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Milestones in microbiology 1546 to 1940


Milestones ln Microbiology 1546 t0 1940 T la d 4 れ d Edited わ ァ Thomas D. Brock University of Wisconsin, Madison ASM PRESS Washington, D. C.

Milestones in microbiology 1546 to 1940


Copyright ◎ 1999 American Society for Microbi010gy 1325 Massachusetts Avenue NW Washington, DC 20005-4171 Original copyright ◎ 1961 Prentice-Hall' lnc. ; renewed by Thomas D. Brock in 1989. AII rights assigned to ASM by Thomas D. Brock in 1998. Library 0 f Congress Cataloging-in-Publication Data Milestones in microbiology: 1546 t0 1940 / translated and edited by Thomas D. Brock. cm. These research papers were published predominantly ⅲ German and French' with a few ⅲ English. ISBN 1 ー 55581-142-6 (pbk. ) 1. Microbiology—History—Sources. QR21. M54 1998 579 ′ .09 ・ 一 DC21 I. Brock, Thomas D. 用 1 Rights Reserved Printed in the United States of America 10 9 8 7 6 5 4 3 2 1 9840858 CIP

Milestones in microbiology 1546 to 1940


MiIestones ln Microbi010gy 1546 t0 1940

Milestones in microbiology 1546 to 1940


HISTORICAL INTRODU(TION into a practical t001 b Ehrlich, it was now possible t0 speci cally cure a per- son suffering from these diseases by lnjectrng intO him some antltoxic serum prepared by earlier immuniza- t10n Of a horse or Other large animal. This led to the development for the first time Of rational cures for infec- t1011S diseases, and was more than any— thing responsible for Ehrlich's later conception Of chemotherapy. The antibiotics era has only been documented here by Fleming's first paper, which should be suffcient t0 ShOW the promise that 、 vas in store. lt would be diffcult indeed to select one or t , 0 additional papers in the field of antibiotics as further milestones along this road. At some future time, a whole book will be needed to docu- ment the historical development Of the antibiotic age. Although the bulk 0f the papers in this collection deal with applied ma- terial, there are a number Of studies which can only be classified as funda- mental in nature. 、 hey were not begun with any practical goal in mind, but only because the investigator was ln- terested in the subject. These have been included here under the heading of GeneraI MicrobioIogy. AIso in this sectlon are several papers dealing With methods, which have been placed here for want Of a better place. The further development of general microbiology is shown here through the papers 0f Winogradsky and Bei- )erinck WhO were primarily interested ln mlcroorganisms as Obj ects Of study, with little regard for practical prob- lems. KIuyver was BeiJerinck's suc- cessor and pioneered the field 0f mi- crobial physiology and biochemistry. The first clear-cut discovery of a gro 、 vth substance WhiCh can be con— sidered tO be a vitamin was made using 7 a mrcroorganism (yeast) as a test or- ganism. Although later work with mammals was necessary tO establish the vitarmns lmportant substances ln nutritlon, the ork With microor- gamsms has always been a great stlmu- lus tO vitamin research. Wildier's early paper on the BiOS substance has been included here. Another whole area of microbiology has been completely omitted in the present collection. ThiS iS the area Of microbial genetics. Although some of the most important developments in microbiology in recent tlmes have OC- curred in this field, it has not yet reached the stage where it can be p , 0 。 000d intelligiblx 0 00d0 , g , ad 。 - ates through isolate articles. ln addi- t10n, there are tWO collections which include most Of the significant work in this area: Lederberg's "Papers in Microbial Genetics ” and Peters' "Clas- SiC Papers in Genetics. ' The articles in the present collection were selected primarily tO show the beginmngs Of the SC1ence Of micro- biology, which arose through practical problems. Today it is becoming in- creasingly evident that microbiology IS at last becoming a true biological SCIence, able tO exist apart from its applications. lt is alSO becomlng in- creasingly clear that bacteria are prob ab the most useful living organisms for studying many processes in thc life of the cell. This is indeed encour- aging. 、 IO SCIence can ever thrive if it is built only on purely practical grounds. lt is to be hoped that if a new collection Of milestones ln mlcro- biology is made 100 years from now, it will amply document the funda- mental aspects 0f microbiology, and ShO 、 V that this science has truly of age ・

Milestones in microbiology 1546 to 1940


Preface 石 か イ な 川 房 0 ん , first published in 1961 , has been widely used as a supplement for bi010 and microbiology courses. 用 - though not a history text, it has also been used as a textbook for courses ⅲ the history of microbiology. I am pleased that the Amer- ican Society for Microbiology has decided to continue to make this book available. I began work on this b00k in 1958 , when I was on the faculty at Western Reserve University (now Case Western Reserve University). At the time I wasjust beginning my career. Although I had a strong interest in the history of microbiology, my working knowledge of the field was quite limited. lndeed, my knowledge of microbiology itself was ね 迂 ly restricted. What I did have was access to the original literature in an excellent history of medicine library, a good work- ing knowledge of German and French, and a willingness-töléön ィ ひ deally, science should be learned from reading original experl- ments rather than textbooks. However, beginning students often XI background. would be: undergraduate students with very little science collected papers. lt was edited with a clear idea ofwho the audience mere "cut and paste" 」 Ob , as were SO many subsequent books of thinking, obviously extraneous material. Thus, this book is not a siderable amount of what was, in the light of contemporary lated the original papers, I carefully edited them, deleting a con- simply a republication of a number of origmal papers. After I trans- lt is important tO emphasize that ハ 石 立 0 れ お 乞 石 c 川 房 0 ん 1 is not microbiology. 石 0 房 0 ん might serve the same purpose ⅲ the teaching of napolis, Md. , and Santa Fe, N. Mex. ). I felt that 石 れ championed since the 1930S so effectively by St. John's College (An- core of the "Great Books" approach to education which has been や 襯 召 厩 & 乞 化 , Harvard University Press, 1957 ) and was at the proach was pioneered by James Conant ( 〃 〃 矼 イ C 召 〃 0 厖 か instilling in the student a feeling for scientific research. This ap- study of a few key historic experiments can go a long way toward foundations Of science are much simpler to understand. A careful ' 復 : & を 、 porary experiments. Those classic experlments that served as the lack the background, chemical or otherwise, to understand contem-

Milestones in microbiology 1546 to 1940


Historical lntroduction ・ TH PRACTICAL AND TH E SCIENTI FIC aspects Of microbiology have been closely woven. Perhaps it is for this reason more than any Other that micro- biology as a el イ of study did not develop until the twentieth century. 、 ・ ineteenth-century workers were chemists or physicians, a few were bot- anists, but none called themselves bac- teriologists or microbiologists. lndeed, even tO this day mcrobiology is often not g as a body of knowledge, in the way botany and zoolo , are, but as an applied SCience. One reason for itS practlcal onenta— uon is that microbiology developed to solve practical problems. The two main areas Of development were in the fields Of fermentation and medicine•, the pure scrence Of microbiology did not eXlSt. Perhaps more than most sclences, the development of microbiology de- pended on the improvement of a tool —the microscope. AIthough there are no papers on the development of the mrcroscope in the present collection (that is really a branch of physics) , the quality 0f all 0f the work reported here reflects the quality of the micro- scopes used. Since bacteria cannot be seen individually 、 廴 & 当 ueu 生 を ye , their eXIStence as individuals can only be known from mcroscopic observa- t10ns. And Since most bactena are very small, ranging in size down to the limits Of resolution of the light micro- scope, their exrstence and character- istics are quite diffcult tO decipher. lndeed, it would be interesting to s ec- ulate on hOW bacteriology might ave developed if the limits of resolution of the microscope were poorer, so that all bacteria would have been invisible. Bacteri010gy could undoubtedly have evolved under these conditions, but surely at a slower rate, and surely in quite a different manner. From the time Leeuwenhoek first saw bacteria, until the time Koch proved a bacterium to be the cause of a disease, there was a 甦 gn. dous de- velopment in the light microscope. From Koch's time, when the Abbé condenser was introduced (latter nine- teenth century) , t0 the present there have been only minor developments, and few of them can be said [ 0 have had a crucial impact on bacteriology. But as he reads the articles, the reader should StOP tO consider in each one: hOW do the observaüons made and conclusions drawn reflect the quality Of the microscope? Many controver- sies may have really been due to the use by the investigators of different nucroscopes. 、 hiS iS a good lesson for tOday: When workers disagree, isn't it usually due tO variations in the methods used? 、 AIthough medicine and fermenta- tion presented the practical problems which stimulated the development of microbiology, the first studies that put the subJect on a scientific basis arose from a problem in pure science. This was the controversy over spontaneous generatlon, a dispute that aroused men 0f 2 philosopffrcal nature, men who studied science for sclence's sake, and consequently led tO the first scientific approaches to microbiology. The early crude ideas of spontaneous generation (mice from 01d rags, mag-

Milestones in microbiology 1546 to 1940


234 Enrichment culture studies With urea bactern 四 ・ 盟 4 切 一 Be 實 切 じ た GENERAL MICROBIOLOGY Beijerinck, M. 、 V. 1 1. Anhäufungsversuche mit Ureumbakterien. C 4 ル 耘 な f. Ba 々 olog , Part Ⅱ , VoI. 7 , pages 33 1. THEPRESENTPHASE INTHEDEVELOPMENT of microbiology can be called the "systematlC" phase, because lll biology, as in every young science, it is first necessary tO describe the ma- terial tO be studied, and tO place it intO an orderly classification. For thiS phase 0f microbiolo the enrichment culture expenment as a special im- portance. The enrichment culture experiment makes it possible tO iSOlate a large variety Of microorgamsms which are adapted tO particular en- vironmental conditions, and tO bring them int0 development side by side in liquid culture media. ln this way it is possible tO set up enrichment cul- tures under conditions which are similar tO those as they exist nature itself, such as in natural fermentations, or in bacterial diseases. But all the uncontrollable variables Of nature can be avoided in enrichment cultures and can be replaced by precisely con- trolled factors. (Footnote: Enrichment culture experiments are Ⅱ 第 ortant not only scientifically, but pe agogically ・ I intend tO publish at a later tlme a summary Of these ideas as ℃ Ⅱ as a description of the bacteriological methods used in my laboratory. ) A 000 。 orthy pr(terty of bo 市 the artificial and natura enrichments iS that the microbes that exist in the enrichment dO not occur ln a single variety, such as iS always the case when a single colony is isolated and 砠 further cultures made from it. ln the enrichment all Of the varieties are present that existed in the material used for inoculum and that can grow under the selected conditions. 、 、 are therefore ab 厄 tO become acquainted with a species not through 2 selected isolated variety, but through a number Of POSSible variatlons Of this species. This latter seems to me to be especially important for taxonomic studies, and it can be stated that every description Of a new species must begin With an enrrchment culture. Because Of our very lmperfect understanding Of the environmental requirements Of the maprity Of mi- crobes, it iS impossible in most enrich- ment culture experiments tO go further than tO bring about a relative lncrease in the numbers Of a desired forr れ , without leading to a complete disap- pearance Of the Other species present. Often this partial enrichment only occurs at a particular stage Of the experiment, whereas earlier and later Other forms predominate. Because Of this, enrichment culture experments can be called perfect" or "imperfect. " ln a perfect experiment, a single spe- iS iSOlated in all its varieties. Only

Milestones in microbiology 1546 to 1940


X11 PREFACE lation. As much as possible, I have tried t0 present a text which lated by me, and I am therefore responsible for any errors ⅲ trans- English translations. 、 /lost (but not all) Of the articles were trans- ing Koch and pasteur. I have saved them this difficulty by providing sian). The language barrier alone will deter most people from read- even fewer in Other European languages such as Dutch and Rus- predominantly in German and French' with a few ⅲ English (and the 19th century. The research papers were therefore published Microbi010gy developed into a science almost exclusively during entertaining as I have. the way t0 the present. I hope readers will find such reading as will read in this volume a wide array 0f historical facts that point interest here. Those curious tO learn hOW microbiology developed working in pure or applied microbiology will find many things 0f read these articles ⅲ their original languages. I think that anyone Further, most microbiologists d0 not possess the language skills t0 presented here, although they are surely aware 0f some 0f them. them have had the occasion or opportunity tO read the articles sional microbiologists will also find this b00k interesting. Few 0f tIOn tO its use by students, it has been 1 れ y experience that profes- These are reasons enough for the present V01u1 れ e. But in addi- of the field. select articles which we know were important for the development and the number of technical words is smaller. And it is possible t0 fragmented into subdisciplines. Their language is generally simplen They were written in earlier times, before microbiology had become historically important papers offer ideal material for the student. completed work, written ⅲ a highly specialized language. But these current research papers. They are generally only fragments 0f a On the other hand, it is difficult for a beginning student t0 read of the way in which current theories have developed out 0f the past. taught as a bOdy Of preexisting factS' and the student is not aware pasteur, and Koch really did. TO a great extent' microbiology is dent tO become aware OfJust what scientists such as Leeuwenhoek' treated in introductory texts, there is little opportunity for the stu- of the science. Although the history 0f microbiology is always are less likely to read any 0f the articles which form the foundations with many Of the papers presented here. But present generations There was a time When most microbiologists were have a level of sophistication unsuitable for young minds. undergraduate students as well as the one at hand' since it might did in 1958. On the other hand, such a new b00k might not serve more about both microbiology and the history of science than I doubtedly write a somewhat different book. Obviously, I know much If I were to prepare this b00k today, 40 years later, I would un-

Milestones in microbiology 1546 to 1940


260 ilation and assimilation, and perhaps may open a way for penetratln more deeply into these problems. A ter all, is quite remarkable that /. み 0 - can grow perfectly well in a mneral medium contmning a few per 00 。 0 0 。 00i , 01 if 0i0 。 g00 is suprlied is thus certain that this bacterium can use ammonra nitrogen for the synthe- sis Of its proteins. But it appears 引 SO that this organism does not grow III the same medium if glucose IS sub- stituted for mannitol, whereas, in the p , 。 。 。 00 of p 0 。 0 gl 。 。 0 , 0 is very rapidly oxidize , and the bacteria then multi ly profusely. Glucose is there- fore ully adequate as an energy-pro- viding substrate. Thus we see hOW t 、 VO substances, each one utilizable in 1 0 、 vn function, can furnish an inade— quate combination as fOOd. Might it perhaps be possible t0 ascribe this un- expected phenomenon tO a difference in the decrease in free energy Of the first stages Of the oxidation Of man- nitol and glucose, respectively, a dif- ference that might cause ammoma nitrogen tO be used for the synthesis Of bacterial protein in the one, but not in the Other case? 、 ル don't know; but we can learn from this example croorgamsms or those exhibitlng unusual chemosynthetic and photosynthetic ml- culture, and industry tO worry about the by applied problems in medicine, agri- When most workers were tOO fascinated microbiology were pioneers lll an era The men of the Delft school of general present. has carried this tradition down tO the his own work and that Of his students, Beijerinck had established, and through tradition of general microbiology that was just the right man 【 0 carry on the nical University at Del t, Holland. He the Chair of Microbiolo y at the Tech- Kluyver , as Bei)erinck's successor tO G N AL MICROBIOLOGY that the occurrence Of a biological oxidation does not depend only on the nature and condition Of the living cells and the presence of an oxidizable sub- strate, but that it is conditioned by very subtle modifications in the com- position Of the medium. lt seems tO be a practicable task tO determrne the successlve lntermediate oxidation stages of physiologically important compounds such as glucose, glycerol, etC. , With numerous orgamsms that exhibit a IOW oxidative capacity. At the same tlme one may attempt tO ascertaln lll a quantltatlve manner the ease with which the varl- ous steps are accomplished. lt is hardly doubtful that such investigations will lncrease our comprehension Of the nature Of microbial metabolism. But by virtue Of the above-mentioned unity it will also be eminently useful for our understanding Of the meta- bOliC activities Of higher organisms which are much less amenable tO ex— perimentation. However this may be, I ho 円 that my discussion may have convinced you that a study Of micro- bial metabolism 0 e many an ln- triguing problem t0 the chemist, par- ticularly in view Of bOth the existing diversity and the manifest unity. due to the influence of Kluyver, both work, and much of the credit for this is rect, in the light of the last 30 years' ThiS view has been shown to be cor- metabolism, even in higher organisms. understanding problems of general this unity was understood, it would help there was an underlying unity, and when diverse organisms 、 vith the vrew that presented here, Kluyver surveyed these lntermediary metabolism. ln the paper much Of our current understanding Of as Kluyver calls them, which has led to oxidative processes Of these specialists, ing Of the nature Of the fermentative and fermentations. Yet it was an understand-

Milestones in microbiology 1546 to 1940


K ル ア 0 ・ U が 4 d 市 ″ ッ 切 the e み 0 〃 可 な - ga な 腕 ゞ 25 3 diversity; quite understandably SO in view Of the fascination Of this type of work. If now I am going to embark on the attempt t0 give you 2 glimpse 0f the unity that can be discovered in m ト crobial metabolism, I do so with the full knowledge that it is but a meagre account I have tO 0 仟 er. That never- theless I have ventured tO SOlicit your attention for this tOPiC finds its ex- planation in the fact that I shall thus have the opportunity to make you realize that this still presents an 111 ト mense field Of endeavour. lt seems tO me that the solution of the problems in this area iS indispensable tO elevate microbiology from its status as a largely descri tive branch Of science t0 a higher p ane, and equally so t0 the application of microbiology t0 many situations. And I am firmly con- vinced that only a close cooperatlon between microbiologists and accom- plished chemists can lead to advances in this respect. Let me then first 0f all show you hOW the unity in the divergent meta- bolic processes 0f the microbes finds expression in the fact that ℃ recog- nlse in them the same general trends that have come tO light as a result Of lnvestigations Of the metabolism Of higher organisms. Without discussing this aspect in detail, the following re- marks m ay se rve tO illustrate this point. Studies 0f the metabolism 0f higher organisms have unequivocally shown that one can always distin ish tWO types Of processes. Part 0 the foodstuffs iS converted intO cell mate- rials, the latter being used either for growth or for replacement 0f degen- erated cellular constituents. Another fraction Of the fOOd appears tO owe its significance largely tO the therein ac- cumulated chemical energy WhiCh iS unleashed by the living cell, thus en- abling it tO carry out energy-requiring functions; this part is ultimately de- graded, generally producing heat. These れ VO processes are differentiated as assimilation and dissimilation. Doubtless the dissimilatory process iS the more essential charactenstic Of life. Whereas other typical vital phe- ・ nomena, such as growth, reproduction, lnternal or external motl()ll, may fre- quently be lacking, dissimilation is ab- sent only in some stages Of latent life, resulting, for example, from dessica- t1011. Consequently it had been estab- lished that in the case of higher or- ganisms the perpetuation Of life is tied tO a contrnuous conversron Of chemical intO Other forms Of energy, and Rubner demonstrated that the first law of thermodynamics applies to the energy transformations in the animal body, while Atwater showed this also for man. AS far as the nature Of the energy- providing reactions IS concerned, La- voisier had already pointed to the sig- nificance Of the respiratory process, え 巳 , the SIOW combustion taking place in living organisms, WhiCh reveals itself through their requirement for oxygen. The validity of this con- cept was emphasized when Rubner experimentally established his princi- ple Of "isodynamic substitution. " He showed that, up tO a point, proteins, fats, and carbohydrates can replace one another, in a weight ratiO that is inversely proportional tO the heat Of combustion Of these substances, in the adult animal for the maintenance Of the same vital functrons; lll Other words that, within certain limits, equality in chemical energy corres- ponds to equality in food value. As early as 1885 Rubner pointed out that in all probability it would be pos- sible that 引 so in the metabolism of mrcroorganisms a conversron could be designated which derives its signifi- cance for the organism entirely from the resultant energy liberation. NOW