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

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

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


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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

of course. states provided that. e. sufficient to shift the energy is greater than (Eoo electron beyond the outermost orbit. the electron is in the orbit corresponding to the lower of these states. If the quantum is larger than Km En. 1216 A.unless its Kn). i. at this moment. Changes between other light states result in the radiation of visible and still others produce radiation in the far infra red.4 x 10.99796 x 10 10 X 6. it may be absorbed and the surplus. .547 x 1C" 27 __ ~ ~~~ 161. An electron so ejected from an atom ionized. and the of its atom itself is said to be As has been stated before. the atom is said to be in an excited state. In this event. is readily calculated from the data this table. if its energy happens to equal the energy difference between any two quantum .16 CHAPTER I The wave length emitted by the hydrogen atom for the energy change in Ex E for example. as _ ~~ This wave length 2. it will not be absorbed -?. The life of an atom in an excited state is of the order of 10~ 7 to 10~ 9 seconds. it is 178 500 A. 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). in the far ultraviolet.. E When it has been raised to a state of higher energy E n by virtue of the absorption of energy. or is . 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. for the shift from the 6th to the 7th orbit. g.ia 1216 xK)" 8 cm. then it may be absorbed.e. After this length of time the atom reverts to its normal state with the resulting emission of radiant energy. Tf its energy is not one of these discrete values. It will be. an electron spends most . existence in the normal state. is called a photoelectron. or hv Ku). hr (K<x> electron away from the atom. 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.

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

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

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

a Hilger quartz double moiioehromator. . the intensity of the unwanted be reduced to practically ZATO. \vave lengths list may is of such filters By so doing. In general. some intensity of radiation of shorter 1 wave length than that desired is transmitted by the instrument. The wave lengths given in the table are those at which the filter ceases 1 ) In particular. 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. relative intensities of the lines in the ultraviolet spectrum of the quartz mercury arc as transmitted by such a monochromator. While monochromators are designed to give high spectral purity of the isolated light. some degree of impurity seems unavoidable. 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). 1 ) Table 4.20 calibrated in CHAPTER wave lengths. 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. Relative Intensities of II g Spectral Lines Uratings are also used to produce spectra of violet light but since their application is will visible 1 and ultra- more strictly confined to special work in spectroseopy. a discussion of their characteristics be omitted.

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

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

which is proportional to the temperature difference of the two junctions. circuit two dissimilar metals or alloys (sec U>a).97 of This. Table 6.97 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. shows plainly why in biological radiations. Figure 16. (J the junctions are maintained at different temperatures. counter. the photocell graphic plate. 10 b in . the photoelectric (1) and the photoelectric. Thermocouples left: a two-metal thermoelement. together with the absorption spectrum of oxygen. a current will flow in the circuit. no experiments mm are carried below 1900 A. The Thermocouple: The composed of thermocouple consists of a fig. The Absorption of Ultraviolet (16. Modern high-sensitivity If thermopiles are frequently built as diagrammed in fig. the detectors of ultraviolet light are: the thermocouple.PHYSICS OF RADIATION 23 distilled water. Table 6 gives the absorption of ultraviolet by 10. right: a modern highsensitivity thermocouple.

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

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

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

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

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

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

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

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

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

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

only the rays between 2000 and ) Radiation passed 2700 A were measured. and by pyrogallic acid in air. In is all larger this. BABTH (1934) could also prove existence* of ultra-violet emission AUDTJBEKT and VAN DooKMAL (1933) tile known of from proteolysis which to give an immeasurably small heat of reaction. Table 10.34 Jn this way. 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. therefore it must be of a wave length shorter than 3500 A. the emanation is greatly increased in the presence of diffuse day light. vessels. trically also measured photo-electhe emission of ultra-violet light by several inorganic oxidations. by the oxidation of alcohol with chromic acid. 1 . not from gla?ss containers. 25 experiments but one. verified by GERLACH (1933) who showed that this radiation appears only from quart/. . This physical proof of light from chemical reactions has been ultra-violet radiation ficiently sensitive instrument. Recently. 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. It was also observed at this time that in some of the reactions.

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

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

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

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

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

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

arc 1

commonly used by

the Russian


.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.



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




of all these



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

frequently combine

of these lines, in addition to

some others



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


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



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

The wave length



this procedure,


they found the spectrum of neutralization

of acid


alkali to consist of three lines

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


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


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


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


arc limited, however, by our



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)


good summary


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).


intensity of ultraviolet radiation

not proportional to the total energy liberated.

from a chemical reaction This was to be



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



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


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 --:<


10 2:i



minimal amount
then be

of radiant energy



g of sucrose would

H.I > 10 23



9.8 < 10~ 12




10 11 ergs

-- 0.042 , 10* cal



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


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


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


with liberation of


be expected to take place spontaneously.

energy, and the process might, therefore, Modern chemistry




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

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


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


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


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
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

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

than that absorbed, because


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




would be an excellent means to measure the energy

of activation.

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

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

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

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

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

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

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

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

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

5 All other enzymic processes which have been tested so far have yielded positive growth stimulation. containing the data obtained by KARPASS and LA. Jnerease in the development of yeast cultures after exposure to the emanation from proteolytic processes Proteolytic profess Pereentual increase of exposed culture over control Egg white with pepsin. 31. - lfi. the irradiated culture grows Table 15.1 %. only one was negative. 15.4% . 10. 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.7%.6 H 2 O ('ells per ec. 37. more rapidly.52 OH AFTER III absorbs the radiation (Table 15). There is the usual lag period of 2 hours. 36. . . Protco- by enzymes yields an ultraviolet emanation which greatly stimulates the growth of yeast as seen in Table 16. Table 16. it is only logical to assume that all living somewhat by the consideration that these organisms radiate.5% 20.6 % %. ex- Development of a culture of Bacterium eoli after posure to emanations from the reaction C 2 4 H 2 + KMnO 4 =. This statement must be modified ultraviolet rays are . Of 12 experiments. of culture exposed through glass .0 egg yolk with pancrcatin fibrin with gastric juiee 30. egg white with pancreatm egg yolk with pepsin .4 %. 25. Since all organisms display processes liberating energy. is also more complicated biochemical processes. 20.K 2 ('O 3 + 2 MnO + 9 (<O 2 H. but after the bacteria once start to grow.NSCHINA (1029).8%.1%.

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

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

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

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

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

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

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

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

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

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

however. 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. When steel. Almost always. it was considered a proof of a raitogenetic effect. while the leaves grew through the hole in the cork. long and absolutely straight. In. this percentage was higher in the irradiated culture than in unexposed controls. usually not over 30 minutes. The number of examples given is not large enough to draw many more conclusions. After this. he spread yeast over the surface of solidified nutrient agar containing glucose. However. When. the symmetrical distribution was disturbed. The original method has since been changed in some details by BAIION (1930) and GUBWITSCH (1932) (see p. sender roots -were taken as controls. 66). The measure was the percentage of yeast cells showing buds.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 63 room at 20 C. and it will be. The yeast was then incubated for from 1 to 2 hours to permit the radiation effect to develop. The exposed roots turn downwards. and theii exposed it to the radiating source for definite short periods. b) The yeast bud method BAIION (1920) suggested that the rate of bud formation of yeast could be used as indicator of mitogenetie radiation. . whether all mitotic The stages were included or only the more conspicuous ones. and they showed a uniform and symmetrical distribution of mitoses. his first experiments. dried and stained. The object of PAIR'S investigation was the establishment of a good method for studying mitogenetie rays with onion roots.5 cm. the yeast was spread on glass slides. When the roots were 1 2. allowed it to grow for from 9 to 15 hours at room temperature. The paper verifies GURWITHOII'S principal experiment. but the microscopic analysis showed no increase in mitosis at the exposed side. the exposed root grew more rapidly than the other roots of the same onion. the starting point for all future work with this type of detector. The onions were never placed in water. the sender root was substituted by a needle of stainless the exposed root also turned downwards. 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 roots developed in the* air through the muslin in two to five days.

e. and the control. he obtained positive results by using a very dilute CuSO 4 solution as oxygen catalyst. he succeeded by placing the acidified pulp atmosphere. Moreover. Table 19 gives some of his experiments. is 9.5% over the control. expressed in pereeiits of the control value.<>4 CHAPTER 19. or excited muscle to radiate strongly while the resting. and this is an increase of 37. sender and detector were J separated by a quartz plate. The numbers indicate the percentages of yeast cells published wit. in air When . Thus. IV Table Effect of Yeast. since the addition of lactic acid alone would not produce radiation. He observed the working. Tt is now customary to record. The induction effect is 37. chemically the pulp of resting muscle to that of working muscle. results as "induction' effect.li buds. i. as increase in buds of the exposed yeast over the control. the increase when the exposed yeast shows 33% buds. 24%. __ 100 (exposed control control) SIEUEKT (1928b) used this method for a number of interesting studies in physiology.. 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). quiet muscle did not do He attempted to produce radiation by changing this (Table 20). in an oxygen Finally. In all his experiments. of Sarcoma.5%.

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

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

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

N type is more sensitive because the old cells act as "amplifiers" (see p. The "mitogeiictic effect" is much greater than in the other method because there are no old. these are probably the largest mitogeiietie effects ever recorded. with very weak radiations. this type of detector is so different from the BAKON type that it may react differently in certain experiments. inactive cells to "dilute'' the counts.68 Table 21. both types are good. sterile raisin agar l ) Raisin extract: 1 pound of chopped. CHAPTER IV Kffeut of Irradiating Yeast Surface Cultures after Incubation for Different Times : Length of Exposure 30 minutes Incubation after Exposure: 30 minutes .5 hrs: 2 hrs [2. the extract 1 liter. On the other hand. A 24 hours old culture in raisin extract 1 ) is flooded over a solidified. 127)..5 hrs.Age of Parent ( 1 I Age of culture when exposed hrs 10. This should make the counting easier. 5 g. with 1 liter of water in steam for 45 minutes. of earliest rejuvenation when exposed. KH PO4 and added. 1 Culture ! hr 1. Probably. 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. As long as they are used merely as detectors to prove the existence of radiations. 6 days . and the cells are far enough apart not to influence each other. . and a limitation in size is not necessary.. pressed made up to 5g. Method by The yeast is TUTIIILL and KAHN (designed for Burgundy yeast): kept throughout the experiment at 30 C. the BARO. In fact. 2 or seeded raisins is is heated off.5.5 hrs Percentage of Buds 24 hours . It is quite permissable to count all buds because the percentage at the beginning is very low.

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

210) and especially of SALKTND (1933) where the percentage of buds decreases while the total cell count increases under the stimulation of mitogenetic radiation. a change would become noticeable if only one certain stage of the cycle should be accelerated. If we take flection. Many observations suggest this. p. they have kept their de12 hours at 25 C while BARON used room tector cultures for 9 . If niitogenotic radiation should speed up the very first stage so much that the "tiny" buds never appeared. It is rather probable that only a certain stage of the cell cycle is affected by mite-genetic rays. the percentage of buds remains 74 75%. This percentage depends only upon the variety of yeast used. This would offer a good explanation for the "false mitogenetic depression" of (rUBWiTScn (1932.8%. However. A change of the growth rate would not affect the bud percentage at all. 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/ . and our present conceptions of the mechanism of cell division do not contradict it. and this lower level would continue as long as radiation accelerated the one particular stage.70 necessarily be CHAPTER TV more cells of the young stages than of the old. the number of buds at the period would drop from 26" out of 35 to 18 out of 27.4 to 66. The reliability of the BAKIXN method has been doubted by NAKAI DZUMI and HCHKEIBKK (1931) who claimed to have followed BARON'S method explicitly. whether the time unit is 30 to 00 minutes (at cellar temperatures) or 10 minutes (under optimal conditions). However. or from 74. Whether it grows rapidly or slowly.

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

the buds of yeast ascertains cell divisions directly while the mitoses in cells. Table.5 ee. The total number of yeast cells (counting each smallest bud as an individual) is determined with a hemacytometer at the start. and it is made discontinuous by moving some object between blood and yeast at intervals of 2 seconds (see p. The number of cells is the best measure of the growth rate. While they are identical in principle. etc. and may be influenced by secondary effects. The exposure lasts 5 minutes. and again after three hours' incubation. are indirect measurements of growth. the respiration. or by hemacytometer count. 116). the yeast bud method indicates a decrease in the growth rate while the total cell count shows an increase (e. 1 . better than any other of the* methods described in this book. 103). they shall bo treated separately in this chapter on Methods. because it onion roots. 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. to the radiation of blood diluted \\itli MgS()4 solution to prevent coagulation. of the exposed yeast suspension is then mixed with an equal amount of beer wort and incubated for three hours at 24 C. an equal amount of a 4% . The hemacytometer has been used by the Russian workers as well as by HEINKMANN (1932) in his studies on blood radiation. An counting by the addition of H a SO4 example of the results and of the method of calculation is given in Table 22.72 c) Detection CHAPTER IV cell by increase in number investigator not familiar with yeasts. There is a difference so fundamental that in a number of experiments. The plate count method has rarely been used. 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. they differ in the method of measuring since yeasts arc so much larger than bacteria. 0. g. This method is used with yeasts as well as with bacteria. cells. It is the same as that used for counting bacteria except that more preferable media are beer wort agar or raisin agar. All cultures are preserved for The control is counted twice. (1) Yeasts: The number of yeast cells in a liquid may be ascertained by plate all the volume of count. For this reason. 34 p. by measuring by comparing the turbidity by means of a nephelometer. both in the control and in the exposed culture. the preceding pages were written.

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

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

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

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

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

2 4. dilute exposure 1 10000 with dilute broth. this has been done by SKWKLITZOWA (Table 24) and Arz.4 and 4. The effect may not become apparent iff he cell concentration is too high (over JOOOOO per cc.8 3. see Table 41 p. cultures 4K hours old still older always responded. diameter. 133. and plate at least every 2 hours for 6 to <S hours.78 CHAPTER The error of the IV method is mentioned in 1934. The best medium was 1 part standard broth plus 9 parts water.9% of the total count. and no rules can be given.e. incubate. it must be considered that the number 183 has no real biological where d is the number t. 1 after the time 1 significance while the other indicates that during the four hours. the increase by irradiation amounted to 25% series are given. . at the start of incubation. 136). By computing the growth rates.1 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(>. 1)1. or in this 1 cc. after a quart /-covered glass dish. is only 27. bacterium and found that it depended primarily upon the age of the culture.4. while the effect in tho same culture. the best results were obtained with 15 to 30 minutes of irradiation 3. However. of an old culture in dilute broth (either in a quartz dish of 4 5 em. The time of exposure depends upon the intenWith a 4 hours old agar surTable 20).3. The growth rate is usually substituted the average by the generation time.1 7.3. The errors are between 3. (see Computation of the Induction Effect: The in- duction effect in these bacterial cultures cau be computed in the same manner as explained on p. we have a really reliable measure. of cells at the beginning. i. this culture had a growth rate higher than the average of 27% the two controls.6. the : ^ ^ i and 28%. computed from the generation time. sity of the source. 64. four detailed counts being 73. or through the bottom.5 and 171. Tho simplest procedure is to irradiate 1 ee. Method: FEUGUSON and RAHN a good mitogenetic effect with (1933) studied the best conditions for cult. 24 hours old cultures never reacted. : from abo\c). face culture (37(-) as sender. while in truth it is exponential. the time required for cell to double. The induction effect calculated from the numbers directly is in one cast (]5 minutes exposure) 183. As a matter of fact. This computation implies that multiplication of bacteria is arithmetical. and tho transmission of ultraviolet by the medium.

8 f-27 This procedure has also been used in Table. It permits no comparison. This means that not all cells had divided in this time. minutes. after diluting the irradiated cultures 10 000 with broth) : Induction Effect 37 168 I83| 111 Generation Times. for the time interval from 47. g.EIIJER does not give the actual numbers of from which he calculated the generation times.5 I Generation Times Induction Effect 45. With Nadxonia. 1 . it seems from his curves that they were computed from less than 100 cells. various lengths of time by an agar surface culture of the same bacterium (The numbers are cells 1 per cc. 64) has no meaning. the especially many investigators. This makes the error very large and a comparison of the growth rates must necessarily result in enormous percentual differences.5 14 ! 41. However. the generation time of yeast for the first 2 hours when the mitogenetic effect is strongest is mostly more than 2 hours. that error of the method.METHODS OF OBSERVING BIOLOGICAL RADIATIONS . and occasion- ally more. 2 --6 hours 52. It could be used with yeasts as well. The "Induction definite Kffect" as usually calculated (p. e. SCHKKIBEII found the variations tiaee/Miromycefi cMipxoirfcuti commonly in duplicate plates of to read) -40%. Thus. In the tables given by iScuitEJBKR (1933). therefore. 24. the error becomes very large if the increase is small. in Table 22. . the deviation of duplicates went as high as 237%. with the probable It is very unfortunate. there is little hope of detecting mitogenetic effects. With such a large error.5 40. record. 79 3 days old culture of Bacterium cult irradiated for Table 26.6 . the Russian scientists. and in one case even 103%. cells Though HCHK.

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

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

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

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

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

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

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

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

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

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

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

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

Though some early workers indicated higher sensitivities. more than 2000 quanta are required to eject one electron. . Most convincing is the experiment that a counter gave a definite increase when exposed to normal blood. 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. In a recent paper. 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. In all cases except the first. are a step in that direction. it seems from later work that this value has never been sur(see of the surfaces used passed if. 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. In his latest paper. KREUCHEN 4 6 from (1935) obtained yields of from 1C to 10 quanta per electron hydrogen-activated zinc and cadmium surfaces. and a number of experiments with water blanks or non-radiant organic materials will be necessary to produce evidence which physically irreproachable. and it seems well to view with caution the positive results claimed for the physical detection experiments so far carried out. in fact. but showed 110 increase when this was KCN. A more careful description of the method of exposure. it ever was reached. In none of the physical measurements of mitogenetic rays is it absolutely certain that these errors have been excluded. is The experiments of SIEBBBT and SEFFERT (1934) who obtained increased counts with several hundred normal blood samples. but no increase with blood from carcinoma patients. 29) by the various investigators.92 Table 30a.

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

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

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

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

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

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

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

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

E. The only one known to the authors is an by their was necessary The temperature of own respiration. the peas rose 0. The only case of near infra-red emanation known to the authors is. The irradiated eggs showed. The cell is alive intensity of this radiation or dead. The authors believe therefore that the reduced fertilization is caused by a photochemical effect. It originates from the radioactive fraction of potassium.METHODS OF OBSERVING BIOLOGICAL RADIATIONS 101 In 25 experiments. Equally rare are observations of an effect of infra-red rays upon living organisms. and is of importance to life An processes. is the same whether the .5 increase of 5. The decrease varied between 18. and really not characteristic of the living cell. namely the bet a -radiation of potassium.06 C.1%. the eggs were fertilized by the usual method. The temperature difference between control and exposed eggs was not more than 0. but of its potassium content. and was not visible. After 15 to 45 minutes exposure. 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 effect passed through glass. it must therefore be of an infra-red nature. BETA-RADIATION entirely different type of radiation should be mentioned only in passing. the suspensions were brought into light.5% and 87. obtained from a Mazda lamp by means of a monochromator. however. They exposed the unfertilized eggs of 2 sea urchin species and of one worm to infrared rays of 8000 to 12 000 A. 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.5 to bring the rate of peroxide decomposition to that produced by peas. but an artificial experiment by NELSON and BIIOOKS (1933). Though it has been proved experimentally. suspension which stood under the quartz vessel uniformly darkened sooner than the other. D. 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.

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

The present chapter initogenetic rays describes some to which are not common all peculiar properties of types of radiations. It travels through space rectilincarly it can be reflected (pp. 39). shows refraction and dispersion (p. the duration of each. disks contained one or several windows of varying width. 59) it . 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. The width of these. much threshold value in mutual yeast irradiation by means of disks The rotating at approximately 3000 revolutions per minute.e. 35). their number. From this angle and from the number of revolutions. initogenetic radiation. It indicates that the minimal . 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 frequency of exposures. A. Table 31 gives the results obtained. 59 and 80) it can be absorbed (p. . of a rotating disk which contains one or several openings or windows. and the total time of actual irradiation. INTERMITTENT IRRADIATION of Very early in the history discovered that the effect could for this purpose is the insertion.. and the rate of rotation of the disk allow one to vary the three items concerned in intermittent intermittently instead of continuously. processes. i. The preceding chapter has revealed that the etic radiation is .

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

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

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

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

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

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

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

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

is conducted to another part and is emitted there. onion root. This was sufficiently to 3 intense to permit the reduction of the window for primary radiation and that for secondary radiation to 1.00140 seconds. This is good agreement with physiological measurements on the rate in of conduction of nerve impulses. side view showing position of primary sender S. LATMANISOWA (1932) has measured the rate of conduction of secondary radiation in the sciatic nerve of the frog. and detector D. CHAPTER V This signifies that a certain time (0. Further experiments showed . The central angle for positive effects r. At right. radiation was observed from the same side of the root which had been exposed to primary radiation.. which means an average increase of 15. of root.5 cm. such as local reactions at the points of absorption and emission.00083 seconds. 2.0022 seconds) must pass before primary radiation falling upon one part of the root. The total time corresponding to The difference. and this is This corresponds. p. at 3000 Figure 41. of root is 0. The accuracy of the in method was thus greatly increased. to 0. to between 40 was hereby increased and 70. the average angle of 55 is 0.5 cm. These values refer to a condition over 2.00306 seconds. except that the 2200 A area of the copper arc was used as primary source. the total time required for transmission through 5 cm.112 results. Then. and the rate of conduction the nerve was found to be 303 meters per second.m. was required for processes other than conduction. the time required for the secondary radiation to pass the additional The rate of conduction is therefore about 30 meters per second. Allowing the same time for transmission through the other 2. the distance was increased to 5 cm. Recently. Kotating disk for measur4 ing the rate of travel of radiation in secondary onion roots. 0. In the experiments with secondary radiation of onion roots.5 cm. The method was exactly the same.00166 seconds.

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

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

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

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

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

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

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

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

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

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

15% more than the control.00042 The probability of being hit in one minute is 0.0252. All hope for proportionality must be given up in this case. uniformly dispersed radiation before each of the yeast cells is likely to be hit by one quantum of ultraviolet light. Since we find the strongest effect under this . P = 0. does a biological measurement of intensity seem at all possible. This may be explained by the recent discovery of WOLFF and HAS (p. and the 40-minute exposure a 30% increase. but senders appears to be about 100 to 1000 quanta per cm 2 per second. Only with a medium which does not produce secondary radiation. tJ. the 10-minute exposure should have produced an increase of approximately 7%. It will require 40 minutes of continuous. 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 entire detector plate begins to emit radiation as soon as it is exposed to a sender. however. If there were proportionality. 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. The pro6x7 is hit in one second. Neither of these other times showed any great effect. either directly or through quartz.00000042 x 1000 = 0. The exposure 20 minutes produced a strong effect.ANALYSIS OF THE MITOGENETIC EFFECT 123 The sender was a 6 hours old yeast of surface culture. 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.

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

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

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

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

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

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

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

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

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

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

This observation had been very puzzling to biologists since tho opposite should be expected. do not start growing at their maximal growth rate. The proof is not complete. The rate of growth of old cultures can be increased for several successive transfers.134 the next experiment. He inoculated increasing amounts of yeast into flasks of sugar -peptone solution. 120). an old culture to a fresh medium. and single-cell cultures frequently die. until it could be explained as a simple stimulating effect by mutual irradiation of the cells.422 0. It is well known that young old bacterial cultures require a longer time to start growing than ones.914 mm. the average diameter as 6. Without accurate statements concerning the number of cells in the sender and in the detector. The volume of one 8 yeast cell is taken as 1 18 ^ .036 0.. Bacteria* and yeasts. PENFOLD (1914) and others that bacteria after being transferred from an old (see p. It has been observed by RAHN (1907). without any irradiation. the stronger must be the effect. with yeast. 612 1224 2436 cell 2 4 8 hrs. were present. The times required to reach the maximal growthrate were: The I) Table with an inoculum of 20 949 750 cells per cc. absolutely correct only with spherical organisms. Lag Period Average cell distance 1 ) 2094975 209498 20950 2095 1 ) .192 0. morphologically and physiologically (2) Reaction Upon Irradiation: when transferred from This results in a very slow growthrate during the hours after transfer. The bacteriologist calls this the lag phase. so that all the decimal gradations from 2095 to 20949750 cells per cc. no definite conclusions can be drawn. The old cells undergo a rejuvenation process. J Jt is .04 ||/~~ / \ cell volume in 100 cc. the quicker the recovery. CHAPTER VII His data show a very consistent increase in the mitogenetic effect through 6 to 7 such transfers. first culture rejuvenate more rapidly if many cells have been transof fresh ferred while with a few cells in a large volume medium. best example is very likely that of HENKICT (1928. rejuvenation is slow. The closer they are together. 8 0. The average distance distance is computed from the equation = ~ cell diameter 74.4 //.086 0. These different inocula resulted in very different lag periods. however.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

the freed leg grew a little more slowly 0.1 more in one series of 33 tadpoles and The actual 1. Removal of the gill. The result of the operation as such was that in the vessels with glass bottoms. or the freeing of the foreleg from the gill cavity where resorption of the gills occurs. retarded growth.5% i tadpoles.8 less in one series of 22 than its mate in the cavity: 6.3 less in more rapidly. and therefore shaded against radiation. the legs if the bottom of resorption.168 CHAPTER VII maximum of blood radiation occurs after completed their metamorphosis. a second series with 44 tadpoles. and that transplantation of gills increased the growth rate of the corresponding foreleg. during successive stages of metamorphosis. it could be shown that a tadpole lying in a quartz -bottomed vessel will show a more rapid growth of Figure 51.5% i 1. while the other leg which served as control remained covered.4% ^ . 4.2 series with 51 tadpoles.1% 1. Intensity of radiation of the resorbed organs of frog tadpoles. gills and intestine have BLACHER and associates (VII) further showed that there was a relation between the resorbed tissue and the developing limbs. the freed legs grew and 4. however. In the quartz -bottomed vessels. in the other it 7. Finally. 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.

Positonly the radiation of the resorbed tissue. too scant to permit tin* of conclusions. are.. .. The triton Pleurodeles Wallii shows similar radiations i (BLACKER etc.4 in the one set gain by irradiation was therefore 10. 120 tomato plants were inoculated with Bact. They do not radiate until about ready to pupate. and an accelerating effect of this radiation upon the rate of development of the organs concerned.THE SIGNIFICANCE OF BIOLOGICAL RADIATIONS ETC. above the tumor in order to induce the development of new shoots. though less intense. the radiation of the resorbed parts ively established had any distinct form-giving effect. HI). Parthenogenesis by Radiation. To test this. RKITEB. Change of Chromosome Number.8 in the other set of experiments.6% and 12. The gain is 7 times that of the average error. He had succeeded in causing cell division in unfertilized frogs' eggs by only a very few percent of the eggs thus treated initiated development. they are suggestive of a new cause for polyploidy. the plants tumefaciens. but in some cases. Experiments were made also with Drosophila larvae (BLACKER etc. were cut off 3 to 5 cm. of course. and the axolotl Ambmtoma tigrinum which retains gills as well as tail throughout its life radiates still less. Seven shoots arose directly in the tumor and of these. While these data drawing 5. and OAROK GURWITSCH'S statement that there is after having verified always radiation around a wound. There is no direct proof that in these cases. is 4. 169 1.1% 1. 5 could be rooted. The maximum intensity is reached 24 hours after pupation. 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. One of them proved to bo tetraploid. After the development of the tumor. 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. very fine needle pricks. BATAILLON obtained tadpoles and even frogs by this method. V). and radiation nearly ceases after 30 hours. It has been repeatedly observed that polyploidy occurs fairly frequently in some individual branches of tomato plants infected with Bacterium tumefaciens.

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

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

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

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

BROMLTGY (1930) has proved that the tail of the salamander Anibi/tttonui tigrimim radiated strongly after being amputated. by LTOSNKR and WORONZOWA (1932 b). 1 minute of exposure to blood sufficed for a similar effect 24 hours after injury. uninjured tadpoles produced a marked effect in 5 minutes. Exposing yeast for 18 minutes to the ground tissue.utt. and a minimum at 24 hours. This agrees with the two maxima in pll of healing wounds as observed by OKUNBFK (1928). and a great WITSCII. 4 tadpoles of the frog Pdobatcs drop in intensity after 4 days. and on the fourth day. This means that at this stage. from the tip of the tail. entire 18 days of the experiment.(>% 41. that radiation one of the necessary While the argument concerning the sedum leaf wounds lias never been settled experimentally. and it stayed at this low level during the . They cut 10mm. FIUCHIMOfutic.2% 2. By this long-continued effect of radiation. VJ1 is but. . and after different times. ground it up and used the pulp to irradiate yeast plates.1% 39.9% 48. 2 minutes were required. an exposure of 15 minutes became necessary. It is not the new tissue which radiates. 12 hours after injury. removed the old tissue which produced the regeneration. Very interesting was the observation by the same authors that the blood of the tadpoles changed its radiation decidedly upon wounding of the tail. The blood of normal. In Table 50.174 besides the physical ones factors. the entire body is affected the wound upon blood and brought into play. The same double maximum could be observed with wounded BLACIIER.4% induction of the injured tissue reaches a It requires 3 hours for radiation to be established. the blood radiated less than half as strongly as that of uninjured tadpoles.4% 50. two distinct maxima are seen at 12 and 36 hours after wounding. the following "induction effects" were obtained: 1 23 5 6 12 24 hours after amputation 6. CHAPTER . but the old cells immediately under the newly-formed tissue.1% 29. The threshold time to distinct was the measure effects mitogenetie necessary produce of intensity. The radiation maximum after 1 2 days. decreases decidely and reaches a second maximum after 5 days.

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

Die mitogenetische Spektralanalyse Spektren des Carcinoms und des Cornealepithels. Die Fortpflanzung des mitogenetischen Erregungszustandes in den Zwiebelwurzeln. 185 pp 120. und mitogenetische Strahlung des 190 ( Bluts. 1932. 84 180 173 Biochem. Z. 180. soc. 1934. 1925.. Roux' Archiv 124.. 159 du systemo nerveux . . Die mitogenetische Sjpektralanalyse IV. 134 HERLANT. de Physiol. . Techn. 1934. M. Z. biol. Z. 701 1935. 1934a. G. Biol. Zentr. Paris. t)ber . Biochem. Springfield. A. 111 1932. et Physicochem. KREUCHEN. M. 152. K. et Teclairago monochromatique dc 1'oeil. Klinische Wochenschr. 246. L'excitation mitogenotique du systeme Tierveux pendant Ann. Cytagenin HEINEMANN. 20 92 HARVEY see TAYLOR. 181.200 GURWITSCH. L'analyse mitogenetique spectrale. Thomas.. Das mitogenetische Verhalten des Bluts Carcinomatoser.. ANIKIN. 178 Krebsforschung 29. II.. Die Fortleitung des mitogenetischen Effekts inLosungen und die Beziehungen zwischen Fermentatigkeit und Strahlung. H. genetischo Spektrum der Nukleinsaurespaltung. C. Physik 15. ttber den Ursprung der mitogenetischen Strahlen. 1931. 152. 1929. 362 65. Quantenausbeuten Lichtzahlern. 1929. bei HAUSSER. . Physico-chemical test for mitogenetic Nature 134. Periodische Anderungen der Permeabilitat beim befruchteten Seeigelei. Z. IV of Exposes 40^ de Physiologie. 211. Das mito. 5. 10.mitogenetische Strahlung". rays. 11. Compt. Roux' Archiv 105. 122. rend. : . Physico-chemical test for mitogenetic GURWITSCH) GURWITSCH) rays. C. . 124 38 . and K. Die mitogenetische -. 731 and S. 1928. 15 HENRICI. Z. L'excitation mitogenetique Strahlung der optischen Pfliigers Archiv 231. Physicochem. 236. 55 . 425 Die Biochem. 39 pp GURWITSCH. adaquater Erregung.Die mitogcnetische Strahhmg des Carcinoma.. LYDIA. 255 Bahn bei 157. 10. ALEANDER. . W. GURWITSCH. ANNA.. 81. 89 ( . biol. 1928. HABERLANDT. 43. 38 and N.. 151 . biol. central. 1918.. Hermann and Cie. . AUTHOR INDEX Page and L. 1153 1934b. Roux' Archiv 113. Morphologic Variation and the Rate of Growth of Bacteria. 470 1929. Z. 1931. SALKIND. 157 Ann.. 142 . 49. Das Cornealepithcl ats Detektor und Sender mitogenetischer Strahlung. f. 246. 220 1932 a. 89 Acta Brevia Neerland. de Physiol. GURWITSCH. 151. 127 . 1166 : 158 II. 1375 72. 226 MALL see JONES. T. Biochem. 1932 b. 357 . 109 1934. \ Vol.

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

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

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

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

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

see also G.



see also

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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".
19, 1


see also

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Intensitat der mito-

Biochem. Z. 227, 386 NAKAIDZUMI. SCHWARZ, 1928, Zur Theorie der mitogenctischen Strahlung.
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1929, Mitogenetische Strahlen.

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The determination of potassium
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L. B.,


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


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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
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1909, t)ber die Oxydationen





Z. Physiol.



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


WASSILIEFF, L. L., 1934,




travail cerebral sur la

radiation mitogenetique



Arch. Sciences Biol. (Russ.)



G. M.

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



Physik WHITE see HILL.


Die Entladungsformen im zylindrischen Zahlrohr. 705




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

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

Strahlen von

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

128, 257

190 132


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,




Bakt. I Orig. 128, 314


146 190

1934 a, VI.

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


37, 1

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

Nature 133, 499

single cells of non-spore

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

Boll. Soc. di

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



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


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

Ruhe gekommenen Herzens durch

die Bestrahlung des


d. ges. Physiol. 215,


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

blood radiation 05, 72. S5,



and cytagcnin 152
after injury




working Mi)

wounding 174

allelocatalysis 137




Ambystoma ombryoH
metamorphosis 109



during methamorphosis 107 in old age 151


radiation 174



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

intensity 151







cancer patients dogs 148, 150



antagonism and radiation 172
antibiosis 172

staiving animals





theory 10
spectra 17

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

atoms, excited
ionized 10


of tadpoles 145

axolotl see



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




.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

menstrual, see menstrual blood
Piotoplafuna-Monotjraphiei) IX:

effect 134,

concentration and mitogcnetic. 13S



112 by radiation fruit flies. 50 chicken embryo. 65 spectrum 35. 113 over-exposure J15. 178 - and polyploidy 169 cytagenin 152. failure of 93 methods finger radiation 97. 77. 80 - - . 150 roots 111. 154 tadpole legs 109 tissue cultures 83 ---- in yeast cells 110 -. 97. 100 184 flocculation of colloids discontinuous irradiation 103 disks. definition 47 - and radiation 43 - chemical radiation 33. 19 dissolution of salt as source of frog eggs as detector 81. radiation intensity 146 yeast buds 71 yeast volume 74 . 91 onion roots 58 - --- [u in in - muscle 154 nerve 112. 76. 160 cotyledon radiation 141 crown gall origin 171. radiating 145. 185 depression of mitogenesis 115 ~. 100 Enchelys radiation 137 radiation 38* error. 169 chain reactions. 89. spectrum 147 erythema 48 exhaustion by irradiation 43. see cornea and radiation human r day light affecting mitogenetic failure in mitogenetic - experiments 93 rays 108 death through irradiation 49. 125. 76. rotating 103. field 6.sources of in bio. 150 muscle radiation 29. 145 coagulation omitting radiation 89. radiation 145 cholesterol and cancer 182. 160 of radiating power 110 eye radiation. J31 secondary 117 see also spectrum 37 filters for light 21 detectors. 143 larvae.radiation 69. 128 radiation 169 embryo enzyme radiation 143. expcri mental. 187 heliotropifctm K eggs as detectors 81.false 70 - . 142 larvae see tadpoles - radiation 50 dog blood radiation 148. general 190 colony si/e method 75 colloid coagulation as detector 89. see also errors false mitogenetic depression 70 fermentation affected by radiation 85 causing radiation 124.of nerves 111. 142 as senders 142 183 affected chromosome number by electric. - 73.210 SUBJECT INDEX drosophila eggs as detectors 81. see drosophila dispersion 2. 161. 105. cornea as detector 84 --. 180 cytophotolysis 182 -. 61 f . with bacteria 78 & 100 conduction of radiation _ J 1 1 Geigcr counter 34.

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

148 bone marrow 64 - cancer 178 chemical reactions 50. 72. spectrum 38 nuclei in onion roots 55. 52 embryos 143 infusoria 133 optic. discovery 55. 110. conducting radiation 154 radiation 65. 61 over-exposure 43. 87 cornea 146 eggs 142 nephelometer measurements 74 nerve conduction 112. . J77. 44 onion root. 51. 115. wave length 49. 115 - monochromator 19 morphologic changes by radiation 86. 80 proteolysis 52 of 43. nuclear stages 129 method 63 radiation 54 . 103 . 96.212 SUBJECT INDEX mitogen?tic 110 field methods: necrobiotic radiation 99 onion root 54 peroxide decomposition 89. enzymes 37. formula of 64 . rays 99 blood 65. . 59. spreading of theory 128 photography 90. 119 stimulation. 77. 160 . 147 - . 119 sender mechanism 139 . 96 82. 59 intensity. see mitogenctic effect yeast buds 66. 140 spectra 140 opalina radiation 133 . 161 effect. spectrum 61 from bacteria 87. protozoa 133 sarcoma 64 tissue cultures 81 tissues 146 detector mechanism 129 . 125. 101 photoelectric counter 90 effect. 99 tissue cultures 81 radiation. 64 plants 138 polarized light 45. cell concentration 134 cause of 122 muscle. 68 yeast growth 72 range 49. secondary radiation onion roots 55 oxidation 29. see intensity properties 59. radiation . 131. secondary radiation 109. analysis J 19 and . 160 metabolism 156 . 70. 156 exhaustion 111. 178 N necrobiotic. larvae 143 from 50 molds menstrual blood 86. 129 nucleic acid. yeast 64. 87 physical nature of. 142 spectrum 40 muscle 65. 59. radiation 157 radiation 154 spectra 156 neutralisation. 59. 113. 142 yeast metabolism 84 mitogenctic depression 70. 61 mitotin 140. 147 neutralisation 50 nuclease action. discovery 55. 161.

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

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

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

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