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

in Chapter IV which discusses the various An approach be predicted from the results of physico-chemical to historical presentation is found methods used. and the important task is to find out in what points the investigations differed. a more conciliatory attitude has become noticeable since it has been shown that mitogonetic radiation not a mysterious force. though several factors responsible for negative results have been discovered. The book deals almost exclusively with mitogenetic rays w hich r exist in the ultraviolet range of the spectrum. With a phenomenon so little understood as these biological radiations. almost all happen to contain negative results. GOKWITSCH had not discovered these emanations 10 years ago. 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). #eta-ray emission from potassium it is biologically imporcell. but the result of biochemical processes. This very fact has been one of the authors' reasons for presenting the more important facts in this book. Many simple chemical reactions have been found to emit weak is ultraviolet rays. of the subject matter is not historical. they would An now investigations. just as strong after The arrangement logical. The objection this country. 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. It results. Both results are correct. to biological radiations has been strongest in but even here.VI of ultraviolet emission FOREWORD from living cells was the inability of some to repeat the positive experiments of others with the same results. ones. . and death as during life. it is not surprising that these apparent contradictions have not as yet been explained in every ease. it is tant. but attempt has been made to show that ultraviolet If radiation from living organisms is nothing at all strange. but not really characteristic of the living is proportional to the potassium content. and of the very few in English. 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.

FOREWORD The book literature is VJf on this subject. The authors are further under great. The authors an? further under great obligations to Mrs. not meant to represent a compilation of all This would have increased its size list of references. to see this field. MARGARET N. during a recent journey to A very brief summary is given at the end. BAKNKS . BARNES for her ceaseless assistance in editing he!]) in tin's book. chapter by chapter. August OTTO RAHX SIDNEY W. and to Miss A. obligation 011 to Professor ALEXANDER CUIIWITHOH for sound advice various points. (1032). This might be more useful in some respects than the customary Table of Contents. One of the authors had occasion. J. FFJKJUSON for her proof-reading. The literature compiled at the end of this book refers only to those greatly. up to the beginning the book by HTKMPKLL. of the entire book. Fthaca. 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. Europe. 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. of 1932.

.

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

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

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

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

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

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

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

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

of course. Such an arrangement is known as an oscillatory circuit. 7. The charge then begins to reverse its direction of flow. 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. The con) denser \\as charged to such a difference of potential that a spark occurred between the balls of the spark gap. not only line of force but in all directions. and S a spark gap. Diagram ot an electromagnetic wave. 7). oscillating rapidly between two .PHYSICS OF RADIATION An along one oscillating charge radiates energy. though the amount of radiant energy sent out in different directions varies. Experiments showing the wave nature of Hertzian waves. Plane ofElectrostatic Field Zero Figure is fi. HEKTZ (1866) caused a charge to oscillate rapidly between ei. for when a spark occurs the charge flows across the as to discharge the condenser. distance from the oscillating charge Mxtor Figure 7. and it continues to recharged with the difference of potential reversed. 1 In fig.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. c represents a condenser. radiated along the line of motion of the charge and the is radiated in a plane normal to the direction of motion.

When the velocity of these waves. cut and the edge viewed under a microscope (see fig. When the. it was immediately suggested that was nothing other than such waves. Experiment showing the wave nature of light. the silver compound was shine perpendicularly unaffected. was found to equal the experimentallyMono . the silver compound was reduced. the direct and the reflected beam interfering to cause standing waves. which arc of the wave length of short radio waves. offered a beautifully direct experiment not only showed that light was a wave motion but way of measuring the wave length. simple experiment performed by LLPPMA^N showed thatbe made to form standing waves and thus must have a wavelike nature. of surface placed normal to the radiation at that point (see . These loops and nodes showed that the energy was being radiated in the form of transverse waves. emulsion was developed. 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. Jight determined velocity of light. 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. at planes in between. Definition of Intensity.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. corresponding to the loops of This 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. only of shorter wave length.chromatic light G -* standing Surface Figure 8.

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

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

CO OC r CO oooooooooooooocoooooooooo : / - -^ A x x y x x .< v > % / >. / CC rH X V N X (M CO I.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^ .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 .

charged body. however. centripetal force upon it by q. For any value of r. .65 X 10~ 24 grams.12 CHAPTER is I by q and r. from each other.5 X 10~ 8 cm. then fig. \ q \ x *> Figure 11. F. it is found experimentits orbital electron . From elementary considerations it would seem that the nucleus with would form a stable system also for values of r other than 0. q. If the mass of q is 1. 11 represents a hydrogen atom. which is thus the unit of positive earth. Electron in a circular orbit traveling about the positive \ * charge q. it will. ' ^\ N Figure. 11). Positive arid negative ative charges at the distance r . a corresponding orbital velocity can be calculated which will bring about the balance of the electrostatic attraction and the centripetal force of the electron. ^ center (see fig. fall toward and the electron into the positively. 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.5xlO~ 8 cm. the distance r js 0. 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. Ef its orbital velocity is adjusted until its just equal to the force of attraction. inversely proportional to the square of the distance. 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. of course.. exerted then the electron will be in stable equilibrium remain- is ing indefinitely in this orbit. q. However. Suppose. electricity.10. An example becomes the nucleus of the atom and the electron represents the hydrogen atom's one orbital electron. and the charge q represents an electron.

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

Figure 12. which are unoccupied 3. The central dot marks the nucleus. about The outer dotted in the circles represent energy levels virtual orbits. amount of energy z would be required to remove either one of the K electrons from the atom. of the A't/ Jb'ig. i. solid circle there is but its showing the various one electron and is binding energy 5 electron volts. the binding energies which arc approximately 35 elec1 tron volts. The next circle represeiitsthe L orbit. which in Na of contains 8 electrons. e. In the last Diagram of the sodium atom shells. 12 is a wholly diagrammatic representation atom.6xlO~ 9 ergs. the first circle is called the K shell and contains two electrons. energy states associated with the orbit occupied by . I Electron Configuration for the Elements from H to Na. The binding energy these is of negative charges about 1000 electron volts or this 1. We have seen that the hydrogen atom can exist in various electron.CHAPTER Table 2. normal Na atom. The Emission and Absorption of Radiation by its Atoms.

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

1216 A. is readily calculated from the data this table.99796 x 10 10 X 6. it is 178 500 A. or is . After this length of time the atom reverts to its normal state with the resulting emission of radiant energy. if its energy happens to equal the energy difference between any two quantum . If the quantum is larger than Km En. states provided that. An electron so ejected from an atom ionized. sufficient to shift the energy is greater than (Eoo electron beyond the outermost orbit. i. Changes between other light states result in the radiation of visible and still others produce radiation in the far infra red. for the shift from the 6th to the 7th orbit. Tf its energy is not one of these discrete values. existence in the normal state. 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. in the far ultraviolet.unless its Kn). e. and the of its atom itself is said to be As has been stated before. it may be absorbed and the surplus. it will not be absorbed -?. 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). 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.e. . the atom is said to be in an excited state. then it may be absorbed. E When it has been raised to a state of higher energy E n by virtue of the absorption of energy.4 x 10.ia 1216 xK)" 8 cm. an electron spends most . or hv Ku). at this moment. as _ ~~ This wave length 2. It will be. the electron is in the orbit corresponding to the lower of these states. The life of an atom in an excited state is of the order of 10~ 7 to 10~ 9 seconds. is called a photoelectron.. In this event.547 x 1C" 27 __ ~ ~~~ 161.16 CHAPTER I The wave length emitted by the hydrogen atom for the energy change in Ex E for example. g. hr (K<x> electron away from the atom. of course.

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

14). is still the reduction of the salt with the deposition of Without presenting the theory of the process. become a chemically just this. molecular. 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 . together with the distribution of energy among these wave lengths. moo visible- A range Figure 14. The most prominent effect is that of light on silver salts as in all photograph ic emulsions. on the absorption of sufficient energy. unstable state. 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. and an incandescent solid emits all wave lengths with varying intensity beyond a certain wave length.18 CHAPTER T In this connection. 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. To realize that such an effect does exist. the effect of radiation upon chemical reactions should be mentioned. and thermal radiation. a molecule may some such spectrum as that of the center strip. Kepresentation of atomic. . E. depending upon the temperature of the solid (see lower spectrum of fig. The result is metallic silver. 14).and this may. a certain energy give rise to (see upper spectrum of fig.

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

. The wave lengths given in the table are those at which the filter ceases 1 ) In particular. the intensity of the unwanted be reduced to practically ZATO. 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. 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). a Hilger quartz double moiioehromator. 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. some degree of impurity seems unavoidable. some intensity of radiation of shorter 1 wave length than that desired is transmitted by the instrument. \vave lengths list may is of such filters By so doing. relative intensities of the lines in the ultraviolet spectrum of the quartz mercury arc as transmitted by such a monochromator.20 calibrated in CHAPTER wave lengths. In general. a discussion of their characteristics be omitted. While monochromators are designed to give high spectral purity of the isolated light. 1 ) Table 4. 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.

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

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

97 of This. 10 b in . circuit two dissimilar metals or alloys (sec U>a). no experiments mm are carried below 1900 A. Figure 16.97 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. The Thermocouple: The composed of thermocouple consists of a fig.PHYSICS OF RADIATION 23 distilled water. the photoelectric (1) and the photoelectric. the photocell graphic plate. Table 6 gives the absorption of ultraviolet by 10. The Absorption of Ultraviolet (16. which is proportional to the temperature difference of the two junctions. the detectors of ultraviolet light are: the thermocouple. Thermocouples left: a two-metal thermoelement. Modern high-sensitivity If thermopiles are frequently built as diagrammed in fig. right: a modern highsensitivity thermocouple. together with the absorption spectrum of oxygen. (J the junctions are maintained at different temperatures. shows plainly why in biological radiations. a current will flow in the circuit. Table 6. counter.

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

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

W . 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. that is remove an electron from the hv>Eoo E . hv. It is rare in this work to find the results of two The investigators coming within more than approximate agreement. ir o for various metals as obtained by different investigators. If light is allowed to fall upon the metal 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. With the best high vacuum technique known today it is impossible to prevent contamination with various atoms. since these data Du BKIDUE). such as oxygen. given. of each quantum be measured must be greater than the energy required to surface. which is a measure of the photoelectric current. 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. The result is tluit 1F for a given metal depends considerably upon its history and the has been freed from gases. to Necessarily. The energy required to remove a photoelectron from a photoelectric surface is called it. Table 7 gives the photoelectric work functions. The threshold wave length in A refers to the longest wave length which will eject photoelectrons from a given surface. 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. The purest surfaces by distilling metals in a high vacuum. . the "work function". photoelectrons are ejected from the atoms of the metal film and are attracted to the central wire. in such cells. (Further be found in Photoelectric Phenomena. chiefly those of the gases prevalent in the air. In a photoelectric cell. This equation applies to isolated atoms. These electrons flowing through the wire cause the galvanometer to deflect. hydrogen and nitrogen. the galvanometer reads zero indicating that no current is flowing in the circuit.20 the CHAPTER I window covered so that no light can enter the cell. the energy. only a few metals being care with which it are prepared .

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

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

Howewer. 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. 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. surface. o minute interval.FeS<). the experience of one of the authors shows that it is difficult to obtain sharply defined counts with this mixture. at.. p. Fig. The source of radiation \\as O 7 -|. O the radiation. More literature is quoted in Chapter IV. At lion the counter is exposed alternately tor 5 ininiit. with which these counters are tilled A\J 11 reduce the sensitivity of the surface. is called the photoelectric t/icltl of the The photoelectric yield is the reciprocal of the effiof a surface. A perfectly efficient surface would yield ciency one photoelectron for every incident quantum.es and shielded from the source of radiation. Yields of surfaces in photoelectric counters have never been as high as those in photoelectric cells because the active gases H. or the average number of quanta required to eject one photoelectron. 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. the riuht a tctam/cd at the left the chemical reaction K 3 2 sartorius muscle of the frou. some inert gas which a counter having a highly will not reduce the WERNER (1935) recommends 75%Ne and 25/ He. etc. (). It should be possible to Jill sensitive surface with sensitivity. "strays" or background radiation. 91. .PHYSICS OF KAD1ATION called L>9 "dark counts". a 5 minute exposure being alternated Avith 5 minutes of shielding.

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

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

that of the tungsten filament in an electric bulb. for 1 cm range of \vave lengths. Radiation which arises as a result of of iron is the temperature of a body radiation. wave lengths Table 9 gives the energy radiated by a tungsten filament at various for the two temperatures 2500 and 3000 0. this radiation is of very long wave length. 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. 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. At still higher temperatures. and in some of the hotter stars to a blue-white eolor. whieh is being heated. the radiation becomes visible when the temperature of the body reaeJies the witness the dull red color neighborhood of 500 C. They are fitted with metal 1 ) Kw(A) is the rate of omission of energy in \\atts. but solely upon its temperature. These are nothing more than discharge tubes filled with neon or a mixture of neon and other gases. the color of the radiation is changed to nearly white. However. Table 9. g. per unit solid angle. e. in a direction perpendicular to the surface. from 1 om 2 of surface. .32 CHAPTER II temperatures. It can be seen from these data that thermal radiators are poor sources of ultraviolet.

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

This physical proof of light from chemical reactions has been ultra-violet radiation ficiently sensitive instrument. . 25 experiments but one. 1 . by the oxidation of alcohol with chromic acid. In is all larger this.34 Jn this way. the emanation is greatly increased in the presence of diffuse day light. not from gla?ss containers. and by pyrogallic acid in air. vessels. Table 10. Recently. therefore it must be of a wave length shorter than 3500 A. It was also observed at this time that in some of the reactions. Ultra -violet radiation from chemical reactions 1 a nionochromator. 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. verified by GERLACH (1933) who showed that this radiation appears only from quart/. 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 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. trically also measured photo-electhe emission of ultra-violet light by several inorganic oxidations. only the rays between 2000 and ) Radiation passed 2700 A were measured.

By blocks. After the irradiation. solution It was by air.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. These radiations are so very weak that only the most sensiHowever. as far as can bo ascertained with this rather crude method. glucose -{. namely pyrogallic acid . photographic plate by a succession of tiuy blocks of nutrient agar on which yeast in the proper physiological condition was growing. 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.SOURCES OF RADIANT ENERGY 35 tive counters will detect them. milogonctic spectrum " 1 excitod muscle: B: tho quartz prism. The organisms most used in these experiments are yeasts the growth rate of which is accelerated by short ultraviolet rays under certain conditions. Kig. The biological aspects of this acceleration will be dis- cussed in Chapter TV. studied three in alkaline types of oxidation. this method. each block receiving rays C knou-n wave length. it would appear that all three oxidations gave the same spectrum. living cells under cer- tain physiological conditions react very promptly upon irradiation in the wave length range 1800 2600 A. Each yeast block was thus exposed to a definite" range of the spectrum which could be determined fairly accurately.. C: agar blocks with yeast on the exposed side. : T d^ "^ the yeast was permitted to grow for a short time order to bring out the growth rate differences.. by KFUNK (1929) to obtain tho spectrum of a frog muscle. KANNEC HESSE it (1931) working with yeast each representing approximately 50 A. 3* . From these results. 22 shows the first attempt. In fact. and to substitute m . and was then compared with the controls. they are so sensitive that it has been possible to use them in place of photographic plates for determining the spectra of such radiations.. in First attempt to obtain a Figure 22.

The limit of error is + 15. 23) the sections divisions of her spectrograph. Figure 23.36 CHAPTER The next advancement was the II division of the spectrum into separate strips of exactly 50 of glass needles and heavy A each. 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. and even the rather crude determination by 50 A strips shows . oxidations possessed certain specific regions besides the general The data of 7 separate experiments are oxidation spectrum. By this simple instrument. reproduced in Table 1 J The various spectra are not quite identical. differences which remained typically constant when the experi- ments were repeated. Table 11. Induction Efiects obtained from 50 ANGSTKOM stups of the Spectra of various Oxidations Detector: Cell Numbers in liquid Yeast Cultures . by means prepared a chamber of which corresponded exactly to the 50 A (fig. POTOZKY (1932).

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

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

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.

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

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

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

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

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

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

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

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

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

15. 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. egg white with pancreatm egg yolk with pepsin .4 %. Since all organisms display processes liberating energy.52 OH AFTER III absorbs the radiation (Table 15).5 All other enzymic processes which have been tested so far have yielded positive growth stimulation.NSCHINA (1029). of culture exposed through glass . This statement must be modified ultraviolet rays are . . containing the data obtained by KARPASS and LA. but after the bacteria once start to grow.4% . 20.7%. 10. it is only logical to assume that all living somewhat by the consideration that these organisms radiate. is also more complicated biochemical processes. .6 H 2 O ('ells per ec. Table 16. 31. - lfi. only one was negative. more rapidly.1%. Protco- by enzymes yields an ultraviolet emanation which greatly stimulates the growth of yeast as seen in Table 16. There is the usual lag period of 2 hours.K 2 ('O 3 + 2 MnO + 9 (<O 2 H. ex- Development of a culture of Bacterium eoli after posure to emanations from the reaction C 2 4 H 2 + KMnO 4 =. 25.0 egg yolk with pancrcatin fibrin with gastric juiee 30. 37. 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.1 %. Of 12 experiments.8%. the irradiated culture grows Table 15.5% 20. 36.6 % %.

and. . and since they may stimulate cell division they may play an extremely important role in the development of all living beings. Which parts of the various animals and plants Attention radiate. Since we must expect ultraviolet radiations from very many biochemical processes. 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.EFFECT OF ULTRAVIOLET RADIATIONS UPON CELLS 53 very readily absorbed. will not pass the skiu man or animals. will be discussed in Chapters IV and VII. for example.

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

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

POTOZKV and Ctontracting muscle (SiEBERT. BBOMLEY) . but not of normal rats \VITSCH) (L. It would scarcely be worthwhile to compile a complete list of organisms found to radiate. sarcoma GUBWITSCH. 24 hours old (ANNA Giwwrrscif) Young tadpole heads. only during the first two days of incubation After establishment of circulation system. intestine of amphibian larvae during metamorphosis (BLACKER. SIEBERT. pulp of tad})ole heads (ANJKTN. Bacterium murimorx (Acz) (SEWKRTZOWA*) Sarcina flara (BARON) Streptococcus fastis (MAGBOU) Yeast (BARON. cotyledons.: tails. 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. gills. GESENIUS. 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). the more important earlier observations.lactic acid -\. KEITEB and GABOB) Spleen of young frogs ( GUBWITSCH) Bone marrow (SIEBERT) Bone marrow and lymph glands of young rats (SUSSMANOWITSCH) ( Resorbed tissue. Radiating organisms and tissues Bacteria Hdclcrium tuwcfacicns.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. radiation of hi the ceases Embryos tkiis amphibia morn la stage (ANIKIN) Plant seedlings: root tips. of chicken. Staphyloeoeci (MAGBOU) JJaciUvti antHiracoidcif. FKANK) Pulp from resting muscle |. GUBWITSCH. and L. SORIN) Blood of man (SIEBEBT. RIOITER and OABOII) Turnip pulp. SIKBHRT. The following list contains compiled by GUBWITSCH (1929). ( J i u- Neoplasms: carcinoma.oxygen Corneal epithelium of starving rats. and GABOK) Blood of frog and rat (GuitwiTscn.

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

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

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

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

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

(4) Omission by GURWITSCH of the microtome sections showing a decreased number of mitoses when they happen to come between. Both papers describe the technique employed very carefully. 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. With yeast and blood as senders. This is no proof that they do not exist. Pressing or rubbing of the roots will increase the number of mitoses. that below this point. of roots (2) Light applied repeatedly in centralization which causes phototropic curving of the root and results in increased mitosis. but rather. the onion root as detector has been substituted by the time-saving yeast methods or by bacterial detectors. and confirmed by WOLFF and KAK (p. and after long continued and pressure were carefully avoided. and they arrive at quite different conclusions. absorption by quartz and air in the spectrograph makes aecurate determinations impossible. this author obtained also negative results. these were held above the ground by simple stands. pressing the opposite effect occurs. (3) Selection of good roots for important experiments which results hi increased mitosis through pressure. It can hardly be doubted from the Lirge amount of speetra analyzed by the Russian school. in a dark . 1932) observed that roots when removed from the water show symmetrical distribution of mitoses. and of less uniform roots when no effect is expected. two extensive recent investigations must be mentioned which concern the question whether onion roots can be used at as detectors. they do not. PAUL fastened the small onions (hazelnut size) by means of gauze to perforated cork stoppers. Since 1928.62 CHAPTER IV given of wave lengths below 1900 A. sections showing an increased number. 40). However. MOTSHEJEWA (1931. which then leads to negative results. that mitogeiictio radiation consists essentially of the wave lengths between 1900 and 2500 A. in a completely covered inoist chamber. no increase* of mitoses was observed upon exposure to another onion root. but all after repeated removal. Most of them do not mention the still more careful work by MAKUARETE PAUL (1933). Realizing the prompt reaction of roots to touching. This careful study has been considered by many critics to be the final proof against mitogcnetic radiation.

After this. However. the roots developed in the* air through the muslin in two to five days. The paper verifies GURWITHOII'S principal experiment. The exposed roots turn downwards. he spread yeast over the surface of solidified nutrient agar containing glucose. Almost always. long and absolutely straight.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 63 room at 20 C. this percentage was higher in the irradiated culture than in unexposed controls. When.5 cm. and theii exposed it to the radiating source for definite short periods. whether all mitotic The stages were included or only the more conspicuous ones. and they showed a uniform and symmetrical distribution of mitoses. In. however. it was considered a proof of a raitogenetic effect. The original method has since been changed in some details by BAIION (1930) and GUBWITSCH (1932) (see p. dried and stained. the exposed root grew more rapidly than the other roots of the same onion. and it will be. but the microscopic analysis showed no increase in mitosis at the exposed side. the yeast was spread on glass slides. usually not over 30 minutes. sender roots -were taken as controls. The onions were never placed in water. The measure was the percentage of yeast cells showing buds. . one could be exposed to a root from another onion without being cut or even touched and without the need of glass tube holders. The object of PAIR'S investigation was the establishment of a good method for studying mitogenetie rays with onion roots. When steel. 66). When the roots were 1 2. 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. while the leaves grew through the hole in the cork. the symmetrical distribution was disturbed. the sender root was substituted by a needle of stainless the exposed root also turned downwards. The yeast was then incubated for from 1 to 2 hours to permit the radiation effect to develop. The number of examples given is not large enough to draw many more conclusions. his first experiments. allowed it to grow for from 9 to 15 hours at room temperature. the starting point for all future work with this type of detector. b) The yeast bud method BAIION (1920) suggested that the rate of bud formation of yeast could be used as indicator of mitogenetie 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). i. quiet muscle did not do He attempted to produce radiation by changing this (Table 20). expressed in pereeiits of the control value. In all his experiments. the increase when the exposed yeast shows 33% buds. sender and detector were J separated by a quartz plate. or excited muscle to radiate strongly while the resting. He observed the working. __ 100 (exposed control control) SIEUEKT (1928b) used this method for a number of interesting studies in physiology. IV Table Effect of Yeast.. The numbers indicate the percentages of yeast cells published wit. in an oxygen Finally. Table 19 gives some of his experiments. is 9. Moreover. as increase in buds of the exposed yeast over the control.5%. Tt is now customary to record. since the addition of lactic acid alone would not produce radiation.<>4 CHAPTER 19.e. The induction effect is 37.5% over the control. results as "induction' effect. he obtained positive results by using a very dilute CuSO 4 solution as oxygen catalyst. and this is an increase of 37. he succeeded by placing the acidified pulp atmosphere. 24%.li buds. in air When . Thus. of Sarcoma. and the control. chemically the pulp of resting muscle to that of working muscle.

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

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

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

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

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

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

at least approx- The Russian investigators have repeatedly stated the imately error of their method. Tin's can be done very easily. knows the error of his methods. A schematic representation of the bud formation of n yeast culture growing at a constant growth rate. the percentage of buds decreased distinctly 1 in 2. though they have not given all the data necessary for others to check their computation. regardless by what method it is computed. can be seen from the fact that in most of their experiments. The claim of these authors that the error of the method has never been considered by the workers is entirely wrong. 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. Every biologist . g.METHODS OF OBSERVING BTOUMJICAL RADIATIONS 71 temperature which may be as low as 13 C in . That their cultures were far too old.Russia. The following computation is made from a study of the effect of upon organic catalysts (19281) KCN 1929). SiiflBERT e. .5 to <S hours. however. the error. from the data on experiments without mitogenctic effect. rarely publishes less than 10 experiments to prove an yone point. 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.

1 . because it onion roots. 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. The hemacytometer has been used by the Russian workers as well as by HEINKMANN (1932) in his studies on blood radiation. by measuring by comparing the turbidity by means of a nephelometer. For this reason. 0. While they are identical in principle. etc. they shall bo treated separately in this chapter on Methods. the respiration. the yeast bud method indicates a decrease in the growth rate while the total cell count shows an increase (e. of the exposed yeast suspension is then mixed with an equal amount of beer wort and incubated for three hours at 24 C. both in the control and in the exposed culture. All cultures are preserved for The control is counted twice. 116).5 ee. 103). better than any other of the* methods described in this book. cells. The plate count method has rarely been used.72 c) Detection CHAPTER IV cell by increase in number investigator not familiar with yeasts. 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. an equal amount of a 4% . The exposure lasts 5 minutes. An counting by the addition of H a SO4 example of the results and of the method of calculation is given in Table 22. and may be influenced by secondary effects. g. and it is made discontinuous by moving some object between blood and yeast at intervals of 2 seconds (see p. the preceding pages were written. There is a difference so fundamental that in a number of experiments. are indirect measurements of growth. the buds of yeast ascertains cell divisions directly while the mitoses in cells. Table. The total number of yeast cells (counting each smallest bud as an individual) is determined with a hemacytometer at the start. This method is used with yeasts as well as with bacteria. or by hemacytometer count. 34 p. (1) Yeasts: The number of yeast cells in a liquid may be ascertained by plate all the volume of count. It is the same as that used for counting bacteria except that more preferable media are beer wort agar or raisin agar. and again after three hours' incubation. The number of cells is the best measure of the growth rate. to the radiation of blood diluted \\itli MgS()4 solution to prevent coagulation. they differ in the method of measuring since yeasts arc so much larger than bacteria.

The measurement of the growth rate of yeast by cell volume has been studied in detail by LITAS (1924).control grows too rapidly. are suitable as detectors. 1932. After exposure. in othor words. of fresh wort. This The . especially in regard to cancer patients. i. and the method requires less time and less eye strain than the hernacytometer method. BUAINKSS (quoted from Ontwrrscu. and the curves and data published by this author show so little deviation only because they are However.*J hours. the yeast is distributed evenly. and that no effect can be expected when the. p. only cultures ]5 (1932) used beer yeast in a 20 hours old (temperature not given) which arc actively fermenting. He also added somo important new facts regarding radiation of the blood of old people and of patients with chronic tonsilitis. Volumetric Method: KALUNDAROFF strong wort (18 22 Balling). e. 17) has adapted the method for the small vnhimina available in 1 mitogenetic work.2 ec. when the control increases to more than double* during the incubation.METHODS OF OBSERVING BIOLOGICAL RADIATIONS Table 22. presented in logarithms instead of actual numbers. an will be shown in Chapter VI 1. also giving quicker results than the plating method. and a definite} amount of the exposed culture is is added to 1 ee. the volume is sufficiently accurate to prove mitogenetic radiation. 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. To avoid lag phase. g. measured by means of a micropipette. 0. period. that the yeast must he in . This means. e. else it would double in less than errors from this source. The accuracy is not greater than with plate counts. and incubated for 4 hours at 2s C.

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

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

rapid growth.76 delivers - CHAPTER ^oU IV of a cc. resulting in a decreased growth rate. WOLFF and HAS irradiated their bacteria in standard broth in a layer of 0.5mm. g. However. covered with quartz. pipettes used standard (Breed) method for the mieroseopie count of bacteria in milk. A fresh suspension of staphylococci in broth. almost doubled their number during the 5th hour. so is the control. using the slide cell method. A most interesting observation was the. the lag becomes shorter and shorter. and exposed (e. of Bacterium coli in a quartz dish I a layer of (U> mm. to milk \. exhaustion of bacteria good mitogenetic effect.6mm. (1931) allowed that a layer of standard nutrient broth 0. irradiated from below over the control. with about 20 000 cells per cc. Hy capillary pipette. After exposure. 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. indicating a return to the normal growth rate (fig.. there was no effect.) can be. a layer of 1 mm. The two experiments in Table 25 show a lag period of about 2 hours in the control Irradiation decreases this period very distinctly. only very small amounts of the culture (about 1 cc. FERGUSON and KAHN in tlie (1933) obtained good results with the ^ i cc. even then part of the bacteria \\ere shaded. thick transmitted only rays above 2500 A. continued irra- diation after the lag phase retards the growth for some time. and at 5 hours. if broth is diluted with 9 parts of water. During during lag phase.. or decreasing distance. by continued irradiation. still transmits some rays as low as 2200 A. This retardation of growth is only temporary most of the irradiated cultures . samples of the exposed culture and control are either plated on ugar. is placed in a glass dish in a very thin layer. 34). and with increasing intensity.rennet in a quart/ tube). ATetliod (the most recent method by WOLFF and HAS. than any of the cultures whose growth was distinctly stimulated. means of a . exposed. (1933) verified this observation. the dish with the bacteria is incubated at 37 (' for 1/5 to 30 minutes. FwimrsoN and KAHJS of a standard broth culture in 1 cc. 011 account of the strong absorption of ultra-violet WOLFF and RAS light by the customary bacteriological media. the control shows more cells per cc. 1933a). The plating of such minute quantities is necessary because. or brought into "slide cells" according to WREGHT.

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

see Table 41 p. and no rules can be given.4.8 3. incubate.78 CHAPTER The error of the IV method is mentioned in 1934. The best medium was 1 part standard broth plus 9 parts water. The induction effect calculated from the numbers directly is in one cast (]5 minutes exposure) 183. while in truth it is exponential. The errors are between 3. it must be considered that the number 183 has no real biological where d is the number t. 133. i. diameter.5 and 171. or in this 1 cc. cultures 4K hours old still older always responded. and tho transmission of ultraviolet by the medium. computed from the generation time.e. is only 27. 64. the increase by irradiation amounted to 25% series are given. at the start of incubation. 1)1. and plate at least every 2 hours for 6 to <S hours. of cells at the beginning.3. This computation implies that multiplication of bacteria is arithmetical. sity of the source.3. 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(>. this culture had a growth rate higher than the average of 27% the two controls. The growth rate is usually substituted the average by the generation time. 1 after the time 1 significance while the other indicates that during the four hours. face culture (37(-) as sender. Tho simplest procedure is to irradiate 1 ee.6. 24 hours old cultures never reacted. (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.1 3. 136). dilute exposure 1 10000 with dilute broth. this has been done by SKWKLITZOWA (Table 24) and Arz. : from abo\c). while the effect in tho same culture. the : ^ ^ i and 28%. The effect may not become apparent iff he cell concentration is too high (over JOOOOO per cc. after a quart /-covered glass dish. four detailed counts being 73. the best results were obtained with 15 to 30 minutes of irradiation 3. bacterium and found that it depended primarily upon the age of the culture. we have a really reliable measure. or through the bottom. The time of exposure depends upon the intenWith a 4 hours old agar surTable 20).1 7.9% of the total count. . However. Method: FEUGUSON and RAHN a good mitogenetic effect with (1933) studied the best conditions for cult. By computing the growth rates. of an old culture in dilute broth (either in a quartz dish of 4 5 em.2 4. As a matter of fact.4 and 4. the time required for cell to double.

cells Though HCHK.8 f-27 This procedure has also been used in Table. with the probable It is very unfortunate. minutes. and in one case even 103%.METHODS OF OBSERVING BIOLOGICAL RADIATIONS . for the time interval from 47. The "Induction definite Kffect" as usually calculated (p.6 . and occasion- ally more. With Nadxonia. various lengths of time by an agar surface culture of the same bacterium (The numbers are cells 1 per cc. the especially many investigators. 64) has no meaning. the generation time of yeast for the first 2 hours when the mitogenetic effect is strongest is mostly more than 2 hours. SCHKKIBEII found the variations tiaee/Miromycefi cMipxoirfcuti commonly in duplicate plates of to read) -40%. However.5 I Generation Times Induction Effect 45. In the tables given by iScuitEJBKR (1933). it seems from his curves that they were computed from less than 100 cells. e. Thus. 1 .5 14 ! 41.EIIJER does not give the actual numbers of from which he calculated the generation times. there is little hope of detecting mitogenetic effects. the deviation of duplicates went as high as 237%. This means that not all cells had divided in this time. the error becomes very large if the increase is small.5 40. the Russian scientists. With such a large error. after diluting the irradiated cultures 10 000 with broth) : Induction Effect 37 168 I83| 111 Generation Times. record. in Table 22. g. 24. 79 3 days old culture of Bacterium cult irradiated for Table 26. therefore. This makes the error very large and a comparison of the growth rates must necessarily result in enormous percentual differences. that error of the method. . 2 --6 hours 52. It could be used with yeasts as well. It permits no comparison.

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

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

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

the effect being 5. as with all other detectors. radiation began after 60 hours with a fibro blast culture from the heart of a chicken embryo. as was shown for yeast and bacterial cultures (pp. those on quartz grew more rapidly as measured by % means of a planimcter. the other half on quartz. and one of the two was irradiated for 48 hours by a beating embryo heart. distance was only Very recently.15. the irradiated culture appeared much denser than the control. CHRUSTSCHOFF believes that radiation is . By placing one half of a culture on glass. which was replaced when necessary. At this time. the average effect was 1. after 3 culture day being without radiation). CHRTTSTSCHOFF used the tissue culture as detector. JAEGER (1930) observed that blood radiation retarded the days (the last was far in tissue cultures. A fibroblast culture (chicken) was divided into halves.91 to actual stimulation by radiation from bacteria was therefore 0. Here. after 12 hours. JULIUS (1935) obtained definite growth stimulation of chick fibroblast cultures. In 56 such pairs. Another set of 48 parrs. even if tbo mm. The 0.17. RAWIDOWICZ (1931) as well as DOLJANSKI (1932) could find no growth of stimulation by mitogciictic radiation. but they did not respond at all to organic radiations. but without irradiation. 61) and 120). With a spleen culture of Amhystoma tigrinum. both were cidtivated in separate drops on the same quartz cover glass. 6* . DOLJANSKI used cultures which reacted promptly upon addition of embryo extract.92 culture. some LASNITZKI and KLEEinvestigators obtained negative results. ho obtained in all 4 experiments a more rapid growth in the culture nearest to the radiating substance. the exposed advance of the control. Next.o^ proving no chemical effect from glass or quartz. The induction effect was recorded as the ratio of increase in the quartz culture over that in the glass 0. but only on poor media.01 _j_ o. gave the ratio 1.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 83 head of a chicken embryo. due to autolysis of nccrotic parts in the culture. and exposing both to radiation from stapbylococcus culture. 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. It may be that a certain stage of development is necessary make the cells sensitive to the mitogenetic stimulus.3 times the probable error.

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

The organism used as detector was a wine yeast. and probably accounts for the depression of GESENIUS tried respiration. mutual irradiation of the cells must play an important role in thew experiments. to blood radiation GESENIUS (1930b) applied this test. free from cells. after the ceptions were only with cases of pernicious anemia.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 85 essentially that of WABBUKO (1923). If a failure occurs. blood radiation decreased the fermentation in 28 out of 30 experiments. He observed further that from patients with most diseases. improvement and found that normal blood 1 always radiated ). While yeast radiation produced no effect. The results can be briefly summarized in the following way: Fermentation in N-CO. e. The role in cancer diagnosis of this loss of radiation will be discussed in Chapter VII. further the iniluencr of radiation upon macerated yeast. and that the consistent exof the technique. The result was a stimulation of fermentation.) . His results agree very well with those of L. GLTEWITSCH and SALKIND and of SIEBERT. > 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. or of RUNNSTROM (1928) for measuring respiration of tissues or tissue pulps. i. This 10 to 100 times the maximal The which can develop in the medium used. 1932. the average depression being 14%. carcinoma and severe sepsis (see fig." (GESENIUS. upon zymase. it is time to test either the yeast or the apparatus. 7 to 10 billion cells per cc. *) "Healthy blood never fails. \ I atmosphere J f68 increase 102 ]02 cxporlmelltg t experiments < 4 decrease 3 | Respiration (oxygeii-uptake) in 2 j |40 decrease increase atmosphere. 40 p. The same retardation of respiration also could be obtained with sea urchin eggs during the early stages of cell division. the blood radiated. leucemia. but a retardation of the oxygen uptake. 153). which was exposed in quartzbottomed dishes to yeast radiation for 4 hours before being tested.

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

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

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

p. 40) regarding Figure 36.8% glucose. i. detector. just over the root tips. has been found on the deterioration of of ultraviolet radiation. the peroxide concentration in this drop is less than that of the control. a volatile substance produced by the sender.5 mm of quartz. Since this book is meant to be limited to biological radiation. e. right: effect through 0. 5/3). or a bundle of several roots. p. 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". the interesting speculations of STEMPELL (1932. under the influence 2 O2 : H One onion root. quite different by STEMPELL (1932. 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. On the opposite side of the quartz plate is placed a drop of 2 O 2 H . the possible biological meaning of the chemical emanations will be omitted.O. Flocculation of Colloidal Solutions: A promising method has been worked out by HEINEMANN (1934. It Ls based into H a O -f. and states that the LIESECJANCS rings at present are the only detector for this substance. is fixed so as to touch the underside of a thin quartz plate. . 1935) who observed that inorganic sols flocculate more readily when exposed to mitogenetic rays. Decomposition of Hydrogen Peroxide Another. The gold sol was prepared in the following way: To 1000 cc. since none of the other detectors for mitogenetic radiation react to it. of a slightly alkaline solution of 0.METHODS OF OBSERVING BIOLOGICAL RADIATIONS STEMPELL (1932. biologically. Gold sol was found to be more satisfactory than iron hydroxides. After long exposure in a moist chamber. He considers this chemical effect to be very important.

and covered with a quartz dish.5 1. None of them are absolutely convincing. the measurements are not as accurate as with physical experiments.90 150 cc. the gold salt clear gold sol. solution is then distributed between two beakers. small amount of solution is added just before the beginning of the experiment to make the gold sol unstable. The following readings were obtained in 15 consecutive minutes. A Nad The is each difference in turbidity between the two beakers is read every minute.5 2 15 NaCl. heating. This is a rule.1 g. in the absence of radiation. the readings remain fairly uniform. The It may reason must be sought in its very weak intensity. The* intensity is so slight that the most sensitive photographic plates and the most elaborate physical instruments have failed to record this radiation. fig. by means of a galvanometer. and those by BRF- NETTI and MAXTA (1930) and by PROTTI (1930). the* arrow indicating the moment when the radiating material control: was applied 3 5 7 to the quartz dish: 1.5 11 IG 2 11 20 2 11 1. and therefore a change of the galvanometer The reading while. This is done in complete darkness. They are placed into a very sensitive photoelectric differential nephelometer. The only photographic records are the ones reproduced in 15 of KEITER and GABOR'S monograph. and the supposedly radiant substance is placed into one of tho*quartz dishes. The best detector in most cases the living is still unsatisfactory since we must allow for considerable individual variation of the detector organisms. Upon is reduced by the glucose to a bright-red. 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. and as organism. of CHAPTER IV a neutralized solution of 0. . and GUBWITSCH has refused them all as proofs of the physical nature of mitogenetic radiation. AuCl 3 are added. TAYLOH and HARVEY (1932) could obtain no effect by exposing plates to frequently renewed fermenting yeast for ninety days.5 22 25 2G 3 3 blood: control: 0000001 1 1 7 8 10 1. Radiation causes a more rapid flocculation.5 1 0001 0^2 2 2 2 19 2 19 2.

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

In a recent paper. it ever was reached. 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. Though some early workers indicated higher sensitivities. 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 no increase with blood from carcinoma patients. In all cases except the first. KREUCHEN 4 6 from (1935) obtained yields of from 1C to 10 quanta per electron hydrogen-activated zinc and cadmium surfaces. it seems from later work that this value has never been sur(see of the surfaces used passed if. and a number of experiments with water blanks or non-radiant organic materials will be necessary to produce evidence which physically irreproachable. 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.92 Table 30a. A more careful description of the method of exposure. In his latest paper. and it seems well to view with caution the positive results claimed for the physical detection experiments so far carried out. are a step in that direction. 29) by the various investigators. but showed 110 increase when this was KCN. Most convincing is the experiment that a counter gave a definite increase when exposed to normal blood. in fact. In none of the physical measurements of mitogenetic rays is it absolutely certain that these errors have been excluded. 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. .

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

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

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

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

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

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

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

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

and is of importance to life An processes. After 15 to 45 minutes exposure.5 increase of 5.5 to bring the rate of peroxide decomposition to that produced by peas. obtained from a Mazda lamp by means of a monochromator. but of its potassium content. They exposed the unfertilized eggs of 2 sea urchin species and of one worm to infrared rays of 8000 to 12 000 A. Though it has been proved experimentally. The authors believe therefore that the reduced fertilization is caused by a photochemical effect. The decrease varied between 18. 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. The only case of near infra-red emanation known to the authors is. The only one known to the authors is an by their was necessary The temperature of own respiration. It originates from the radioactive fraction of potassium. is the same whether the . the eggs were fertilized by the usual method. it must therefore be of an infra-red nature. but an artificial experiment by NELSON and BIIOOKS (1933).METHODS OF OBSERVING BIOLOGICAL RADIATIONS 101 In 25 experiments. BETA-RADIATION entirely different type of radiation should be mentioned only in passing. the suspensions were brought into light. the peas rose 0.1%. The temperature difference between control and exposed eggs was not more than 0. namely the bet a -radiation of potassium. suspension which stood under the quartz vessel uniformly darkened sooner than the other. and really not characteristic of the living cell.5% and 87. Equally rare are observations of an effect of infra-red rays upon living organisms. however. The cell is alive intensity of this radiation or dead. D. The effect passed through glass.06 C. The irradiated eggs showed. it will not be discussed extensively here because this type of radiation is utterly unlike the mitogenotic and related is rays. in each of the 9 experiments a distinct decrease in the percentage fertilization. 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. E. and was not visible.

the systolic contraction". In 34 experiments. Nevertheless. rubidium. heart beat soon ceased again and could frequently be brought back a third time by to new exposure beta rays. this radiation is biologically important. After removal of the radioactive substance.64x 10~~ 6 ergs per second. started again in solution after about half an hour's irradiation by mesothorium (in glass) or radium (through mica). 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. uranium. METHODS OP OBSERVING BIOLOGICAL ETC. transform the potential energy of the heart muscle in response to node and bundle impulses into the enormously greater manifestation of kinetic energy. may tion. He states that "one may readily conceive that the free energy of the beta particles can be cumulative and.102 CHAPTER IV. 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. and be the chief reason for the indispensibility of potassium in ZWAABDEMAKEB (1921) was the first to test living organisms. . thorium. radium or ionium. reaching a maximum.

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

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

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

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

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

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

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

The from starving animals which are free from glycogeii. 55% 8 mm. 43% 9 mm. did not radiate. supports this view. 80% 6 mm. distant. 44). secondary radiation seemed to be glycolytic.110 ail CHAPTER V fact that livers these cases. 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. that one hour later.1 ram. has produced 41% less buds than the Exhaustion has also been demonstrated with chemical solutions (see p. The increase in buds was 65% at the irradiated zone. but as far as 10mm. When tho percentage of buds was counted. 10 mm. 79% 4 5 mm. 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. It will be shown later that the tips of onion roots are only secondary senders. of spreading. A very recent illustration for such exhaustion has been given LATMANLSOWA by (1932) on the secondary radiation from nerves . 2 100% mm. effect after 30 minutes. it was 33% 25% respectively Experiments with larger irradiated surfaces gave the same amount 12 mm. 87% 3 mm. there was a distinct increase not only in the irradiated region. Even then. this cannot be generalized because the secondary radiation from nucleic acid is not glycolytic. However. it unirradiated control. the primary rays are produced in the onion bulb. the organisms will still produce secondary radiation under the influence of an arc light spectrum. GTTRWITSCH irradiated such a culture through a slit 0. but not after 40 Tho same authors could also show that the production of Freshly-cut secondary radiation exhausted the plant rapidly. 80% 7 mm.. but not 30 minutes later. by oxidation. POTOZKY and ZOOLINA (1928) found a positive 45 minutes. Yeast cells may help to radiate imme- diately after being washed. This spreading of the mitogenetic effect can be plainly shgwii with densely grown agar surface cultures of yeast. This exhausts the yeast so much. 86% mm. primary radiations can be spread and transmitted to distant parts of the plant or animal body. from the border of the irradiated area. wide. By this mechanism. Another experiment with starving yeast cells throw some light on this phenomenon.

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

m. of root. 2. p. onion root. the average angle of 55 is 0.00306 seconds. The central angle for positive effects r.5 cm. Allowing the same time for transmission through the other 2. side view showing position of primary sender S. was required for processes other than conduction.5 cm. at 3000 Figure 41. Further experiments showed . Kotating disk for measur4 ing the rate of travel of radiation in secondary onion roots. This was sufficiently to 3 intense to permit the reduction of the window for primary radiation and that for secondary radiation to 1. CHAPTER V This signifies that a certain time (0. except that the 2200 A area of the copper arc was used as primary source. At right.00140 seconds. the time required for the secondary radiation to pass the additional The rate of conduction is therefore about 30 meters per second. of root is 0. The method was exactly the same. The accuracy of the in method was thus greatly increased. is conducted to another part and is emitted there. and the rate of conduction the nerve was found to be 303 meters per second.00166 seconds. the distance was increased to 5 cm.112 results. the total time required for transmission through 5 cm.. such as local reactions at the points of absorption and emission. Then. which means an average increase of 15. These values refer to a condition over 2. This is good agreement with physiological measurements on the rate in of conduction of nerve impulses. Recently. to between 40 was hereby increased and 70. and this is This corresponds. LATMANISOWA (1932) has measured the rate of conduction of secondary radiation in the sciatic nerve of the frog. to 0. In the experiments with secondary radiation of onion roots. and detector D.0022 seconds) must pass before primary radiation falling upon one part of the root. 0. radiation was observed from the same side of the root which had been exposed to primary radiation. The total time corresponding to The difference.00083 seconds.5 cm.

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

been pointed out above (p. and thus will become polarized. However. Table 46 p. . It is not possible. and recently also by WOLFF and RAS. effect" The customary way of comparing intensities is to compare the minima] time of exposure (threshold time) required to give definite mitogenetie effects. Examples may be found in Table 12 p. All previous measurements of intensities have become practically meaningless since that mitogenetie rays may (1934 a) showed easily become polarized. This method has been used repeatedly by the Russian workers. This does not affect the intensity of the biological radiation at all. and Table 49 p. 24) that. the reciprocity law (double intensity means half as long exposure) does not hold with very low intensities. If the number of impacts induced by biological radiations is expressed in percentage of the stray radiations of the surroundings. 79.114 CHAPTER V least part of this radiation will be reflected from the cell walls. 145. and that WOLFF and RAS polarized rays have an enormously much stronger biological effect than the ordinary radiations of this type. at the present moment. One very simple reason for this is the usual method of recording the results. latter evident method of recording resu]ts becomes most applied to physical measurements. there are physical reasons to warn It has against quantitative conclusions from threshold times. 44. expressed in percentages of the cannot possibly be used as quantitative measure. 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. to foresee all the consequences of such polarisation. with photographic plates. 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. The "mitogenetie as used especially by the Russian investigators has no It is not surprising that it was quantitative value whatever. The "induction effect" as the increase in the exposed yeast over that of the control. as explained on p. never possible to use it for measurements of intensities. D.

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

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

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

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

The customary method is to . he succeeded in proving it with onion roots. and all other effects of secondary importance. these were the large first detectors. Hence. he concluded that this factor could not be chemical. and since he made a them.T proved this radiation to be emitted also from purely chemical oxidations. From a certain rhythm observed in the division of the sperm cells of amphibia. GURWITSGH has always distinguished between the "Ureffekt". number the most of experiments with familiar of all detectors. 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.CHAPTER VI ANALYSIS OF THE MITOGENETIC EFFECT At the present time. GURWITSCH (1922) predicted a factor which controlled cell division. but must be of a physical nature. This chapter does not offer one theory. Mitogcnetic radiation was considered at first merely from the cytological viewpoint. they are to him probably However. and in 1923. and in plant roots after special treatment. The number of not offering of mitoses in different roots oven from the same bulb varies greatly. we lack a clear conception of the working mechanism of these rays. the mechanism by which short ultra- violet rays affect living cells is not understood. All of GURWITSCH'S speculations and explanations start with the results obtained with onion roots. From the mode of action. but presents a number of attempts to account for the various phenomena observed. These rays were not discovered by chance. the primary effect which is the increase in the number of mitoses. they have the great disadvantage perfect controls. the physico-chemical viewpoint entered into consideration. as an emanation produced somehow in the very complicated process of cell division. when SJEBEJK.

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

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

for a short time. 66) suggests this strongly. This need not necessarily involve a mechanism different from that assumed It may well be that in the ageing the additional. This may also bo the cause of the mitogenetic effect in onion roots (see p. The description of the physiological condition of BAKON'S yeast plate (p. and so does HEINEMANN'S hemacytomcter method (p. B.122 set CHAPTER VI potential necessary for normal cell functions the candle which then keeps on burning as long as the cell feeds normally). the percentage of buds was not at all proportional to the length of irradiation time. namely immediately before entering the resting stage. sufficiently in the rejuvenation process. where mutual cell influences are practically excluded. cell. Or. the cells. When freshly prepared detector plates were exposed for different lengths of time. under the more favorable conditions of rejuvenation. 72). This may be the same mechanism which. TUTHILL and RAHN. properly dosed energy from the sender prevents a certain phase of the ageing process. cell division might give us the clue for the mitogenetic There seems to be another stage where cells respond. at the point of going to rest. The volumetric method as described by KALENDAROFF (p. it must be kept in mind that so far. the later stage of active cell division does not seem to be greatly influenced by these rays. Whatever up the reduction (light be the explanation. BAKON'S yeast plate. or the But even with the yeast plate of volumetric yeast method. long to permit one more cell division. is stimulated so greatly. We could not expect this with detectors involving secondary radiation by old cells. such as the onion root. the following percentages of buds were found (after 2 hours of incubation) : . 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. 129). are stimulated to at least one more cell division by irradiation. possibly. Apparently. It appears that the difference between rejuvenation and active effect. the cell wall becomes transparent to these rays only at one certain stage of development. 73) appears to make use of this stage.

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

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

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

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

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

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

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

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

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

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

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

J Jt is . 612 1224 2436 cell 2 4 8 hrs. the average diameter as 6.036 0. The volume of one 8 yeast cell is taken as 1 18 ^ . Without accurate statements concerning the number of cells in the sender and in the detector. Lag Period Average cell distance 1 ) 2094975 209498 20950 2095 1 ) . the stronger must be the effect. CHAPTER VII His data show a very consistent increase in the mitogenetic effect through 6 to 7 such transfers. The rate of growth of old cultures can be increased for several successive transfers.192 0. This observation had been very puzzling to biologists since tho opposite should be expected. 120).422 0. so that all the decimal gradations from 2095 to 20949750 cells per cc. the quicker the recovery. The times required to reach the maximal growthrate were: The I) Table with an inoculum of 20 949 750 cells per cc.086 0. rejuvenation is slow. and single-cell cultures frequently die.914 mm. no definite conclusions can be drawn. an old culture to a fresh medium. however. The proof is not complete.134 the next experiment. The old cells undergo a rejuvenation process. absolutely correct only with spherical organisms.04 ||/~~ / \ cell volume in 100 cc. These different inocula resulted in very different lag periods. do not start growing at their maximal growth rate. morphologically and physiologically (2) Reaction Upon Irradiation: when transferred from This results in a very slow growthrate during the hours after transfer. 8 0. without any irradiation. first culture rejuvenate more rapidly if many cells have been transof fresh ferred while with a few cells in a large volume medium. Bacteria* and yeasts. until it could be explained as a simple stimulating effect by mutual irradiation of the cells. It is well known that young old bacterial cultures require a longer time to start growing than ones. The closer they are together. best example is very likely that of HENKICT (1928. It has been observed by RAHN (1907). PENFOLD (1914) and others that bacteria after being transferred from an old (see p. with yeast. The average distance distance is computed from the equation = ~ cell diameter 74.4 //. He inoculated increasing amounts of yeast into flasks of sugar -peptone solution. The bacteriologist calls this the lag phase. were present..

59). as a result. and. ETC.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 39. After having formation in fresh yeast cultures begins sooner is BAKON larger (Table 39). the lag phase became independent of the cell concentration. ftelatin Table 40. is shortened by the mutual irradiation of shown that bud when the inoculum (1930) added another. muto -induction was prevented. 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. and while all other conditions of life remained the same (gelatin contains no nutrients for yeast). he showed that is this difference disappears when gelatin added (Table 40). 135 Growth Acceleration by Mutual Irradiation with Saccharoniyces ellipsoideus To these proofs that the lag phase (jells. .

? ^ r} A A <M CO ^* <O 00 .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^ .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 - .

the only difference being their distances Table 41 presents the data from one another after dilution. . 10000 and 1000000 times. Table 42 gives the progeny derived in 44 hours from single cells viEnchelya in hay infusion drops of various sizes. 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. i.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 42. came from the same 1 cc. 1 cc.15 cc. the inhibiting volume is 0. we must conclude that a single cell of culture A in 1. In large drops. This is cultures usually borne out by experience. and if extrapolation is permissible. verifies this comparably. it differs by the fact that the size of the drops has a great influence.. with culture B. the absence of a mitogenetic effect when the cells are too close together. appears a duplication of the lag phase of bacteria. it from Vioo cc f ^ nc concentrations. and then diluted in broth The cells in all three cultures 100. when grown in different cell The slower growth due to greater dilution is The more important result is plainly evident in the controls. ROBERTSON mentions that such show no growth. 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. 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). ETC. While the growth is much slower. (3) to fungi. of a culture of Bacterium coli was irradiated for 30 minutes. would not grow at all. - e. shows the development of the progeny arising kl culture.5 cc.

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

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

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

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

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

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

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

165). insects.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 46. and cease to radiate 90 hours Quite different is the behavior of the chicken embryo. 168). 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. 146 Effect of various parts of the embryos of the axolotl upon onion roots Of the analyzed. 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.Monographien 10 . positive results were obtained with the zone around the embryo. However. during the second and of incubation. only the larvae of Drosopkila have until shortly before after this (see also They do not begin to radiate pupation. p. 169). Pro to plasma . ROT: IX R a h n : may also be mentioned here (see p. The morpho- changes in sea urchin larvae observed by MAC. ETC. liquefied (d) Embryos as Detectors.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

1 more in one series of 33 tadpoles and The actual 1. In the quartz -bottomed vessels. gills and intestine have BLACHER and associates (VII) further showed that there was a relation between the resorbed tissue and the developing limbs. a second series with 44 tadpoles. it could be shown that a tadpole lying in a quartz -bottomed vessel will show a more rapid growth of Figure 51.8 less in one series of 22 than its mate in the cavity: 6. 4.3 less in more rapidly. and therefore shaded against radiation.2 series with 51 tadpoles. Intensity of radiation of the resorbed organs of frog tadpoles.5% i tadpoles. while the other leg which served as control remained covered. The result of the operation as such was that in the vessels with glass bottoms.168 CHAPTER VII maximum of blood radiation occurs after completed their metamorphosis. Finally. the legs if the bottom of resorption. the freed leg grew a little more slowly 0.1% 1.4% ^ . however. and that transplantation of gills increased the growth rate of the corresponding foreleg. retarded growth.5% i 1. or the freeing of the foreleg from the gill cavity where resorption of the gills occurs. the freed legs grew and 4. in the other it 7. during successive stages of metamorphosis. 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. Removal of the gill.

He had succeeded in causing cell division in unfertilized frogs' eggs by only a very few percent of the eggs thus treated initiated development. the plants tumefaciens. and radiation nearly ceases after 30 hours.8 in the other set of experiments. but in some cases. Seven shoots arose directly in the tumor and of these. V). and OAROK GURWITSCH'S statement that there is after having verified always radiation around a wound. and an accelerating effect of this radiation upon the rate of development of the organs concerned. To test this. BATAILLON obtained tadpoles and even frogs by this method.. RKITEB. of course. above the tumor in order to induce the development of new shoots. and the axolotl Ambmtoma tigrinum which retains gills as well as tail throughout its life radiates still less. HI). They do not radiate until about ready to pupate. are. One of them proved to bo tetraploid. The triton Pleurodeles Wallii shows similar radiations i (BLACKER etc. Change of Chromosome Number. Positonly the radiation of the resorbed tissue. Parthenogenesis by Radiation. The maximum intensity is reached 24 hours after pupation. After the development of the tumor. 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.1% 1. 5 could be rooted. The gain is 7 times that of the average error. 120 tomato plants were inoculated with Bact. were cut off 3 to 5 cm. though less intense. very fine needle pricks. too scant to permit tin* of conclusions. 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. 169 1. While these data drawing 5. . Experiments were made also with Drosophila larvae (BLACKER etc. is 4.6% and 12. There is no direct proof that in these cases. they are suggestive of a new cause for polyploidy. It has been repeatedly observed that polyploidy occurs fairly frequently in some individual branches of tomato plants infected with Bacterium tumefaciens. the radiation of the resorbed parts ively established had any distinct form-giving effect.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC..4 in the one set gain by irradiation was therefore 10.

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

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

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

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

that radiation one of the necessary While the argument concerning the sedum leaf wounds lias never been settled experimentally. but the old cells immediately under the newly-formed tissue. and a minimum at 24 hours. The threshold time to distinct was the measure effects mitogenetie necessary produce of intensity. . BROMLTGY (1930) has proved that the tail of the salamander Anibi/tttonui tigrimim radiated strongly after being amputated.4% 50. FIUCHIMOfutic. 4 tadpoles of the frog Pdobatcs drop in intensity after 4 days. and it stayed at this low level during the .2% 2.9% 48.(>% 41. VJ1 is but. 2 minutes were required. Exposing yeast for 18 minutes to the ground tissue. They cut 10mm.174 besides the physical ones factors. the entire body is affected the wound upon blood and brought into play. The radiation maximum after 1 2 days. and after different times. The same double maximum could be observed with wounded BLACIIER. The blood of normal. 1 minute of exposure to blood sufficed for a similar effect 24 hours after injury. CHAPTER . by LTOSNKR and WORONZOWA (1932 b). from the tip of the tail. an exposure of 15 minutes became necessary. the blood radiated less than half as strongly as that of uninjured tadpoles. It is not the new tissue which radiates.utt.1% 29. 12 hours after injury. the following "induction effects" were obtained: 1 23 5 6 12 24 hours after amputation 6. In Table 50. and a great WITSCII.1% 39. ground it up and used the pulp to irradiate yeast plates.4% induction of the injured tissue reaches a It requires 3 hours for radiation to be established. This means that at this stage. Very interesting was the observation by the same authors that the blood of the tadpoles changed its radiation decidedly upon wounding of the tail. removed the old tissue which produced the regeneration. decreases decidely and reaches a second maximum after 5 days. entire 18 days of the experiment. By this long-continued effect of radiation. uninjured tadpoles produced a marked effect in 5 minutes. two distinct maxima are seen at 12 and 36 hours after wounding. and on the fourth day. This agrees with the two maxima in pll of healing wounds as observed by OKUNBFK (1928).

leaving earthworms. method of measuring the amount of regeneration. More than 500 tadpoles were used for Into their tails were cut triangular wounds. method of wound- the tadpole tails for irradiation from below. at right. and found that the first mitogenetie effect could not be detected until 20 hours after indicting the wound. figure. At ing left. The rate of healing was measured as shown in the same 52. LiosNER and WORONZBWA (1932 a) succeeded in proving this by producing a more rapid regeneration of wounds by biological irradiation. fig. Kadiation SAMABAJEFF was weak throughout. but was noticeable even after days. bringing about healing. ETC. under a low power lens. with experiments them in earth. as in this proof. and it is here recorded in ft of new .THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS Table 50. IRICKIMOWITSOH. 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. Figure 52. 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.

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

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

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

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

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

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

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

61% 3. i.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. not a single animal had developped a tumor. It would be a splendid agreement if this range which is known to stimulate cell division could also cells. exposed to ultraviolet light. avoiding tho hottest sun. Then. The author has not been able to ascertain whether ROFFO has tested the mitogenetic range of rays by itself. there is a decrease of 25 to 40%.07% 1. sunlight as well as ultraviolet light increased the cholesterol content of the exposed parts of the skin. 183 cancerous lesions to be much higher than the normal skin of the same patient. Within 4 months. The mode be shown to produce abnormal cell division of the epithelial The fact that sunlight can do this speaks very much against .. which was exclusively on the naked parts of the body (ears and eyes mainly. 52% of all rats had developped cancer. development and the histology of the rat cancers corresponded exactly with that of human cancers. e. ROFFO (1934 a) studied the "heliotropism" of cholesterol very thoroughly. According KAWAGUCHJ to MALCYNSKI (1930). (1930) had shown that the cholesterol content of the skin increases when it is exposed to sun light.50. as average of 5000 cases: skin of tho face skin of the back of the scalp skin of the foot hand 95. 700 rats were kept in sunlight daily for not more than 6 hours. of an intensity of 14 erythemal units. every one of these rats showed tumors. and many of them had several tumors. also twice on the hind feet and once oil the none). He found the frequency of skin cancer to be distributed in the following way. ROFFO (1934b) studied the tendency for cancer development in white. the range between 1800 and 2600 A. beginning with 5 minutes per day and gradually increasing the exposure to 6 hours. With white rats. This agrees very well with the above-mentioned frequency of skin cancer on different parts of the human of body. He proved it to take place only in living organisms.02% 0. rats. After 10 to 11 months. The wavelength of the ultraviolet was above 2300 A. Of the control rats receiving light from a tungsten filament lamp. ultraviolet irradiation of healthy persons increases the blood cholesterol while with cancer patients. 150 rats wen. X-rays or radium rays produced the opposite effect.

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

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

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

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

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

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

KANNEGIESSEK and SOLOWJEFF. 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. TUTHILL and RAHN have also published two sets of counts of yeast buds from many This has been done (p. omit the work of SIEBEJRT who published more detailed experiments than any other investigator in this field. Other error limits can be found on pp. 58) different parts of the same culture. GIJRWITSCH ciates have stated repeatedly that with the yeast and his assobud method. 71. 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. When the error or the reliability of the method not stated as such. The reliability of the yeast measurement by volume can be estimated from the data by BILUG. NAKAIDZUMI and SCHBEIBEB. working with the yeast bud method. 34 (BAETH) and 168 (BLACKER). 78). the mitogenetic effects were more than 3 times the experimental error. 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. 92). This explains the difference of error in the onion root experiments which was (see p. HEINEMANN has stated that in his method of counting yeast with the hemacytometer. The frequently made statement that the biological investigators do not state the error of their methods is not in accordance facts. WOLFF and RAS have frequently given all individual counts of bacteria for one experiment (p.190 OUTLOOK or even in the same laboratory with different investigators. some weak papers are quoted extensively while some of the best proof in favor of mitogenetic radiation is completely omitted. . even physical measurements show great differences (see Table 30a p. Any serious criticism should start with the most reliable and best founded papers. it is usually possible to compute the error from those experiments where no mitogenetic effects were obtained. 83 (JULIUS). they consider any increase less than 15% over the control to be experimental error. 58). 10% with some investigators and 50% with When it comes to such delicate instruments others as the GEIGER counter. It is somewhat surprising to find that in critical summaries.

sornea.ornea are affected by the season. that the reader does not or lad it 3% is 25% of know whether the controls buds which would have given a suggestion Therefore. mper largely lost. 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. It might be argued that the burden of the proof lies with he opponents since the mitogeneticists claim to have proved heir point. and the roots as well as the number of mitoses in the . or the yeast volume measured. the value of the least of the condition of the yeast. the is term "yeast" means very little. Another justifiable argument against the weight of some jublished papers is the lack of precision in the description of the nethod. 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. the publication of the complete records vould give the reader a conception of the reliability of any >bserved effects. As an example may be mentioned the paper by SALKIND PONOMARBWA (1934) which might be very important for the However.OUTLOOK On 191 the other hand. .he chemist whose methods are really standardized publishes merely the formula of his new compound.he authors do not mention the age of the yeast culture. it is of inite. physiological explanation of the mitogenetic effect. but such an attitude would not be very helpful in lolving the real problem. the critics have good reason to disregard which give no precise account of methods or results. nor ind he medium on which ffect so it was grown they give only the induction . statements as "8 10 hours at room temperature" are too indestate. Since the error in biological experiments varies yith the investigator. even when the standard deviation has not been lot Computed. . Even /ell a good deal about the performance of the experiment. but only in a definite physiological Such greatest importance to describe all details. the percentage of buds. Since the biological detectors do not respond at all imcs to mitogenetic rays. but also the actual inalytical data. We are dealing with a very complex .

a settlement of the question of mitogenetic radiation will be accomplished in a very short time. and both sides should do all they can to bring about a real understanding of the facts. . 93).192 OUTLOOK phenomenon. reasons (p. and the authors hope sincerely that by pointing out the mistakes and misunderstandings. 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.

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

ARNOLD. Das Wachstum der Gewebskulturen in vitro und die GuRWiTsCH-Strahlung. 184. d. Das Monotoxinproblem und die mitogenetischen Strahlen. Zentr.. fra fotografia a la eccitatione il Cultori delle CHARITON. 49. 95. DUGGAR.. 1931. R. 129 35. Nachweis mitogenetischer Strahlung durch beschleunigtes Wachstum der Bakterien. and OTTO RAHN. Irradiation of plant viruses and of microorganisms with monochromatic light.) 36. G. and W. W. Roux' Archiv 121. HOLLAENDER... 1932. A. BRUNETTI.. 1929. (Russ. Sulla delle radiazione del Gurwitsch. Bact. and A. 411 60 50. Ges. RODIONOW. R. J. Roux' Archiv 126. Archiv Sciences Biol. MAXIA. 1930. E. Journ. Das mitogenetische Reizminimum und -maximum und die Wellenlange mitogenetischer Strahlen. 121. 137 Zum FRANK. exper. 634 138 and S. of Research 7. Mikrobiol. 674 76. 321 29. K.198 BROOKS see AUTHOR INDEX Page NELSON. A separation of the reactions in Journ. STAIR and J. deutsch. DOLJANSKI. Archiv f. M. e Nat.. CHRISTIANSEN. 15. 39. 249.. G. 1929. Die Quellen der mitogenetischen Strahlen in Gewebekulturen. 391 107 FELDMAN. hot. 83 A balanced thermocouple and filter method for ultraviolet radiomotry. Archiv f. Die gegenseitige Beeinflussung der Seeigeleier als mitogenetischer Effekt betrachtet. 1930. Cornell University 1933. XJber die Ursachen des Gewebewaclistums in vitro. 4.. FRANK and N. Gen. H. W. i>u BRIDGE sec HUGHES. Physiol.. FERGUSON. COBLENTZ. 185 CHRUSTSCHOFF. 723 48 37 DECKER. KUREPINA. Naturwisson- 90 schaften 18. HOGUE. 109 . Morphological changes in yeasts induced by 96. KANNEGIESSER. 145 DIETL see POLANO. Phila- delphia. 161. tJber die Wellenlange und Intensitat mitogenetischer Strahlung. with practical applications. 1932. Biochem. Neoplasms of Domesticated Animals. 1932. 34. G.219 83 48 EMERSON. W. and C. Science Med. M. 60 and M. 207 VAN DOORMAL SCO AUDUBERT. 47. B. Cagliari 2 J. t)ber die Scharfe mitogenetischer Spektrallinien. 78. 203 R. Saunders 177 161 1932. Z. Thesis. 367 86. 1932. biological radiation. Ber. 126. G. 1934.. Physikalische Untersuchung mitogenetischer Strahlung der Muskeln und einiger Oxydationsmodelle. 1934. 1930. W. Biol.. Atti Soc. photosynthesis by means of intermittent light. I. Journ. L. 1930. 91. 27. Bureau of Standards. Zellforschung 9.

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

central. 220 1932 a. 1929.. 11. Techn. 1375 72. Zentr. ANIKIN. t)ber . Z. bei HAUSSER. 1918. rend. Thomas. Z. K. Physik 15. 226 MALL see JONES. et Teclairago monochromatique dc 1'oeil. Roux' Archiv 113. 43. Morphologic Variation and the Rate of Growth of Bacteria. 1929. biol. Hermann and Cie. 1925. 185 pp 120. and K. SALKIND. . 159 du systemo nerveux .. Paris. 731 and S. G. LYDIA. genetischo Spektrum der Nukleinsaurespaltung. 10. 1934a. 236. L'excitation mitogenotique du systeme Tierveux pendant Ann. 1932 b. L'excitation mitogenetique Strahlung der optischen Pfliigers Archiv 231.. Springfield. Periodische Anderungen der Permeabilitat beim befruchteten Seeigelei. 127 . ttber den Ursprung der mitogenetischen Strahlen. rays. Das mito. M. 701 1935. 89 ( . Roux' Archiv 105. .Die mitogcnetische Strahhmg des Carcinoma. 152. f. Z. 1153 1934b. Roux' Archiv 124. 151. 84 180 173 Biochem.. 181. 81. . Die mitogenetische Spektralanalyse Spektren des Carcinoms und des Cornealepithels. 1934. C. . 362 65.. Biochem.. GURWITSCH. de Physiol. H. 134 HERLANT. 1931. Compt. A. 255 Bahn bei 157.. Quantenausbeuten Lichtzahlern. Das Cornealepithcl ats Detektor und Sender mitogenetischer Strahlung. Physicochem. AUTHOR INDEX Page and L. 1931. 55 . C. 109 1934. 38 and N. 142 . 124 38 . Z. 246. ANNA. 152. GURWITSCH.. 470 1929. 211. : . II. 246. 10. 1932.200 GURWITSCH. 111 1932. 180. 178 Krebsforschung 29.mitogenetische Strahlung". 49. KREUCHEN. T. L'analyse mitogenetique spectrale. IV of Exposes 40^ de Physiologie. 89 Acta Brevia Neerland. Z. Die Fortleitung des mitogenetischen Effekts inLosungen und die Beziehungen zwischen Fermentatigkeit und Strahlung. ALEANDER. . HABERLANDT.. Z. \ Vol. 39 pp GURWITSCH. Das mitogenetische Verhalten des Bluts Carcinomatoser. biol. Physico-chemical test for mitogenetic GURWITSCH) GURWITSCH) rays. 1928. Biochem. et Physicochem. Die Fortpflanzung des mitogenetischen Erregungszustandes in den Zwiebelwurzeln. 20 92 HARVEY see TAYLOR. biol. 15 HENRICI. 1934. W. Cytagenin HEINEMANN. 157 Ann. 357 . und mitogenetische Strahlung des 190 ( Bluts. adaquater Erregung. Klinische Wochenschr. 425 Die Biochem. Biol. 5. de Physiol. Die mitogenetische -. Die mitogenetische Sjpektralanalyse IV.. 151 . Physico-chemical test for mitogenetic Nature 134. M. 1166 : 158 II. 1928. . soc. 122.

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

1180 Journ. Journ. 86 . 188 photoelectric counter tube. . 549 und J. 159 LASNITZKI.. 330 . 802 . d. U. Bot. D. Journ. AUTHOR INDEX Page KENDALL.. E. 22. v<g. Naturwissenschaften 21. Parabiose der Nerven Folge mitogenetischer Bestrahlung.. LUBLIN. W. Radiations emises par path. 413 95. method. Ann. Origin of a tetraploid shoot from 169 the region of a tumor on tomato. 19. 113. deutsch. 1934.. Ber. Compt. W. J. Krebsgenotischen Induktion von Warmbluterzellen. 178 178 1927 b. f. Compt. de Physiol. 14. Ges. forschung 34.. and Exp. 185 J.S. physical 73 MACHT. MACHT. 1928. L. 186. see also * LATMANISOWA. General Physiol. 1927 c. KUREPINA see G. f. and M. 1930. 905 171.. 232 Loos. 367 1933. Untersuchungen iiber mitogeiietisehe Strahlen. K. 1932. Nekrobiotische Strahlen. LANSCHINA see KARPASS. 10. D. 91 LUCAS. Action a distance du Bacterium tumefaciens sur loppement de 1'oeuf d'oursin. Influence of 83 visible and ultraviolet rays upon the . Recherches sur le les radiations . FRANK. Zur Frage der mitoZ.. and E. . 67.. 1927a. 1934. 22. rend. 1932a.. Sur la parabiose mitoge'n&ique et de Physicochem. Nekrobiotische Strahlen I. 547 . I. rend. 1923/24. Bull. Search for mitogenetic radiation electric by means of the photo41. 843 bios. W. M \GROU.. 243 Protoplasma 92. KLEE-RAWIDOWICZ. bot. 1924. Physik 94.202 KOSTOFF. Am. . Physikalische und biologische Untersuchungen uber mitogenetische Strahlung. Ther. LORENZ.. 1311 . 1933. . BATEMAN. 184. Protoplasma 20.. mitoge*n6tiques. Z.. 99 99 99 1932 b. MAGROU. A. 1935.. agr. 95. H. wissensch.. 1934. Bacterium tumefaciens. 611 57 LIOSNER see BLACKER.. Rev. le deve- . A phyto-pharmacological study of menstrual toxin. Jahrb. and D. 184. W. . 144 KREUCHEN. 190 HAUSER. desNerven. 57. Pfliigors Archiv 231. 28. Science 76. 265 als 159 159 1933. . see LUBIN S. et ent. 1932. stability of living matter. The fractionation of Chem. Health Reports 48. 112. Rayons mitoge*netiques et gen6se des tumours.. 50. Investigation of mitogenetic radiation by means of a 91. Die mitogenetische Sekundarstrahlung 110. G. . and J. 1931. d'Histologie appliqu6 4. Biol. W. 178 . H. B. f. Messung geringer Lichtintensitaten mit Hilfe 92 von Zahlrohren. 141 du nerf. 518 LEPESCHKJN. . 244 . 253 . of Pharm. Botanik 72. 17.

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

49 153 1934 b. Comm all Sed Scient. PROTTI. No 99 RAHN. see Physik. Journ. 1932. Osped. Zentr. 1929. 249. AUTHOR INDEX Page and K. also: Eifetti citolitici da radiazioni biologiche (citofotolisi). 1931. 16.204 POLANO. DESSAUER . S.TEWSKY. The fermentometer. Physiology of Bacteria. ZOGLINA. 131 different . Wachstum der Bakt. SALKIND and and I. Ober Beeinflussung des mitogenetischeri Effekts durch sichtbares Licht. 16. Wochenschrift 1385 95 detaillierte PONOMAREWA. f.Zehn Jahre Forschung auf dem . IT. Krebs91 forschung 35. das . . 199 Philadelphia. 211. 387 . f. 1930. Zum Problem der mitogenetischen Strahlung. 1934a. Blakiston.) 35. L'emoinnesto intramuscolare.. Biol. BARNES. 92 RAS WOLFF.. OTTO. Die mitogenetische Strahlung des Bluts und der Gewebe von Wirbellosen. ttber einen empfindlichen Lichtzahler. 418 134 125 1929. . 255 see romyces cerevisiae. . 282 . Dio mitogenetischen Spektra der Oxydationsreaktionen. ZOGLINA. 352 149 see also BRAUNSTEIN and SALKIND. 217. Gen. 71. Biochem. 32. 91 1932. 579 125 see also BARNES and FERGUSON and TUTHILL. 36 Biochem. An experimental comparison of criteria of death in yeast. see PROKOFIEWA KLENITZKY. II fenomeno (4). 424 POTOZKY. 1933. Impressioni fotografiche di radiazioni ematiche ottenuti attraverso Civile di Venezia . DIETL. Sciences Biol... N. . 121 . 90 151 1931. 1931. 427 pp. Physiol. Das Verhalten ciniger Tumoren gegeniiber dem SacchaArch.. Z. A. Bact. 140 pp. 1928. physikalisch-medizinischen Grenzgebiet". Roux' Archiv 114. . 50. and M. Centr. 712 108 . . Mitogenetische Strahlung des Bluts. (Russ. Die Einwirkung der Hautabsonderung der Menstruierenden auf die Hefegarung. and I. La 182 Ricerca Scientifica 2. 1930. 37 Biochem. 1934. in FR. G. Das glykolytischc Spektrum. 1 109 . Biochem. B. 1932. della emoradiazione applicato alia dinica. Milano. J. t)ber don EinfluB der Stoffwechselprodukte auf Bakterien. 178 146 1. t)ber mitogenetische Sekundarstrahlung aus abgeschnittenen Zwiebelwurzeln. Miinchencr Mod. 18. il quarzo. Z. Z. bei O. Journ. Abt. Radiobiologia 1 . 239. 1924. Z. 1906. 1930. Leipzig 1931 Z.. RA. Z. ZOGLINA. Ulrico Hoepli.

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

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

see drosophila dispersion 2. 161. rotating 103. 160 of radiating power 110 eye radiation. 160 cotyledon radiation 141 crown gall origin 171. 187 heliotropifctm K eggs as detectors 81. 150 muscle radiation 29. radiation 145 cholesterol and cancer 182. 91 onion roots 58 - --- [u in in - muscle 154 nerve 112. 100 184 flocculation of colloids discontinuous irradiation 103 disks. 19 dissolution of salt as source of frog eggs as detector 81.210 SUBJECT INDEX drosophila eggs as detectors 81. 65 spectrum 35. - 73. 61 f . 50 chicken embryo. 178 - and polyploidy 169 cytagenin 152. 150 roots 111. 89.sources of in bio. field 6. 97. 143 larvae. 76. 145 coagulation omitting radiation 89. 125. J31 secondary 117 see also spectrum 37 filters for light 21 detectors.radiation 69.of nerves 111. see also errors false mitogenetic depression 70 fermentation affected by radiation 85 causing radiation 124. 100 Enchelys radiation 137 radiation 38* error. 76. 169 chain reactions. 77. radiation intensity 146 yeast buds 71 yeast volume 74 . expcri mental. 112 by radiation fruit flies. 154 tadpole legs 109 tissue cultures 83 ---- in yeast cells 110 -. 128 radiation 169 embryo enzyme radiation 143. spectrum 147 erythema 48 exhaustion by irradiation 43. 142 as senders 142 183 affected chromosome number by electric. 80 - - . general 190 colony si/e method 75 colloid coagulation as detector 89. 180 cytophotolysis 182 -. failure of 93 methods finger radiation 97. 185 depression of mitogenesis 115 ~. with bacteria 78 & 100 conduction of radiation _ J 1 1 Geigcr counter 34. see cornea and radiation human r day light affecting mitogenetic failure in mitogenetic - experiments 93 rays 108 death through irradiation 49. 113 over-exposure J15.false 70 - . 105. 142 larvae see tadpoles - radiation 50 dog blood radiation 148. definition 47 - and radiation 43 - chemical radiation 33. cornea as detector 84 --. radiating 145.

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

156 exhaustion 111. 96 82. see intensity properties 59.212 SUBJECT INDEX mitogen?tic 110 field methods: necrobiotic radiation 99 onion root 54 peroxide decomposition 89. spectrum 61 from bacteria 87. 115. . 161 effect. 44 onion root. 101 photoelectric counter 90 effect. 125. 99 tissue cultures 81 radiation. 70. 96. yeast 64. 148 bone marrow 64 - cancer 178 chemical reactions 50. spreading of theory 128 photography 90. protozoa 133 sarcoma 64 tissue cultures 81 tissues 146 detector mechanism 129 . 87 physical nature of. . 142 yeast metabolism 84 mitogenctic depression 70. rays 99 blood 65. see mitogenctic effect yeast buds 66. 147 - . 178 N necrobiotic. 147 neutralisation 50 nuclease action. secondary radiation onion roots 55 oxidation 29. cell concentration 134 cause of 122 muscle. 59. radiation 157 radiation 154 spectra 156 neutralisation. 87 cornea 146 eggs 142 nephelometer measurements 74 nerve conduction 112. 59. conducting radiation 154 radiation 65. 131. discovery 55. 61 mitotin 140. 160 . 59 intensity. secondary radiation 109. spectrum 38 nuclei in onion roots 55. analysis J 19 and . 77. 119 stimulation. 129 nucleic acid. 68 yeast growth 72 range 49. larvae 143 from 50 molds menstrual blood 86. 80 proteolysis 52 of 43. enzymes 37. J77. 119 sender mechanism 139 . 113. 72. 52 embryos 143 infusoria 133 optic. 51. nuclear stages 129 method 63 radiation 54 . 103 . 115 - monochromator 19 morphologic changes by radiation 86. 161. 64 plants 138 polarized light 45. 160 metabolism 156 . wave length 49. 59. discovery 55. radiation . 140 spectra 140 opalina radiation 133 . 61 over-exposure 43. 142 spectrum 40 muscle 65. 110. formula of 64 .

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

tumors caused by radiation 171 of plants 178 see also cancer . 161 sarcoma. atomic 17 molecular 17 . 114. . and radiation. 25 . definition 43 . absorption 23. and photographic plate blood rad- as senders 83 septicemia decreasing iation 152 solar radiation limits radiation 146. 39 muscle 61. 40 cornea 147 fermentation 37 frog muscle 35. see intensity. 147 -. 37 saliva affecting yeasts 96. and thres- tissue cultures as detectors 81 hold. 164 by infra-red 101 storage of radiation 126 sunflower.sucrase 37. of tungsten 32 of amylase 37. 175 rate of travel in nerves 112 in r0 ots 111 for production of effect 104. 112 spectrum of mercury arc 20 maltase 37. definition 178 . - of cells 108 of solutions 43 . 124 T tadpoles affected by radiation 167 radiation of different parts 145 wound healing with 175 . sea urchin eggs as detectors 81 aa senders 142 - solar 22 retarded 86. blood 150 affecting organisms 48 bunsen burner flame 40 cancer tissue 179 -- photographic plates 24 causing cancer 183 chemical reactions 37.onion - roots ]40 oxidation 36. . 61 glycolysis 37 * from chemical reactions 29. . 98. effect on intensity measurements 123. 123. 39 turbidity measurements 74 U ultraviolet rays.nerves 156 neutralisation 40 nuclease 39 -- . mitogenetic Ureffekt 119 . mitogenetic 35 thermal. see Helianthus larvae changed by irradiation symbiosis and radiation 170 secondary depression 117 radiation. see mitogenetic effect. sedum leaves 141 123 sensitivity. 167 tonsillitis decreasing blood radiation 22 152 spectrometers 19 spectrum. phosphate cleavage 38 * proteolysis 38 39 a radiation 64 -. 33 from organisms.214 reversal SUBJECT INDEX of effect with photo- graphic rhythm 107 plates 116 in cell division 119 rhythmic interruption of radiation rotating disks 103. thermocouples 23 threshold time as measure of intensity 44.

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

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