of gastric digestion. while the cell volume of bacteria is too small to be measured with sufficient accuracy in the earlier stages of growth. 'Fable 23 shows some results obtained with tins method by it BILLH. Attention may be called to the description of a simple nephelo meter by RICHARDS and JAHN (1933). exposed to the various spectral regions of the radiation produced by gastrie digestion of serum albumin. The nephelometer can be used for bacteria as well as for yeasts.74 CHAPTER IV Table 23. the methods are reviewed and analyzed by STRAUSS (1929).. p. of 20% ir 2( SO4 and .

. was the (2) Bacteria: mitogeiietic radiation i . by the same species. the agar plate count water. either of the same species. greater. Table 24 gives some of the results obtained by the latter. BARON and Ars did not recommend the growth rate of bacteria as a universal indicator for mitogeiietic radiation. (Jells through Quartz Time start per on bio millimeter ontrol irradialed Effect Another way of estimating the amount of growth in \east cultures has been suggested by BARON (1930) who compared the This method has been size of yeast colonies in hanging drops. or of yeast.Distance. slightly modified by BORODIN ( 934) who photographed the colonies and measured their area with a planimeter. Bacillus mescntericus 75 Irradiated Continuously by Yeast at 12 mm. they took their samples with a WKKJHT pipette which . This These was done most successfully by WOLFF and K\s (1931) authors worked with different species.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 24. and the number of cells was determined by the customary method of bacteriological Instead of making dilutions in technique. who irradiated liquid cultures of Kacillns murvmors with agar cultures. 1 The stimulation of bacterial growth by had already been observed by BARON (192(>) and by SKWTCRTZOWA in 1929. The data were verified by Ars (1931). and found that the e effect by "miito-induetion".

if broth is diluted with 9 parts of water. FERGUSON and KAHN in tlie (1933) obtained good results with the ^ i cc. A most interesting observation was the. even then part of the bacteria \\ere shaded. of Bacterium coli in a quartz dish I a layer of (U> mm.76 delivers - CHAPTER ^oU IV of a cc. exhaustion of bacteria good mitogenetic effect. Hy capillary pipette. and with increasing intensity. rapid growth. 1933a). the lag becomes shorter and shorter. During during lag phase.6mm.) can be. The two experiments in Table 25 show a lag period of about 2 hours in the control Irradiation decreases this period very distinctly. and at 5 hours.. (1931) allowed that a layer of standard nutrient broth 0. there was no effect. After exposure. indicating a return to the normal growth rate (fig. only very small amounts of the culture (about 1 cc. The plating of such minute quantities is necessary because. or decreasing distance. However. while a culture in broth diluted showed no increase 10 showed a 1 : WOI/FF and HAS pointed out that only the definite results could be obtained. or brought into "slide cells" according to WREGHT. ATetliod (the most recent method by WOLFF and HAS. covered with quartz. is placed in a glass dish in a very thin layer. to milk \. a layer of 1 mm. FwimrsoN and KAHJS of a standard broth culture in 1 cc. A fresh suspension of staphylococci in broth. irradiated from below over the control. pipettes used standard (Breed) method for the mieroseopie count of bacteria in milk. thick transmitted only rays above 2500 A.rennet in a quart/ tube). g. continued irra- diation after the lag phase retards the growth for some time. exposed. still transmits some rays as low as 2200 A. This retardation of growth is only temporary most of the irradiated cultures . almost doubled their number during the 5th hour. the dish with the bacteria is incubated at 37 (' for 1/5 to 30 minutes. by continued irradiation. 011 account of the strong absorption of ultra-violet WOLFF and RAS light by the customary bacteriological media. means of a . than any of the cultures whose growth was distinctly stimulated. WOLFF and HAS irradiated their bacteria in standard broth in a layer of 0. resulting in a decreased growth rate.5mm. with about 20 000 cells per cc. and exposed (e. samples of the exposed culture and control are either plated on ugar.. so is the control. using the slide cell method. 34). the control shows more cells per cc. (1933) verified this observation.

of Ntapliylocoerits nitrcim. or eventually decrease of growth rate (see p. 115). Developpemcait of two liquid staphylococcus cultures of which one was exposed continuously to the radiation from a staphylococcus agar plate. 77 The Effect of Different Intensities of Coiitinous Radiation through Quartz upon the Rate of Crowlli of NiaphyIOMCCHS aweus Sender: Agar surface culture.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 25. consider over-exposure the most common cause of over-exposure either produces no effect at all. . failure. ItSMO WOLFF and HAS Figure 34. at various distances The umrradiated controls never show oro\\th in this shojt time A\hile the irradiated cells do.

this culture had a growth rate higher than the average of 27% the two controls. This computation implies that multiplication of bacteria is arithmetical. The best medium was 1 part standard broth plus 9 parts water. while in truth it is exponential. sity of the source. 133.8 3. diameter. or in this 1 cc. 64. 24 hours old cultures never reacted. The growth rate is usually substituted the average by the generation time.9% of the total count. this has been done by SKWKLITZOWA (Table 24) and Arz.4. 136). is only 27. face culture (37(-) as sender. cultures 4K hours old still older always responded.1 7. . i. However.5 and 171. the time required for cell to double. dilute exposure 1 10000 with dilute broth. Method: FEUGUSON and RAHN a good mitogenetic effect with (1933) studied the best conditions for cult. of cells at the beginning. we have a really reliable measure.6. while the effect in tho same culture. As a matter of fact.e. Tho simplest procedure is to irradiate 1 ee. bacterium and found that it depended primarily upon the age of the culture. The errors are between 3. the : ^ ^ i and 28%. four detailed counts being 73. it must be considered that the number 183 has no real biological where d is the number t. and plate at least every 2 hours for 6 to <S hours.3. 1 after the time 1 significance while the other indicates that during the four hours. : from abo\c). By computing the growth rates. the best results were obtained with 15 to 30 minutes of irradiation 3.4 and 4. The effect may not become apparent iff he cell concentration is too high (over JOOOOO per cc. It is computed from the formula t OT =zr X -- log 2 log b log a and b the number Both methods have been applied in Table 2(>.78 CHAPTER The error of the IV method is mentioned in 1934. of an old culture in dilute broth (either in a quartz dish of 4 5 em. see Table 41 p. after a quart /-covered glass dish.1 3.3. (see Computation of the Induction Effect: The in- duction effect in these bacterial cultures cau be computed in the same manner as explained on p.2 4. 1)1. incubate. computed from the generation time. and no rules can be given. and tho transmission of ultraviolet by the medium. or through the bottom. at the start of incubation. The induction effect calculated from the numbers directly is in one cast (]5 minutes exposure) 183. the increase by irradiation amounted to 25% series are given. The time of exposure depends upon the intenWith a 4 hours old agar surTable 20).

the error becomes very large if the increase is small. It could be used with yeasts as well.EIIJER does not give the actual numbers of from which he calculated the generation times. cells Though HCHK. after diluting the irradiated cultures 10 000 with broth) : Induction Effect 37 168 I83| 111 Generation Times. With Nadxonia. This means that not all cells had divided in this time. In the tables given by iScuitEJBKR (1933).5 14 ! 41.5 I Generation Times Induction Effect 45. the Russian scientists.8 f-27 This procedure has also been used in Table. Thus. for the time interval from 47. The "Induction definite Kffect" as usually calculated (p. 1 . the especially many investigators. 64) has no meaning. with the probable It is very unfortunate.6 . SCHKKIBEII found the variations tiaee/Miromycefi cMipxoirfcuti commonly in duplicate plates of to read) -40%. With such a large error. However. and occasion- ally more. 24.METHODS OF OBSERVING BIOLOGICAL RADIATIONS . and in one case even 103%. in Table 22. it seems from his curves that they were computed from less than 100 cells. 79 3 days old culture of Bacterium cult irradiated for Table 26. the generation time of yeast for the first 2 hours when the mitogenetic effect is strongest is mostly more than 2 hours. e. therefore. It permits no comparison. . the deviation of duplicates went as high as 237%. that error of the method. g. minutes. various lengths of time by an agar surface culture of the same bacterium (The numbers are cells 1 per cc. This makes the error very large and a comparison of the growth rates must necessarily result in enormous percentual differences.5 40. there is little hope of detecting mitogenetic effects. 2 --6 hours 52. record.

the control and also for the same culture before the beginning of the experiment. The critics point out very justly that such relative numbers are not convincing. observed that the radiation of the detector culture may be reflected.) were given for the exposed culture. and is not always present. 45). It would add a great deal to the general recognition of biological radiation if the actual data obtained (numbers of cells. . a few others have been employed occasionally. percentage of buds etc. d) Detection by cell division in larger organisms The three detectors mentioned above are the only ones that have been commonly used to prove the existence of mitogenetic radiation. This reflection may be the cause of effect many failures to observe mitogenetic rays. germination was more rapid than when the cover consisted of black paper or sterile agar. WOLFF and HAS (1933 c) mention that they react slowly and require about ten times as long an exposure as staphylococcus cultures. but must be guarded against in this technique and probably in most others. The effect varies in magnitude. When the dish was covered with a glass cover or a quartz plafc.80 CHAPTER IV obtained effects merely by giving the "Induction Effect". and may thus give a mitogenetic effect even in the controls which received no radiation from outside. in some unpublished experiments. of mitoses. Mention is made of the reaction of mold spores upon mitoThe first publication of actual effects is probably genetic rays. that by SCHOUTEN (1933). The strong can be explained by polarisation of the rays through reflection (see p. but not often. Spores of Aspergillus niger were spread on an agar surface. FERGUSON and RASN.

rate with which they divide. Table 27. Too long an exposure retarded the de- The wave length is unusual as in all publications by REITEK and GABOB (see p.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 81 In the animal kingdom. SALKTND. Tissue cultures would appear to be an interesting subject for the study of this radiation. Not all species are equally well adapted as detectors. of sea urchins were found to be quite good detectors. BEITER and GABOB (1928) showed that frog eggs when irradiated with the spectral line 3340 A developped more rapidly into tadpoles than the controls. Table 27 shows Mitogenetic Effect upon the Eggs of the Sea Urchin Paracentrotus lividus The morphological changes of the larvae brought about by irradiation of sea urchin eggs will be discussed later (p. In a short paper. 164) since a different principle is involved. having been exposed to the radiation from bacterial cultures (see also p. 60). The first investigation was started without the knowledge of GUBWITSCH'S discovery. others. The some being much more sensitive than some data by ZIBPOLO (1930). can be easily seen under the microscope. the eggs of the smaller animals have been used occasionally to demonstrate the mitogenotic effect. 144). Protoplasma-Monographien IX: Rahn 6 . POTOZKX" The eggs The and ZoGLfNA (1930) were the first to show that biological radiation from growing yeast or contracting muscle will increase the rate of development of the eggs. Italian school of mitogcnetioists has also used sea urchins repeatedly. and the percentage of eggs in each of the different stages is a good indication of the growth rate. WOLFF and RAS (1934b) showed that eggs of the fruit fly Drosophila melanogaster hatch more rapidly after velopment.

Daily growth increments of three cultures of chicken embryo. the effect of the glass was found to be that of shading.82 CHAPTER IV GUILLEEY (1928) observed during some experiments on the tissue cultures. When a glass slide was used to separate the cultures. with embryo extract. This can be explained only as radiant energy which stimulates growth. and inserting small glass strips between some of them. acts essentially as a source of radiation. Fig. but were changed when all throo were continued in the same dish. While the most rapidly-growing culture usually maintained its growth rate. grown separately for 8 days. It could also be shown that the radiation was reflected from metal mirrors. the effect sometimes continued. 35 shows the relative daily increase of three cultures from the heart of chicken embryos. However. These observations suggested to GUILLEBY the possibility embryo extract which is necessary for the growth of He tissue cultures. that two or three growth -promoting agents for cultures in the same dish influenced one another. irradiated a number of cultures in a dish. but the result was negative. Even when a strip of solid medium was completely removed between two cultures. which were transplanted at different ages of the embryo. and therefore had different characteristic growth rates. and permitted diffusion. . from one side. were grown in separate dishes. even if the slide Further did not touch the bottom. These remained constant as long as the cultures 6 8 10 12 ft days Figure 35. that of the more slowly-growing ones was distinctly increased. experiments showed that the effect spreads re ctili nearly by placing several cultures in the same dish. Incisions in the solid medium which separated the cultures and prevented diffusion from one to the other did not prevent the mutual stimulation. with a that the . the influence ceased. then united in the same dish.

both were cidtivated in separate drops on the same quartz cover glass. those on quartz grew more rapidly as measured by % means of a planimcter. after 12 hours. Next. JAEGER (1930) observed that blood radiation retarded the days (the last was far in tissue cultures. DOLJANSKI used cultures which reacted promptly upon addition of embryo extract. The 0. the exposed advance of the control. It may be that a certain stage of development is necessary make the cells sensitive to the mitogenetic stimulus. radiation began after 60 hours with a fibro blast culture from the heart of a chicken embryo. At this time. ho obtained in all 4 experiments a more rapid growth in the culture nearest to the radiating substance. Here. A fibroblast culture (chicken) was divided into halves. as was shown for yeast and bacterial cultures (pp. In 56 such pairs.17. The induction effect was recorded as the ratio of increase in the quartz culture over that in the glass 0. 61) and 120). but without irradiation. RAWIDOWICZ (1931) as well as DOLJANSKI (1932) could find no growth of stimulation by mitogciictic radiation. as with all other detectors.01 _j_ o. CHRUSTSCHOFF believes that radiation is . after 3 culture day being without radiation). the average effect was 1. which was replaced when necessary. the irradiated culture appeared much denser than the control.91 to actual stimulation by radiation from bacteria was therefore 0. distance was only Very recently. Another set of 48 parrs.o^ proving no chemical effect from glass or quartz. some LASNITZKI and KLEEinvestigators obtained negative results.92 culture. gave the ratio 1. even if tbo mm. but they did not respond at all to organic radiations. and one of the two was irradiated for 48 hours by a beating embryo heart.15. CHRTTSTSCHOFF used the tissue culture as detector. The second paper on this subject was that of CHRUSTSCHOFF (1930) who observed that growing tissue cultures began to radiate some little time after transplantation from the original tissue. 6* . and exposing both to radiation from stapbylococcus culture. By placing one half of a culture on glass. the effect being 5.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 83 head of a chicken embryo. With a spleen culture of Amhystoma tigrinum.3 times the probable error. but only on poor media. due to autolysis of nccrotic parts in the culture. JULIUS (1935) obtained definite growth stimulation of chick fibroblast cultures. the other half on quartz.

after irradiation. the animal's physical light sources. 3 of the head into an immovable position. the two corneae of the same animal always show it irradiated. so that one eye can be and the other used as control. Mitogenetic Effects produced in the Oorneal Epithelium 4 minutes exposure of the left eye to the of vertebrates by 3 spectral line 2030 A A very good detector is the corncal epithelium of verte- brates.84 CHAFTER IV Table 28. Method: For With sufficient. an exposure of 3 4 minutes was head being held in the hands of the experimcntor. very nearly the same number of mitoses. or perhaps preceded by changes in metabolism. Ordi4 hours time was given for the manifestation fixed for 40 minutes in of the effect. *) upon the rate of respiration The technique employed was culture. The number of mitoses varies with the physioincreases rapidly with good nourishment. e) Detection by changes in yeast metabolism G-ESENius (1930 a) concluded that such decisive changes as the acceleration of the growth rate of yeast must be accompanied. stained with hamalaun. The results in Table 28 show very strong mitogenetic effects. logical state . therefore. The cornea was 70% alcohol 4 5% acetic acid. dropping in rats during starvation from approximately 2000 to about 50. He studied. and clarified in glyeerol. The source of radiation was a yeast . which necessitated the tying down narily. the influence of irradiation and of fermentation of yeast. according to LYDIA GURWITSCH and ANIKIN (1928). Fortunately. exposure had to be continued for 20 minutes. It is the only easily accessible tissue of the grown animal showing frequent mitosis. biological sources.

He observed further that from patients with most diseases. leucemia. after the ceptions were only with cases of pernicious anemia. improvement and found that normal blood 1 always radiated ). the average depression being 14%. mutual irradiation of the cells must play an important role in thew experiments. His results agree very well with those of L. blood radiation decreased the fermentation in 28 out of 30 experiments. the blood radiated. If a failure occurs. The role in cancer diagnosis of this loss of radiation will be discussed in Chapter VII. free from cells. *) "Healthy blood never fails. e. The results can be briefly summarized in the following way: Fermentation in N-CO. further the iniluencr of radiation upon macerated yeast. and that the consistent exof the technique. 1932. upon zymase. While yeast radiation produced no effect. carcinoma and severe sepsis (see fig. 7 to 10 billion cells per cc. GLTEWITSCH and SALKIND and of SIEBERT. and probably accounts for the depression of GESENIUS tried respiration. The same retardation of respiration also could be obtained with sea urchin eggs during the early stages of cell division.) .METHODS OF OBSERVING BIOLOGICAL RADIATIONS 85 essentially that of WABBUKO (1923). i. which was exposed in quartzbottomed dishes to yeast radiation for 4 hours before being tested. 153). \ I atmosphere J f68 increase 102 ]02 cxporlmelltg t experiments < 4 decrease 3 | Respiration (oxygeii-uptake) in 2 j |40 decrease increase atmosphere. to blood radiation GESENIUS (1930b) applied this test. it is time to test either the yeast or the apparatus. This 10 to 100 times the maximal The which can develop in the medium used. > 54 experiments 1 [ without sugar J he limits of error [l3 within t cells The number population of yeast in these tests is was very large. The result was a stimulation of fermentation. 40 p." (GESENIUS. but a retardation of the oxygen uptake. The organism used as detector was a wine yeast. or of RUNNSTROM (1928) for measuring respiration of tissues or tissue pulps.

who exposed and even from chemical reactions. later experiments. The first observations of this kind were those by and M. In each case. but not through glass. They found that though a layer of glass prevents the effect. and if further the difference in oxidation-reduction potential between the two liquids is very great. these authors conclude that they are dealing with an electric effect. the author and his associates have regularly observed similar morphological changes. they elongated and produced hyphae or. they did not grow at all. the greatest care was taken in later experiments effect any chemical influences. When the electric insulation was prevented by a metallic connection. Upon various criticisms. . the was transmitted through quartz. Table 29 shows a summary of the results of these experiments. While the MAGROTJS originally explained this morphological change through mitogenetic rays. During the last two years. and must blood. the MAGKOTJS do not give it preference to the ultraviolet radiation theory. with enormously distended vacuoles or. but died. This explanation is tentative. MAGKOU the eggs of the sea urchin Paracentrotus (1928) lividus to radiation from bacteria. . In their recent publications. From this and similar experiments. 49 p. the larvae remain normal. yeasts or other organisms. offered another possible explanation. CHRISTIANSEN observed very strange morphological changes in yeasts and in bacteria brought about by menstrual The effect passed through quartz coverslips. together with REISS (1931). The larvae become abnormal if the source of radiation is separated from the medium of the sea urchin eggs by a very good electric insulator.86 CHAPTER IV f) Morphological changes by biological radiation is Quite different from the previously described manifestations a decided morphological change in the irradiated organisms. two layers of glass with a coat of paraffine between them permit the effect to pass. They obtained quite abnormal. more or less spherical larvae while the normal larvae possess a very characteristic conical form (see fig. The yeast cells either became large and spherical. 165). J. In 1929. and the purely physical nature of the effect cannot possibly be (1931) to prevent doubted. therefore be considered as the result of biological radiation. .

METHODS OF OBSEKVING BIOLOGICAL RADIATIONS Table 29 87 MAGROU'S Results with Biological Irradiation of Sea Urchin Larvae .

In later publications. young seeds. ) a clean glass plate r 4 inches. After solidification. bichromate. Silver cbromate is gradually precipitated in concentric rings \\icii spread over the entire plate in about 24 hours. STEMPELL'S since. However. The precipitant diffuses gradually and the* precipitate is deposited in concentric 0. we could never observe the true Of the branching of cells which OJIKISTTANNEN has described. 2 drops of a 20% AgN<) 3 solution are placed on tin* center of the plate by means of a fine pipette. the ring formation while small doses intensify it. lie observed II LIESEGANU that the LiESEdANcj rings an* disturbed by biological radiation. gelatin. STEMPKLL could prove disturbance of the rings when chemical influences were completely eliminated. tubes as closely as possible over the gelatin surface. rings. I of eliminate gelatin (12g. exposed places. a drop of another solution is placed causing a precipitate with the salts in into the gelatin. g) Physico-chemical detectors ings: It was recognized by NT KM TELL that a physico-chemical detector would carry much more weight than biological ones for the proof of mitogenetie radiation. the tendency was not a shortening but rather a lengthening of the cells. Strong ultraviolet from a quartz mercury vapor lamp thrown through a narrow slit upon the gelatin intensifies the ring formation at the.4 g. Method: 2 ammonium water -| 1 co. various parts of plants. with drop of 3" aqueous p\Togallic acid) are poured hot upon ec. KiOcc. were essentially identical with those observed by CHRIHTIANSKN. the gelatin.. immediately before use. The light situation is rather complicated. however. These rings appear when. it first experiments (1929) with onions were not 1 could be shown that the ally -mustard oil of the accepted onion caused a chemical disturbance of the LIESKUANU rings. while leaves had little or no effect (see fig. and Chapter VII). while weak light. this was so pronounced with some Mycoderinas that their growth strongly resembled that of mold mycelium. 3. being mixed. on a gelatin gel containing certain salts. seedlings and pollen were the most effective. water. 102. Onion . oil acts in the opposite way a large dose of onion. at the edge of the slit. 48 p. oil gas lessens . decreases it. The uniformity of these rings is disturbed by radiating material placed in quart/. however.88 CHAPTER IV The effects in beer and wine yeasts produced by saliva. When irradiated by plants. the roots.

detector. p. 46) states that the disturbance of 89 CIANG rings is usually brought about by a combined action of radiation and chemical effect of a "gas". and states that the LIESECJANCS rings at present are the only detector for this substance. He considers this chemical effect to be very important.5 mm of quartz. . The gold sol was prepared in the following way: To 1000 cc. 5/3). right: effect through 0. After long exposure in a moist chamber. Flocculation of Colloidal Solutions: A promising method has been worked out by HEINEMANN (1934. quite different by STEMPELL (1932. a volatile substance produced by the sender. p. It Ls based into H a O -f. has been found on the deterioration of of ultraviolet radiation. the possible biological meaning of the chemical emanations will be omitted. or a bundle of several roots. Gold sol was found to be more satisfactory than iron hydroxides. of a slightly alkaline solution of 0. 40) regarding Figure 36. just over the root tips. Decomposition of Hydrogen Peroxide Another.METHODS OF OBSERVING BIOLOGICAL RADIATIONS STEMPELL (1932. since none of the other detectors for mitogenetic radiation react to it. 1935) who observed that inorganic sols flocculate more readily when exposed to mitogenetic rays.8% glucose. is fixed so as to touch the underside of a thin quartz plate. On the opposite side of the quartz plate is placed a drop of 2 O 2 H . e. the peroxide concentration in this drop is less than that of the control. biologically. under the influence 2 O2 : H One onion root. The effect upon LIES EG A NG rings of onion base pulp in metal tubes with a slit whose position is indicated by the black line left: effect through cellophane. Since this book is meant to be limited to biological radiation. i. the interesting speculations of STEMPELL (1932. p.O.

TAYLOH and HARVEY (1932) could obtain no effect by exposing plates to frequently renewed fermenting yeast for ninety days. small amount of solution is added just before the beginning of the experiment to make the gold sol unstable.5 1 0001 0^2 2 2 2 19 2 19 2.1 g. and as organism. dissolving: 0$ 5 10 10 21 h) Measurement by physical instruments be surprising that radiation by organisms has not been recognized and proved conclusively long before this. fig. Upon is reduced by the glucose to a bright-red.5 2 15 NaCl. the* arrow indicating the moment when the radiating material control: was applied 3 5 7 to the quartz dish: 1. and those by BRF- NETTI and MAXTA (1930) and by PROTTI (1930). heating. in the absence of radiation. and therefore a change of the galvanometer The reading while. .5 1. This is a rule. Radiation causes a more rapid flocculation. and covered with a quartz dish.90 150 cc. The best detector in most cases the living is still unsatisfactory since we must allow for considerable individual variation of the detector organisms. They are placed into a very sensitive photoelectric differential nephelometer. AuCl 3 are added. The It may reason must be sought in its very weak intensity. the readings remain fairly uniform. the gold salt clear gold sol. This is done in complete darkness. solution is then distributed between two beakers. None of them are absolutely convincing.5 22 25 2G 3 3 blood: control: 0000001 1 1 7 8 10 1. by means of a galvanometer. The following readings were obtained in 15 consecutive minutes.5 11 IG 2 11 20 2 11 1. and GUBWITSCH has refused them all as proofs of the physical nature of mitogenetic radiation. The* intensity is so slight that the most sensitive photographic plates and the most elaborate physical instruments have failed to record this radiation. and the supposedly radiant substance is placed into one of tho*quartz dishes. the measurements are not as accurate as with physical experiments. of CHAPTER IV a neutralized solution of 0. A Nad The is each difference in turbidity between the two beakers is read every minute. The only photographic records are the ones reproduced in 15 of KEITER and GABOR'S monograph.

29). at least. muscle was tetanized by means of an induction coil directly in front of the window.30 and Fig. even though slight. 21. Table 30. may definitely change the counting rate even if there is no radiation present. 28). These experiments were repeated successfully by (1930) with working muscle (see Table. RAJEWSKY succeeded in obtaining direct physical proof of this radiation by means of a photo-electric counter (see His data with onion roots and carcinoma are shown in p. changing its resistance and thus altering the counting rate.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 30. the water vapor from the material. LOKENTZ (1933. Later experiments of this nature are those by BARTH (1934) and SIEBERT and SEFFEHT (1934). or opening a shutter between the counter and source. FRANK and RODJONOW : . will condense upon the quartz of the counter. at least not by physicists. the charges which are inevitably present on the outside of the quartz tube containing the biological material are almost sure to change the counting rate. To enumerate (1) If the biological material is not in a closed quartz container. Some experimenters have even placed their moist material directly upon the quartz window which will certainly change the counting rate. (2) If the water vapor is carefully kept away from the counter. p. However. 1934) could show that bringing the "radiating material near the counter. 91 Measurements of Mitogcnetic Radiation by Means of the GETGER Counter In 1929. (3) In one case. these results have not been accepted generally.

are a step in that direction. CHAPTER IV Photo-electric yields obtained for ultraviolet light To separate the effect of extremely low intensity from these other effects is very difficult even when their existence is realized. Most convincing is the experiment that a counter gave a definite increase when exposed to normal blood. KREUCHEN 4 6 from (1935) obtained yields of from 1C to 10 quanta per electron hydrogen-activated zinc and cadmium surfaces. it ever was reached. but showed 110 increase when this was KCN. it seems from later work that this value has never been sur(see of the surfaces used passed if. in fact. but no increase with blood from carcinoma patients. In his latest paper. Many more extended removed and replaced by blood | experiments of this general nature will be necessary to establish finally the radiant nature of the biological effects. more than 2000 quanta are required to eject one electron. is The experiments of SIEBBBT and SEFFERT (1934) who obtained increased counts with several hundred normal blood samples. A more careful description of the method of exposure. and it seems well to view with caution the positive results claimed for the physical detection experiments so far carried out. Though some early workers indicated higher sensitivities.92 Table 30a. In a recent paper. . and a number of experiments with water blanks or non-radiant organic materials will be necessary to produce evidence which physically irreproachable. In none of the physical measurements of mitogenetic rays is it absolutely certain that these errors have been excluded. 29) by the various investigators. In all cases except the first. KREUCHEN and BATEMAN (1934) reviewed the field of physical detection and present their results in a table Table 30 a) which gives the photoelectric yield (see p.

but by discussing this point with the various investigators in this field. and it was impossible to produce even . For the reproduction. for the wave length 2300 A). failures in proving radiation Unaccounted It must be stated with fail detectors sometimes for unknown perfect frankness that biological reasons. This cause became evident through the fact that all other associates using the same culture on the same day obtained negative results. much larger intensities are required (about 6. Twice it happened that all the experiments of one day proved to be negative though the culture had reacted promptly on the previous day. It was found that the sensitivity culture had changed. FRANK and RODINOW observed higher values. Most of the sudden failures of cultures to react have. These authors believe that a change in the opposite direction may also take place. 54 failed on account of a poor quality of the yeast culture which was used as detector.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 03 It would be of great advantage if some surface having a Prehigher efficiency for the ultraviolet could be obtained. and that a longer exposure was necessary. or even for a number of weeks. in OUKWITSOH'S laboratory. with the working frog muscle and heart. Professor GUKWITSCH has told the author that in his experience such a condition usually remained for several days. (10 to 100 quanta/cm 2 /sec. estimated from to offer encouraging results. WOLFF and RAS (1933 c) working with Staphylococci had a similar experience.6 of 10~ 10 to 10~ 9 xlO 5 quanta. practically all seem to have had the same experience. GOLLSHEWA (1933) mentions that out of 373 experiments with blood radiation. they obtained with pulp from muscle. values up to 2000 quanta/cm 2 /sec. not been published. No reason for the abnormality of the yeast culture is mentioned. experiments the intensity and found it for onion roots and for 2 carcinoma tissue to be of the magnitude erg/cm /sec. of the various niitogenetic phenomena with physical sources of light. 1932). i) according to STEMPELL. liminary experiments by one of the authors using magnesium surfaces sensitized by oxygen seem lu's RAJBWSKY of this radiation. Acs (1932) claims to have inof the creased sensitivity by selection. Probably all investigators working with biological detectors have been worried by such failures. Some of them have published short remarks.

may it the contrary. and he resorted to the GEIGER electron counter as a more dependable detector (see p. sunspots. to a change of sensitivity of the detector culture WOLFF and HAS). and they come and go at irregular As a rule. terrestrial magnetism. to a retarding effect by human radiation (RAHN). But with him. and thereby may On bring about a better understanding of mitogenetic effects. too. These occasional failures have nothing to do with the error method. The author himself has also had long periods of negative results in his laborafailure tion (see p. it help to explain the cause of these failures. This induced him to look for physico-chemical methods of detecProfessor WERNER SIEBERT'S many succesful with a experiments yeast detector have been mentioned in practically every chapter of this book. g. At the present. it does not seem wise to belittle this experience. we do not know whether the culture. ( The cause of this disturbance might be more readily traced and overcome by the cooperation of various laboratories. it might be that the senders do not function under the prevailing detector. When mitogenetic effects are observed. as condition. it is only an assumption that the change has influenced the If the cause should prove to be of such general nature weather. cosmic rays. 92). the investigators do not is still intervals. e. with yeast as detector. 181) Frankfurt and in London. not even the testing of a large number of different yeast cultures. the yeast suddenly ceased to react. the experimenter or the environmental laboratory conditions have changed.94 CHAPTER IV Eventually the culture reacted Doctor HEINEMANN. they are outside the limits of error. after a very successful the simplest mitogenetic effect. by calling attention to it. discuss these periods of failure because there considerable doubt among physicists and some biologists concerning the existence of the mitogenetic phenomena. normally again. and none of the various attempts to obtain normal reactions proved successful. in fact. tory. to climatic changes or some other cause. diagnosis of cancer in by the absence of blood radiation (see p. Whether is due to disturbance by short radio waves (suggestion by GURWITSCH). and the emphasis increase this doubt. is not known. 89). of such periodical failures might While this can not be denied. suddenly experienced a complete lack of reaction.. The failures might be compared to of the .

Experiments to prove this have been successful with some . of hours of light per day (if decided The 1 consistent observation of this disturbance by most not ah ) investigators who have obtained large series of positive results.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 95 the experience of expert florists that sometimes. detailed investigation woman technician A by CHRISTIANSEN (1929) led to the discovery that the effect from menstrual blood passed through . This was not caused by poor seed nor wrong soil. and the term "menotoxin" for the hypothetical compound causing it is in general use. MACHT and LUBIN. POLAND and DIETL. as bacteriologist of a dairy laboratory in Germany. 1921. 1920. it was ultimately found that this abnormality occurred during the menstrual period of the in charge of the cultures. no great stress can be laid upon the results obtained on any one day. CHRISTIANSEN. points out a common error in some criticisms. but would not increase the proof or disproof of biological radiation. B. in- vestigators (ScHiOK. It has been claimed that many simultaneous parallel experiments prove more* than similar experiments spread over a longer period of time. that Hewers in their hands wilt readily. certain plants refuse to bloom in the greenhouse. the belief in this effect seems to have been rather prevalent among medical men. BOHMER. Still. It is believed an that bread dough kneaded by them will not rise. that food preserved by them will not keep. As long as it is not known why the occasional failures occur. observed that the pure cultures used for dairy starters occasionally developed poorly and abnormally. It is INJURIOUS HUMAN RADIATION from old "superstition" that a harmful emanation comes the body or the hands of menstruating women. 1924). and the multiplication of experiments on such days would only reveal the error of the method as such. g. but remained unexplained for a long time until it was found that the number this. KRETJCHEN and theirs is is BATEMANN (1934) state that one series of That equivalent to 140 single experiments by GURWITHCH. After eliminating all other causes. It might be a day where the detector fails. 1927) FRANK. E. 1921. a mistake. 1927 and gave 110 results with others (&ANGKU.

he showed that wine made with a pure culture yeast by a menstruating woman fermented feebly.signs indicate growth. the controls nearly always grew normally 37). and no growth of yeast.in droplet cultures transferred hy the same person. stronger in summer than in winter. CHAPTER IV and must. and since one woman who had been treated with ultraviolet light in winter. be considered a radiation.96 quartz. Further. The droplet culture on the coverglass requires that the certain of woman coverglass be held in the fingers while the droplets are a pen dipped into the yeast suspension. called attention to the fact that the He FERGUSON (1932) during an investigation of morphological changes in yeast by plant radiation. he thought difference very probable that the seasonal in solar irradiation. or killed them. while the negative ones had been obtained in winter. Aside CHRISTIANSEN did not go into the nature of the effect. continued to show from this proof. observed occasionally that for short periods. H. Alternation of growth it strong radiation. The blood produced cither abnormal morphological changes The effect was much in yeasts and bacteria. but not with radiations from the body. (fig. therefore. o signs in black bars indicate no growth. The periods of no growth through summer and while in winter. growth alternated with those student. was brought about by the variation above-mentioned investigators obtaining positive results had made their experiments in summer. covcrglass cultures of one yeast did not grow when made by one fall. He worked with menstrual blood and saliva. made with . while the control showed a vigorous normal fermentation. July 1931 August 13M February March June 1331 July Figure 37.

but have not been able to prevent it. drops of yea/t'7 raisin / \ . regardless of the fact that this radiation does not reach further than a few inches. and that only one A A especially sensitive species of yeast could be killed in this way.METHODS OF OBS FRYING BIOLOGICAL RADIATIONS 97 The above-mentioned experiments of CHRISTIANSEN made it probable that this failure was due to human radiation. this method. the authors are in no way responsible for this kind of publicity. he frequently killed yeast through quartz in 15 minutes. he did not fool quite woll. finger of the face. After a summer vacation. With this person. thickness was placed between finger and yeast (fig. This test has been applied to a number of people. during 3 successive days of sinus infection. a quartz plate of 2 mm. it reminutes to kill the This student menstruat- by 1. fourth case was one of the authors. and it was found that this radiation occurs only very rarely. i. Frotoplasma-Monographien IX: Rahn 7 . the fingertip was held closely over the yeast by the support of a glass ring (fig. for about fi months after recovery. In another experiment. and some experiments proved that an emanation from the fingertips of this person killed yeast in 5 minutes while others making the same test with the same culture produced no marked effects. Never before or afterwards did this person show any effect upon the yeast. he felt perfectly normal. ^g. radiation also occurred from the tip of the nose and from the region of the eye. quired yeast. In this experiment. but this was not always the case. 38). During this timo. It was most pronounced with a man who had recently recovered from herpes zoster ^ ^^J^ $d f $* rir >9 ^overglaL Fimro38 Methods of testing radiation. and the ease suggests an abnormal parallel to CHRISTIANSEN'S oKservations. 1 ) was a hypo-thyroid patient tested only once. and his power of radiation was gone. third case *) This has been interpreted by imaginative but uncritical newspaper reporters as a scientific proof of the "Evil Eye". 38).5 ed rarely and irregularly.

This radiation be shown later is quite likely duo to a skin excretion. Two other females stimulated yeast growth. the typical saliva reaction could not be produced through quartz. with Saccharomyces mycoderma punctisporus. a manner of growth strikingly resembling mycelium. It is It was observed not certain. This yeast does not cause fermenOther yeasts showed slight retardation. saliva was mixed with the raisin extract in which the yeast cultivated. and one female member. isolated complete killing observed. and it must be left to a later investigation to solve this problem. and pratieally independent It is typical An of the diet. as will (p. not even with half and half raisin extract. the other hand. but never wan tation. This harmful effect for the individual. this did not exclude a chemical effect. and. were never convincing. from the scum of a fruit juice. granuof beer and wine yeasts to spherical cells of increased killing effect many tents. This seems to exclude any chemical effect. such as absence The pictures of granulation.98 CHAPTER IV The experiments were always made with droplet cultures through quartz. saliva of lated cells was frequently observed with saliva. Another The size. 185). the droplets were prepared by the same person who had been found over a 2-year period never to radiate. or cease to do so after a few cell divisions. as shown in fig. The only organism which reacted upon this radiation was Saccharomyces mycoderma punctisporus GUILLIERMOND. in droplet cultures which were mounted imme- When was saliva diately above the saliva. In all more recent work. 38. however. investigation of the saliva reactions of several members of a family showed the above-mentioned injurious effect with male members of the family. persons changes the oval or elliptical. produced elongated forms. only partial effects could be obtained. it produced no abnormal cells. with greatly distended vacuoles and homogeneous cell conOften. the organisms either do not grow at all. is not typical for all saliva. with yeast on one side and saliva On on the other. or tendency to become spherical. that the effect is one of radiation. .

though parallel experiments were made also with leaves of Jtilodea. when cell constituents break down. the suspension exposed to dying proved to have received some radiation. The gray color of the silverprotein precipitate with living yeast was supposed to be brought about by ultraviolet rays emitted by the when yeast a mixture of solutions of KBr and AgNO 8 Living yeast with ether caused a dark discoloration. with petals of flowers. very sensitive photographic plates (EASTMAN SPEEDWAY) cut in small strips. were submerged directly into the yeast suspension. Ho con- cludes (1932b) that weak intensities of ultraviolet must help in the synthesis of cell constituents. while strong intensities made cells less The two effects are independent of each other. least. the yeast died within 12 minutes. erythrocytes) is increased by irradiation with weak stable. emitted. and the suspension was gray. After exposure with continuous shaking in the dark room. the dying colls. AgBr cells developer. the energy absorbed during synthesis must be released again. and that vice versa. the mixture remained light-colored. suspensions were removed and mixed with a photographic In all experiments. Part of the plate was covered with filter paper which excluded physical. ultraviolet light. but not chemical effects by soluble substances. but turned gray upon exposure to light. the plates upon development remained light. With living . After 10 to 25 minutes exposure to yeast previously killed by ether.METHODS OF OBSERVING BIOLOGICAL RADIATIONS C. but when the yeast had been killed by ether before the AgBr mixture was added. 99 NECBOBIOTIC BAYS LBPESCHKIN had observed (1932 a) that the stability of living matter (plant cells. g. When silver nitrate was added to a living yeast later suspension in the dark room. A similar difference could be observed was suspended in . and also into those with living yeast plus ether. Then. and with suspensions of Bacillus subtilis. e. When the yeast had been killed by ether or by heat before the silver salt was added. AgBr suspensions in small quartz tubes were inserted into tubes with dead yeast. the suspension remained white. Since these experiments did not exclude chemical effects. of Pajmver. The test organism was nearly always yeast. partly at radiation. as ultraviolet The experimental proof is given in considerable detail in a paper (1933).

the living yeast. but the uninjured cells next to the wound. PESCHKLN apparently feels this. e. the effect from the same mixture with ether i. though there was a slight from the fermentation upon the AgBr. effect 10% sugar. he believes it possible that of vital "necrobiotic rays" might also be emitted from a decomposition compounds in living cells which might be imagincable during very rapid physiological processes. From the absorption of these rays by glass and by gelatin. and they tested it by coagulating egg white by alcohol in quartz or glass vessels which were placed over AgBr-suspensions as in LEPESCHKIN'S experiments. except for the little strip shaded by the filter paper. They conceived the idea that the "necrobiotic rays" were emitted from the coagulation of proteins. LE- PESCHKIN was not familiar with the latest literature on this subject. autolysis and of wounds as proof for his contention. they were also light. In the case of wounds. However. He emphasizes the radiation of necrobiotie processes e. lengths. and after exposure. LEPESCHKIN estimates their wave length to be largely between 1800 and 2300 A. He mentions respiration as a possibility.100 CHAPTER IV However. with a very weak emission of greater wave This agrees fairly well with the range of mitogenetie From the above experiments. it is not the injured cells which radiate. the dying cells affected the plates so that during developing they turned dark. However. SUCHOW and SUCHOWA (1934) have perhaps found the link between LEPESCHKIN'S and GURWITSCH'S explanation. it would be necessary to consider almost all exothermic reactions as neerobiotic processes in order to combine the two types of radiation. The experiment was carried out in the dark. rays were stronger than those from actively fermenting yeast cells. This proved to LEPESCHKIN'S satisfaction that the effect was physical and not chemical. g. it seems as if the necrobiotie rays. the two . LEPESOHKIN then ventures further to state that many of the mitogcnetic phenomena are in reality due to necrobiotie rays. with dying cells was much and added stronger. when ether was added to yeast. The radiation spectra of the various chemical processes are ample proof that dying cells LEare not necessary for the production of mitogenetic rays. Evidently. LEPESCIIKIN could obtain an indication of an effect upon AgBr suspensions if he used living beer yeast instead of baker's yeast.

is the same whether the . The cell is alive intensity of this radiation or dead. the suspensions were brought into light. The effect passed through glass. D. it must therefore be of an infra-red nature. The authors believe therefore that the reduced fertilization is caused by a photochemical effect. however. and was not visible. it will not be discussed extensively here because this type of radiation is utterly unlike the mitogenotic and related is rays. the eggs were fertilized by the usual method. It originates from the radioactive fraction of potassium. a series of observations by STEMPELL (1931) that sprouting peas will increase distinctly the rate of spontaneous decomposition of a saturated solution of HO 2 2. BETA-RADIATION entirely different type of radiation should be mentioned only in passing. in each of the 9 experiments a distinct decrease in the percentage fertilization. suspension which stood under the quartz vessel uniformly darkened sooner than the other. The only one known to the authors is an by their was necessary The temperature of own respiration. Though it has been proved experimentally. but an artificial experiment by NELSON and BIIOOKS (1933). obtained from a Mazda lamp by means of a monochromator. namely the bet a -radiation of potassium. and really not characteristic of the living cell. and is of importance to life An processes. the peas rose 0.5 increase of 5. Equally rare are observations of an effect of infra-red rays upon living organisms.06 C.5% and 87. The temperature difference between control and exposed eggs was not more than 0. The only case of near infra-red emanation known to the authors is.5 to bring the rate of peroxide decomposition to that produced by peas.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 101 In 25 experiments. The decrease varied between 18.1%. The irradiated eggs showed. They exposed the unfertilized eggs of 2 sea urchin species and of one worm to infrared rays of 8000 to 12 000 A. E. but of its potassium content. It INFRA-RED RADIATION might appear rather probable that some organisms would emit infra-red rays since some are capable of producing visible and ultraviolet rays. After 15 to 45 minutes exposure.

thorium. he could show that frog hearts which had ceased the same to beat in RINGEB'S solution minus potassium. total energy SCOTT (1931) determined the from potassium of the average human heart to be 9. transform the potential energy of the heart muscle in response to node and bundle impulses into the enormously greater manifestation of kinetic energy. . In 34 experiments. experimentally the physiological importance of potassium radiaHe succeeded (1926) in keeping isolated frog hearts beating by substituting the potassium of RINGER'S solution by radioactive equivalents. He states that "one may readily conceive that the free energy of the beta particles can be cumulative and.102 CHAPTER IV. rubidium. this radiation is biologically important. Nevertheless. started again in solution after about half an hour's irradiation by mesothorium (in glass) or radium (through mica). heart beat soon ceased again and could frequently be brought back a third time by to new exposure beta rays. METHODS OP OBSERVING BIOLOGICAL ETC. After removal of the radioactive substance. uranium. the systolic contraction". may tion. radium or ionium.64x 10~~ 6 ergs per second. reaching a maximum. and be the chief reason for the indispensibility of potassium in ZWAABDEMAKEB (1921) was the first to test living organisms.

The width of the window is measured by the central angle (fig. it was ho intensified by irradiating The customary method between sender and detector. From this angle and from the number of revolutions. initogenetic radiation. 35). The present chapter initogenetic rays describes some to which are not common all peculiar properties of types of radiations. the duration of each. processes. The preceding chapter has revealed that the etic radiation is .CHAPTER V SPECIAL PROPERTIES OF M1TOGENETIC RADIATION socalled mitogena real radiation according to tho strict physical definition of the word. i.. A. Table 31 gives the results obtained. 59) it . of a rotating disk which contains one or several openings or windows. 39). and the total time of actual irradiation. disks contained one or several windows of varying width. and the rate of rotation of the disk allow one to vary the three items concerned in intermittent intermittently instead of continuously. 59 and 80) it can be absorbed (p. It travels through space rectilincarly it can be reflected (pp. It indicates that the minimal . .e. The width of these. the frequency of exposures. shows refraction and dispersion (p. their number. much threshold value in mutual yeast irradiation by means of disks The rotating at approximately 3000 revolutions per minute. the duration of each interval and the frequency of interruption can be calculated. INTERMITTENT IRRADIATION of Very early in the history discovered that the effect could for this purpose is the insertion. The most striking result with intermittent irradiation is the shorter total time of exposure necessary to bring about OrmwiTSCH (1!)32) determined the distinct mitogenetic effects.

104 CHAPTER V O Op O ppp o ooo" o p rH G^ L-" i-H C^l i~H r-l C .

12. CHLOamount measured the (1919) BELLA. 13 seconds are sufficient for induction.5 to 13 seconds When the same experiment was tried with being sufficient. 256) repeated the same experiments with a half -disk rotating at 1 Or. at right: side view showing the position two agar of the blocks with the yeast sides each facing other. Figure 39. When the frequency is between 800 per second. ZOGLINA (quoted from 1932. The results are given in Table 32. it a distinct effect. This meant uniform intervals of 1 exposure and irradiation of / 2o^ second each. The following per- GUKWITSCH centages in increase of buds were obtained with a total exposiiro time of 60 seconds: 60 7 15 -|-56 5 +4 +36 +28 +64 +28 The intermittent increase in efficiency of a photochemical reaction by irradiation has its analogy in the increase of photo- by intermittent exposure. A rotating disk was used which was in principle like that of fig. since it signifies an approach to continuous irradiation. p. The rhythmic interruption required 6 to 8 minutes to produce of radiation had decreased the threshold time to about l/30th of the amount required with continuous exposure. 39. WARBURG of CO 2 absorbed by an alga. 15 and 10 seconds of total exposure are usually insufficient. but the times for light and dark were made synthesis of green plants equal. p. for intermit- tent muto-induction. uninterrupted irradiation. during 15 minutes of actual exposure. either continuously or discoiitinuously. The frequency of interruption has some bearing threshold value. An increase up to . upon the and 100 However. Rotating disk for in- termittent radiation. when it falls to 50 per second. and 30 seconds are necessary to show a mitogenetic effect.m.SPECIAL PROPERTIES OF M1TOGKNET1C RADIATION 105 actual exposure must be more than 10 seconds. This is to bo expected.

106 CHAPTER V -d so CO CO CO -t CO >O iO - 00 OS .

found a 30-fold increase in the threshold value while the greatest difference between light and dark periods was only 1:0. 1932. their concenHis intention to investigate is only comparatively small. but also with very strong sources which produce either no effects or depressions (see p. It is not at all certain that this observation is really anal- ogous to the increase of the mitogeiietic effect. in more detail the latter. assimilation is more rapid at the first moments of exposure because more substances have tration accumulated ready for photosynthesis while later. Besides. WARBURG considers two possible explanations: either assimilation continues for some time after darkening (e. A rhythmical interruption of the radiation seems to be essential for mitogenctic induction. The greater susceptibility also While mutual induction permits transmission over longer distances. however. where weak intensities of radiation could not be induced to produce stronger photosynthesis by intermittent exposure (Table 32). we cannot be certain that these threshold times are reliable measures of intensity of radiation (see p. 115). 200). GURWITSCH. there seems to be no difference between strong and weak intensities. p. 114). a paper by EM EKSON and ARNOLD (1932) came to our notice. with continuous exposure. more probable case seems never to have materialised. of yeast has its limits at 3 -4cm. On the other hand. The greatly intensified susceptibility of tho living detectors by rhythmic discontinuity of radiation shows that by this method. This holds true not only with very weak senders. very good results can be obtained over 15 cm. or. weak intensities 1 ). Parallel experiments were made with l ) After the manuscript was finished.SPECIAL PROPERTIES OF MITOGENETIC RADIATION practically the double but no difference with 107 amount was observed with strong light. radiations can be detected which otherwise produce no effect whatever when applied continuously. . They were able to increase photosynthesis 400% by intermittent irradiation whereas WARBURG (1919) could only double it. These facts differ greatly from WARBURG'S observations with algae. WARBURG could double the amount of photosynthesis by distributing the same total radiant energy over twice as long a period. with intermittent irradiation (GuRwrrson. through some short storage of energy). g.

inductioii of yeast. and whether tested in light or dark. Diffused lignt is entirely sufficient. varying in size from 2. measuring the percentage increase of buds on yeast Many affect other grown on agar blocks. All experiments wero made by mutoby exposing yeast to yeast. .108 CHAPTER V total two disks rotating at the same speed.5 to 30. In 1930. and for the pulp of the onion base. as well as for the pulp of a number of plant tissues. Influence of Daylight upon the Yeast as Sender and as . o. the detector and the light during exposure. the other disk contained openings. POTOZKY gave several series of experiments showing that the same holds true also for yeast. p. The experiments were made by the Baron technique. Yeast grown in daylight yeast. Only with the regular spacing were positive results obtained (GuRWiTSCH 1932. B. could hardly be accounted for by any of the two explanations of WARBURG'S. This has been observed for onion roots cut off from the onion bulb. both with a width of 75.Detector of Mitogeiietic Effects Obtained by Muto-Induction Three factors wore varied: the sender yeast. INFLUENCE OF DIFFUSED DAYLIGHT of the common "senders" of mitogenetie radiation organisms only when in daylight. This. i. 261). Yeast grown in the dark had no effect upon yeast. whether grown in the light or the dark. Table 33. in irregular distribution. the 75 were distributed uniformly. window In one disk.

phenomenon characteristic of the living cells only.SPECIAL PROPERTIES OF MITOGKNETIC RADIATION showed distinct radiation 109 and mitogenetic effect upon the detector yeast. in nerves. and WOLFF and HAS (1933.been transmitted through the root by reflection without having been absorbed completely. 34). by means a GEIGEK counter. Secondary radiation was observed in muscle. or at least with part of it. e. SECONDARY RADIATION original experiments by GTJKWITSCH had shown that only the meristem. when severed from the bulb. There was only one alternative loft the radiation Irom the outside induced the exposed cells to produce some radiation of their own these "secondary rays" again induced the neighboring colls to radiate. until in 1932. will emit a radiation when it is exposed to ultraviolet light. in liver. This explanation was proved by many variations of the For some time. This ''secondary" radiation of the root ceases when the "primary" radiation docs. GTJRWITSCH found it to occur also in nucleic acid solutions. after being cut : . Yeast grown in the dark regained the property of radiation whether this had been grown in but only when the exposure was made after remaining in daylight for about two hours. Many illuminating details have been worked out by POTOZKY and ZOCLTNA (1928). Even the root tips radiated The only when connected with the lost their radiation They completely bulb. regardless of light or dark. in suspensions of yeast. of protozoa. were inactive. This accounts probably for a number of negative results by some experimentors. in daylight. In . GURWITSCH discovered that a from the bulb. of bacteria. as o. of FKANK and RODIONOW have shown. where the cells had waned to multiply. A chemical effect could hardly be passed along so rapidly. 1934) proved it to be primarily a photochemical phenomenon. the growing tissue near the tips of onion roots radiated while the older parts of the root. ALEXANDER. ANNA and LYDTA GUUWITSCH and others. oxidations. and so the effect was passed along root. arid L. it was believed to bo a original experiment. g. the root without losing in intensity. that light affects greatly radiation from some chemical Cr 2 O 7 f FeSO 4 (p. In search for an explanation. A. It seemed quite impossible that the original rays as such could have. i. 2 K C.

55% 8 mm. This exhausts the yeast so much. by oxidation. it unirradiated control.. but as far as 10mm. 44). However. 9 The ability of roots to produce and conduct secondary radiation is limited to a short time after the severing of tho root from the bulb. The increase in buds was 65% at the irradiated zone. 43% 9 mm. 86% mm. It will be shown later that the tips of onion roots are only secondary senders. of spreading. GTTRWITSCH irradiated such a culture through a slit 0. there was a distinct increase not only in the irradiated region. When tho percentage of buds was counted. the primary rays are produced in the onion bulb. Another experiment with starving yeast cells throw some light on this phenomenon. did not radiate. secondary radiation seemed to be glycolytic. this cannot be generalized because the secondary radiation from nucleic acid is not glycolytic. 80% 7 mm. Yeast cells may help to radiate imme- diately after being washed. but not after 40 Tho same authors could also show that the production of Freshly-cut secondary radiation exhausted the plant rapidly. but not 30 minutes later. 87% 3 mm.1 ram. The from starving animals which are free from glycogeii. A very recent illustration for such exhaustion has been given LATMANLSOWA by (1932) on the secondary radiation from nerves . primary radiations can be spread and transmitted to distant parts of the plant or animal body. This spreading of the mitogenetic effect can be plainly shgwii with densely grown agar surface cultures of yeast. supports this view. the organisms will still produce secondary radiation under the influence of an arc light spectrum. distant. effect after 30 minutes. Even then. 2 100% mm.110 ail CHAPTER V fact that livers these cases. 79% 4 5 mm. By this mechanism. that one hour later. has produced 41% less buds than the Exhaustion has also been demonstrated with chemical solutions (see p. 10 mm. from the border of the irradiated area. wide. POTOZKY and ZOOLINA (1928) found a positive 45 minutes. roots which gave strong secondary effects during the first 5 minutes of irradiation showed no reaction after 10 more minutes of exposure to monochromatic light of 2020 A. and at a distance of 1 mm. it was 33% 25% respectively Experiments with larger irradiated surfaces gave the same amount 12 mm. 80% 6 mm.

30 minutes.m. These two slits were so arranged on the rotating disk that after the primary rays had fallen upon the root. too. 111 The sciatic nerve of a frog was Another yeast block. 40A shows that the induction effect of every 5 minutes. but decreases after 5 minutes. ANNA GTJKWITSCH (1931) made some accurate measurements with onion roots. one nearer the center through (fig. A: the 40 minutes. only the angles between 25 and 50 gave positive. the disk had to be turned through 50 before the secondary rays from the meristem could fall upon the detector. however.. is given 10 minutes rest after which irradiation is continued. This central angle was varied from 20 to 85. is not characteristic of nerves It can be duplicated with cell-free solutions (p. and will become much more readily exhausted This phenomenon. the nerve is given a ''rest" for 10 minutes. rays secondary Figure 40. which the primary rays (from a yeast culture) fell upon the older part of the root.SPECIAL PROPERTIES OF MITOGENETIC RADIATION after mitogenetic irradiation. But the nerve is still than the only. 40 B). showing exhaustion of the nervo. suggested measuring the rate of travel. serving as was placed near the irradiated part of the nerve so that was exposed only to secondary radiation from the nerve. it Fig. and does not appear any more upon continued irradiation. and another towards the periphery of the disk through which the radiation from the meristem of the root fell upon the detector. This block was changed detector. irradiated by a yeast culture. is very strong at first. but not to primary radiation from the yeast. minutes minutes and after approximately . If. With a definite speed of rotation of 3000 r. by removal of the source of irradiation. 43). "tired" first. it will recover sufficiently to react again upon renewed irradiation (fig. p. The observation that secondary radiation could be passed on over considerable distances. Secondary radiation from a nervo exposed to continuous yeast first irradiation. exhausted after 35 minutes. After some preliminary experiments by ALEXANDER GUKWTTSCH. it has disappeared. . B: a nerve. 41). by means of a rotating disk This had two windows..

The accuracy of the in method was thus greatly increased. at 3000 Figure 41. and this is This corresponds.. This was sufficiently to 3 intense to permit the reduction of the window for primary radiation and that for secondary radiation to 1. The method was exactly the same. of root. At right. CHAPTER V This signifies that a certain time (0. is conducted to another part and is emitted there. This is good agreement with physiological measurements on the rate in of conduction of nerve impulses. Recently. the time required for the secondary radiation to pass the additional The rate of conduction is therefore about 30 meters per second. In the experiments with secondary radiation of onion roots. LATMANISOWA (1932) has measured the rate of conduction of secondary radiation in the sciatic nerve of the frog. except that the 2200 A area of the copper arc was used as primary source.5 cm.0022 seconds) must pass before primary radiation falling upon one part of the root. to between 40 was hereby increased and 70. Then. radiation was observed from the same side of the root which had been exposed to primary radiation. Allowing the same time for transmission through the other 2. p. the distance was increased to 5 cm. and detector D. to 0. The total time corresponding to The difference. and the rate of conduction the nerve was found to be 303 meters per second. onion root. Kotating disk for measur4 ing the rate of travel of radiation in secondary onion roots. which means an average increase of 15.00140 seconds.112 results. The central angle for positive effects r. of root is 0.00166 seconds. the total time required for transmission through 5 cm.5 cm. These values refer to a condition over 2.00306 seconds. Further experiments showed . was required for processes other than conduction.00083 seconds.m. 0. such as local reactions at the points of absorption and emission. side view showing position of primary sender S.5 cm. 2. the average angle of 55 is 0.

they were separated and tested as senders one was tested at once. None produced an increase in cultures lose their .SPECIAL PROPERTIES OF MITOGENETIC RADIATION 113 ease. it seems highly probable that at Pro to plasma -Monographien IX: Rahn . When mitogenetic rays fall upon any cell. effect upon new detector After 15 minutes muto-induction and 15 mi- nutes recovery After 15 minutes muto-induction and 30 minutes recovery . 45). as the following data show: Mutual induction during 15 minutes . 300) that radiating power rather readily when they are them- selves exposed to mitogenetic rays. before it was found that in certain cell-free solutions. the generation time of which must have been at least one hour under the condition of the experiment. Perhaps 30 minutes is too short a time for recovery of yeast. This does not alter the explanations materially. but that that the radiation effect was transferred longitudinally with great no conduction occurred transversely across the root to the opposite side. 1-8% . polarized mitogenetic rays have an enormously greater biological effect. the same effect can be obtained (see p. however. 37% 2% 0% increase over control After 15 minutes rnuto-induction.. strong radiation has been obtained from the unexposed side (LATMANISOWA 1932). 8 . In the nerve.. After 15 minutes. 44). ^our yeast agar blocks were placed so as to irradiate one another. and that apparently. common light does. however. the others after 16 and 30 minutes. It is brought about by some unknown influence of ultraviolet rays which induce certain complex organic substances. It only means that secondary radiation need not be connected with life processes. p. budding. Even young. The first mutual induction was plainly noticeable. actively radiating power when exposed to their own wave lengths. WOLFF and RAS point out that mitogenetic rays become polarized by reflection like chemical reactions in cells or In their latest publication (1934 a).. Practically all these facts had been discovered... This phenomenon itself can bo at least partially explained by the experiences with secondary radiation (see p. An important phenomenon for the explanation of mitogenetic effects is the observation cells or tissues lose this (GuRWiTSCH 1932.

with photographic plates. the reciprocity law (double intensity means half as long exposure) does not hold with very low intensities. and thus will become polarized. It is not possible. Table 46 p. 145. If the number of impacts induced by biological radiations is expressed in percentage of the stray radiations of the surroundings. Examples may be found in Table 12 p. D. and recently also by WOLFF and RAS. This does not affect the intensity of the biological radiation at all. 79. never possible to use it for measurements of intensities. The "induction effect" as the increase in the exposed yeast over that of the control. 157. It INTENSITY OF THE MITOGENETIC EFFECT the effect has already been stated repeatedly that the intensity of is not proportional to the intensity of the radiation. effect" The customary way of comparing intensities is to compare the minima] time of exposure (threshold time) required to give definite mitogenetie effects. and Table 49 p. to foresee all the consequences of such polarisation. at the present moment.114 CHAPTER V least part of this radiation will be reflected from the cell walls. and that WOLFF and RAS polarized rays have an enormously much stronger biological effect than the ordinary radiations of this type. The "mitogenetie as used especially by the Russian investigators has no It is not surprising that it was quantitative value whatever. 24) that. as explained on p. latter evident method of recording resu]ts becomes most applied to physical measurements. This method has been used repeatedly by the Russian workers. there are physical reasons to warn It has against quantitative conclusions from threshold times. it is utterly error of this The when meaningless from a quantitative viewpoint because this stray radiation (the background radiation) can be greatly altered by shielding the instrument with iron or lead. been pointed out above (p. One very simple reason for this is the usual method of recording the results. 44. expressed in percentages of the cannot possibly be used as quantitative measure. However. All previous measurements of intensities have become practically meaningless since that mitogenetie rays may (1934 a) showed easily become polarized. .

Real retardation by biological radiation can be measured only by decrease in the growth rate. that with roots as detectors. As early as 1928. in the same experiments. to unicellular detectors. we have 110 real controls. It must be remembered. is larger than that of the controls. shaded one. a smaller increase than in the control can only signify a retardation 8* . Strong physical light produced tho same depression in a few minutes. The measurement of the actual number of cells permits of only one interpretation. The yeast bud method appears to be the one by which "depression" is observed most easily. or it may mean no effect radiation) through over-exposure. and prevent or retard mitosis. really. or both. She observed a decrease of mitoses in the exposed side of the root as compared with tho opposite. 69).5 minutes and longer. radiation. a difference between tho two sides of the root may mean stimulation on one side. however. or retardation on the other side. there may have been retardation through over-exposure. the actual number of cells. But it is just these retardations by the yeast bud method which are frequently contradicted. self -contradictory a detectors sometimes give opposite results. however. warns against hasty conclusions. together with control roots exposed only during the last 2. by parallel measurements of the actual cell increase. 115 RETARDATION THROUGH RADIATION It seems quite probable that an overdose of radiation might produce the opposite effect of mitogenesis. i. Table 34 shows SALKIND'S experiments (1933) with rat blood With exposures of 2. This was interpreted as "exhaustion" by too much radiation. STLSSMANOWITSCH irradiated onion roots biologically for 12 hours and longer. In this case of over-exposure.SPECIAL PROPERTIES OF MITOGENETIC RADIATION E. e. and stimulation (through secondary on the shaded side.5 to 3 hours. This can only mean that the yeast bud technique fails to indicate the true growth rate (see p. the percentage of buds showed a decrease against tho controls. GUKWITSCH called this is phenomenon mitogenetic de- term. We must therefore turn to other detectors which permit absolute controls. but the circumstance that different pression which. recorded rather frequently.. it should be Observations of this kind have been "depressed mitogenesis".

as well in the experiment of Table 34. WOLFF and RAS it the normal reaction after continued irraanalysis of this phenomenon. no mitogenetic effects were obtained. 77) consider diation. after stimulation followed depression. SALKIND (1933) senders. A more detailed investigation revealed a certain periodicity. again stimulation could be observed. Such cases are also reported. the depression did not increase. "Reversal by Light: With a we get a latent image which on development yields a negative. the image becomes positive instead of negative when and it is developed. If the exposure is lengthened considerably. In his observed that with prolonged irradiation. CHAPTER V Induction Effects from Intermittent Radiation of Rat Blood. This was the case with physical as well as biological In each instance. but after depression. probable that the cycle may be repeated indefinitely. (Table 35). and by the Increase in Total Cells of the growth rate. could be found in SALKIND'S data. although . No definite periodicity GUKWITSCH as well as WOLFF and RAS maxima this observation of several (1933c) have verified at widely different exposure times while between these maxima. while still further exposure will produce a second negative. if radiation continued. as Measured by the Relative Increase in Yeast Buds. irradiation was applied intermittently. (see p. A striking parallel exists between this effect and that of the photographic plate. as may be seen by the following quotation short exposure to light from NEBLETTE (1930).116 Table 34.

An experiment was started with two yeast agar blocks mounted 6 cm." GURWITSOH (1932. until after 5 to 8 minutes. on the movable substage of a microscope. apart. but was delayed for a short time. 117 Periodicity of the Mitogenetic Effect Measured by the Increase in Cell Numbers with Yeast owing to the enormous exposures required. the effect was not at once harmful. no one has been able to go past the second negative stage. 219) gives some examples whore. p. This has been demonstrated most simply in experiments on mutual yeast irradiation (GuRWiTSCH 1932. 263). p. the two blocks were made to approach one another. F.SPECIAL PROPERTIES OF MITOGENETIC RADIATION Table 35. He calls this "secondary depression". ADAPTATION TO GRADUAL INCREASES IN INTENSITY When the intensity of radiation is gradually increased from below the threshold to a value which would produce a strong effect under usual conditions of exposure. Very slowly. This distance is too far to produce a mitogenetic effect. after too long an exposure. no induction takes place. The reactions which result in reversal are still obscure. acceleration being noticeable followed by a distinct retardation. they remained in this position for some . they were very close together .

and gradually shaded them again. This disk was mounted so that in rotation. SPECIAL PROPERTIES ETC. set CHAPTER V.118 time. it gradually exposed the two agar blocks to each other. This was sufficient to prevent induction. the yeast of the equally long exposed. . The total irradiation time corresponded to that of another with the same yeast culture which had been placed in the final position at the start. but gradually nearing agar blocks paralleled the controls. the regular mitogenetic effect is observed. The time required for this slow approach must be about 5 to 6 minutes. When it is reduced to 3 minutes. While this latter set showed increases of 40 to 50% over the controls. The same phenomenon was obtained when an elliptical disk was rotated between two yeast agar blocks.

number the most of experiments with familiar of all detectors. From a certain rhythm observed in the division of the sperm cells of amphibia. they are to him probably However. he succeeded in proving it with onion roots. Mitogcnetic radiation was considered at first merely from the cytological viewpoint. the mechanism by which short ultra- violet rays affect living cells is not understood. when SJEBEJK. The number of not offering of mitoses in different roots oven from the same bulb varies greatly. as an emanation produced somehow in the very complicated process of cell division. and all other effects of secondary importance. they have the great disadvantage perfect controls. Hence. but must be of a physical nature. these were the large first detectors. This chapter does not offer one theory. the primary effect which is the increase in the number of mitoses. From the mode of action. These rays were not discovered by chance.CHAPTER VI ANALYSIS OF THE MITOGENETIC EFFECT At the present time. GURWITSGH has always distinguished between the "Ureffekt".T proved this radiation to be emitted also from purely chemical oxidations. GURWITSCH (1922) predicted a factor which controlled cell division. the physico-chemical viewpoint entered into consideration. All of GURWITSCH'S speculations and explanations start with the results obtained with onion roots. we lack a clear conception of the working mechanism of these rays. and in plant roots after special treatment. he concluded that this factor could not be chemical. and in 1923. but presents a number of attempts to account for the various phenomena observed. and since he made a them. The theories which were developed during the "biological stage" of the discovery have never been fitted completely into the physico-chemical facts observed later. Only after 1928. The customary method is to .

or changed. unicellular forms as the first objects for an attempt to interpret the primary mitogenetic effect. The best method for this purpose is the yeast bud method* by TV THILL and RAHN (p. . multicellular organisms. before old can divide again. This In period of adjustment is called the lag phase (see p.120 CHAPTER VI we cannot be use the shaded. g. but are familiar with yeasts and bacteria. 68) where all cells are of the same age no secondary radiation from older cells complicates the analysis. 67). able to multiply at the normal rate. The old cells need from one to several hours before they are "rejuvenated". but also for cultures of yeasts and bacteria (see e. old cells can be induced to cell division is by wounding. e. This holds not only for the larger plants and animals. (3) (4) The minimal The harmful A. rejuvenation brought about by transferring the old cells to a fresh medium. THE NECESSITY OP A PARTICULAR PHYSIOLOGICAL STAGE This neces&ity will not appear improbable to a cytologist. this factors. Any interpretation of the mitogenetic effect should account at least for the most remarkable facts observed. In fact.t they are affected. but at all certain that irradiation of one side does not influence the cells on the opposite side as well. or neoplasma in animals. The following have been selected as the most important: (1) (2) The necessity of a particular physiological stage of the cell. HBNKICI. 1928). effect of over -exposure. and the percentage of buds is a true measure of the rate of cell division. When any cell changes from the stage of active cell division to the resting stage. The relation between the intensities of ra-diation and of effect. intensity required for an effect. With unicellular organisms. i. cells is caused by some external or internal These factors must be removed. The authors of this book who have made no experiments with onion roots. unexposed side of the same root as control. prefer to start with these simple. REITER and GABOR claim tha. resting cells of the same tissue. or by unknown outside stimuli as in the case of gall formation in plants. The cells of growing tissues are morphologically and chemically quite different from the old. .

the lag phase. 68) where the yeast produced buds very rapidly when exposed within half an hour after being transferred to the fresh nutrient medium. Perhaps. The stage of active cell division.ANALYSIS OF THE MITOGENETIC EFFECT 121 It would not appear probable that a resting cell can be induced to a new cell division by a short. the ultraviolet. not during the phase of constant growth rate. it does not seem likely that the factors which induce ageing could be removed by irradiation. whether exposed as such or diluted 1 10 000. but not later. such cultures should rcart cells irradiate one another. as a rule. cytological. They explain it by the assumption that the rapidly multiplying and being so close together. WOLFF and RAS (1932) make the unrestricted statement that mitogenetic effects can be obtained only during the lag phase. weak irradiation. 24 hours old. p. e. Very striking are the results of TUTHILL and RAHN (Table 21. might . their stronger than that from any external source. while older cultures : gave very pronounced effects. they should also react to an external source when they arc widely The latter was tried dispersed so that the cells are far apart. With yeasts as well as with bacteria. This is borne out by experiment. chemical or physical. but failed to respond an hour later. but the explanation by WOLFF and RAS is doubtful. and consequently the intensity of radiation. without success by FEKGUSON and RAHJS (1933). Mitogenetic effects are not. Their experiments support this claim. the stage of rejuvenation. which is necessarily weakened by distance and absorption. a certain chemical process in the rejuvenating cell is greatly stimu- The explanation may be It lated . observed with old cells left in an old environment. mitotic one be but that stage can take advantage of the may energy introduced into the cell by this radiation. is one of strong response. does not appear very favorable either for mitogenetic effects. is own radiation to outside irradiation at low temperatures where the rate of metabolism.chemically the ageing process of a cell. i. g. though the control had not as yet started to produce buds. is weak . The fact that actively dividing cells do not respond readily to mitogenetic rays is thus verified. never reacted upon irradiation. Though we do not really understand physico. If this explanation were correct. There seems to be one vstage during rejuvenation when the cells are most susceptible. by means of a chain reaction. e. Cultures of Bacterium coli.

The volumetric method as described by KALENDAROFF (p. This may be the same mechanism which.122 set CHAPTER VI potential necessary for normal cell functions the candle which then keeps on burning as long as the cell feeds normally). cell division might give us the clue for the mitogenetic There seems to be another stage where cells respond. TUTHILL and RAHN. long to permit one more cell division. such as the onion root. Apparently. The description of the physiological condition of BAKON'S yeast plate (p. B. namely immediately before entering the resting stage. possibly. When freshly prepared detector plates were exposed for different lengths of time. cell. for a short time. the following percentages of buds were found (after 2 hours of incubation) : . the cells. It appears that the difference between rejuvenation and active effect. it must be kept in mind that so far. the percentage of buds was not at all proportional to the length of irradiation time. We could not expect this with detectors involving secondary radiation by old cells. RELATION BETWEEN THE INTENSITIES OF RADIATION AND OF EFFECT been one of the most annoying puzzles of mitogenetic experiments that there seemed to be no proportionality between It has cause and effect even when polarisation is excluded. at the point of going to rest. This need not necessarily involve a mechanism different from that assumed It may well be that in the ageing the additional. Or. 73) appears to make use of this stage. sufficiently in the rejuvenation process. where mutual cell influences are practically excluded. the cell wall becomes transparent to these rays only at one certain stage of development. 72). under the more favorable conditions of rejuvenation. or the But even with the yeast plate of volumetric yeast method. This may also bo the cause of the mitogenetic effect in onion roots (see p. 129). Whatever up the reduction (light be the explanation. is stimulated so greatly. and so does HEINEMANN'S hemacytomcter method (p. BAKON'S yeast plate. properly dosed energy from the sender prevents a certain phase of the ageing process. 66) suggests this strongly. are stimulated to at least one more cell division by irradiation. the later stage of active cell division does not seem to be greatly influenced by these rays.

P = 0. This may be explained by the recent discovery of WOLFF and HAS (p. but that of TUTHTLL and is bability that a yeast cell of RAKN has single cells. Neither of these other times showed any great effect. The intensity of secondary radiation does not depend so much upon that of the primary source as upon the chemical composition of the medium. If there were proportionality. the 10-minute exposure should have produced an increase of approximately 7%. however. either directly or through quartz. but senders appears to be about 100 to 1000 quanta per cm 2 per second. The entire detector plate begins to emit radiation as soon as it is exposed to a sender. 15% more than the control. Since we find the strongest effect under this . tJ.00000042 x 1000 = 0. All hope for proportionality must be given up in this case. THE MINIMAL INTENSITY REQUIRED FOR AN EFFECT measurements of the intensity of mitogeuetic rays are the order of magnitude of the strongest All very inaccurate. does a biological measurement of intensity seem at all possible. The detector plate by BAKON is completely covered with cells.00042 The probability of being hit in one minute is 0.ANALYSIS OF THE MITOGENETIC EFFECT 123 The sender was a 6 hours old yeast of surface culture. The exposure 20 minutes produced a strong effect.0252. assuming JLI 1000 quanta/cm 2 /sec. uniformly dispersed radiation before each of the yeast cells is likely to be hit by one quantum of ultraviolet light. Only with a medium which does not produce secondary radiation. The pro6x7 is hit in one second. It will require 40 minutes of continuous. and the 40-minute exposure a 30% increase. 43) that nutrient media produce secondary radiation when they have been in contact with microorganisms.

it seems safe to assume that protoplasm generally will respond in this way. e. the absorption of one quantum of ultraviolet of fairly definite wave length. 37). 1930. This induces a state of excitement. many processes going on automatically and exothermically. will produce secondary radiation. He claimed. but they may be explained in some other way. Irradiation will then set the entire mass of agar radiating. it can be well imagined that such a release will start many wheels turning. One definite mitogenetic wave lengths. and it is quite probable that a more rapid cell division may be brought about. until cell division is in slightly different terms. Possibly. and the number of quanta thus produced. This is. since certain solutions such as blood serum. and seems to assume oven now that no cell division is possible completed. how can A (p. it arrangement at 20 minutes would seem that one to produce the mitogenetic effect. yield living is of the cell at the proper cytological stage. however. If all namely of 1900. makes the above explanation too simple. if the cell absorbs one or a number of quanta of ultraviolet. the assumption of a single quantum acting has become unnecessary. energy of fact. without this external. Considering the systematic arrangement of all molecules in the cell. or absorbed by the cells. cells yeast cells produce this wave be stimulated by the same wave lengths from .124 CHAPTER VI (see above). or broth in which bacteria have lived or are living. Then. i. the entire cell begins to radiate. even improbable. It is known that yeast cells. within the cell. 1910. ultraviolet stimulus. not visibly. There has been a good deal of speculation as to the mechanism by which one single quantum could affect the cell so greatly. cannot be estimated. The upon which the yeast is spread will gradually become transformed into a secondary sender by the very presence of the yeast. Every fermenting yeast cell liberates. but measurably. 1950 and 2170 length. or onion root cells. or pulp of upon the cell raisin agar secondary radiation. provided that the cell tissues. GURWITSCH'S original conception of the mechanism of the mitogenetic effect. It would appear that a proof against single -cell cultures of bacteria and yeasts were this assumption. quantum per cell is sufficient However. the synthetic powers work to a certain morphological and physiological culmination which can be released only by a very accurately measured impulse.

effect I). but that only those cells which . But the assumption is not justified. the stimulating effect which the first quantum had produced. or excitation. They also will become exhausted upon long -continued exposure (p. Whatever the explanation. RAHN (1928) and RAHN and BARNES (1932) found that old yeast cells. Nothing is known about the chemistry involved. 43) that too long exposiu-e destroys the ability of a solution to produce secondary radiation. fermented strongly within sugar solution. it has been shown that recovery is slow. appeared to be counteracted by the ab- The harmful sorption of a second quantum. 110). but the assumption of an equilibrium. Now we realize that the first as well as the second quantum are probably multiplied manyfold within and outside the It cell. After a day or two of rest. Moreover. If we A could make the cells show no metabolism. at least emit no ultraviolet. have already seen that very likely they are all capable of secondary radiation. stage of most active multiplication which is also that of most active metabolism. compressed baker's yeast as well as beer yeast stored for several weeks at low temperature. 10 minutes after being placed in it is certain that the mitogenetio does not occur merely through the increase in energy content of the cell. fite best into our present conceptions of life functions. If we assume that all cells are brought to a state of radiation. We Something similar to these effects may happen in the cell. or the assumption that during the period of sensitivity. then the mitogenetic effect could be easily explained. this property returns.ANALYSIS OF THE MITOGENET1G EFFECT an external source ? 125 We may return to the first of our fundamental facts that only at a certain cytological stage. Before this was found. disturbed by irradiation and slowly re -established after discontinuance. THE HARMFUL EFFECT OF OVER-EXPOSURE effect of over-exposure is more easily understood by the secondary radiation of the cell and of the medium. cells will react to No reaction has been observed at the mitogenetic radiation. very definite and strong response was obtained at the first stage of the rejuvenation process. has been shown (p.

Since exhaustion of solutions lasts for a day or two (p. Table 36. it would be difficult to explain the periodical alternation of stimulation and depression observed with long. etc. p. and eventually a temporary interruption of mitosis. irradiated by an agar culture of Bacterium coli for 30 minutes .126 CHAPTER VI are at the proper cytological stage can respond to this stimulus by dividing more rapidly. 44) strong positive : 5 minutes exposure. 7 or 8 minutes. This is doubtless caused by the uniform age of all cells in this kind of detector while BARON'S yeast plate with effect with cells of many different physiological stage*. The time during which mitogenetic effects can be observed seems to vary with the detector. The data of WOLFF and RAS (Table 25. 6. 116). 43) and exhaustion of cells for hours (p. p. E.continued irradiation by SALKIND (p. and still show stimulation of growth when transferred to a fresh medium (Table 36). on account of exhaustion of certain chemicals in the cells. 110). none whatever with 4. by the prolonged secondary radiation. the cells of other stages will soon become exhausted. 3-day old culture of Bacterium coll. has a more gradual range of response and tolerance. The sharpest limitations observed aro those by WOLFF and RAS (Table 12. This would mean at first a normal rate of cell division. 77) also seem to indicate recovery from depression though irradiation is continued. STORAGE OP MITOGENETIC CHARGES (1933) observed that bacterial cells FERGUSON and R\HN could be kept in their old environment for 2 hours after exposure to mitogenetic rays.

cells are lying closely side by side. though absorbing ultraviolet. may be observed of 10mm. We can only assume that the cells were changed chemically. However. After that. which then. structure of the BARON yeast plate. mitogenetic effects cells. The limited space of this book does not permit detailed quotation. Some cells. secondary radiation. F. This eventually help to locate the exact process by the mitogenetic impact. On removed from the exposed quanta will penetrate into p. 66. they are exhausted and remain inactive. on its part. here. these secondary younger and stimulate them to bud formation. has been described in detail on p. especially since the complexity of this detector leaves too many possible explanations. rejuvenation set in at once. They can emit only a definite (though unknown) amount of radiation. The absorption of one quantum is sufficient to induce secondary radiation in a surface cell provided that the cell is not too old. However.ANALYSIS OF THE MITOGENETIC EFFECT 127 A The storage of energy as such appears out of the question. decreases therefore gradually during exposure. with old on the surface. analysis of mitogenotic effects in the BAKON yeast plates. that the unknown process of rejuvenation was released. the intensity decreases very rapidly with the distance from this cell. but could not materialize under unfavorable environmental conditions . . continued internal secondary radiation is also impossible. since in this detector. MECHANISM OF THE BARON YEAST DETECTOR GUEWITSCH devotes 50 pages to the In his monograph. 110. multicellular detectors. The old surface cells react upon irradiation by producing beyond the stage of cell division. Secondary radiation consists in the emission of a number of quanta. for the duration The number of these secondary senders of the experiment. and with normally-dividing cells at the bottom. since it is a good parallel to we shall at least give a brief summary The complex cells. Since the emission takes place in all directions. as soon as this situation was observation released may altered. a quantum emitted through secondary radiation may be cell absorbed by another reactive new secondary quanta. emits it has been shown that in these yeast plates.

that if two or more quanta strike the surface of the detector at The mitogenetic stimulus the same moment. we look at the mitogenetic effect as a photochemical one. but hardly in three- G. Bacteria and yeasts in the detectors mentioned have a very large amount of food at their immediate multicellular detectors which command. Thus. Such interference is imagineable in a onedimensional system. cell electro-magnetic quanta striking a effect. pro- duce no effect . though the total energy cell is increased. there is less equalization. even after adequate stimulation. the potential lacking. and therefore a relatively stronger mitogenetic effect must be expected from biological necessary to initiate will occur division is senders. obtained a strong mitogenetic effect. while in biological sources. We would not expect the reaction between hydrogen and chlorine (p. Such interference seems rather improbable if and below at the same time. g. Miss FERGUSON. from all sides will not produce a mitogenotic consists in the one-sided discharge GUBWITSCH assumes (release) of a neighboring secondary sender. the food may be insufficient for continuing at a subsequent normal rate of cell division. 46) to be suspended if the gas mixture were irradiated from two opposite sides. . MITOGENETIC EFFECTS IN MULTICELLULAIl ORGANISMS is There one essential difference between the cells of unicellular and must be kept in mind to prevent misleading generalizations. sequently. in the latter case. By irradiating a liquid bacterial culture in quartz from above in an unpublished experiment. e. the peripheries of the streams of secondary quanta may be partly superposed and thus by interference. the food supply is limited. a nerve fiber. dimensional media. e. g. there may not be enough food readily available to permit rapid cell division.128 CHAPTER VI GURWJTSCH considers the "mitogenetic field" similar to an He believes that a uniform stream of field. or after a premature mitosis. whereas in tissues. in onion roots or in the cornea. Conradiation comes from many cells unevenly distributed. Such "equalization" more commonly with physical sources of ultraviolet because the light is more uniform. The two radiations did not cancel.

the shaded zone indicates the sensitive root. They distinguished the a-type. the authors raised two questions: does the exposed side of the root differ from the Pro to plasma . The latter have based their intrepretation upon a cyto- logical analysis of the onion root which deserves attention because it suggests a close analogy of the root with the BARON yeast The root can be divided into transverse cross sections be designated by the number of successive cells from each section to the tip. e. a 2 ripe nuclei. Distribution of various cells in the onion root. The first newly-born type could be subdivided into 3 nuclear stages a^ The various mitotic stages. 65) and by REITER and GABOR do not agree. part of the root. 3 distribution of these types is shown in Table 37 and fig. born cells. at newlyft for the four types in different parts of the root. Below: cross section through the with its vascular bundle . In the region which is 125 or more cells distant from the : = = The zone dividing stages arc very rare. the resting cells amounting to less than half of the dividing types. 42. measured by the number of cells. Above: distribution cell -elongated. The interpretations of the onion root effect by GURWITSCH (p. To analyze the How muoh effect. a3 mitotio stages. which may Figure 42.Monographien IX : Rahn 9 . In the root tip. resting cells. short activelygrowing cells. cells the successive the sides.ANALYSIS OF THE MITOGENETIC EFFECT The one 129 multicellular detector that has been studied cyto- logically is the onion root. practically all cells are resting . This number is fairly uniform whether plate. division. i. and the /?-type. resting cells. nuclei. tip. elongated. the abcissa represents the distance from the tip. no colls of the All cells arc in active /J-type. rays is The most are counted in the center of the root or along sensitive region in respect to mitogeiietic cells approximately 50 to 70 from the tip. REITER and GABOR studied the distribution of cell types along the root. no resting cells are found. as far as about 25 cell layers upwards. a2 ripe nuclei. a = short dividing colls. of mitogenetic sensitivity contains cells of all types.

and this stimulus is transmitted by secondary radiation of the old cells to the young. . without irradiation. ANALYSIS OF THE MITOGENETIC EFFECT Distribution of the Nuclear Stages Table 37. shaded side differ ? and ? How much does the exposed side from the normal The latter question could be answered only by analogy. Jt will be seen later that the onion root is irradiated continuously from above. by the bulb. of sectioning and counting. Both consist of many cells.130 CHAPTER VI. opposite. all cells The conclusion was that "under the method There described. On the opposite side of the root. but only in about half of all his (GuRWiTscn's) and also of ROSSMANN'R experiments. GUKWITSCH (1932) does not agree with this interpretation. dividing ones at the bottom. and points out that a decrease of mitoses in the opposite side of the root has been observed occasionally. He criticizes the method of counting "ripe nuclei" only. both have the oldest. more cells go into the resting stage than would normally do so". is a certain similarity between the onion root thus closely-packed on top. He doubts the possibility of accurately distinguishing between nuclei of resting and dividing cells. and produce again ripe nuclei while normally. non-dividing cells and very young. a certain percentage would go into the resting stage. growing cells in the root tip. and the BARON yeast plate. influence of mitogenetio born during the experiment remain in the actively-dividing stage. and this can be accounted for by the irradiation.

The loft-hand curves represent the multiplication of the bacteria and the gradual accumulation of lactic acid from sugar.CHAPTER VII THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS IN BIOLOGY. (1) UNICELLULAR ORGANISMS It this Emission at Different Physiological Stages: had been emphasized throughout book that intense mito- genetie radiation lias been observed almost exclusively in young. or only rather weakly. The bacteria starting with 38 000 cells per cc. MEDICINE AND AGRICULTURE A. while fullgrown ones do not radiate at all. proteolysis. as it is apparent that there can be no other important source of radiation. to the intensity of radiation of the culture. can have emitted a noticeable degree of mitogenetic radiation only between 12 and 24 hours. the radiation per cell might have been quite as strong or stronger. though more than a billion per cc. Fig. while afterwards the cells. Under optimal conditions of food and temperature. 9* . glycolysis. by glycolysis. and the other resulting from general cell metabolism. 43 shows the development of a culture of Streptococcus lactis at 21 C (RAira. p. 1932. we must distinguish between two sources of radiation. aetively growing tissues or cell cultures. have ceased to produce acid. both these functions should cease almost entirely within 24 hours after transfer. According to GURWTSCII.. such as oxidation. Before that time. A culture of yeasts or bacteria should be a good sender as long as either the cell division is rapid.. but the number of cells was too small. 401). and therefore to radiate. or the metabolism remains active. The right-hand curves show the increase* These increases must be proportional in each 3-hour interval. the one emitted at the moment of nuclear division.

the theoretical deductions are essentially verified WOLFF and RAS (1932) that an agar surface approximately 18 hours culture of Staphylococcus aureus remains actively radiating until old. medium is continuously changed by property as secondary sender may change. by the observation of However. Development of a culture Full line: of lactose of Streptococcus lactis in milk. cells. i.grown. the period of may also be longer. must follow the culture lines of 43. radiation inoculation. c. This a priori deduction is made doubtful by the secondary radiation of the medium in which the bacteria grow* The actual radiation of the bacteria themselves fig. At lower temperatures. was decided in the affirmative by a simple experiment of the authors with bakers' yeast suspended in sugar . do not result in an inhibiting product active radiation With processes which like acid. since the nutrient its the bacteria. cells Age of culture in hours.132 CHAPTER With a heavier VII flooding the surface of bacteria. nor in metabolic products.j dotted lino: mg left: total cells in 100 cc. e. and sides. In- tensity of secondary radiation may not be proportional to intensity. e. is i. when there is no further appreciable increase in cells. number of decomposed per cc. Whether metabolism without cell division can produce mitogezietic rays. When the cultures are lull. g. primary and be- Figure 43. and lactose consumed during these 3 hour periods. the emission by the entire may not. the number of is less. by an agar plate with a suspension of yeasts or begins sooner because the "active mass". total lactose fermented. the intensity but the phase of active radiation is prolonged. greater. right: cell increase for each successive 3-hour period. the culture ceases to radiate.

+4. No increase in emission could be detected during regeneration. If dissected. +19. p. . from Paramaecium . are given in Table 38. the well-known fresh-water polyp. however. 133 Mitogenetio Effects from Protozoa in Glucose Solution solution. During the first hoiir. i. many hours would be required to overcome the lag. the irradiated culture. interesting set of data on the radiation of Hydra fiisca. but produce a secondary radiation. even if slight growth were to occur. it required about 15 minutes before The mitogcnotic effects observed radiation became noticeable. ETC. +47 +21 +15. +20. the hypostom and the budding zone radiate while the other parts do not. 64) the following 0. as sender for . +38. A very interesting claim has been made by Acs (1932). +3. +1.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 38. e. +40 spectral light . tions because too This yeast cannot grow under the experimental condimany cells are present. and SAMARA JEFF has been given by BLAOHER that a pulp of the entire found (1930). obtained: Zoglina from Opalina irradiated 1932. no radiation from cell division An is possible while alcoholic fermentation starts at once.. +12 by blood frog heart . +3. According to (quoted after GURWITSCH mitogenetic effects were. irradiated Radiation could also be obtained from protozoa when glucoso was added to the culture.. They organism radiates.. . +22. +6. therefore. Ho succeeded in increasing the mitogenetic effect of muto-induction of bacterial cultures (Bacterium typhi-murium) by using the detector of one day. +50 3 +2. +13. The common infusoria do not appear to radiate under normal conditions.

The rate of growth of old cultures can be increased for several successive transfers. morphologically and physiologically (2) Reaction Upon Irradiation: when transferred from This results in a very slow growthrate during the hours after transfer.. He inoculated increasing amounts of yeast into flasks of sugar -peptone solution. so that all the decimal gradations from 2095 to 20949750 cells per cc. The proof is not complete. Without accurate statements concerning the number of cells in the sender and in the detector. The bacteriologist calls this the lag phase. the stronger must be the effect. rejuvenation is slow. The average distance distance is computed from the equation = ~ cell diameter 74.914 mm.192 0. These different inocula resulted in very different lag periods. without any irradiation. It is well known that young old bacterial cultures require a longer time to start growing than ones. do not start growing at their maximal growth rate. absolutely correct only with spherical organisms.086 0. an old culture to a fresh medium. with yeast. J Jt is . 120). 8 0. best example is very likely that of HENKICT (1928. until it could be explained as a simple stimulating effect by mutual irradiation of the cells. were present.134 the next experiment. the average diameter as 6. and single-cell cultures frequently die. however. CHAPTER VII His data show a very consistent increase in the mitogenetic effect through 6 to 7 such transfers. This observation had been very puzzling to biologists since tho opposite should be expected. The old cells undergo a rejuvenation process. 612 1224 2436 cell 2 4 8 hrs. PENFOLD (1914) and others that bacteria after being transferred from an old (see p. The times required to reach the maximal growthrate were: The I) Table with an inoculum of 20 949 750 cells per cc.4 //. The volume of one 8 yeast cell is taken as 1 18 ^ . Lag Period Average cell distance 1 ) 2094975 209498 20950 2095 1 ) . Bacteria* and yeasts.04 ||/~~ / \ cell volume in 100 cc. The closer they are together.422 0. the quicker the recovery.036 0. first culture rejuvenate more rapidly if many cells have been transof fresh ferred while with a few cells in a large volume medium. It has been observed by RAHN (1907). no definite conclusions can be drawn.

as a result. 135 Growth Acceleration by Mutual Irradiation with Saccharoniyces ellipsoideus To these proofs that the lag phase (jells. he showed that is this difference disappears when gelatin added (Table 40). muto -induction was prevented. . and. and while all other conditions of life remained the same (gelatin contains no nutrients for yeast). After having formation in fresh yeast cultures begins sooner is BAKON larger (Table 39). 59). ftelatin Table 40. the lag phase became independent of the cell concentration. Prevention of Growth Acceleration by Absorbing the Mitogenetic Kays by Means of Gelatin Number of Buds per 100 Cells Inoculum start after 3 hours after 5 hours 7 hours after after 8 hours after 10 hours after 12 hours absorbs mitogenetic rays very completely (see p. is shortened by the mutual irradiation of shown that bud when the inoculum (1930) added another.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 39. ETC.

? ^ r} A A <M CO ^* <O 00 .136 CHAPTER VII *-* ^ a o | 2 o a 1 l F t o I i I o t oo d O W !| S -go rH rM S - .a O rH O H O S ^ TJ 0) i 5 Hi ^3S ic i- a a s a a s ^ ^ efl i I O O co O CO S8 "8 00 rH R^ .

appears a duplication of the lag phase of bacteria. came from the same 1 cc. verifies this comparably. with culture B. 137 Offspring in 44 hours of the protozoon Enchelys farcinem in hay infusion drops of different sizes The following experiment by FERGUSON and RAHN (1933) from a different angle. it from Vioo cc f ^ nc concentrations. the inhibiting volume is 0. In large drops. single individuals of Enchelys /arcinem multiplied but very slowly or not at all in small drops of hay infusion while the rate was quite large when two this or more individuals were in the same drop. 1 cc. and then diluted in broth The cells in all three cultures 100. would not grow at all. 10000 and 1000000 times. This is cultures usually borne out by experience. we must conclude that a single cell of culture A in 1. While the growth is much slower. Allelocatalysis: The lag phase of growth is not limited Quite striking examples of lag in protozoa have been This investigator found that reported by ROBERTSON (1924). it differs by the fact that the size of the drops has a great influence. shows the development of the progeny arising kl culture. the only difference being their distances Table 41 presents the data from one another after dilution. of a culture of Bacterium coli was irradiated for 30 minutes. ROBERTSON mentions that such show no growth. . Table 42 gives the progeny derived in 44 hours from single cells viEnchelya in hay infusion drops of various sizes.5 cc. the absence of a mitogenetic effect when the cells are too close together. - e. (3) to fungi.15 cc. when grown in different cell The slower growth due to greater dilution is The more important result is plainly evident in the controls.. and if extrapolation is permissible. i.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 42. ETC.

It is possible to reconcile the influence of the drop size with mitogeiietic radiation. but onion root. The ailelocatalytic effect is not limited to unicellular organisms. HIGHER PLANTS first The first mitogenetic sender and the It is detector was the established beyond doubt that the tip of the onion root radiates. small drops are an important factor in success (WRIGHT. In the isolation of single bacterial cells by the micro -pipette. because cells can In a small drop. B. A very pretty exemple of mutual stimulation has been observed by FRANK and KITREPINA (1930) with the eggs of a sea urchin. development went much more rapidly than if there were only a very few eggs per drop (see Table 43). This was verified by NATTMANN (1919) who observed that 5 yeast cells would die in a medium where 50 produced growth. most of it points q\iite distinctly in the direction of growth. a good share influence one another ^mutually. It will be shown later that sea urchin eggs at this stage of development are good senders as well as good detectors. With an increase in drop size. the observation that in the same medium. necessary to cause This theory of "alleloeatalysis" has not been accell division. However. a small inoculum did not reproduce while a large one did. among other things. considerable part of it must leave the cell. of this emitted radiation is reflected from the air surface and finds its A way back to the cell before it is absorbed. upon ROBERTSON assumes that a "catalyst" and that a certain concentration of this produced by the catalyst. absorption is greater and the probability of the rays being reflected back to the cell is smaller. 1929). With 10 to 20 eggs per drop of sea water. This radiation is not entirely diffuse. WILDIER'S "bios" (1901) is essentially identical with ROBERTSON'S The "bios" theory was based. there is no definite experimental proof that alleloeatalysis is a radiation phenomenon. a chemical mutual influence was not excluded. cepted generally. The ability to grow in a small drop indicates that the cell has been able to produce sufficient radiation to induce mitosis. though similar phenomena are known. All media absorb mitogenetic rays fairly readily. However. distances become greater.138 CHAPTER VII is is cell. . here too.

Radiation ceases at once when the root is severed from the bulb. and has been used as a strong source of mitogenetic rays in the early experiAfter approximately half an hour. however.) This suggests that the substances which are the source of radiation are centralized in the onion bulb. radiates strongly. however. The spectrum is in all of the pulp is purely oxidative. the heated pulp furnishing the compound to be oxidized and the exhausted pulp supplying . and especially in its base. 38). radiation ceases.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 43. Heated pulp mixed with exhausted pulp starts to radiate anew. some negative results obtained with onion roots are very likely due to inaccurate direction of the mitogeiietic beam. ments. continue to radiate after being cut from the plant they also continue to grow for a con. Onion roots radiate only as long as they are connected with the bulb. (The? roots of the sunflower [Hi'lianthus]. 139 Development of Sea Urchin Eggs Experiment II According to GTKWITSCH. Heat destroys the radiating power which strongly suggests that an enzyme the active part (Table p. or at least with a part of the bulb. ETC. and its cessation known is probability due to the completed oxidation of some uncompound by the oxidases of the root. siderable time after being severed. When the base is cut from the bulb. it radiates weakly. The pulp of the base.

The new terms might better be discontinued because two they are likely to be considered as introducing a mysterious new element into life processes. Probably. and cannot therefore be caused by the same process as that of the onion base. The bulb spectrum is oxidative. CHAPTER VII Radiation Spectra of different parts of the onion the oxidase. The conduction of mitogenetic rays through normal tissue over a distance of . the tip will radiate. It seems regardless of the wavelength of primary radiation. the oxidase is located in the onion base. and the new oxidizable material is transported to continuously from the leaves through the vascular system of the plant. but not for the emission from the root tips. however. The "secondary radiation" of the roots (sec p. This accounts for the radiation of the bulb. therefore most probable to assume that the normal radiation of the intact root tip is really a secondary radiation induced by the oxidation processes in the onion base. The normal root tip radiates glycolytically. a small part of the base remains on the root. If this part is narcotized with chloral hydrate. for these GUBWITSCH has used the words essential factors. there can be scarcely any doubt that mitotase is an oxidase. This question has been decided it by spectral analysis (Table 44). mitotin and mitotase Considering the oxidative spectrum of the pulp.140 Table 44. the root tips do not radiate. 110) is also glycolytic. it seems likely that a relation between these two radiations might exist. This conclusion is biologically very important. If. If the base is cut off. Since the base of the onion bulb as well as the tip of the root radiates. the tips do not radiate.

The facts as well as the interpretations have been discussed in great detail in preceding chapters. After 48 hours. There are innumerable data on experiments with onion roots. . vein led to the discovery that radiation arises from the vascular How mitogenetie radiation is conducted from the vascular system to the growing parts of the plant. 1929). portions from leptome are inactive (KISLIAK-STATKEAYITSCH. A However. The potato when cut. The radiation of wounds in plant together with the wounds of animals on will tissues will p. very interesting investigation concerning the radiation of sunflower seedlings has been carried out by FRANK and SALKIND (1926). 173. but after 18 hours. Very little experimentation has been done with other plant and though mitogenetie radiation started with plant tissues. be? discussed The plant tumors be discussed on p. A priori. without great loss of energy. absorption appear the only way to explain the absence of radiation sides of the sunflower cotyledons. is unknown. Failure with one cotyledon which showed an abnormal curvature of the central system. only the very center of the cotyledon edge radiated. but only from the leptome fascicles. 1927). Since neither of these two experiments were carried out aseptically. The pulp of turnips radiates when 24 hours old (ANNA GUKWTTSCH) the pulp from fleduni leaves does not radiate when fresh. Just as important is the reaction of plants to mitogenetie rays. they cannot be considered as exact proofs of radiation. radiates free Some other evidence supports this explanation. It was found that radiation can be obtained from the root tips.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS several inches of is ETC. we should expect these crushed tissues to radiate because they must contain oxhlase. from the plumulae (the first young leaves) and from the cotyledons. It seems improbable to asvsume total reflection from the vascular walls. So far. and no other part of these organs. we know much more about radiations from animals than from plants. radiation has disappeared (GURWITSCII. 141 bound to affect our conception of growth and growth stimuli considerably. and yeasts growing in the pulp. Either this or complete from the tissues. it begins and continues until 24 hours old. 178. the discussion on higher plants has been limited to the emission of rays. and no accurate record was made of the number of bacteria . the meristem.

species. other part of grown plants or seedlings has ever been We cannot as yet form any opinion about the bearing of mitogenetic radiation to total growth. and not within the egg.142 CHAPTER VII No tried as detector. they are also senders. WARBURG (1909). FBANK and SALKIND (1927) observed that with eggs of the arctic sea urchin Strongylocentrotus Drobachensis. C. upon the "mitotin" which also diffuses out. EGGS AND EMBRYONIC STAGES OF HIGHER ANIMALS as well as Eggs as Menders. diffuses and acts rays. At The first cleavage this time. 80). while they have been shown to react upon mitogenetic rays. experimenting with the Mediterranean species Strongylocentrotus lividus which produces the first cleavage furrow in 40 minutes. had observed a large increase in the rate These of respiration of the egg 10 minutes after fertilization. radiation begins again. 10 minutes would correspond to about 40 minutes in the arctic The increase in oxygen consumption begins about 20 30 minutes earlier than radiation. there is no noticeable emission of rays. the amphiaster stage is reached. The chemical reaction furnishing the radiant energy would thus take place on the egg surface. ZIKPOLO'S data (1930) on the same subject (b) . becomes activated on the egg surface. and continues for about 1 hour (Table 45). GUKWITSCH (1932. as far as they have been investigated. furrow appears after 2 hours and 45 minutes. In the case of mold spores (see p. it occurs approximately 1 hour later. Half an hour after the first division. HERLANT (1918) observed a great increase in permeability 2 minutes after fertilization. to the formcontrolling factors and to the reproductive mechanism of higher plants. radiation does not (a) begin immediately after fertilization. 99) assumes that a prophase of the "mitotase" from the egg plasma. For 1 hour before and for 30 minutes after the first cell division. Eggs as Detectors: Fertilized eggs not only send out MAXIA showed in 1929 that sea urchin eggs can be stimulated in their rate of development by mitogenetic rays. p. but also respond to radiation. and muto-induction effects have been obtained. The eggs of animals are strong senders good detectors.

in quartz tubes. the medullar plates were dissected.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 45. a greater percentage of furrowed eggs was observed in 23 out of 20 experiments. and vice versa. crab hearts or the hemolymph of crabs. (c) Embryonic and Larval Stages earlier investigation of this the embryos of the axolotl as Senders: The is most detailed (1926) with kind that by ANIKIN (Ambystoma tigrinum). POTOZKY and ZOGLTNA (1930) proved the same for the eggs of the protoannelids tirtccocirrus jjapillocercus and Protodrilm hobrezkii. They were exposed for 15 to 30 minutes to a broth culture of Staphylococcus aureufi) 3 hours old. The eggs emitted nutogenetic rays during their development. always more irradiated eggs had hatched than unirradiated ones. From very young embryos. and the percentage of hatching eggs at the was ascertained in equal time intervals. 143 Effect of Sea Urchin Eggs at Different Stages after Fertilization upon Onion Boots are given in Table 27 p. the eggs of Drosophila melanoyaster are good detectors. ETC. with open rnedullar furrows. 82. . ground to pulp and used as sender. their development was accelerated by the rays from isolated frog hearts. According to WOLFF and RAS (1934b). as example of alleloeatalysis. Table 45 a shows that same moment. The mutual stimulation of sea urchin eggs has already been mentioned on p. 137. After an exposure to these radiations for 5 to 10 minutes. SALKINP.

The brain pulp of the fullgrown animal did not (see however p.144 Table 45 a. long. . pore seems to radiate.. and that the center of radiation appears to be in the head. and it was found that the brain. 158). .. just previous to hatching. The results are shown in Table 46.. CHAPTER Kate VII Effect of a Culture of Staphylococcus aurevs upon the of Hatching of the Eggs of Drosophila The same was done with the rest of the embryo. Then. and of REITER and GABOB. As detector. after the removal of the surrounding mucus. 30% 63% 54% 37% 50% 30% 37% 28% 48% 30% 49% 34% induction 32% . With slightly larger embryos. radiated. Among the invertebrates.. the entire blasto- detector root. living embryos. Swimming Larvae Trochophore Stage . that frog tadpoles radiate only until about 1 cm. onion roots were used. the brain could be dissected out. see p. Concerning the radiation of certain organs during metamorphosis. 167. During gastrulation. the embryos of Saccocirms radiate during the entire stage of their development.. Only the mcdullar plate radiated. This agrees with the earlier statements of GTTBWITSCH. . found the following induction effects: Blastula Stage Gastrula Stage SALKIND (1931) . but none of the other tissues. the vegetative hemisphere did not radiate. were placed in a glass tube so that either only the ventral or only the dorsal side faced the Even at the morula stage.. .

168).Monographien 10 . during the second and of incubation. insects. The only example of embryos or larvae as detectors of mitogenetic radiation is that of BLACK wit and associates who could make the fore leg of a tadpole grow third day more rapidly by exposure to radiation logical (see p. ROT: IX R a h n : may also be mentioned here (see p.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 46. only the larvae of Drosopkila have until shortly before after this (see also They do not begin to radiate pupation. However. Pro to plasma . 165). ETC. liquefied (d) Embryos as Detectors. The morpho- changes in sea urchin larvae observed by MAC. and cease to radiate 90 hours Quite different is the behavior of the chicken embryo. 169). positive results were obtained with the zone around the embryo. SOKIN and KTSLIAK-STATKEWITSCH (1928) working with entire embryos two days old and also testing the brains of older embryos could find no evidence of radiation. p. 146 Effect of various parts of the embryos of the axolotl upon onion roots Of the analyzed.

however. . Other actively reacting digestive enzymes are likely to be good senders. suffice to absorb completely a mitogenetic radiation of normal intensity. 45). testicles. The very strong absorption of these short-waved rays has GURWITSCH found that already been emphasized repeatedly. The cornea of the eye is also a good sender. The working muscle. TISSUES VII OP ADULT ANIMALS Tissues as Senders: Until recently. being a con- tinually renewed tissue. GTJBWITSCH has recently favored the peptic digestion as a strong and fairly constant source.146 CHAPTER D. planations for the differences between radiating and non-radiating tissues. and with the resting muscle. Probably. the seat of active proteolysis. but would hardly be considered as tissues. about 10 times as strong as blood. equally thin films of substances capable of enzymatic cleavage. workers with lymph glands. POTOZKY. such as lecithin. in the consider radiation to be produced largely by the chemical reactions cells. are also capable of secondary radiation (see p. contains no membrane. organ is muscle would radiate. skin. WOLFF and RAS (1933c) consider it to be one of the strongest sources. most of the tissues of If we adult animals had been thought to be non-radiating. proved to be very active. particularly when obtained with tissue pulp. and they will pass on radiation not as a beam. especially with organs in situ. extremely thin films of oil or related substances. but in all dimensions Thus we observe strong radiation in blood which of the film. chapters. ovaries. autolyzing vertebrates. it would seem that all tissues should radiate quite GUBWITSOH (1934) has given some convincing exstrongly. negative results With all had been obtained by earlier liver. are now considered of little significance since better methods. Carcinus ma-enas and a species of Pachygrapsus they found the pulp from gills and from testicles . SALKIND and ZOGLIKA (1930) studied the tissues of two crabs. inactive. and in nerves which contain plenty of These two will be treated separately in the next two lecithin. on account of their importance. This while the hepatopancreas radiated distinctly. films of practically only one molecule thickness. showed distinct radiation. On the other hand. Negative results.

GUKWITSCH had been greatty puzzled in his earlier work by the observation that the very distinct radiation from the rabbit's eye disappeared during starvation. SIEBEKT'S results have been given. is radiation main source of muscle an oxidation process. p. with a much larger total energy output. however. 1932. p. the pH changed from 6. spectral analysis brought a simple is explanation glycolytic. 147 On p. In pulp from working muscle.85. radiation also was present. The normal radiation and ceases during starvation because of lack of sugar. the pH changed from 5. In growing organs. while glycolysis.82 during the experiment. showing that the muscle radiates only during work. The role of radiation in wounds will be treated in one of the following chapters.THE SIGNIFICANCE OP BIOLOGICAL RADIATIONS ETC. the muscle in situ radiates strongly while working. 65. the cells lie so 10* . A good example of applied spectral analysis should be mentioned in this connection. emits only a very small fraction of it as mitogenetic rays. (GuRWiTSCH 1932. 41). partly of unknown origin. It must be remembered that there is no quantitative relation between the total energy liberated by a chemical reaction. proteolysis of the tissues becomes necessary for the continuation of life processes. 67). and . but weakly when The resting. energy output is negligible for the muscle work emits most of spectral analysis indicates that the not glycolysis. 149). and this produces a proteolytic spectrum- Tissues as Detectors: Only a few instances are known where fullgrown animals or their tissues reacted upon mitogenetic rays. the stimulation of the growth not so readily proved.33 to 5. This refers only to the pulp. and that pulp from resting muscles does not radiate. According to recent experiments by FRANK and KBBPS (quoted from GUKWITSCH. the results were the opposite. By dropping the organ into liquid air and grinding it while frozen. tired muscle does. With rate is multicellular organisms. partly it is its energy in radiant form. After prolonged starvation. while that from active. and radiation was absent while with rested muscle.96 to 5. and the ultraviolet radiation emitted (see It may well be that some proteolytic process whose p. years later. Several but reappeared after 8 days of continuous starving. this result was caused by an irritation of the muscle during grinding.

is The only animal tissue that has been actually used as detector the cornea. Another example Further illustrations the cornea (see p. Blood radiates quite strongly. E. and exposing a detector to the blood radiation through the inner wall of the blood vessels which is transparent to these rays (spectrum see fig. weak gly- Blood radiates within the veins or arteries as could be shown by removing the tissues around the veins or arteries. p. dog's blood which possesses only vory colysLs gives very fluctuating results. 124). very likely can only be harmful. 167). however. cancers arise through mitogenetie radiation will be discussed in later chapters. am the effects of resorbed tissue in different stages of (see p. The onion root is usually considered the classical differently is example though the data have been interpreted somewhat by REITER and GABOB. . outside. birds and amphibia. BLOOD RADIATION all work on blood radiation has been given by W. 44). According to GURWITSCH. the only exception being extreme old age and a very few diseases which will be discussed later. and also the hemolymph of the crabs Care in us and Pacliygrahsus and of the clam Mytilt/s edulis has been tested. especially purpose. 144) radiations as well as of purely chemical means to achieve its This and the possibility that neoplasms. 84).T (1934) in the second volume of . it yields vory good results (see p. if they are not absorbed completely before they reach the region of growth.(see p. SIEBER.. KANNEUJESSER and KAZWA (quoted from GTJRWTTSCH.148 CHAPTER VII closely together that the optimal intensity of radiation may Emanations from already be furnished by the organ itself. A few instances arc known. These latter results together amphibian metamorphosis with those obtained with em- suggest very strongly that the developmental mechanism controlling size and form makes use of ultraviolet bryos (p. and strong mitogenetic effects have always been observed. Ac- A comprehensive review of cording to 1932. The blood of various mammals. even in adult animals and men. 84). 129). where the rate of cell division is accelerated.Handbuch der allgemeinen Hamatologie".

under continuous stirring. The mitogenetic effects obtained with the blood of those animals were without addition with 2% glucose 2. GUBWITSCH'S method (1932. alcohol) because traces of disinfectants appear to inhibit glycolysis. 149 Outside the blood vessel. especially in diagnostic medicine. but can be brought back by adding glucose. Drying must be accomplished very rapidly to preserve full radiating power. and softened in a tiny shallow dish with 6 to 6 drops of distilled water. p. the blood does not radiate.5 5 2 21 44 36 39 These strong effects could be obtained only by adding much more sugar than is normally contained in the blood. blood loses its radiating power within 10 to 15 minutes.continued work. the dried sample should be kept dark. . it is drawn off with a pipette and used at once for irradiation. 132) investigated a number of laborers before and after the day's work. BBALNESS (quoted from GUR- WITSCH. not more than 1 to 1. 121) shall be mentioned here: disinfection of the skin The blood should be drawn without previous (iodine. or it is hemolyzed with distilled water (PoTOZKY and ZOGLINA. but normal radiation after 2 hours of rest. This must be considered when making blood tests for diagnostic purposes. 72. 1929). p. It is possible to dry blood on filter paper and thus preserve its amount radiating power for 2 to 3 days. POTOZKY and ZOGLINA starved rats until they had lost about 30% of their body weight. Since this might be of practical importance. Table 47 gives a few of his data which show consistently no radiation immediately after long. For radiation experiments. A very important observation is the disappearance of blood radiation after continued work.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. In making the test. The spot is cut into very small pieces.5 minutes should elapse. radiation can be restored by adding glucose. HELNEMANN'S method been mentioned on p. The drops are spread widely on the filter paper to prevent coagulation and thick layers. In hemolyzed blood. 1932. detector and everything else must be prepared before water is put upon the dry blood. blood is mixed with an equal of a 4% MgSO 4 solution to prevent clotting. From the adding of the water to the beginning of exposure. this holds also for the hemolymph of crabs. for detecting blood radiation has With starving mammals. As soon as the water becomes dark red.

CHAPTER VII Mitogenetic Effects from the Blood of Factory Workers before and after Work WASSJLIEFF (1934) repeated the investigation with mental work The. though not quantiIntravenous insulin injection tative. In addition to the starvation experiments just 2000 The main source mentioned. (calculation). in addition. of blood radiation with mammals appears to be glycolysis. It has practically all the lines charac- teristic for glycolysis. This suggests glycolysis. and it could be demonstrated that muscular work connected with calculation caused the decrease in radiation while the mental work as such does not affect it. Of these latter. reduced glycolysis and radiation to zero. The spectrum of the radiation of rabbit blood from the streaming blood in the vena saphena is shown in fig. which is affected by . first results showed a decrease in radiation. Rat blood treated with heparin loses its radiation by NaF. addition of glucose (probably in overdose) interfered with radiation. measuring the glycolytic power and the radiation of each sample. but this happened before psychic analysis showed mental fatigue. 194050. the lines new lines.160 Table 47. proteolysis. relation between the two. 47). creatin hydrolysis. KANNEGIESSEK and KAWZA made extensive experiments with dog's blood. some 2010 have also been found in nerve spectra (fig. and oxidation. 196070 and and. phosphatid hydrolysis. In some eases. 192030. 44 as measured by GOLISCHEWA (1933). but not by KCN. There was a good.

A somewhat different way was used by HEINBMANN and SEYDERHELM animals. no hemoglobin) oxidation Of the various fractions of the blood. Below: the combined spectrum of 5 common biological processes (oxiC 9 protcolysis -. some blood of old people thus treated radiated again. 151 its NaF.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. radiate distinctly but not The spectrum very strongly. Distinct clinical evidence of "rejuvenation" by this treatment has been claimed. even after several months (p. and this result was confirmed by HEINEMANN (1932). and in the blood of crabs (which contains is the deciding reaction. is greatly changed after of amphibia. and the of radiation in blood during senility. loses power promptly in the presence of 0. according to KLENJTZKY (1932). polynuclear as well as lymphocytes. but in hemolyzed blood. 43). Table 48 gives of PROTTI'S results. In his reason why ultraviolet light was beneficial to (1932) succeeded in isolating a SEYDERHELM compound . radiation. the metamorphosis During wounding it varies enormously in the different stages of development. however. PROTTI (1931) observed a great decline or complete absence The intensity of blood radiation (see p. however. indicates glycolysis.Ar . The leucocytes. but loses this power rapidly. Hemolymph of crabs. effort to find the to renew blood radiation in old persons. and probably proteolysis. This suggests that in normal rat blood. In consequence PROTTI injected blood from young persons into old ones. and so does hemolyzed rat blood. 174).0001 molar KCN. to produce secondary 1 I n i n \mt\m u \m\ w Figure 44. Above: spectrum of streaming blood in the artery of a rabbit. oxidation and phosphatase action. but not by KCN. It continues. the serum radiates immediately after the blood is taken from the animal. O f glycolysis -^ G. glycolysis is the dominant source. creatinin hydrolysis dation phosphatid hydrolysis =_ 7*).

The only diseases from the blood which showed this conspicuous absence were leucemia. 45). With patients suffering from anemia. When injected into patients 70 to 80 years old. without blood radiation. 1932). and severe with septicemia high fever (also poisoning with nitro-benzene). . and repeated tests showed no decrease. cytagenin produced either no effect or a slight depression effect during the first days. is L. but not in pernicious anemia in which the bone marrow no longer produces blood cells. When this substance was injected into healthy persons. CHAPTER VII Mitogenetic Effects Produced by the Blood of Old Persons After Injection of Blood from Young Persons from the blood corpuscles which he called cytayenin. dead. it increased blood radiation (fig. GUBWITSCH and SALKIND the persistence of blood radiation during (1929) obtained radiation of tuberculous guinea pigs until almost immebefore death. is It loses is power before the animal reversible. Diabetes.152 Table 48. osteomyelitis and ulcer of diately the stomach did not decrease blood radiation. was possible in all cases of secondary anemia. the blood began to radiate. The blood this still of asphyxiated animals does not radiate. How long these experiments were continued. lues. it washed the newly-formed blood corpuscles out of the bone marrow and This produced a normal blood picture (HEINEMANN. HEINEMANN did not state. but after 1 to 2 weeks. even when the process Quite remarkable illness.

white after injection. ETC. non-radiating group. upon a carcinoma patient. / and // are normal healthy persons. in some cases.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS and cancer. Injections 11 / 2 3 / 83 10 11 Jim At Figure 45. . and scarlatina. and are Decrease of Respiration Decrease of respiration of yeast irradiated with the blood of healthy and sick persons. 46 shows the results obtained by GESENIUS (1930) who measured blood by the decrease spiration (see radiation yeast reSince p. loft: black indicates radiation before injection. l^ig. III is a very old person. 163 This was verified by SIEBERT of radiation in severe cases of sepsis. 84). Figure 46. and by GESENIUS tonsilitis to this (1930). it can be used for the diagnosis of cancer (see p. A very comprehensive review of his extended research on blood radiation has been given by PBOTTI (1934 a). radiation appeared again soon after the removal of the tonsils. of only very few easily recognized diseases prevent blood radiation. Increase of blood radiation by injection of cytagenin. All measurements refer to apparently healthy persons. At right: the effect of repeated injections IV is a carcinoma patient. who observed absence pneumonia. HEINEMANN (1932) added chronic. 180).

animals. the . the stronger radiation of tall. the effect does not last more than one hour. longitudinally at the rate of approximately 30 meters per second. In the same individual.. at a distance was much stronger than at 10 mm. hypo-radiant and hyper -radiant" types can be distinguished. the effect from a place 20 mm away. further by inhalation of oxygen. also by a trip to the sea shore. Later. The resting sartorius muscle of a frog when irradiated biologically at a definite place for 10 seconds emitted a strong mito- geiietic radiation of 20 mm. radiation increases 1 to 2 hours after each meal. Irradiation of the older. These results. Irradiation of the tip produced The impulse was conducted radiation from the older tissue. 66). made by During the winter months. . NERVE RADIATION AND THE CONDUCTION OF STIMULI IN ORGANISMS The results obtained with secondary radiation led GURWITSCH to the idea that possibly secondary radiation might be a controlling factor in the conduction of stimuli in plants and Let us recapitulate briefly the facts obtained with onion roots (see p. all reacting in a parallel manner upon the same changes. upper part of the severed root produced radiation from the tip. PROTTI states that even among healthy individuals. he demonstrates the decrease of radiation with age. 140). It is by high altitudes. together with the discovery of secondary radiation in nerve tissue. In fact. but there was no conduction to the diametrically opposite side of the root. FRANK observed that the same held true for muscles. En a large number of tables and graphs. and decreases with physical fatigue. it is slightly lower. inspired in GURWTTSCH the bold thought that conduction of stimuli in nerves and muscles may be ac- . P. "normoradiant.154 CHAPTER VII the yeast bud method by means of the hemoradiometer (p. slender people as compared with short individuals with a tendency for stoutness. and the slight increase of male over female blood. intensity had increased by conduction. though in this latter case. Menstruation and increased pregnancy have characteristic curves.

Irritation of the nervo it was accomplished either by cutting into with platinum minute (faradisation). It is evident that the chemical analysis of is One them can give only a rather incomplete picture of the chemistry of nerve stimulation. or mechanically. The rather surprising fact was revealed that different kinds of irritation produce slightly different spectra. but he could also study its spectrum by using intermittent exposure. 47. apart. 73) as detector. . the possibility of studying. on account of several suggestive facts. GUKWITSCH himself (1932 a) says that this it theory may appear bold. (traumatisation). or by electrical tetanization 4mm. 145) and those by KEITJCR and GABOK (1928) with the sciatic nerve of the frog gave negative results. FRANK and GOLDEN BEKO (1931) who obtained the pickerel. the metabolism of the nerve in the resting and in the excited stage. They were confirmed by WASSJLIEW. The result of the spectral analysis of nerve radiation has. revealed some chemical processes which had not as yet been discovered by chemical analysis. intermittently. while radiation can be observed without injury. The minimal total exposure required to bring about mitogenetic effects was approximately 6 minutes with the resting nerve and 2 3 minutes with the excited one. All experiments were conducted with This intermittent exposure by the rotating disk (see p. 105). as a matter of fact.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. by moans of the mitogenetic spectrum. but also eliminated the possibility of secondary radiation from the nerve induced by the yeast detector. KAJLENDAUOFF (1932) not only proved that the sciatic nerve of the frog radiated distinctly. The older experiments by ANIKINT with the brain of adult salamanders (see p. The detailed spectra are shown in fig. The spectrum was determined with resting as well as with irritated nerves. The views considerable concerning nerve radiation have undergone change with the improvement of the methods. positive results only with the olfactory nerve of By using the yeast volume (p. because it involves destruction of the nerve. but it seems justified to approach experimentally. 155 complished or aided by secondary mitogenetic radiation. by hitting with a light hammer. 8 12 shocks per electrodes 3 not only increased the intensity of the effect. This statement is possibly too blunt. but it expresses in a few words the ultimate aim.

and de-aminisation of amino acid A few were observed also which cannot at present be accounted for. The spectra (of which each 10 A strip is the result of at least three determinations) show five well-known chemical pro- cesses: oxidation. as a shift of the preceding line to the right. . this J^ine 2410 way. This may be due to a slight shift in the cali- bration of the instrument. 1930-40 and 195060. or rather left. excited by illumination of the eye Figure 47. excited by incision 20 mm. glyeolysis. electrically excited nerve. 20 mm. Fn mechanical and electric irritation. phosphatase action. from combined spectrum 6 common bio- chemical reactions 44) optical nerve. the line 2000 10 in more than half of the spectra and 2370 80 in 3 of the spectra cannot be explained by the same error of calibration. cleavage of resting nerve pulp of resting nerve nerve. from point of electrical excitation point of mechanical excitation of (fig. 20 might also be accounted for in an error in the calibration to the However.156 Further. most of those concerned with do-aminisation are missing. mechanically excited nerve. The spectra of the sciatic nerve of the frog under different The lowest line represents radiation from the optical nerve in consitu. are missing in most of the nerve spectra while they all have the neighboring lines 194050 and 196070. ditions. removed from the zone of *) Two lines of the glycolytic spectrum. and in the electricallystimulated nerve. 20 mm. differs CHAPTER VII it was found that radiation at the point of irritation from that of the same nerve a short distance away. Strange is the absence of certain lines lines. 1 ) These five chemical processes agree with chemical investigations on nerve metabolism. the glycolytic regions are absent while* in the spectrum of the same nerve. creatinm phosphate.

1. It agrees essentially with those by KALENDAROFF. small window. After decapitation and removal of the lower jaw. the part of nerve conduction. and that even the type of excitation may change the spectrum. The radiation from this was tested in the darkroom while directing A a beam of light on one of the eyes. As soon as light was directed onto the eyes of the frog.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS irritation. This difference in the spectra raised the question of 'adequate" nerve stimulation. number of yeast cells by count was the method used to prove the Increase in suits direct microscopic radiation.. The creatinin lines are absent in mechanical stimulation. ETC. but 24 hours later. These fine results induced ANNA OURAVITSOH (1934 a) to approach the more fundamental question whether this radiation of the optic nerve after illumination of the eye was only a simple secondary radiation. The established lines are doubtless correct.5x2 mm. good example of this has been ' A furnished by ANNA GURWITSCH (1932) with the optic nerve of the frog. It has also some unknown lines agreeing with KALENDAROFF'S. The spectrum is shown in the lowest line of Figure 47. The re- were always positive while controls without illumination of the eye showed consistently 110 effect. the* optic nerve was laid bare by dissection from the roof of the mouth. the lobes arid hemispheres radiated strongly while the medulla and spinal cord did not change. and kept these parts covered with the skin. optic lobes. It might be well to remember here GURWITSCH\S statement that these spectra represent "minimal spectra". Those corres- ponding to proteolysis (de-aminisation) are present only in part. or represented an important functional She laid bare the optic nerve. all nerves radiated strongly. 157 they are present. The spectra of nerve conduction lack entirely the phosphatase lines. is sufficient to permit access to the chiasina and tracti optici with part of the optic nerve. . the medulla oblongata and the spinal cord of living For several frogs. These experiments have revealed that there are differences in the metabolism of the resting and the excited nerve. but there may be other lines too weak to be recorded by the detectors used. lobes and hemispheres did not radiate while medulla and spinal cord usually continued to radiate weakly. hours after the operation.

Addition of the glycolytic component (1900 1920 A from a monochromator) to nerve radiation produced good radiation in lobes and hemispheres. however. the last mentioned part was missing. they did not react promptly upon illumination This suggested an interference between traumatic of the eye. and and with blue all lines were present. The spectrum of the radiation of the lobes contains always glycolytic lines even when radiation has been brought about by exposure of the chiasma to rays from the. while the hemispheres showed no effect. the lobes reacted strongly upon illumination of the eye when the current was applied. oxidation and phosphate light. but their relative intensity was varied. It had been shown at this time (p. cleavage. intensity of the colored lights used and the wave lengths of the colors are not mentioned. but indicates the of an unknown independant chemical reaction in the optic lobes. there were no proteolytic lines. and after that. therefore. radiation ceased completely for a short time. Before the circuit was closed. the metabolism of the nerve was not changed completely. and normal reaction upon optic irritation. With red proteolysis. 45) that secondary radiation usually is a resonant radiation responding only to the lengths characteristic for it. The . Probably. induced strong radiation in both.158 CHAPTER VII When the optic lobes were still in the excited stage from the operation. It mere secondary radiation. ANNA GUBWJTSCH (1934b) obtained different spectra from the hemispheres of the frog brain when the eyes were illuminated with different colors. the reaction upon light was weak and irregular. The explanaby irritating electrically the sciatic nerve. the normal strong effect appeared. not a release which are very different from those of glycolysis. oxidation of pyroalso gallic acid is. light. The extent of radiation in a nervous system during illumination permits thus the experimental approach to the problem of localisation of functions in the brain. radiation tion could be verified . after ceasing the irritation. Yeast radiation. In another paper. ANNA GUKWITSCH studied the reactions produced by various spectra applied to the chiasma wave of the optic system. Green light produced the lines of glycolysis. but continuing illumination. The emission from an electrically irritated nerve made the lobes radiate weakly.

5 10 15 20 25 13 5 Induction Effect at 18cm. are also the experiments by LATMANISOWA on the "fatigue" of the nerve by continued irradiation with strong light (see p. Very suggestive. Table 49. It could further be shown that continued exposure to light produces a long after-effect if the head is cut from the animal. . 112) and this agrees. The result was as follows : Length of Exposure Induction Effect at 6cm. but there no appreciable difference in the intensity of the effect. distance) requires a longer time to pro- effect. though not positive proof. 14 is 4 4 30 -16 35 seconds 6 4 The weaker duce the light (18 cm. . however. e. radiation ceases when the eye is darkened (see Table 49). varying the intensity from 9 to 1) and determining the necessary exposure time. in the living animal. (i. LATMANISOWA (1932) found this to nerve correlate accurate radiation be 30^3 meters per second (see p. within the limits of error. Later publications by LATMANISOWA added another impressive (1933. 1934) have fact to prove the role of mitogenetic . 111). with the rate of conduction of nerve stimuli. Mitogenetic Effect of the Optic Nerve after illumination and darkening of the eye Another important advancement in the effort to conduction with mitogenetic radiation is the measurement of the velocity of progress of secondary in the sciatic nerve. 159 was tested by ANNA distance between light and eye from 6 to 18 cm. 1 3 19 5 22.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS The effect of the intensity of GUBWITSCH (1932) by varying the light ETC.

By exposing the sciatic nerve. The nerve outside of the irradiated zone reacted normally at the same time when the exposed part of the same nerve showed parabiosis. but soon the muscle reactions became weaker and weaker. Perhaps. causing a stronger muscle contraction with a stronger impulse. Though a number of physiologists oppose this idea.160 CHAPTER VII rays in nerve conduction. For about 2 hours. and the place between irradiated. and then. the nerve showed all symp- toms of parabiosis. the same success. it does not seem impossible that the two observed facts may be some time be combined to produce a more comprehensive explanation of the nerve mechanism. The one example of true adequate stimulation are the above-mentioned experiments by ANNA GrrrtWiTSCH with the optic nerve. Two them was electrodes touched the nerve. Finally. few minutes. or a corresponding eifect. with the attached muscle. and its reactions became normal again. the nerve reacted nor- mally upon electric impulses. . It seems too early to speculate on the relation between the mitogenetic radiation of nerves and the action current. This experiment has been repeated more than 40 times with Controls with gastric juice without protein were not affected. a * such combination of definite wave lengths is necessary to produce effects. the typical parabiotic stage set in. the contraction being stronger when the impulse was weaker. and it took two hours after the removal of the mitogenetic source before the nerve had recovered. This may be due to the absence of an 'adequate" stimulation. . After this time. by irradiating a nerve. the emission seems to be much stronger than that of the normal nerve. It has not been possible as yet to produce muscle contraction. the nerve became at first more excitable. in a moist chamber for about 2 hours to the radiation from protein digestion. the nerve ceased to react altogether. The author quotes a paper by LAPITZKI who obtained in a the same effect by rays from a mercury vapor lamp The nerve at the parabiotic stage has not ceased to radiate on the contrary.

ETC. At times. Yeasts and Bacteria: CHRISTIANSEN (]{)28) striking morphological changes in yeast cells as well as bacteria under the influence of radiations emanating from menstrual blood. 1. produced they shall by a short observed note on unicellular organisms. 1932). Particularly the large spherical cells with tremendously extended vacuoles and with complete loss of granulation are considered It has not been possible. there was a decided Still other cultures showed for- Similar morphological changes could be produced by exposure* normal persons (FERGUSON. however. tendency to grow into hyphae. the blood was strong enough to kill microorganisms more commonly. of yeast to saliva of apparently In other cases. While the classical by be biological radiation is that of sea urchin larvae preceded. With Bacterium coli. it is evidently not a purely chemical effect of some saliva constituent because the yeasts grew quite normally in mixtures of equal volumes of saliva and raisin extract. Thus. 161 MORPHOLOGICAL EFFECTS example of morphological effects 4 . but the same effects could be obtained when a quartz coverglass protected the test organisms perfectly against vapors from the blood or from outside. to typical "saliva cells". these cultures must have obtained more saliva constituents than could possibly distil over from the Protoplasma-Monographien IX: Rahn 11 . Frequently. . Streptococcus luctis and cremoris did not appear to be changed morphologically. leaving the protoplasm only as a very thin layer around the cell membrane. produce these same forms by interposing a quartz eoverglass between saliva and yeast. the non -motile cells immediately above the drop of blood were 3 to 5 times as long as the farther removed cells. Bacterium vulgare lost its pellicie formation Laclobacillus bvlyaricus grew to long threads without cell division Oidium formed no oidia. tho physical nature of this phenomenon is not proved. .THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS G. Just as striking were the changes in yeasts. retardation of growth was accompanied by an enormous expansion of vacuolcs. it affected their cell forms. On the other hand. for the sake of logical arrangement. mation of giant cells. . It seems that most of these experiments were* carried out without exclusion of chemical effects from volatile substances of the blood.

Normal growth of tiacclwrowycett Mycoderma Above: 100 times. puttctisporui . below: 500 timca.162 CHAPTER Vll Figure 48 a.

103 Figure 48 b. below: 600 times. Growth of irradiated tiaccliarontyces Mycodcnna Above: 100 times.THE SIGNIFICANCE OJB 1 BIOLOGICAL RADIATIONS ETC. 11* . -punctisporus.

when exposed continously to mitogenetic rays from various sources. fig. the weakest effects. the effect of plants upon the morphology of yeasts was studied. . will change morphologiyeast cells. develop into very abnormal larvae. It has also been stated there that recently. Mrs. Seed embryos. cillus In the aiithor's laboratory. pollen.1(>4 CHAPTER VII saliva droj) to the yeast culture. but scarcely ever the complete set of morphological changes. tiactfiaro'tnyces and also produced The most sensitive yeast was M ycoderma putu'tisporus GuiLLfEimoND which was frequently greatly elongated so that it appeared under the low power like a mold colony. 2. but to an electric effect. the steam distillates of carrots or potatoes. Tt was found that most parts of the various plants stimulated yeast growth considerably. 49. as well as those by CHRISTIANSEN. Home of the wine yeasts also showed changes of cell form and size. nor did the juice of crushed carrots produce any change. killing were observed. great morphological changes. effects . When no effects the sender was poisoned chemically. and roots produced the strongest leaves. Perhaps we are dealing here with a combined chemical and physical effect. it has been suggested that this is not due to a real radiation. cally the giving them the appearance of "saliva types". when added to the culture medium in large amounts did not affect the morphology. all cell activity. BARNES has isolated a bawhich through a quartz coverglass. These results could not usually be repeated with the interposition of a quartz coverglass between yeast and sender. However. true branching was never observed. 48 shows the change produced by exposure to a young agar culture of the same species. Sea Urchin Larvae* It has already been discussed on 86 that sea urchin eggs. By the same method. Mcrophotographs of these forms are shown in p. These experiments. were made in hanging droplets on a coverglass which was sealed to a moist ehamher slide with vaseline or lanolin. the "sender" was at the bottom of the cavity. such as STEMPELL (1932) assumed to be rather common in nature. 48). "Only the one or the other symptom appeared occasionally (see fig. However. and the growth resembled more that of a MoniJia Fig.

Control. but fertilized with sperm exposed for 45 minutes to the same solution as 11. Control. . F/1V. untreated. 11. Eggs of the same origin as ///.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS J. Paracentrotm I. This #iu's a simple explanation of the morphological changes by the inito- * // Figure 49. Same to origin and age as /. larvae of the sea urchin. lividus. but exposed continuously through quartz an acid solution of phenosaffranino reduced by KHSO 3 normal sperm. fertilized with . arid (1931) have studied the abnormal larvae have come to the conclusion that they are primarily due to overproduction of the mesenchyme. IV. 165 and M. while the ectoderm and endoderm appear normal (fig. 50). MAUKOU histologically. 111.

ect = mouth. mes -= mestmchyme The most remarkable fact is perhaps the observation that the irradiation of the sperm alone or of the unfertilized ovum alone will suffice to bring about the same morphological changes. while the mosenchyrno grows entirely out of bounds though it is really protected against radiation by the outer cell layers. Figure 50. affected (there may blast. the mesenchymatic cells. tumefadens. Mid oca -^ endoderm. . the radiation received by the single egg or sperm cell affects the entire future development of the larva. and the stimulus received by such a cell affects only one part of the offspring. J est Cross section through sea urchin larvae. there is considerable difference in the reaction of ectoderm and endoderm cells on one side. Right: larva exposed to the radiation of ttact. = ectoderm. . Left: the normal larva. Thus. i. intestine. oesophagus. e. stomach.166 CHAPTER VII genetic effect. The former are not essentially be some retardation eventually). However. and mesen- chymatic cells on the other. Hast int blastophore.

All organs radiate only during their resorption. The second pronounced . the main shortening results are occurring at the time the forelegs develop. Radiation arises from the resorbed The riewly- forrned fore or hind legs do not radiate. IV forelegs arc out. forelegs not visible through the skin. the tail begins. no parallelism. elbows of forelegs distend the skin.THE SIGNIFICANCE OP BIOLOGICAL RADIATIONS ETC. followed closely by the intestine which is shortened considerably. e. strongest blood radiation coincides with the stage where intestine* and tail are all near the maximum of radiation. g. Metamorphosis of Amphibia. The gills initiate the process. there is Otherwise. metamorphosis completed. When the gills are almost completely resorbed. The role of mitogenetic radiation in the metamorphosis of amphibia has been studied extensively by BLACKER and his associates. The shown in fig. but they did not radiate. bring about a distinct mitogenetic effect. tail has full length : Va: Vb: Vc: VI about one-fourth of the tail is resorbed about one-half of the tail is resorbed about three-fourths of the tail is resorbed the entire tail is : resorbed. The following developmental stages are distinguished: : Stage I II hind legs differentiated hind legs scarcely motile Ilia: hind legs actively motile. 167 3. the back of the skill. belly rounded. : Illb: belly becomes loan. Other parts of the tadpole were tried. 51. In a later paper. The intensity of radiation was estimated by the amount of the mitogenetic effect (yeast bud method) which is not a particularly good measure. relative intensity: normal one-sixth normal 20 times normal normal 10 times normal normal Ilia: 30 minutes Illb: 15 seconds IV VI : 5 minutes 5 minutes Vb: 30 seconds : The gills. BLACHER and LIOSNER (1932) estimated the intensity of blood radiation of Jiana ridibunda during metaThey ascertained the minimal time necessary to morphosis. at Stage The results were II : 5 minutes. tissues. The most detailed investigation concerned the development of tadpoles.

during successive stages of metamorphosis. In the quartz -bottomed vessels.1 more in one series of 33 tadpoles and The actual 1. Finally. the legs if the bottom of resorption. The result of the operation as such was that in the vessels with glass bottoms. it could be shown that a tadpole lying in a quartz -bottomed vessel will show a more rapid growth of Figure 51. gills and intestine have BLACHER and associates (VII) further showed that there was a relation between the resorbed tissue and the developing limbs.3 less in more rapidly.5% i 1.5% i tadpoles. 4. while the other leg which served as control remained covered.8 less in one series of 22 than its mate in the cavity: 6. Removal of the gill. a second series with 44 tadpoles. and that transplantation of gills increased the growth rate of the corresponding foreleg. however.168 CHAPTER VII maximum of blood radiation occurs after completed their metamorphosis.1% 1. and therefore shaded against radiation.2 series with 51 tadpoles.4% ^ . Intensity of radiation of the resorbed organs of frog tadpoles. The placed over pulp of tissues in the process foreleg to be irradiated was freed from its is cavity by a cut in the covering skin. in the other it 7. or the freeing of the foreleg from the gill cavity where resorption of the gills occurs. retarded growth. the freed legs grew and 4. the freed leg grew a little more slowly 0.

1% 1. the plants tumefaciens. 169 1. KOSTOFF and KENDALL (1932) conceived the idea that polyploidy might be somehow linked with the strong rnitogeiietic radiation which is known to be emitted by this bacterium. are. After the development of the tumor. of course. To test this. Parthenogenesis by Radiation. too scant to permit tin* of conclusions. they are suggestive of a new cause for polyploidy.. above the tumor in order to induce the development of new shoots. 5 could be rooted.6% and 12. HI). and OAROK GURWITSCH'S statement that there is after having verified always radiation around a wound. They do not radiate until about ready to pupate. is 4. Change of Chromosome Number. He had succeeded in causing cell division in unfertilized frogs' eggs by only a very few percent of the eggs thus treated initiated development. though less intense. BATAILLON obtained tadpoles and even frogs by this method. and an accelerating effect of this radiation upon the rate of development of the organs concerned. Positonly the radiation of the resorbed tissue. The triton Pleurodeles Wallii shows similar radiations i (BLACKER etc. were cut off 3 to 5 cm. and radiation nearly ceases after 30 hours. The maximum intensity is reached 24 hours after pupation. 120 tomato plants were inoculated with Bact. refer to the theory of BATAILLON that the developmental mechanism of an egg is released by the wound produced through the entrance of the spermatozoon.. Seven shoots arose directly in the tumor and of these.8 in the other set of experiments.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. The gain is 7 times that of the average error. It has been repeatedly observed that polyploidy occurs fairly frequently in some individual branches of tomato plants infected with Bacterium tumefaciens. One of them proved to bo tetraploid. but in some cases. V). . Experiments were made also with Drosophila larvae (BLACKER etc. There is no direct proof that in these cases. While these data drawing 5. RKITEB.4 in the one set gain by irradiation was therefore 10. very fine needle pricks. and the axolotl Ambmtoma tigrinum which retains gills as well as tail throughout its life radiates still less. the radiation of the resorbed parts ively established had any distinct form-giving effect.

we have been able to observe for the most part It has this one type. 60). the controls. only 2 unfertilized eggs of the salamander of could bo induced to develop. one to the 8 -cell stage. been shown that as a rule. as well as in all experiments with longer or shorter irradiation. mitogenetic radiation has as yet scarcely been entered. 3 could be induced to start cell division by irradiation with monochromatic light of 3340 A. the first division being completed 3 hours after the 5-minute period of irradiation. It PARASITISM AND ANTAGONISM seems very likely that biological radiations play an important role in the mutual relationship not only of cells. except that it was stated that each group contained about 5 eggs. Even with this wave length. In. cell division of spores. It may be that only for reasons of technique. They exposed unfertilized eggs of salamanders and frogs. radiation alone. 173). In a later experiment with "a few hundred" unfertilized frogs' eggs. only those 3340 A proved efficient (Attention may be called again to the circumstance that REITEB and GABOR have always claimed that only the rays around 3340 A are mitogenetic. A few isolated facts stand out plainly. Of these "induced eggs' '. should produce the same result. Possibly. obtained with great precautions from the females. without wounding. to various sources of ultraviolet. H. SYMBIOSIS. this conception is wrong. the eggs died in a short time. A repetition caused only one such egg to develop. The radiation from yeast stimulates the many bacteria. biological radiations appear to be largely stimulating. but not proved.170 CHAPTER VII REITEB and GABOB speculated that this effect was in all probability due to the mitogenetic radiation of the wound (see wound radiation p. The number of eggs exposed is not given. and the rate of germination of mold Similar effects with multicellular organisms are imagineable. The radiation from an onion root will stimulate cell division in another root under laboratorv . and they reasoned that in this event. From the material mentioned in the preceding chapters. or with different wave lengths. see p. unicellular organisms further each other's growth. or beneficial. Of all the wave lengths tried. one developed to 32 cells. but This field of also of cell groups and multicellular organisms.

But it must be admitted that we have no Parasitic existence It is or animals. tmm>faclens. On the other hand. The abnormal proliferations on the leaves and stems of plants due to insect eggs (commonly called galls) are usually attributed to chemical irritation by the larva. for some reason. or cell division. of specific radiations has very little experimental evidence for its support. (see g. 1 by irritation is limited to a very few species of plants customary to assume that for chemical reasons. but it does not seem impossible that the young larvae radiate strongly during their early period of growth. or.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. but not to rats could not be explained by different radiations from the two hosts. it was not attempted to prove that a tumor would be formed even if chemical effects by bacteria were excluded from action through a wall of quartz. there has never been observed any specific action of definite wave lengths upon definite organisms. the plant or animal. this assumption would imply very specific radiations: either beneficial. The assumption that . MAUROU (\U27c) observed that cell division in the tumor was not limited to the immediate presence of the bacterial cells. The role of radiation in parasitism is strongly suggested. providing that 28 e. which favor one speeies. antagonistic radiations which prevent growth. However. but that can hardly occur under normal conditions of plant growth. chemical explanation either. processes which are common to most plants and animals. However. and cause abnormal proliferation of tissue through ultraviolet rays. more probably. p. All experiments so far point to the conclusion that any radiation 4 between 1900 and 2600 A can stimulate the division of any cell. all other plants and animals are prevented from living as parasites.specific. oxidation. fig. The assumption. at our present state of knowledge. all well-known radiations arise from entirely unspecific biochemical processes such as proteolysis. permit physiological . conditions 49). the parasite is not affected by the harmful radiation. with the soil absorbing all radiation. While we are still far from knowing all sources of mitogenetic rays.. unicellular or part of a tissue. The pathogenicity of the typhoid bacterium to man. but none of its nearest relatives. etc. but not proved in the case of plant tumors caused by ttact. of all species except the parasite. 171 conditions. radiatioiis also play a part offers itseK naturally. glycolysis.

Different from the specific harmful radiation which injures one species. i. pyocyanca 42 and 48% retarded. 184) which are linked with certain pathological conditions. in 5 experiments. could be killed. using B. . which was not retarded. are the generally harmful human radiations observed by CHRISTIANSEN and by BAK^ES and RAHN (see p. the harm is probably done by an overdose and not by any widely different specific wave lengths since such sources as chemical oxidation. Acs used old. a more detailed speetroseopic investigation of such types might add greatly to our understanding of the significance of biological radiations. or yeast with which were 6 S hours staphylococci or streptococci. glycolysis by . but then* is no evidence that parasites are more tolerant to radiations of this kind than While the assumption of specific related. or not at The example of the sea urchin larvae (p. to 2 hours by Ps. ftaccharomyces Mycodcrma jmnctisporus. but stimulates other species. In this case. but stimulated 42 and 50%. while other yeast species were retarded. 1 Antagonistic Radiations: The only good case of specific antagonistic radiations is the investigation of Acs (1933) who experimented with microorganisms known to antagonize each other when grown simultaneously in the same culture. finlhrricift for 1 gave growth retardations of 22 In two experiments. for his experiments cultures at the stage of rapid growth. Since such selectivity of antagonistic radiation cannot be explained by our present knowledge. 86 all affected. and 164) is quite characteristic of a harmful effect produced by a primarily beneficial radiation. in 14 experiments. such as Bacillus anthracix and Psc udow onax pyocyanca. he obtained e. Yeast radiation was found to retard staphylococci very distinctly. and also streptococci in the one experiment made.172 Jt is CHAPTER VII known. The radiation may be truly specific only one species of yeast. and exposed the one pure culture to the radiation from the other. ant/tracts as sender. it has as yet no experimental foundation. and found Px. pyocyanca Irradiation of B. distinct retardation of growth. he reversed to 136%. that an overdose of rays will not and might even retard growth. non-parasitic species. stimulate. hi this way. radiations would be a very convenient explanation for some problems in parasitism. the arrangement. of course. The same organism was used simultaneously to irradiate Racillus rnfhnors.

HABERLAND (1929) attempted sedum leaf pulp would cause dry. as with grown animals or plants. revert to multiplying cells. not "heal". They divide. di- leaving the wound surface wet with cell debris. that would do pulp of scdum leaves did not radiate when fresh. that there are chemical requirements . the cells show no signs of division kept in a moist chamber. using yeast as detector.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. In most cases. They will begin to if smeared with juice of crushed leaves. of cells it came from the cell pulp. will They can be torn without rupturing the cells separate. but so after 18 to 24 hours. the torn leaf will also reproduce if the leaf is cut instead of carefully torn. THE HEALING OF WOUNDS When wounds begin to heal. and protcolysis by staphyloeocci gave the same results. is e. losing this power again after 48 hours. Such i. this is usually accomplished by mitosis of the cells nearest to the wound. to discover whether radiation by a healing or renewed mitosis of a that leaf pulp did not induce any division from the injured leaf and concluded that the wound hormones act chemically and not He found when held at short distances physically. GURWTTSCH (1929) expressed his opinion that such new and that very probably vision of old cells could not take place without a mitogenetic stimulus. but young leaves of fledum s-pectahile and Esrhcverta a surface if sec nmla of the mesophyll. He states that in his opinion. this would mean that cells which have already come to a resting stage. This resembles somewhat the situation of old yeast or bacterial cells transferred to a fresh medium. however. torn leaf. Tt was precisely at this stage that we found mitogenetic radiation to be most effective upon bacteria or yeasts. 173 yeasts or streptococci. and leave a dry surface. GURWITSCH (1929) found. I. HABJOKLAND had found that full-grown. The first discussion concerning the possible role of biological radiation in the healing of wounds occurred in 1929. Something like a rejuvenation process of the old resting cells is necessary before they can divide again. the mesetich vine being the only ones which were stimulated out of proportion. The main cause of the harmful effect in this case cells is the difference in sensitivity or reactivity. radiation alone will not cause the healing process in this case .

(>% 41. . decreases decidely and reaches a second maximum after 5 days. The same double maximum could be observed with wounded BLACIIER.9% 48. entire 18 days of the experiment. 4 tadpoles of the frog Pdobatcs drop in intensity after 4 days.utt. The threshold time to distinct was the measure effects mitogenetie necessary produce of intensity. that radiation one of the necessary While the argument concerning the sedum leaf wounds lias never been settled experimentally. uninjured tadpoles produced a marked effect in 5 minutes. ground it up and used the pulp to irradiate yeast plates. The blood of normal. and after different times. the blood radiated less than half as strongly as that of uninjured tadpoles. FIUCHIMOfutic. The radiation maximum after 1 2 days. by LTOSNKR and WORONZOWA (1932 b).2% 2. By this long-continued effect of radiation. the entire body is affected the wound upon blood and brought into play. from the tip of the tail. It is not the new tissue which radiates. BROMLTGY (1930) has proved that the tail of the salamander Anibi/tttonui tigrimim radiated strongly after being amputated. 12 hours after injury. and a minimum at 24 hours. Very interesting was the observation by the same authors that the blood of the tadpoles changed its radiation decidedly upon wounding of the tail. This agrees with the two maxima in pll of healing wounds as observed by OKUNBFK (1928). VJ1 is but. They cut 10mm. removed the old tissue which produced the regeneration. and it stayed at this low level during the . the following "induction effects" were obtained: 1 23 5 6 12 24 hours after amputation 6.174 besides the physical ones factors. Exposing yeast for 18 minutes to the ground tissue. two distinct maxima are seen at 12 and 36 hours after wounding. CHAPTER .4% 50.4% induction of the injured tissue reaches a It requires 3 hours for radiation to be established. In Table 50. and a great WITSCII. but the old cells immediately under the newly-formed tissue.1% 29. an exposure of 15 minutes became necessary. 1 minute of exposure to blood sufficed for a similar effect 24 hours after injury. This means that at this stage. 2 minutes were required.1% 39. and on the fourth day.

method of wound- the tadpole tails for irradiation from below. and found that the first mitogenetie effect could not be detected until 20 hours after indicting the wound. 175 Induction Effects Produced in Yeast by Irradiation with Old Tissue Bordering the Regeneration of Wounded Tadpole Tails (Pdobates fuscus) (1932) repeated the tadpole and salamander He cut them through. leaving earthworms. and it is here recorded in ft of new . ETC. LiosNER and WORONZBWA (1932 a) succeeded in proving this by producing a more rapid regeneration of wounds by biological irradiation. bringing about healing. At ing left. Kadiation SAMABAJEFF was weak throughout. as in this proof. Considering that regeneration in wounds is accompanied by a radiation of the injured radiation tissues it was essential in seemed probable that this BLACK KB. method of measuring the amount of regeneration. under a low power lens. but was noticeable even after days. fig. The rate of healing was measured as shown in the same 52. figure.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 50. Figure 52. at right. More than 500 tadpoles were used for Into their tails were cut triangular wounds. IRICKIMOWITSOH. with experiments them in earth.

for the tadpoles were left untreated for 24 hours and then exposed 24 hours. CHAPTER Amount VI T of Regeneration in the Wounds of Tadpole Tails. This pulp is known to radiate strongly. and the wounds on the under side of the tail were thus exposed through quartz. of which a few are reproduced in Table 51. there was no difference when the tad- poles were exposed for 24 hours immediately after wounding. showed that in the control**. while the wounds on the upper side were not. There was also no difference when the tadpoles were left without treatment for 48 hours and then exposed for 24 hoiirs.176 Table 51. no difference . When all the upper (non-irradiated) wounds of the exposed tadpoles were compared of regeneration with those of the controls. However. If. Irradiation produced in all series of experiments a distinctly more rapid healing of the irradiated wounds. The tadpoles were placed in dishes with quartz bottoms which rested on a pulp of tadpole tails and intestine. the upper wounds had healed 26% less than the controls. A comparison of the lower wounds showed an increase in healing of 33% when irradiated at once. this statement needs some modification as a careful checking of the individual wounds showed. These experiments. The controls were in similar dishes with glass bottoms. there was 110 difference in the rate between the upper and the under side of the tail. however. but when irradiation began after 24 hours. Irradiated from Below growth from the deepest part of the wound.

and lacking orderly structural arrangement. utero. hay fever. the cells resembling those of the parent cell from which they derived. that the continuous removal of pus and dead tissue cells by the maggots induces more rapid healing. may of wound treatment. THE CANCER PROBLEM is not very accurately defined. manifesting an effect which is only apparently beneficial. Irradiation 48 hours after wounding again produces a true stimulation. NORTH (1909) and GILTNEB and HIMMELBEBGEB (1912) applied cultures of Lactobacillua bulgaricw to inflamed mucous membranes (gonorrhoea. there is a tivitis. partly at least. One is the beneficial influence of the presence of LactobaciDi." Protoplasma-Monogrraphien IX: Bahn 12 . The proof that the rate of healing of ated by mitogenetic rays. The other method is the rapid healing of wounds infested with maggots of flies. Two explained by mitogenetic rays.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS - ETC." "Carcinoma is preferable to the older term cancer as designating tumors of epithelial origin that are malignant. yet serving no useful function. There is a suggestion of the same double maximum which we have already encountered. a neoplasm may be defined us an autonomous proliferation of cells. mitogenetic effect upon regeneration by the larvae. stimulating them. The good success is ascribed to the lactic acid. J. conjunc- wounds can be accelerbe of importance for the future treatments might be. Sincr the authors do not feel competent to pass judgment upon medical definitions. they have adopted the following of FELDMAN'S The word cancer (1932): "With certain reservations. It is meant to indicate the most common form of malignant tumor. irradiation 24 hours after wounding does not stimulate these. While doubtless the present explanation is correct. but retards the others. it may well be that in addition. which grow continuously and without restraint. No explanation has been attempted. non-inflammatory. 177 This signifies that irradiation immediately after injury affects only the irradiated wounds.vaginal affection of cows after abortion) and to suppurating wounds with very good healing results. From the physician's point of view all tumors arc neoplasms.

A The if nucleic acid spectrum is also found in carcinoma itself the exposure is sufficiently long. A. which are not capable of other occurs primarily in the necrotic parts of cancerous tissue By comparing the spectra glycolysis. g. the carcinoma problem in man and animals was attacked from various angles. and by various methods. REITEK. the most definite facts of mitogenetic radiation. (3) the origin of cancer. SIEBERT. while the benign do not. and b) of plant The first study on the relation between tumors and mitogenetic was the investigation by the MAOROUS (1927 a tumors caused by the crown gall organism." It has already been mentioned repeatedly in previous chapters that contrary to the very weak radiations of the normal tissues of grown animals. cell division had taken place quite removed from the location of the bacteria. the malignant tumors radiate strongly.178 CHAPTER VII "Sarcoma refers to a tumor is nective tissue elements. that consisting of immature conclinically or histologically malignant. e. They proved.STATKEWITSOH (1929) had found that there were two kinds of carcinoma radiations. (1) the radiation of cancerous tissues. it could be shown that the one was plainly a glycolytic radiation while the lines emitted by the necrotic parts of the tumors agreed with those of proteolysis (see Table 52). one requires glucose in order to emit rays. (2) the radiation of blood of cancer patients. as a of diagnosis. and L. GESENIUS. that cultures of the bacterium radiated. Soon afterwards. Bacterium tumefaciens. while the vestigators. It is one of STBMPELL. of these two different types with the known spectra. physical effect They did not produce experimentally plant tumors by radiation. An analysis with strips of 50 to 60 was sufficient to prove the difference. however. but did not exclude chemical effects. they also showed (1927c) that in the tumors. means Radiation of Cancerous Tissues: The strong radiation of cancerous tissue as contrasted with the non-radiating normal tissue has been established in a number of cases by various in- GITRWITSCH. It is also an established fact that only malignant tumors radiate. GTJRWITSCH (1929) and KTSLTAK . the proper time for the production of a glycolytic spectrum is too short for that of nucleic . with onion roots as detectors. and GABOB. rays. This made a probable.

46. Absence of Blood Radiation: The second outstanding and thoroughly established fact is the absence of blood radiation in cancer patients (see fig. hypernephroma (benign kidney tumor). and myoma (benign muscle tumor). states that patients with teratomes (monstrosities) and probably with mixed tumors never show blood radiation. GESENIUS (1932). retina or auditory nerve). 153).THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 52. 12* . p. even in severe anemia caused by bleeding of the myoma. 179 Analysis of the (the Two Spectra of Carcinoma effects) numbers indicate induction acid. glioma (benign tumor of the brain. HEYFETZ (1933) determined the time They inoculated rats with tumor same strong tissue shows the *) This seems strange since the sarcoma radiation as carcinoma tissue. in summarizing 3 years of clinical experience. while normal radiation is observed in all cases of sarcoma 1 ). and BRAUNSTELN when blood radiation decreases. ETC. This difference in intensity of various reactions occurring simultaneously adds greatly to the difficulties of spectral analysis.

but were normal again after 9 days. showed no blood radiation after 2. glycolysis and sugar content were not affected. . 45. a tumor the size of a barleycorn had developed. KIND observed that the blood of cancer patients suppressed the radiation of normal blood upon mixing the two. that in pernicious anemia. When its drops to cytagenin is discontinued. injected with cell-free tumor extract. There is really much more to this negative induction effect of cancer blood than merely the absence of radiation. 153). radiation begins again after liver diet. p. Rats. emission of rays negative level in 1 to 3 days. In short intervals. Injection of cytagenin (see fig. they determined the decrease in blood sugar. HEINEMANN (1932) very definitely found growth retardation when he used the actual rate of cell increase as a measure (Table 22. even the complete removal of the tumor and radium treatment will not restore normal blood radiation. LYDIA GURWITSCH and SALfig. While the latter decreased distinctly from the fifth day. while in carcinoma. since the earliest GESENITTS reports (1932) observations as a diagnostic means. p. 3. 45) caused a temporary increase of blood radiation. 73 and As early as 1920. Cancer Diagnosis: Blood radiation is so conspicuous even with patients suffering from many kinds of severe illnesses that its absence in cancer has been considered. and 4 days. and the mitogenetic radiation (Table 53).cells through an incision of the skin of the back. after 6 days.

most careful clinical investigation had given no reason to suspect tumors (e. blood radiation has been found infrequently connected with severe carcinoma. For this . It remains absent after removal of the tumor. though the preceding. During the last year. only such cases in which diag- nosis was uncertain were investigated. Since practically all diseases producing decreased mitogenetic radiation in blood are readily recognized. 84). the opposite takes place. blood radiation is a valuable diagnostic means when used as a part of the clinical examination. These facts plainly suggest that the growth-stimulating source of radiation is removed from the blood and concentrates in the neoplasm." Very encouraging is further the summary by GESENIUS (1932) of 3 years experience in the Berlin University clinics. Germany. However. g. further observations with carcinoma suspects justified the conclusion that a positive mitogenetic effect excluded the possibility of a tumor with certainty. a small carcinoma in the upper rectum which caused no direct pain). a negative effect changing promptly to positive after cytagenin injection. caused us repeatedly to search for a tumor and to prove it beyond doubt. The method consisted in the decrease of yeast respiration by blood radiation (see p. cancer diagnosis by The best results this method has been studied systematically. this simple assumption does not agree with our explanation of these radiations as originating from biochemical reactions. On the other hand. and has rarely been absent in patients where autopsy revealed absence of carcinoma. It has just been shown that sugar content and the rate of glycolysis in the blood of cancer patients are not greatly altered. KLENITZKY (1934) observed the return of blood radiation when every last trace of cancerous tissue had been removed. Loss of radiation is independent of the size and location of the tumor.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. 181 Recently. HEINEMANN from "On the one side. OriginofCancer:It has been stated repeatedly that normal adult tissues radiate very weakly while normal blood radiates strongly. and cannot be used as a test for the success of treatment. obtained are very likely those recorded by one of the hospitals in Frankfurt. in cancer. the new growth radiating strongly while the blood ceases to do so. Occasional exceptions have been observed. Nevertheless.

from about 50 to about 10. the discovery of their effect does not really explain the formation of cancers. MORAVEK and others that cholesterol stimulates the growth of malignant tumors. together with the abovemcntioned "cytophotolysis". the tumors grew more rapidly in the starved mice whose blood radiation had decreased. For the first 15 days. eventually resulting in a neoplasm. on the day of injection. The same was the case with intravenous injections. not usually Somewhat different is PROTTI'S conception (19346) who assumes that a cellular disorder may arise when blood radiation becomes very low. MACKENZIE. injections of small amounts of yeast into the tumor stimulated its growth while large amounts retarded it. The same happened when the neoplasmic cells were separated from the yeast cells by a quartz plate. as far as is known. been produced by frequent application of Since the compounds found so are not normal or pathological products of the human body. but to radiation.182 CHAPTER VII is reason. but may give valuable suggestions towards the solution of the problem. With the adeno-carcinoma of mice. that the results are not due to enzymes. this explanation of the origin of cancer stressed. ROFFO (1933) found the cholesterol content of the skin near cancerous or pre- . one lot was being fed normally. More important is perhaps the discovery that cholesterol metabolism is in some way connected with cancer. leaving finally a cavity with thin fibrous walls. PROTTI calls this "cytophotolysis". Yeast heated to 60 produced no effect which suggests. Repeated injections of yeast suspensions into the GallieraSarcoma of rats caused a liquefaction of the tumor. Cancers far have certain chemicals to the skin. It has been claimed by SHAW. It must be remembered that cancer is most frequent in old age when blood radiation has a tendency to become very low. PBOTTI observed further that a mixture of neoplasmic cells and yeast cells in vitro resulted in a destruction of the neoplasmic cells while cells from normal tissues were not influenced by yeast. without pus formation. while the other was on a starving ration. Intravenous injections produced no effect upon the tumors. He proved his point by injecting a cell suspension of adeno-carcinoma into two lots of 12 mice each.

After 10 to 11 months. beginning with 5 minutes per day and gradually increasing the exposure to 6 hours. It would be a splendid agreement if this range which is known to stimulate cell division could also cells. 183 cancerous lesions to be much higher than the normal skin of the same patient. This agrees very well with the above-mentioned frequency of skin cancer on different parts of the human of body. Within 4 months. Of the control rats receiving light from a tungsten filament lamp. He proved it to take place only in living organisms. The author has not been able to ascertain whether ROFFO has tested the mitogenetic range of rays by itself. which was exclusively on the naked parts of the body (ears and eyes mainly. avoiding tho hottest sun.61% 3. X-rays or radium rays produced the opposite effect. and many of them had several tumors. i. not a single animal had developped a tumor. there is a decrease of 25 to 40%. With white rats. ROFFO (1934 a) studied the "heliotropism" of cholesterol very thoroughly. development and the histology of the rat cancers corresponded exactly with that of human cancers. 700 rats were kept in sunlight daily for not more than 6 hours. every one of these rats showed tumors. Then. ROFFO (1934b) studied the tendency for cancer development in white. The wavelength of the ultraviolet was above 2300 A.. e. (1930) had shown that the cholesterol content of the skin increases when it is exposed to sun light. ultraviolet irradiation of healthy persons increases the blood cholesterol while with cancer patients. rats. exposed to ultraviolet light. of an intensity of 14 erythemal units. 52% of all rats had developped cancer.50.02% 0. He found the frequency of skin cancer to be distributed in the following way. sunlight as well as ultraviolet light increased the cholesterol content of the exposed parts of the skin.07% 1. also twice on the hind feet and once oil the none). The mode be shown to produce abnormal cell division of the epithelial The fact that sunlight can do this speaks very much against . 150 rats wen. as average of 5000 cases: skin of tho face skin of the back of the scalp skin of the foot hand 95.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. According KAWAGUCHJ to MALCYNSKI (1930). the range between 1800 and 2600 A.

Then. three types of radiation from organisms have been mentioned which are not identical with mitogenetic radiation. Quite commonly known : are readily in their collecting flowers for the perfume factories of France. menstruating women are excluded from and bacteria become abnormal when handled and the common belief that bread dough will not rise normally. since CHRISTIANSEN proved that wine fermentation was distinctly cultures of yeasts by them (see p. Attention efficient in may be called here to the effect of radiation of exist It is impossible to sa^y whether cells. One was entirely different. it would be purely accidental. and they produce distinct biological the reactions brought about effects. namely the TJeto-radiation of the potassium in the cells. However. and does not really belong in this group. 95).cholesterol upon yeast any relation might problem. Details may be found in the paper by MACHT and LX^BIN (1923 1924).cholesterol and the cancer oxy. such as have been shown to produce mitogenetic effects. though apparently stronger than mitogenetic rays. might possibly be present (at least in Buenos Aires where these experiments were made). between oxy. and does not have any definite biological purpose or effect as far as has been ascertained. and that pickles and sauerkraut packed by them will not keep. This type. K. The much stronger effect of artificial ultraviolet containing also the shorter waves suggests that the shorter waves are more bringing about abnormal cell division. The third group are the harmful human radiations. and should it produce biological effects. there was necrobiotic radiation. very small amounts. but it is absorbed by the surrounding medium. does not sound improbable. pure by menstruating women flowers wilt hands. But it cannot be considered proved that the effect . 96). affected (see p. It is an emission of energy typical for the chemical changes connected with death. is emitted by dying cells. mushroom beds are said to be utterly ruined by their mere presence.184 it CHAPTER VII since sunlight contains no wavelengths shorter than 2700 A. The facts as such can be considered fairly definitely established. RADIATIONS OTHER THAN MITOGENETIC In Chapter IV.

on account of the exposure could not be started until one day later. the yeast was killed by emanation from the finger tips. and effect upon the yeast. cannot be stated at present. likely that the radiation comes from some reaction of the cholesterol or oxycholesterol. of the oxycholesterol an error was made in the separation from the other reagents. 4 killed Mycoderma during the first day when exposed continuously through quartz. The results were quite definitely positive. through a thick quartz plate excluding chemical influences. was emulsified with a little water to make oxidation possible. 185 men physical origin. produced by a compound called meno- toxin. must be either oxy cholesterol or a closely related compound. and we would think above In the above experiments the oxycholesterol all of oxidations. In this case. this product usually has lost its power. Probably the same type of harmful radiation has been observed in a very few cases of illness (p. The more common conception among medical is that it is chemical. With Batch No.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS is of ETC. but not on the second day. at least. Why this radiation is harmful instead of stimulating. by this time. 97) . have but rarely been able to obtain good results with wilting flowers. Out of 5 batches of oxycholesterol made at different times. The colorimetric tests mentioned for oxycholesterol can be obtained long after radiation has become too weak to be detected biologically. The intensity of this effect varies greatly with the individual. not probable that a compound as such radiates. the radiated. The simplest assumption would be that the death of the Mycoderma cells is due to over-exposure because in Since it is it appears more . it has not been decided as yet whether the effect is physical or We chemical. of Harmful Human Radiations: MACHT came to the conclusion that the u meno- 1924) toxin". This induced BARNES and RAHN (1933) to test whether this compound e. compound in the blood during menstruation responsible for the various phenomena just mentioned. The Source and LUBIN (1923 i. CHRISTIANSEN found the effect to be much stronger in summer than in winter. and even from the face. Ill which had no this. Attention should be called here to the observation by HILL (1933) that certain persons kill bacteria on agar plates when placing their fingers right upon the seeded plate.

complete killing of all cells is not usually observed even in over-exposure. An exception is the fat in the finger nails and toe nails. the largest . The other alternative that we are dealing with different wave lengths does not seem very probable either. Even its chemical composition is not certain. p. and can therefore be hardly anything but ultraviolet. and no exceptions are known. While a good deal of attention has been paid in recent years to cholesterol metabolism. of cholesterol Oxycholesterol in the Body: A study of the distribution and oxycholesterol in the fats of the human skin. The only more recent investigation on oxycholesterol in the body is that by PFEIFFEK (1931). An analysis of the spectrum may give the explanation. by UNNA and GOLODETZ (1909) shows that oxycholesterol is not present in the normal cellular fats but only in secreted fats (Table 54). and only rarely was stimulation found. CHAPTER VII Effect of Oxy-cholesterol upon through fused Quartz Mycoderma experiments the yeast was exposed continuously to this raHowever. 49).186 Table 53a. the oxycholesterol radiation passes through quartz but not through glass. All wave lengths between about 1800 and 2600 A give the same mitogenetic effect all diation. According to his analyses. very little is known about oxycholesterol in the body. This agrees quite well with our observation that radiation was obtained from hands (and feet) and from the face where sebacious glands exist. (see fig. but not from the chest where they are very rare.

THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 54. While previous authors. on account of the uncertain chemical composition. ETC. The relation between this substance in the tissues and radiation is therefore not definitely established. erythrocytes with 0. In the preceding chapter.29%). . estimated the oxycholesterol colorimetrically. followed by the liver (0. adrenals and bile Lowest are the (about 0. amount of oxy cholesterol.cholesterol in connection. The results may therefore not be comparable.003%. The above results suggest that an investigation about the role of oxy. e. next are bone when It has been stated already that oxyeholesterol radiation ceased the colorimetric tests were still very strong. with the cholesterol metabolism in cancer might yield interesting results. 187- The Cholesterol and Oxycholesterol Content of Human Skin Fat is found marrow. in the brain (0. g. UNNA and GOLODETZ.09%). attention has been called to the close relation between cholesterol and cancer.44%). PFEIFFER determined it by the difference in melting points between cholesterol and oxyeholesterol. on the basis of total solids.

OUTLOOK The gist who bioJogical part of this book has been written by a biolois convinced. Many others have repeated these experiments. we are dealing with an entirely new phenomenon. MOISSEJEWA. or have been contradicted by Most of the critics dismiss other. and consequently cannot predict which changes of technique might increase or decrease the effect. and with very sensitive detectors. g. in the course of 12 it lies years. and obtained no mitogenetic effect. He has realized that it is difficult to prove it because we are dealing with an extremely weak effect. . Some such attempts have been made (e. Such claim is unscientific as has already been pointed out in the foreword. The only way to disprove any theory is to obtain the same results. The fault equally with the two groups of contestants. Several of the latter group have claimed therefore that they have disproved biological radiation. more recent investigations. those for and those against radiation. the question of the existence of this radiation. following directions as exactly as they were given. The facts are these GXJBWITSCH and a number of his pupils and also many other investigators have presented a very large amount of experimental data to show that mitogenetic radiation exists. : the question with the simple statement that all so-called mitogenetic effects are within the limits of experimental error. It does not speak well for the present status of science that has not been possible to settle definitely. that mitogenetic radiation exists. LORENZ) but they have not been carried far enough. Above all. from his own experiences as well as from the study of literature. and to show that they are due to another cause.

It is evident that this point can be settled only by biological experiments. one is the existence of the biological effect. In biology.OUTLOOK 189 Let us realize from the beginning that the differences of opinion center around two essentially different points. e. PAUL of example painstaking (1933) A roots. After all. will not make the effect less important for biology. It may be that both are affected. the variety of onion). it depends very much upon the choice of the organism (e. it requires a thorough understanding of their physiology to interprete differences of growth rate. the explanation is secondary. is with onion the work M. This objection may also be made to other tissues. Biological measurements are not at all simple. but the absence of radiation does not disprove the biological effect. While it is easy to make yeasts or bacteria grow in the customary culture media. the error of may be widely different in different laboratories . g. The error in biological experiments is not as absolutely fixed as in physical or chemical methods. where large numbers are used. the controls are not perfect. The effect is A the important thing. while the theories in deciding the existence of the mitogenetic be sought largely in the sensitivity of the methods. g. the uniformity of environment before and during the experiment. When higher effect are to come and go. the uniformity of the material. in an analysis where it can be stated reliably for all laboratories that the accuracy of the method is 0. Several authors have questioned the use of one side of the onion root as control for the other side. only the facts remain permanent in science. and errors have been made in this respect by those opposing biological radiation as well as by those convinced of it. e. Physical measurements can tell only whether or not it is caused by radiation. With unicellular organisms. As a the same method result of the various factors creeping in. and the other is its interdifferent interpretation pretation as an ultraviolet radiation. g.005 g. the ability of the experimentor fine to recognize and avoid disturbing or secondary influences. the controls are as reliable as they can possibly be in any biological experiment. the cornea. the treatment of the organisms before the beginning of the experiment. The difficulties organisms are used as detectors.

Any serious criticism should start with the most reliable and best founded papers. the mitogenetic effects were more than 3 times the experimental error. RREUCHEN and BATEMAN also mention neither SIEBERT'S papers nor the extensive work of WOLFF and RAS with bacteria which is the best material with this detector. 83 (JULIUS). 58) different parts of the same culture. omit the work of SIEBEJRT who published more detailed experiments than any other investigator in this field. It is only natural that any new development in science will attract speculative minds who generalize from a few experiments and come to conclusions which are not shared by the more conservative workers in this field. even physical measurements show great differences (see Table 30a p. KANNEGIESSEK and SOLOWJEFF. 34 (BAETH) and 168 (BLACKER). 10% with some investigators and 50% with When it comes to such delicate instruments others as the GEIGER counter. HEINEMANN has stated that in his method of counting yeast with the hemacytometer. GIJRWITSCH ciates have stated repeatedly that with the yeast and his assobud method. WOLFF and RAS have frequently given all individual counts of bacteria for one experiment (p. This explains the difference of error in the onion root experiments which was (see p. they consider any increase less than 15% over the control to be experimental error. 92). 58). Other error limits can be found on pp. TUTHILL and RAHN have also published two sets of counts of yeast buds from many This has been done (p.190 OUTLOOK or even in the same laboratory with different investigators. NAKAIDZUMI and SCHBEIBEB. with the is experiments by SCHWEMMLE for all earlier onion root and the authors gave some similar calculations for the yeast bud method on p. The reliability of the yeast measurement by volume can be estimated from the data by BILUG. some weak papers are quoted extensively while some of the best proof in favor of mitogenetic radiation is completely omitted. 71. it is usually possible to compute the error from those experiments where no mitogenetic effects were obtained. 78). When the error or the reliability of the method not stated as such. working with the yeast bud method. . It is somewhat surprising to find that in critical summaries. The frequently made statement that the biological investigators do not state the error of their methods is not in accordance facts.

that the reader does not or lad it 3% is 25% of know whether the controls buds which would have given a suggestion Therefore. It might be argued that the burden of the proof lies with he opponents since the mitogeneticists claim to have proved heir point. .OUTLOOK On 191 the other hand.ornea are affected by the season. but only in a definite physiological Such greatest importance to describe all details. Another justifiable argument against the weight of some jublished papers is the lack of precision in the description of the nethod. physiological explanation of the mitogenetic effect. and even such terms "onion root" or "cornea of a frog" should be specified in much nore detail since there are many different kinds of onions or rogs. Even /ell a good deal about the performance of the experiment. the critics have good reason to disregard which give no precise account of methods or results. Since the biological detectors do not respond at all imcs to mitogenetic rays. the is term "yeast" means very little. rapers Probably the main reason why mitogenetic effects are still doubted las been the recording of results by merely giving the "Induction Effect" without mention of the experimental data from which The actual number of mitoses in a >he effect was computed. As an example may be mentioned the paper by SALKIND PONOMARBWA (1934) which might be very important for the However.he chemist whose methods are really standardized publishes merely the formula of his new compound. sornea. and the roots as well as the number of mitoses in the . but such an attitude would not be very helpful in lolving the real problem. mper largely lost. but also the actual inalytical data. nor ind he medium on which ffect so it was grown they give only the induction . it is of inite. Since the error in biological experiments varies yith the investigator. even when the standard deviation has not been lot Computed. statements as "8 10 hours at room temperature" are too indestate. the value of the least of the condition of the yeast. or the yeast volume measured. We are dealing with a very complex . . the percentage of buds. the publication of the complete records vould give the reader a conception of the reliability of any >bserved effects.he authors do not mention the age of the yeast culture.

a settlement of the question of mitogenetic radiation will be accomplished in a very short time. and the authors hope sincerely that by pointing out the mistakes and misunderstandings. 93).192 OUTLOOK phenomenon. The complexity is greatly increased by the occasional failure of the phenomenon for unknown Errors have been made by the defendants of both sides of the argument. . reasons (p. and both sides should do all they can to bring about a real understanding of the facts.

coll division processes. Die mitogenetische Strahlung des Regenerate . I. 138 BLACKER. BROMLEY. 197 Page BLACHBK and N. 363 and W. Roux' Archiv 127. 364 167 A. fusca als Quelle mitogenetischer Ausstrahlungen. Menotoxin und Hefegarung.AUTHOR INDEX II. M-rays macro-effect and planimetric drop culture method. 230 and N. 255. Biochem. D. D. Die Glycolyse und die mitogenetische Strahlung dos Bluts bei experimentellem Carci- 179 noma. Mitogenet. 121.. A. Mitogenetische Ausstrahlungen bei IV. N. 249. E. L. LIOSNER and W. 1932 a. N. Biol. 124. Mitogenetischc Ausstrahlungen wahrend der Metamorphose der Urodelen. Energy emanation during . 1930. 128. 119 1934.. W. Mitogenetische Ausstrahlungen wahrend der Metamorphose bei Drosophila melanogaster. 79 IT I. als M. Die Organisationszentren von Hydra . 175 and A. Mitogenet. POTOZKY. Zentr. and L. 1932. Blutstrahlung der Amphibien wahrend der Metamorphose. Internat. Z. 270 and B. und des Bluts der Kaulquappen wahrend der Regeneration. 174 Roux' Archiv 127. 128. L. L. ges.. W. Z. -. D. WORONZOWA. Der Zerfall der Kreatinphosphorsaure als 38 mitogenetische Strahlungsquelle. Biochem. K. 448 Deutsch. der Schwanzregeneration der Urodelen. SAMARAJEW. 1927. Der EinfluB der primaren Verheilung der Wunde auf die Entstehung in ihr. O. and P. A. Plant Physiol. J. 266 VI. A. Der EinfluB der primaren Vorheilung der Wunde auf die Entstehung mitogenetischer Ausstrahlungen in 174 Roux' Archiv 128. G. L. 38 BROMLEY. 1930. . SAMARA JEW. 1932. IRICHIMOWITSCH. LIOSNER and M. gerichtliche Medizin 10. HEYFETZ. W. Biochem. 128. HOLZMANN. 1933. I. EinfluB der mitogenetischen Strahlen auf die Goschwindig175 keit der Regeneration. Oxydationsreaktionen als Quelle mito36 genetischer Strahlung. 95 57 75 BORODIN. 128. N. 339 1932 b. . Die mitogonetischen Ausstrahlungen Stimulus des Wachstums der Vorderbeine bei der Metamorphose von Rana temporaries. Ausstrahlungen bei der Regeneration des Kaulquappenschwanzes. G. N. 240 HOLZMANN.. Venice BRAUNSTEIN. 60. V. A. Roux' Archiv 127. 5. Congress of Electro-Radio-Biology. WORONZOWA. 624 BOIIMER. Z. and 0. 1932. . see also BLACKER. SEVERIN. BKOMLEY. 274 mitogenetischer Ausstrahlungen VII. 133 d. Z. J. LIOSNER. D. BLACKER. A. f. W. BROMLEY. 1930. 274 ihr. 259.

KANNEGIESSER. Physikalische Untersuchung mitogenetischer Strahlung der Muskeln und einiger Oxydationsmodelle. I. 411 60 50. Naturwisson- 90 schaften 18. Neoplasms of Domesticated Animals. 60 and M. and A. 34.. and W. R.) 36. Ges. G. Archiv f.198 BROOKS see AUTHOR INDEX Page NELSON. Morphological changes in yeasts induced by 96. Physiol. COBLENTZ. Cagliari 2 J. 1932. Archiv Sciences Biol. deutsch. d.. 109 . 1934. photosynthesis by means of intermittent light. MAXIA.. 47. 1929. Nachweis mitogenetischer Strahlung durch beschleunigtes Wachstum der Bakterien. W. W. Biochem. 1930. Saunders 177 161 1932. K. 95. Atti Soc. 1934. 126. G. Journ. t)ber die Scharfe mitogenetischer Spektrallinien. CHRISTIANSEN.. W. 1930. 27.. 367 86. FERGUSON. 1929. Zentr. 15. Die Quellen der mitogenetischen Strahlen in Gewebekulturen. 39. W.. B.. Sulla delle radiazione del Gurwitsch. Bact. HOLLAENDER. FRANK and N. HOGUE.. M. 121. and OTTO RAHN. e Nat. Bureau of Standards. Das mitogenetische Reizminimum und -maximum und die Wellenlange mitogenetischer Strahlen. ARNOLD. DOLJANSKI. M. 321 29. hot. Zellforschung 9. 203 R. and C. Science Med. DUGGAR. R. 49. fra fotografia a la eccitatione il Cultori delle CHARITON. RODIONOW. 185 CHRUSTSCHOFF. Phila- delphia. A separation of the reactions in Journ. 1930. (Russ. G. 1932. 723 48 37 DECKER. 83 A balanced thermocouple and filter method for ultraviolet radiomotry. 634 138 and S. 161. Cornell University 1933. tJber die Wellenlange und Intensitat mitogenetischer Strahlung. 1930. E. 1931.219 83 48 EMERSON. G.. Biol. Das Monotoxinproblem und die mitogenetischen Strahlen. Ber. Roux' Archiv 121. 249. XJber die Ursachen des Gewebewaclistums in vitro. 207 VAN DOORMAL SCO AUDUBERT. STAIR and J. biological radiation. 1932. with practical applications. L. 129 35.. Thesis. 674 76. Das Wachstum der Gewebskulturen in vitro und die GuRWiTsCH-Strahlung. Journ.. Roux' Archiv 126. BRUNETTI. i>u BRIDGE sec HUGHES. 78. 137 Zum FRANK. Die gegenseitige Beeinflussung der Seeigeleier als mitogenetischer Effekt betrachtet. 1932. Irradiation of plant viruses and of microorganisms with monochromatic light. A. 91. Mikrobiol. 184. Gen. KUREPINA. 391 107 FELDMAN. 4. H. 145 DIETL see POLANO. Z. J. Archiv f. of Research 7. exper.

117. M. W. Monatsschrift fur Kinderheilkunde 21. 85. 1930a. 149 . 25. Biol. Physikalisches zum Problem mitogenetischen Gesellsch. B 114. 1932 a. Die mitogenetische Strahlung. u. 49. Cold Spring Harbor Symposia IT. 66. Venice 45. .. Menotoxine in der Frauenmilch. 108. GABOR see REITER. Miinchen 42. 155 Pflugers Archiv 231. and FRANK. Die mitogenetische Strahlung aus den Blattern von Sedum. 103. . 113. 1926. 474 95 FRICKE. 626 142 S. Congress of Electro-Radio-Biology.. 312 GOLISCHEWA. Die Natur des spezifischen Erregers der Zellteilung. 311 82 GURWITSOH. Archiv 52. 328 . Die mitogenetische Strahlung des markhaltigen Nerven. 63. G. . Proc. Roy. 11 Roux' 54.AUTHOR INDEX FRANK. f. 1932. 146 . t)ber Ursachen der Zellteilung. 177 Pathol. Archiv 100. 220. 257 . and L. 70. HIMMELBERGEK. W. I. P. Z. 150 Blutstrahlung am lebenden Tier. 199 Page SALKIND. Z.. 142. II. Biochem. Z. 42 FRIEDRICH SCHREIBER. Mitogenetische Strahlung der Seeigeleier. 153 1932. 1929 a. 225. H. 696 . 1933. . 1921. 119 1923. . Roux' . 449 66. 1933. SALKIND. 1922. Die Quellcn der mitogenetischen Strahlung im Pflanzenkeimling. Comp. Morphol. GESENIITS. Berlin. 1933. GATES see RIVERS. 179. Biochem. The chemical-physical foundation for the biological effects of x-rays. Protoplasma 6. 33 181 GILTNER. 147. and Ther. Die mitogenetische Spektralanalyse der 93. 1 Strahlung. 107. R.. tlber den derzeitigen Stand des Problems der mitogenetischen Strahlen. Biochem. 4 GUILLERY. tiber Bedingungen des Wachstums auf Grund von Untersuchungen an Gewebskulturen. Apparent mitogenetic inactivity of 92 active cells.. Blutstrahlung (2). Roux' Archiv 110. 167 . 173 1929 b. 127. see GERLACH. 1929. J. 141 and S. 380pp. 180.. 449 141. 119 . Roux' Archiv 108. The use of lactic acid cultures in combating infections of mucous membranes. Physiol. J. Radiobiologia 1 85. 1934. OUELLET. 241 . 73. 234 1934. and C. t)ber Stoffwechselwirkungen von GURWITSCHStrahlen. 34 84 1930 b. . Der gegenwartige Stand des mitogenetischen Problems. GREY. 59. Zentr. 1912. 1927.. . Virchows Archiv 270. ALEXANDER. . 60 . Internat. 74. Springer. 52 GOLODETZ see UNNA.. 260. und Carcinom-Diagnostik. Soc. K. tTber die GuRWiTSCH-Strahlung menschlichen Bluts und ihre Bedeutung fur die Carcinom-Diagnostik.. HUGO..

genetischo Spektrum der Nukleinsaurespaltung. f. 1928. et Physicochem. 1929. 701 1935. GURWITSCH.. G. 1166 : 158 II. Das mitogenetische Verhalten des Bluts Carcinomatoser. 211. LYDIA. A.. 731 and S. 122. de Physiol. und mitogenetische Strahlung des 190 ( Bluts.. Klinische Wochenschr. Die mitogenetische Spektralanalyse Spektren des Carcinoms und des Cornealepithels. 38 and N. 151. T. 142 . biol. 1925. C. Das mito.Die mitogcnetische Strahhmg des Carcinoma.. Roux' Archiv 124. 362 65. SALKIND. 357 . . 152. 1934a. 425 Die Biochem.mitogenetische Strahlung". Cytagenin HEINEMANN. . : . ANIKIN. 124 38 . Roux' Archiv 105. 159 du systemo nerveux . biol. Z. 1932. Morphologic Variation and the Rate of Growth of Bacteria. Biol. 43. Z. Z. Die Fortleitung des mitogenetischen Effekts inLosungen und die Beziehungen zwischen Fermentatigkeit und Strahlung. Quantenausbeuten Lichtzahlern. . de Physiol. 226 MALL see JONES. rays. rend. 246. Zentr. t)ber . 11. Die Fortpflanzung des mitogenetischen Erregungszustandes in den Zwiebelwurzeln. AUTHOR INDEX Page and L. Hermann and Cie. ALEANDER. 255 Bahn bei 157. Z. Biochem. 49. W. 15 HENRICI. ttber den Ursprung der mitogenetischen Strahlen. Z. 5. 157 Ann. 1928. .200 GURWITSCH. 84 180 173 Biochem. GURWITSCH.. L'excitation mitogenotique du systeme Tierveux pendant Ann. 89 Acta Brevia Neerland. 127 . 55 . Compt. 1153 1934b. Physico-chemical test for mitogenetic Nature 134. 1934. 1931. 181. IV of Exposes 40^ de Physiologie. 10. Springfield. soc. 246. M. Biochem. 81. 1931. Roux' Archiv 113. 39 pp GURWITSCH. ANNA. and K. 470 1929. Periodische Anderungen der Permeabilitat beim befruchteten Seeigelei. 185 pp 120. \ Vol. 109 1934. H. KREUCHEN. Physik 15. 1932 b. Paris. 220 1932 a. C. Die mitogenetische Sjpektralanalyse IV. HABERLANDT.. . adaquater Erregung.. L'analyse mitogenetique spectrale.. 20 92 HARVEY see TAYLOR. II. central. 236. biol. 152. 180.. Physico-chemical test for mitogenetic GURWITSCH) GURWITSCH) rays. 134 HERLANT. 1929. 178 Krebsforschung 29. 111 1932. 1934. Techn. 1918. 1375 72. M. L'excitation mitogenetique Strahlung der optischen Pfliigers Archiv 231. Z. Das Cornealepithcl ats Detektor und Sender mitogenetischer Strahlung. et Teclairago monochromatique dc 1'oeil.. 89 ( . . 10. K. 151 . Thomas. Die mitogenetische -. Physicochem. bei HAUSSER.

1935. On relation between time and intenJ. 0. Action of normal skin on bacteria. 236. sity in photographic exposure (IV). Archives of Surgery 26. McGraw-Hill.. 46 . 1934. 460 . 415 and CHARITON. L)er Einflufi der Blutstrahlung auf die Gowebsf. L. DE. 337 KAWAGUCIII. KENDALL see KOSTOFF KISLIAK-STATKEWISCH. N.. (Russ. Pfliigers Archiv 231. Die Spektralanalyse der Strahlung des Nervcn im Ruhezustande und bei kiinstlicher 73. Internat. 354 1930. see also BILLIG Erregung. tissue cultures ? mitogenetic rays have any influence on Acta Brevia Neerland. G. Z. 252. . and M. Die nu'togenetische 155 Spektralanalyse I. . L'analyse mitogenetique de la desaggregation des polysaccharides. K. 1932. see 185 HOLLAENDER 866 DlTGGAR. 122. S. 239 1931. LANSCHINA. Zellforschung 12.. 1932. 145 Kartoffeleptoms. 39 KOROSI. J. 201 Page BRAUNSTEIN. KLEE-RAWIDOWICZ KLENITZKY. PROKOFIEWA. 35 KARPASS. York. Roux' Archiv 109. and L. Am. KANNEGIESSER. tJber den EinfluB der Lichtstrahleii auf don 183 Gesamt-Cholesteringehalt der Haut. 211 Arch. A. Z. 218 181 spectrale de la radiation and E. Biochem. Z. 13.. 126 1934. Krebs178 forschung 29. markhaltigcn S. 283 . 1930. 214 see also SORIN. Z. 24 83 JULIUS. Z. L'action a distance de la levure sur I.)35. HOLZMANN see BLACKER. Photoelectric Phenomena. 1933. Congress of Electro-Radio-Biology. A.. 1927. M. Die mitogenetische Strahlung des Carcinoms. 83 JAHN see RICHARDS. Sciences Biol.. A... L. 1926. 1932. JONES. C. Mitogenetische Strahlung 52 bei Eiweifiverdauung. 1934. 20. WHITE. J. N. Die mitogenetische Strahlung der weifien Blut151 Biochem. Die mitogenetische Btrahlung des 141. 531 pp see 26 IRICHIMOWITSCH C. Le retablissement de la radiation mitogenetique du sang apres 1'extirpation d'une tumeur cancereuse. kulturen. . 5.AUTHOR INDEX HEYFETZ HILL. (Russ. sec LASNITZKY. 1929. Venice les sucres.. BLACKER. H.. HUGHES. Sciences Biol.) 35. Biochem. and V. New JAEGER.. Arch. Z. and E. H. Soc. Opt. 215. A. DU BRIDGE. HALL. Biochem. 51 Do KALENDAROFF. 901 HlMMELBERGEB See GlLTNER. 1929. elemente. W. 221 232 . f. M.

d'Histologie appliqu6 4. Action a distance du Bacterium tumefaciens sur loppement de 1'oeuf d'oursin. . E. 185 J. Die mitogenetische Sekundarstrahlung 110.. see also * LATMANISOWA. 518 LEPESCHKJN. The fractionation of Chem. f.202 KOSTOFF. Untersuchungen iiber mitogeiietisehe Strahlen. physical 73 MACHT. 1927a. Science 76. Biol. Botanik 72. f. 905 171. Search for mitogenetic radiation electric by means of the photo41. L. Z. Krebsgenotischen Induktion von Warmbluterzellen. stability of living matter. Bull.. Parabiose der Nerven Folge mitogenetischer Bestrahlung. 113.. 265 als 159 159 1933. 186. Journ. Physikalische und biologische Untersuchungen uber mitogenetische Strahlung. 95. d. 190 HAUSER. 22. LUBLIN. I. 1934. MAGROU.. Messung geringer Lichtintensitaten mit Hilfe 92 von Zahlrohren. 159 LASNITZKI.. 243 Protoplasma 92. A. Health Reports 48. 99 99 99 1932 b. J. 1930. BATEMAN. H.. . rend. and Exp. W. f. . Journ. 178 178 1927 b. . 184. 802 . Pfliigors Archiv 231. 50. Ges. 112. 367 1933. wissensch. B. . Rev. 1933. Zur Frage der mitoZ. 611 57 LIOSNER see BLACKER. 19. Ther. M \GROU. de Physiol. 1928. Influence of 83 visible and ultraviolet rays upon the . LORENZ.. and M. and E. 547 . and D. Nekrobiotische Strahlen. Radiations emises par path. 57. General Physiol. 1924. 91 LUCAS. 1932. et ent. Origin of a tetraploid shoot from 169 the region of a tumor on tomato. Compt. 1931. H.. mitoge*n6tiques. 178 . Recherches sur le les radiations . W. 10. Protoplasma 20. Ann. 1932a. v<g. Physik 94.. Bot... 144 KREUCHEN. desNerven. 1923/24. Naturwissenschaften 21. deutsch. le deve- . D. 1935. Sur la parabiose mitoge'n&ique et de Physicochem. see LUBIN S. Ber. . rend. W. . 253 .. 1934. AUTHOR INDEX Page KENDALL. Bacterium tumefaciens. KLEE-RAWIDOWICZ. MACHT.. 232 Loos. KUREPINA see G. Investigation of mitogenetic radiation by means of a 91.S. 188 photoelectric counter tube. . 22. 1934.. D. 141 du nerf.. W. 244 . 413 95.. FRANK.. and J. 843 bios. Compt. method. 1927 c. 28. W. 549 und J. agr. U. K. of Pharm. 1180 Journ. 14. 17. Jahrb. Rayons mitoge*netiques et gen6se des tumours. 86 . 330 . Nekrobiotische Strahlen I. forschung 34. 1932. bot. G. 184. Am. 67. A phyto-pharmacological study of menstrual toxin. 1311 . LANSCHINA see KARPASS.

265. 1914. Klinische Wochen936 183 MAXIA. Biochem. 35870. 62.. Compt. 190 NAKAJIMA see WRIGHT.. Zoologie 14. Soc. 1929. 30. fl. tJber den EinfluB der einmaligen Bestrahlung nichtkrebsigen schrift 9. G. Com. 763 see also MOISSEJEWA. Zur Theoric der mitogenetischen Strahlung. mittels S. Z. 1930. 155 BRUNETTI. W. Opt. 222. 231. Biol. Biochem. 195. B. and BROOKS. 1931. 235. 214.. Radiobiologia 1 (1). 133 62. 1931. Z. M. March 27 OKUNEFF. 1909. 188 NAKAIDZUMI. rend. 237. M. New York. N. 192831. Action a distance et embryog&iese. Action a distance sur le developpement do 165 . 1931. REISS. 232. 1928. G. MEES. 1919. On the nature of bacterial lag.. 421 OUELLET see GREY. 1. Untersuchungen uber das mitogemrtisehe Strahlungsproblem 11. 457 . and Med. MAEOOU. K. Van Nost116 198 pp NELSON. SCHREIBER. 609 Kssai d'interpretation.. 86 MALCZYNSKI. Die Lebenstatigkeit von SproBpilzen in minoralisohen Nahrlosungen. Hyg. Annales des Sciences naturelles: 86. Z. E. C. 86 Poeuf d'oursin. Paris 193. 439. Ober einige physico-chemische Erscheinungen 174 wahrend der Regeneration. Tallasografico It. 203 Page and M.. 1931. Biochem. Z. Z. 53 and 210. 239.. f. 215 134 PFEIFFBR. J. Medical Record. K. S. 189 PEXFOLD. Biochem. 243. Effect of infra-red light on subsequent fertilisation of the eggs of certain marine invertebrates. N \UMANN. Reports of 300 cases treated with lactic acid 177 bacteria. 2nd ed. 97. Photography. 149 1932. C. Mem. J. 21.AUTHOR INDEX MAROOU... 186 230.. 236. Exp.. Acad.. und H. 67. 14. 142 R. Biochem. C. 251. I. Soc. E. 32 et P.. PAUL. HI. techn.). 201. 1933. 1933. C. Journ. 1931. 241. Biochem. 1007 C. Quarzlampe auf den Cholesteringehalt im Bluto der und krebskranken Personen. Proc. Z. II. Roux' Archiv 128. Am. rand. Z. 108 57. Die Cholesterine im Strukturverbande des Protoplasmas. 101 NORTH. Intensificazione delle segmentazione din ova di Paracentrotus lividus sotto Pinfluenza di radiazione mitogcnetiche. Action a distance de facteurs biologiques et chiniiqucs sur le developpoment de Toeuf d'ourein (Paracentrotus lividus L. Photographic plates for use in spectroscopy 24 and astronomy. 7. 220. 1 138 NKBLETTE. M. 424. J. Zwiebelwurzeln als Detektoreii bei Untersuchungen uber mitogenetische Strahlung. Biol. 1930.

Mitogenetische Strahlung des Bluts. Wachstum der Bakt. (Russ. Journ. 249. and I. in FR. f.) 35. . 1932. 140 pp. 49 153 1934 b. 427 pp. No 99 RAHN. Roux' Archiv 114. Zum Problem der mitogenetischen Strahlung. . 418 134 125 1929. 121 . The fermentometer. ttber einen empfindlichen Lichtzahler. Bact. 1924. La 182 Ricerca Scientifica 2. see Physik. 90 151 1931. 712 108 . Comm all Sed Scient. 199 Philadelphia. 424 POTOZKY. Die Einwirkung der Hautabsonderung der Menstruierenden auf die Hefegarung. t)ber don EinfluB der Stoffwechselprodukte auf Bakterien. Biochem. Physiol. Krebs91 forschung 35. see PROKOFIEWA KLENITZKY. 579 125 see also BARNES and FERGUSON and TUTHILL. RA. della emoradiazione applicato alia dinica. Radiobiologia 1 . Leipzig 1931 Z. 16. Z. Das Verhalten ciniger Tumoren gegeniiber dem SacchaArch. Z. . Biol. . ZOGLINA.. An experimental comparison of criteria of death in yeast. 1930. Z. 91 1932. 1930. . 1930. ZOGLINA. bei O. Osped. 131 different . Biochem. 1933. 16. Z. 239. 178 146 1. Ulrico Hoepli.. A. PROTTI. 1934a. J. . Impressioni fotografiche di radiazioni ematiche ottenuti attraverso Civile di Venezia . G. 36 Biochem. ZOGLINA. 1906. 255 see romyces cerevisiae. 32. 1931.. Zentr. Journ. Physiology of Bacteria. 92 RAS WOLFF. t)ber mitogenetische Sekundarstrahlung aus abgeschnittenen Zwiebelwurzeln.TEWSKY. Gen. Die mitogenetische Strahlung des Bluts und der Gewebe von Wirbellosen. physikalisch-medizinischen Grenzgebiet". B. Z. DIETL. Das glykolytischc Spektrum. 211. Centr. SALKIND and and I. Dio mitogenetischen Spektra der Oxydationsreaktionen. 1934. BARNES. S. 37 Biochem. Sciences Biol. 217.. Wochenschrift 1385 95 detaillierte PONOMAREWA. L'emoinnesto intramuscolare. 387 . DESSAUER . 1931. das . il quarzo. Miinchencr Mod. II fenomeno (4). N. Blakiston. 1932. 352 149 see also BRAUNSTEIN and SALKIND. 1 109 . AUTHOR INDEX Page and K. and M.. f. 282 . 1928. 50. OTTO.204 POLANO. Abt. 18. Ober Beeinflussung des mitogenetischeri Effekts durch sichtbares Licht..Zehn Jahre Forschung auf dem . IT. Milano. 71. 1929. also: Eifetti citolitici da radiazioni biologiche (citofotolisi).

1933. heft Wissenschaftl. RICHARDS. Exp. 1934a. 1932. 1. 42 I. Heliotropisme de la cholesterine. and T. bei dor Entwicklungs- erregung des Seeigelcies. 1934. Die mitogenetische Beeinflussung der Eier von Protodrilus und Saccocirrus. 143 121. . genetischen Strahlen.. virus. 1924. 467 . (4). 169 Berlin. 57. 128. 1933. Electro. f. 9. aus Siemons-Konzern. Gynakol. GATES. Roux' Archiv 81. The nature of the factors which determine the Austral. p. Bacteriol. L. 1928. 630 I. 1 alkoholischcii Garung. 137 137 151 RODIONOW see G. MAX. (Congress of Electro. SALKIND. I. Ibid. 346 58 RUBNER. 95 819 S. Struktur und Atmung. A. JAHN. 144 A. W. 183 HOSSMANN. W. Untersuchungen tiber die Thcorie der mito57. 113 RIVERS.. 535 es ein Menstruationsgift t 40 Zentr. 445 85 RUYSSEN. Suppl. Med. PONOMAREWA. 1912. 1. T.. 120. 1921.. Der unmittelbare EinfluB der mitoRadio191 genetischen Strahlen auf den Verlauf der Zellteilung. Journ. SAENGER. 129. L. 148.. GAB OR.. duration of the period of lag in cultures of infusoria. Biol. Roux' Archiv 113. M. L'emission do rayons de Gurwitsoh par les reactions chimicjues entre gaz. Die Ernahrungsphysiologie der Hefczclle bci der Archiv f. 385 and G. 1928. und J.Radio-Biology.. Beitrage zur Analyse der mitogenetischen Effekte. Sci. 1934b. Veroffentl. 1930.. T.. 81. R. . J. Action cancerigene des irradiations solaires.FRANK. Venice B. Venice .. TAYLOR. Physiol. 1928. POTOZKY and ZOGLINA. Springer . 26. Acta zoolog. 45 B.. . a critique of the 69 The Biol. 1928.Mitogenetic Rays" yeast detector method. 1. 59. 206 Page and D. der Zellteilung und Strahlung. 0. 41 RUNNSTROM. 19. J. Ultraviolet Journ. 1933. Gibt 45. RBISS see MARGOU. 378 1931. and F. . Belgiquo.Radio-Biology. H. Die mitogeiietische Strahlung der Larve Roux' Archiv 70. 116 von Saccocirrus.. 1912. Heliotropism of cholesterol in relation to skin cancer. Exper. 182 Congress of 183 I. of Cancer 17. 47. liiternat. Roy. ROFFO. Sciences. 114. Am. and Med. 90. 1933. Journ. 5e serie. 74 Journ. Internat. H. I. A photoelectric ncphelometer. Acad..AUTHOR INDEX REITER. biologia 3 .. 105 Influence of washing etc. light and vaccine 48 ROBERTSON. Bulletin 63. Roux' Archiv 124. Sonderd. 155.

206
SALKIND
see also G.

AUTHOR INDEX
Page

FRANK, L. GURWITSCH and POTOZKY.

SAMARAJEW, W. N.,
see also

1932, Die mitogenetische Ausstrahlung bei der Regeneration des Regenwurms. Roux' Archiv 126, 633 ... 175

BLACKER.
95

SCHICK, B., 1920, Das Menstruationsgift. Wiener klin. Wochenschr. 33,

395 SCHOUTEN, S. L., 1933, GuRWiTSOH-Strahlen and Pilzsporcnkeimung. Acta Brevia Neerland. 3, 68 SCHREIBER, H., 1933, Zur Theorie der ,,mitogenetischen Strahlung".
Protoplasma
19, 1

80
79

und W. FRIED RICH,
see also

1930, tlber Nachweis

und

Intensitat der mito-

Biochem. Z. 227, 386 NAKAIDZUMI. SCHWARZ, 1928, Zur Theorie der mitogenctischen Strahlung.
genetischen Strahlung.
Zentralbl. 48, 302

92
'

Biol.

57, Biol. Zentralbl. 49, 57,

58
190

SCHWEMMLE,
421
SCOTT, L.

J.,

1929, Mitogenetische Strahlen.

0., 1931,

The determination of potassium
influence of the

in cardiac muscle

and the presumable
rythm.

beta-radiations on the

Journ. of Clinical Med. 10, 745

102

SEVERIN

see

BRAUNSTEJN.
L. B.,
1929,

SEWERTZOWA,
212

Ober den

EirifluB der

mitogenetischen
75,

Strahlen auf die Vermehrung der Bakterien.

Biol. Zentralbl. 49,

78

SEYDERHELM, R.,

1932, t)ber cinen durch ultraviolcttc Bestrahlung Kliaktivierbaren, antianamisch wirkenden Stoff im Blut.

151 nischeWochenschr.il, 628 SIEBERT, W. W., 1928a, t)ber die mitogenetische Strahlung des Arbeitsmuskels und einiger anderer Gewebe. Biochem. Z. 202, 115 64, 147
,

1928b, t)ber die Ursachen der mitogenetischen Strahlung. chem. Z. 202, 133

Bio64,

71

,

1929, Aktionsstrahlung des Muskels und Wachstumswirkung des elektrodynamischen Feldes. Biochem. Z. 215, 152
1930, Die mitogenetische Strahlung des Bluts und des Hams gesunder und kranker Menschen. Biochem. Z. 226, 253 65, 71,

71

,

153

,

1934, Die
allg.

,,

mitogenetische" Strahlung des Bluts in Handb. d.
II, 2, 1339.

Hamatologie Bd.

Berlin,

Urban und Schwarzen148

berg

und H. SEFFERT,

1933, Physikalischer Nachweis der GURWITSCHNaturwissenStrahlung mit Hilfe eines Differenzverfahrens. schaften 21, 193 1934, Zur Frage des physikalischen Nachweises der Strahlung. Archiv Sciences Biol. (Russ.) 35, 177
,

91

,

GURWITSCH9J

AUTHOR INDEX
SOLOWJEFF
SORIN,
see BILLIG.

207
Page

A. N., and M. KISLIAK-STATKBWITSCH, 1928, Cber initogonetische Induktion in den fruhen Entwicklungsstadien des

Hiihnerembryo.

Roux' Archiv 113, 724

145

STEMPELL, W., 1929, Nachweis der von frischom Zwiebelsohlenbrei ausgesandten Strahlen durch Stoning der LiESEGANGschen
Ringbildung.
,

Biol. Zentralbl. 49,

607

88

1931,

Das Wasserstoffsuperoxyd als Detektor fur Organismen89, 101 strahlung und Organismengasung. Protoplasma 12, 538
.

,

1932, Die unsichtbare Strahlung der Lebewesen.

Jena, Gustav
59, 89, 93,

Fischer, 88

pp

164

STRAUSS, W., 1929, Objektive Nephelonietrie mittels des MoLLschen Triibungsmessers, deinonstriert am Beispiel der Bakterienzahlung.
Centr.
f.

Bakt.,

I.

Abt. Orig. 115, 228

74

Sucirow, K., and M. SUCHOWA, 1934, t)bcr Coagulationsstrahlung. 100 Archiv Sciences Biol. (Russ.) 35, 307 SUSSMANOWITSCII, M., 1928, Erschopfung durch raitogenotische Induktion.

Roux' Archiv

113, 753

115

TAYLOR, G. W., and E. N. HARVEY, 1931, The theory of mitogenetic
radiation.
see also
Biol. Bulletin 61,

280

57,

58

RICHARDS. 90

TAYLOR, H. S., 1931, A Treatise on Physical Chemistry. New York, D. van Nostrand Co 46, TUTHILL, J. B., and OTTO RAHN, 1933, Zum Nachweis mitogenetigcher
Strahlung durch Hefesprossung.

Archiv

f.

Mikrobiol.

4,

565
190

66, 120, 121,

UNNA,

P. G.,

and L. GOLODETZ, 1909, Die Hautfette. Biochem.

Z. 20,

469

186

WAGNER,

N., 1927, t)ber den von A. GURWITSCH entdeckten spezifischen Erreger der Zellteilung (mitogenetische Strahlen). Biol. Zentralbl. 47, 670 57,
1909, t)ber die Oxydationen

58
142

WARBURG, OTTO,
Chem.
,

im

Ei.

Z. Physiol.

60,

443

1919, t)ber die Geschwindigkeit der photochemischen Kohlensaurezersetzung in lebenden Zellen. Biochem. Z. 100, 230

105

WASSILIEFF, L. L., 1934,
35,
,

De

Pinfluence

du

travail cerebral sur la

radiation mitogenetique

du

sang.

Arch. Sciences Biol. (Russ.)

104

150

G. M.

E. E. GOLDENBERG, 1931, Versuche uber die mitogenetische Strahlung der Nerven. Biol. Zentralbl. 51, 225 155
S., 1935,

FRANK und

WERNER,
Z.
f.

Physik WHITE see HILL.

92,

Die Entladungsformen im zylindrischen Zahlrohr. 705

29

208

AUTHOR INDEX
Page

WILDIERS, E., 1901, Une nouvelle substance indispensible au developpeinent de la levure. La Cellule 18, 313 138

WOLFF, L. K., and G. RAS,
geiietischcn

Strahlen von

1931, Einige Untersuchungen iiber die mitoGURWITSCH. Centr. Bakt. I Orig.
75, 76,

128, 257
,

190 132
76
109

,

1932, tfber mitogenetische Strahlen: II.

Biochem. Z. 250, 305
39, 121,

,
,

,

,

1933 a, Uber mitogenetische Strahlen: III. Biochem. Z. 259, 210 1933 b, t)ber mitogenetische Strahlen: IV. ttber Sekundar43,

,

strahlung. Centr. f. Bakt. I Orig. 128, 306 1933 c, V.: Ober die Mothodik zumNachweis der ,

GUHWITSCH80, 93, 116,

Strahlen.
,
,

Centr.
:

f.

Bakt. I Orig. 128, 314

....

146 190
143

1934 a, VI.

Le rayonnement secondaire. K. Akad. Wetensch/
45, 109, 113, 114, 123,

Amsterdam
,
,

37, 1

1934b, Effects of mitogcnetic rays upon eggs of Drosophila.
81,

Nature 133, 499

WORONZOWA see BLACHER. WRIGHT, W. H., and H. NAKAJIMA,
from
10
single cells of non-spore

1929, The growing of pure cultures forming bacteria. Journ. Bact. 17,

138
Boll. Soc. di

ZIRPOLO, G., Ricerche sulle radiazionc mitogcnetiche. Naturalist! Napoli 42, Atti 169

81,

142

POTOZKY and SALKIND. ZWAARDEMAKER, H., 1921, t)ber die Bedeutung der Radioaktivitat
ZOGLINA
see
,

fur

102 das tierische Leben. Ergebnisse d. Physiol. 19, 326 1926/27, t)ber das Erwachen des durch Kaliumentziehung zur

Ruhe gekommenen Herzens durch
Archiv
f.

die Bestrahlung des

Radiums.
102

d. ges. Physiol. 215,

460

SUBJECT INDEX
A
absorption of energy 2, 4, 16 of ultraviolet by water 23
gelatin 25, 59 adaptation to radiation ]J7
-

blood radiation 05, 72. S5,
-

I

IS

and cytagcnin 152
after injury
-

174

--

by

working Mi)

wounding 174
,

allelocatalysis 137

cause

<>f

150

Ambystoma ombryoH
metamorphosis 109

143,

145

during methamorphosis 107 in old age 151
in

wound

radiation 174

disease

152

amphibia, metamorphosis 1(57 amylase spectrum 37, 30 analysis of mitogeuetie effect J 19 anemia treatment witlicylagcnm 152 antagonistic rays GO

intensity 151

methods
-

72,

1

19
17!)

oi
-

-

cancer patients dogs 148, 150

man
rats

148
110
I

antagonism and radiation 172
antibiosis 172

staiving animals

(!)

atomic
-

radifitioii

32

theory 10
spectra 17
lit

spectrum 15O serum radiation 151 bone marro\v radiation 04
brain radiation 140.

atoms, excited
ionized 10

158

of tadpoles 145

axolotl see

ambystoma
C
It

-

bacteria as detectors 75, 70, 134 as senders 87, 131, 161, 177, 17S
,

17t> Cancer, blood radiation in caused by irradiation 183 - cells destroyed by \east 182

morphological change by
diation 161

inn-

definition

178

.Baron method 03, 06, 127 beta-rays from organisms 101 affecting organisms 102
bios 138

diagnosis 18O origin 1st

problem 177
,

radiation of tissue 178
,

blood drying for radiation test 149
grafting 151
,

spectrum 179 carcinoma see cancer
cell

menstrual, see menstrual blood
Piotoplafuna-Monotjraphiei) IX:

effect 134,

concentration and mitogcnetic. 13S

Rahn

14

80 - - . 100 Enchelys radiation 137 radiation 38* error. 125. see cornea and radiation human r day light affecting mitogenetic failure in mitogenetic - experiments 93 rays 108 death through irradiation 49. 143 larvae. 76. 142 larvae see tadpoles - radiation 50 dog blood radiation 148. 112 by radiation fruit flies. spectrum 147 erythema 48 exhaustion by irradiation 43. rotating 103. - 73. J31 secondary 117 see also spectrum 37 filters for light 21 detectors. radiation intensity 146 yeast buds 71 yeast volume 74 . definition 47 - and radiation 43 - chemical radiation 33. 105. 178 - and polyploidy 169 cytagenin 152. 142 as senders 142 183 affected chromosome number by electric. expcri mental. 77. 160 cotyledon radiation 141 crown gall origin 171. with bacteria 78 & 100 conduction of radiation _ J 1 1 Geigcr counter 34. 185 depression of mitogenesis 115 ~.210 SUBJECT INDEX drosophila eggs as detectors 81. general 190 colony si/e method 75 colloid coagulation as detector 89. radiating 145. 150 roots 111. radiation 145 cholesterol and cancer 182. 91 onion roots 58 - --- [u in in - muscle 154 nerve 112.false 70 - . field 6. 128 radiation 169 embryo enzyme radiation 143. 145 coagulation omitting radiation 89. 160 of radiating power 110 eye radiation. 50 chicken embryo. 113 over-exposure J15. 89. 154 tadpole legs 109 tissue cultures 83 ---- in yeast cells 110 -.radiation 69. 161. 180 cytophotolysis 182 -. 97.sources of in bio. 19 dissolution of salt as source of frog eggs as detector 81. 65 spectrum 35. 100 184 flocculation of colloids discontinuous irradiation 103 disks. cornea as detector 84 --. 187 heliotropifctm K eggs as detectors 81. 76. see also errors false mitogenetic depression 70 fermentation affected by radiation 85 causing radiation 124. 61 f . failure of 93 methods finger radiation 97. see drosophila dispersion 2. 169 chain reactions. 150 muscle radiation 29.of nerves 111.

149. 191 infra-red rays affecting organisms cornea 84 finger radiation 97 flocculation 89 101 from organisms 101 injury. see wounds egg development 81 Geiger counter 90 through over-exposure 49. 63. 66 1 14 blood drying 149 colloid coagulation formula 64 89 significance 79. 128 intermittent irradiation 103 generation time formula 78 glucose causing radiation in blood 148. 165. 184 metamorphosis affected by radiation 167 hydra. 39 mechanism of mitogenesis 119 menotoxin 95. 123 Geiger counter measurements 29. 95. f biological 79. 78 Baron induction effect and intensity 79. 185 hemoradiometer 66. 135 of secondary radiation 46. 169 leucemia decreasing blood radiation 152 leucocyte radiation 151 light affecting mitogenetic rays 108 filters stimulation. 161. see threshold of optic nerve 169 opaque for ultraviolet mitogenetio 25. 96. physical 23. 124 interference 1. 154 human radiation 95. see cancer maltase spectrum 37. 180 in protozoa 133 lag phase 67. 97. . 125. radiation 86. increase by selection 134 Liesegang rings 88 metabolic changes 84 mold spores 80 morphologic changes 86 14* . 120. 121 glycolysis spectrum 37 gradual increase in intensity 117 growth rate computation 78 larvae radiating 145. 185 intensity of radiation. 91 gelatin minimal. 185 menstrual blood. 114.SUBJECT INDEX intensity galls and radiation 171 gas effect from organisms 211 of radiation. 93 89 . 123. measure- ment. radiating 133 methods of detection: bacterial increase 76. 59 effects preventing 59. . definition 8 . see mitogenetic effect H 21 H O -decomposition 2 a Liesegang ring method 88 by mitogenetic rays 89 liquid yeast culture method 69 by infra-red rays 101 healing of wounds 175 beat controlled ^ heart radiation 102 M beta- by Helianthus roots radiating 139 seedlings radiating 141 heliotropism of cholesterol 183 malignant tumors. 115. 35.

spectrum 38 nuclei in onion roots 55. discovery 55. 156 exhaustion 111. 70. larvae 143 from 50 molds menstrual blood 86. . radiation 157 radiation 154 spectra 156 neutralisation. 87 physical nature of. J77. 148 bone marrow 64 - cancer 178 chemical reactions 50. see mitogenctic effect yeast buds 66. 115 - monochromator 19 morphologic changes by radiation 86. secondary radiation 109. 125. 119 stimulation. discovery 55. spreading of theory 128 photography 90. 147 neutralisation 50 nuclease action. 59 intensity.212 SUBJECT INDEX mitogen?tic 110 field methods: necrobiotic radiation 99 onion root 54 peroxide decomposition 89. wave length 49. 59. 110. 161. analysis J 19 and . nuclear stages 129 method 63 radiation 54 . 61 mitotin 140. protozoa 133 sarcoma 64 tissue cultures 81 tissues 146 detector mechanism 129 . 87 cornea 146 eggs 142 nephelometer measurements 74 nerve conduction 112. 77. 131. 113. 160 metabolism 156 . see intensity properties 59. 115. 178 N necrobiotic. formula of 64 . 129 nucleic acid. 160 . 59. . 161 effect. 96 82. 68 yeast growth 72 range 49. 103 . 44 onion root. 101 photoelectric counter 90 effect. 59. 140 spectra 140 opalina radiation 133 . 147 - . 52 embryos 143 infusoria 133 optic. cell concentration 134 cause of 122 muscle. 64 plants 138 polarized light 45. enzymes 37. spectrum 61 from bacteria 87. 119 sender mechanism 139 . 142 spectrum 40 muscle 65. conducting radiation 154 radiation 65. 99 tissue cultures 81 radiation. 51. secondary radiation onion roots 55 oxidation 29. 142 yeast metabolism 84 mitogenctic depression 70. yeast 64. radiation . 96. 72. 61 over-exposure 43. 80 proteolysis 52 of 43. rays 99 blood 65.

discovery of 55. atomic 32 chemical 29. 178 polarization 113. see secondary radiation solar 22 measurement 28 -. 124 theory of radiation 9 15. 52 persons by blood grafting 151 resonant radiation 45 in brain 158 spectrum 38 protozoa radiation 133. 33. 137 resorption of tissues 167 respiration of yeast and radiation 84 Q quantum .tube counter 28. 184 . 59. 115. . 113 reduction potential 86. 64 reduction potential 86. 160 . 125 reversal of effect with organisms 1 single producing effects 46. 91 parthenogenesis 169 polyploidy 169 tumors 171. 80. 158 yield 29. 37 definition 1 oxy cholesterol radiation 185 distribution in human body 187 . definition 16 . 122 spectrum 36. 213 U Radiation. photographic detection 90. energies 11 retardation through irradiation 70. reversal of effect 116 . human 95. tumors 169. 99 plate. 161 _- 169 potato radiation 141 parasitism 171 phosphate cleavage spectrum 38 photoelectric cell 25 - . infra-red 101 P parabiosis through radiation 160 Paracentrotus. . 92 photoelcctron. sensitivity 24 iation photosynthesis 105 physical nature of mitogenetic effect 59 plant radiation 138 - radium rays and organisms rate of conduction in nerves in roots 111 101 1 1 2 affecting morphology 96. see ultraviolet rad- . thermal 32 ultraviolet. secondary. 164 reflection 45. 114 recovery from over-exposure 43. see sea urchin . etic effect) mitogenetic (see also mitogen54 causing antagonism 172 cancer 183 healing of Paramecium radiation 133 parasitism and radiation 171 parthenogenesis through radiation wounds 175 morphologic changes 86. .SUBJECT INDEX oxidation causing radiation 29. 119 resonant 45. 120. . 80 as source of error 80 refraction 2 polyploidy through radiation 169 potassium radiation affecting heart rejuvenation of cells 67. 121 beat 102 proteolysis radiation 34. 171. definition 2. 50 . 4 by 80 polarized mitogenetic rays 45. 183 work function 27 . of mitogenetic rays 45. 122 reflection. 51.

sea urchin eggs as detectors 81 aa senders 142 - solar 22 retarded 86. effect on intensity measurements 123.sucrase 37. see intensity. 39 muscle 61.214 reversal SUBJECT INDEX of effect with photo- graphic rhythm 107 plates 116 in cell division 119 rhythmic interruption of radiation rotating disks 103. sedum leaves 141 123 sensitivity. and photographic plate blood rad- as senders 83 septicemia decreasing iation 152 solar radiation limits radiation 146. - of cells 108 of solutions 43 . absorption 23. 40 cornea 147 fermentation 37 frog muscle 35. . 61 glycolysis 37 * from chemical reactions 29. see Helianthus larvae changed by irradiation symbiosis and radiation 170 secondary depression 117 radiation. 161 sarcoma. 123.onion - roots ]40 oxidation 36. 124 T tadpoles affected by radiation 167 radiation of different parts 145 wound healing with 175 . of tungsten 32 of amylase 37. mitogenetic 35 thermal. 175 rate of travel in nerves 112 in r0 ots 111 for production of effect 104. 33 from organisms. . 114. see mitogenetic effect. and radiation. 112 spectrum of mercury arc 20 maltase 37. 98. definition 43 . definition 178 . mitogenetic Ureffekt 119 . atomic 17 molecular 17 . 164 by infra-red 101 storage of radiation 126 sunflower. thermocouples 23 threshold time as measure of intensity 44. 39 turbidity measurements 74 U ultraviolet rays. 25 . tumors caused by radiation 171 of plants 178 see also cancer . . 167 tonsillitis decreasing blood radiation 22 152 spectrometers 19 spectrum. phosphate cleavage 38 * proteolysis 38 39 a radiation 64 -. 147 -. 37 saliva affecting yeasts 96. blood 150 affecting organisms 48 bunsen burner flame 40 cancer tissue 179 -- photographic plates 24 causing cancer 183 chemical reactions 37.nerves 156 neutralisation 40 nuclease 39 -- . and thres- tissue cultures as detectors 81 hold.

. 161 radiation 64 affecting cancer cells 182 x-rays 10 volume measurements 73 .SUBJECT INDEX 1 volumetric method 73 215 W work function. 69 changed by radiation 86. photoelectric 27 wound healing by radiation 175 . yeast bud method 63. 161 radiation 173 treatment with bacteria 177 fly maggots 177 by plants 164 by saliva 96. 95. metabolism affected by rays 84 morphology affected by blood 96. 66. liquid culture method 69 blood radiation 174 _. 161 . 68.

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