A Life with Yeast Molecular Biology - Prof. Dr. Horst Feldmann

V.P. Skulachev and G. Semenza (Eds.) 

Stories of Success – Personal Recollections. XI 

(Comprehensive Biochemistry Vol. 46) r 2008 Elsevier B.V. All rights reserved. 

DOI: 10.1016/S0069-8032(08)00004-1 275 

Chapter 4 

A Life with Yeast Molecular Biology 

HORST FELDMANN 

Adolf-Butenandt-Institute, Ludwig-Maximilians-University, Munich, 

Molecular Biology. Schillerstrasse 44, D-80336 München, Germany 

E-mail: horst.feldmann@med.uni-muenchen.de 

Horst Wilhelm Albert Feldmann

276 

Abstract 

Born 1932, I was brought up in Stettin. I was trained as an 

organic chemist at Cologne University, before I entered the 

Institute of Genetics there in 1962 to work on tRNA structure and 

function. From 1967 to my retirement in 1997 I worked at the 

Adolf-Butenandt-Institute (Munich University) on several 

aspects of molecular biology of the budding yeast, for example 

tRNA genes, Ty elements, the yeast genome project, and 

programmed proteolysis. I include facets of my engagement in 

national and international organizations. 

Keywords: Yeast; tRNA; Ty; Yeast genome; Programmed proteolysis; FEBS; Spetses 

Within the recent 3 years, several anniversaries have been 

celebrated – the 40th anniversaries of the European Molecular 

Biology Organization (EMBO) and the Federation of European 

Biochemical Societies (FEBS) both of which were founded in 1964 – 

and that of the Spetses Summer Schools on Molecular and Cellular 

Biology in 2006. In 2005, the Cologne Institute of Genetics had 

organized a workshop to remember the ‘‘Early History’’ of this 

institution, the first of its kind in Germany opened in 1961. Finally, 

in 2006 many of the participants in the International Yeast 

Sequencing Genome Project gathered in Brussels to memorize the 

deciphering of the first eukaryotic genome in 1996. These venues 

remind me that I myself by now have devoted nearly all of my 

scientific life to Molecular Biology, especially that of a small 

unicellular ‘‘model’’ organism, the yeast Saccharomyces cerevisiae. 

But it took my first 30 years to bring me to this attractive field. 

My Early Years and Education 

H. FELDMANN 

I was born in Stettin, on a date the impact of which should 

determine the further political development in Germany and with 

all its consequences imprint the future of the next decades worldwide. 

On March 13, 1932 – my birthday – Paul von Hindenburg 

was re-elected Reichspräsident (German President). I keep an 

edition of the Frankfurter Zeitung (Frankfort News) of this day 

reporting on the massive and impudent election campaign, Hitler, 

Goebbels, and the Nazis had set up to push Hitler into the

A LIFE WITH YEAST MOLECULAR BIOLOGY 277 

position of Reichspräsident. This time the Nazis lost because the 

majority of the Germans highly respected Hindenburg and 

trusted that he was the one to save German unity and 

independence (actually, Hindenburg’s nomination had to be 

confirmed on April 10, 1932). But already several months later, 

Hindenburg lost any confidence in Heinrich Brüning, than 

Chancellor of the Republic, who had tried to overcome inflation 

and growing underemployment by emergency decree. Thus, 

Brüning’s demission marked the end of the Weimar Republic, 

and Hitler was appointed Reich Chancellor on January 30, 1933. 

My father, Wilhelm Feldmann, born in Siegen (Westfalia) was 

the eldest of five children. My grandparents run a bakery, and my 

father visited a technical school in Siegen to be trained as an 

engineer for countryside cultivation, water management as well 

as canal and river engineering. He found a position as a civil 

servant in a government facility in Stettin responsible for these 

affairs and married my mother, Hertha Mansen in 1931. My 

maternal grandfather’s family originated from Schleswig 

Holstein, and his genealogy is the only one in my family which I 

could trace back to the 17th century. As an employee of the 

German National Railway my grandfather Albert Mansen had 

been moved to Stettin, where he married Agnes Bohm, whose 

relatives lived in Stettin and Berlin. 

I still keep good reminiscences of Stettin as a wonderful town 

and would be able to draw a city map with street names and 

important buildings in German (such a map is no longer 

available) though I spent only 11 years of my youth there. My 

parents and I lived in an apartment of a tenement quarter built 

during the Gründerzeit and in the early years of the 20th century, 

in the same style as those quarters that became characteristic for 

Berlin and other fast growing towns around Germany. Fortunately, 

our domicile was located not far from the center, and my 

grandfather very often took me for long walks downtown 

explaining to me the meaning and history of buildings, monuments, 

and other attractive places. My favorite place was the 

famous ‘‘Haken-Terrasse’’ (a terrace and park named after a 

Stettin mayor) at the banks of Oder river surrounding a huge 

building that harbored the departmental administration and the 

shipping museum. From here, one could overlook the whole area: 

the harbor with boats of all kinds and steamers moored to the 

quays, the busy traffic on the Oder and its side arms, and, on the

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opposite banks, the free port area governed by huge cranes, docks, 

and shipyards. At home I transformed these impressions into 

drawings of ships, the essentials of which I knew from memory 

already at the age of four, or I constructed cranes with my metal 

construction set. My parents argued that with these skills and 

interests I would become a shipbuilding engineer. 

The harbor – already busy during the time when Stettin was a 

member of the Hanseatic league – at the end of the 19th century 

had developed into the most important port of the Baltic Sea and 

German trading place with the Scandinavian and Baltic countries. 

This became only possible after the strong fortifications 

had been torn down, so that the town could expand. New 

modern quarters and huge parks were created, which together 

with its natural lovely hinterland made Stettin a ‘‘green town,’’ 

offering numerous possibilities for relaxing tours. During summer 

time, we used to undertake week-end trips to one of the many 

seaside resorts at the Baltic Sea, which were served by regular 

pleasure steamers from Stettin. Sometimes, we visited my aunt 

and uncle in Berlin, and so I saw the Olympics in 1936 for 

one day. 

In 1938, I entered primary school, where we started writing in 

Sütterlinschrift; which was changed to Latin lettering during my 

third year at school. But I still can read and write this Gothic type 

which already my parents had been taught at school. Surprisingly, 

it was the Nazis who introduced the change. Other of their 

measures turned out to be less harmless, and I was at an age to 

realize some of the threatening developments, such as the 

increasing prosecution of the Jewish population and foreigners. 

For our daily life, the most far-reaching effect was Hitler’s 

rigorous expansion policy: in September 1939 he broke the nonaggression 

pact and started war. Progressively, all goods and food 

became scarce, but the extending war also began to exert a 

subliminal influence on our lives as schoolboys: we were obliged to 

collect warm clothing or money for the Winterhilfswerk (winter 

relief organization), or to collect waste paper and other waste 

material, even gnawed bones that were used for the production of 

soap. We had to set up silk moth colonies and provide mulberry 

leaves to feed them. Though this was given the veneer of a playful 

competition, my parents and all the more my grand parents were 

alerted. When the local Hitlerjugend forced me to visit their 

‘‘social evenings’’ held in a dump and dreary cellar – a measure


that I hated from the very first moment – my mother succeeded 

that I was exempted from this dismal obligation. 

The consequences of the war became worse for us during 1942, 

the year I entered secondary school, because air raids had reached 

Stettin. Measures against the bomb attacks were noticeable long 

before. Cellars had been transformed into air raid shelters and 

public air bunkers had been put up in public parks. Anti-aircraft 

units were installed all around the town and we could follow their 

practice to fix air planes by spotlights during night. I vividly 

remember that one nice morning in September 1942 one of my 

classmates appeared with singed trousers telling us that their 

house had been bombed last night, everything had burnt down 

and that this was left as his sole property. The attacks increased 

in 1943: we had to spend nearly every night in the shelter of our 

house. The most frightening missiles were the air bombs: coming 

down, they produced a sharp whistle. One started counting to ten, 

and if one was still alive after, the bomb had struck elsewhere. 

Finally, the authorities decided to evacuate all children to the 

countryside. Since my father, due to his occupation, had been 

exempted from military service and instead had been moved to 

West Prussia, my mother insisted that the whole family should 

take up residence in Kulm, a small town at the Vistula. So we 

went there in autumn 1943 and after some gipsy life obtained a 

newly built apartment. It was a quiet and splendid time for me. 

I attended an inter-denominational school in Kulm, where boys 

and girls were co-educated together with young Germanized 

Poles, and found my first love, a tender girl with curled blond 

hair. This happy period ended abruptly in bitter cold January 

1945. The Russian army was on the advance and we could already 

hear the near-by artillery fire. The Nazis, rigorously prepared to 

defend Kulm began, like all around Germany, to conscript all 

youngsters as well as any old men to the Volkssturm (German 

territorial army) endowing them with bazookas. My mother’s 

reaction was to pack a few small suitcases. (My father, for obvious 

reasons, had to stay behind.) I was to carry my satchel with some 

‘‘important’’ belongings of my own, our silver stowed in a small 

suitcase in one hand and my violin given to me by my grandfather 

in the other hand. Where to turn? All nearby bridges across the 

Vistula had been conquered by the Russians and railway 

connections had been interrupted. Fortunately, the Vistula was 

completely covered with ice, so that our doctor was able to drive

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H. FELDMANN 

us by car across the frozen river, where in a small town further to 

the west we waited for the ‘‘last train’’ towards Danzig. When it 

arrived it was already full of refugees. By the crowd storming the 

wagons I was pushed under the slowly rolling train but my 

mother pulled me out safely, only my violin case was hit by one of 

the wheels and has retained a deep notch until today. 

From Danzig we could make our way to Stettin to meet our 

relatives, but as the town had been heavily bombed the city was 

nearly 90% destroyed, and we did not get permission to stay. 

Finally, my mother decided to turn northwest, to cross the Danish 

border and to reach Sønderborg, where my grandfather’s cousin 

lived. As Denmark was occupied by the German army, more than 

200,000 refugees from the Eastern territories had been transported 

to Denmark. Many like us were allowed to stay in private 

quarters, until in May 1945 the Germans had to surrender to the 

British, an event that occurred without firing a shot. 

My grandparents, my mother, and I were then accommodated 

at the Sønderborg Masonic lodge which served as an internment 

camp. We lived together with some 40 people in the ceremonial 

hall, the walls painted black with a blue ceiling decorated with 

stars – but no daylight. Given this mystic atmosphere, the elderly 

ladies used to practice occultism behind black curtains. In early 

1946, all German refugees were concentrated in larger camps and 

we were transferred to one near Sønderborg, a former large shack 

camp of the German marine. With some 40 people, we had to 

share a tiny hut heated by one cast-iron oven that was also used 

for cooking, no nearby showers or toilets were available. A young 

girl of 16 in the bed next to me died from tuberculosis. As I fell 

seriously ill, our family was moved to a smaller camp (the former 

German school in Broager), which was sort of return to more 

freedom. There were some 40 children who could enjoy a huge 

playground and go for swimming. I organized a children’s circus, 

staged two of Grimm’s tales and founded a harmonica ensemble. 

I found much leisure to do handicrafts and to read books 

I discovered in the attic, such as Eddington’s ‘‘Popular Astronomy,’’ 

teaching books in algebra and geometry, physics, and 

chemistry. I had an opportunity to learn French and English, an 

elderly woman teacher taking care of me. 

In February 1947, my father was able to return to his parent’s 

house in Siegen, which however had been nearly completely 

destroyed during the war. We were allowed to leave Denmark and


to join him. We were fortunate to find one intact room in this 

house and, more importantly, an intact garden to grow all kinds 

of vegetables scarcely available otherwise (food ration cards were 

still in use). I managed to re-enter the local secondary school, and 

my grandfather taught me how to play the violin. My free time I 

spent with my friend Dieter Zimelka, who was handicapped by 

reduced verbal and physical capabilities due to spastic lesions, 

which he compensated by intelligence and will. We undertook 

cycle tours, collected plants for a herbarium, and were both 

interested in chemistry. After he had passed school with the best 

marks, he studied engineering at Aachen Technical University. 

We kept contact by correspondence and mutual visits until he 

died at the age of 47. 

In 1949, my father found a position in Düsseldorf and we moved 

into a new modest apartment there. Three years later, I finished 

school at the Max-Planck-Gymnasium in Düsseldorf and had to 

think about my future occupation. On the one hand, I was 

attracted by chemistry. During my time at school, I had been 

allowed to set up experiments for our lessons in physics and 

chemistry, and I even had performed some risky experiments at 

home. One day I stunned my mother, when I tried to produce 

bromine, which however ruined all nutrients in our kitchen: 

unfortunately, the bromine had crystallized in the cooling device 

and caused an explosion of the whole device. On the other hand, I 

developed a foible for architecture. From my father I had learned 

some skills in trigonometric techniques, civil engineering, and 

how to draw plans. Above all, I felt that architecture was an ideal 

subject for creativity. (Later, I experienced that chemistry could 

do the same!) Finally, it was decided that I should enrol for 

chemistry at Cologne University. The argument was that I could 

live at home and travel to Cologne every day by train; the main 

station was near our domicile. Unfortunately, chemistry as a 

subject was over-crowded. A major bottle-neck was to allocate so 

many newcomers (about three hundred candidates had accumulated 

among which were many late returnees from prisoners-ofwar 

camps) to a lab space to begin practical work. So I 

concentrated on lectures in chemistry, physics, and philosophy. 

Together with one of my new companions, Max-Dieter, I enrolled 

for mathematical courses which I followed for four terms. In my 

second year, I passed a test and was admitted to the first lab 

course starting with an endless number of inorganic analyses, for

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which we had to provide our personal equipment and all 

necessary chemicals ourselves, on top of the usual university 

charge. The good news was that once one had successfully 

finished a given program after passing an oral exam, there was a 

guarantee to take up the next course, which culminated in the 

analyses of an ‘‘exploded drugstore.’’ The ultimate goal – before 

we were admitted to the intermediate diploma – was to correctly 

work out the quantitative analysis of a piece of mineral within 2 

days. The only unforeseen difficulty that arose for me and a few 

other candidates was that our assistant had overlooked that the 

institute was completely closed the first day (a local holiday), so 

that we had to find our way in and out through a window in the 

basement. 

Despite the very rigid schedule, I found time for some 

interesting ventures. After our third term, Max-Dieter and I 

participated in a vacation program across Italy organized by the 

Italian student association, my first ‘‘voluntary’’ trip to a foreign 

country. By train we traveled via Munich to Rome and continued 

to Naples. There our German group met with students from Italy, 

France, Finland, and England. In the evening we embarked on a 

small boat to sail to the volcanic island of Stromboli, where we 

were accommodated for 10 days in an open air camp near the 

picturesque, black shore. At this time, Stromboli was nearly 

empty – few inhabitants, no tourists. The painful aftermath of the 

big eruption 5 years earlier were still visible: destroyed and left 

houses, uncultivated gardens, and wild shrubs. A commemorative 

plaque on one of the houses documents that Roberto Rossellini 

spent some time on the island with Ingrid Bergman to produce his 

film Stromboli (1949). The most fascinating enterprise was a 

guided night-tour to the peak of the volcano where we enjoyed a 

marvellous sunrise. We continued by sailing towards Sicily 

touching most of the scenic Eolic islands and spent a week’s time 

in Messina and Taormina. 

In my fifth term, I was elected member of the student 

representatives (Cologne Student Council) and my job became 

to take care of ‘‘social and cultural affaires.’’ Among other 

activities, I was able to invite well-known literary cabarets, 

readings, the University of Michigan student choir, a Jamaican (!) 

steel band, and to arrange for an exhibition of Cologne student 

paintings. A most successful activity was to organize cheap bus 

tours to Paris. Transport plus a stay in a small hotel for 6 days


was offered for 40 (!) Deutsche Mark per person. During my time, 

some 20 Cologne students got a chance to participate in a meeting 

with French students in northern France. We were put up in a 

small hotel located in the dunes of the Channel, enjoyed excellent 

meals (with four courses!) twice a day, met for political 

discussions, and were invited for sight-seeing and official 

receptions in Lille, Calais, Boulogne-sur-mer, and Dunquerque. 

An incredible story happened at my time as a member of the 

Student Council. The university financed a secretary to help in 

our common office. We hired a young lady saying she had escaped 

from Deutsche Demokratische Republik (DDR). After a year or so 

she left to take a governmental post in Bonn. It turned out, 

however, that she was a high-ranked spy who had used her initial 

job as a springboard to infiltrate the Ministry of Defence in Bonn. 

Like many students, I regularly took a job during the vacations. 

I worked in my father’s office, at the central chemical laboratory 

of Henkel Company in Düsseldorf or the chemical laboratory of 

Leybold (vacuum technologies) in Cologne. Henkel at that time 

was developing long-chain fatty alcohols esterified to sulfonic acid 

as new detergents for washing powder. They kept it secret, and 

only much later I realized what I had been working with. Several 

times I volunteered at the municipal gardening bureau in Neuss, 

where I was responsible for designing children’s playgrounds, and 

in the office of an architect in Cologne. In between I earned 

money by designing graphical advertisement for shops, for the 

Cologne tram, or privately. At one of such occasions as a working 

student I met Hildegard Beissel, who was enrolled as a student of 

economics at Cologne University, and we married after my 

diploma in 1960. Her brother Heribert Beissel, then a young 

conductor, arranged for a contact with the Bonn Theatre, and 

together with the Bonn ballet we staged three pieces, Mozart’s 

Bastien and Bastienne; Negro Spirituals; and Prokofieff’s Peter 

and the Wolfe. The material for the simplistic scenery I built in 

my 9 m 2 Cologne mansard. 

After my intermediate diploma, I had to return to serious life in 

the lab and to follow the strictly regulated training as an organic 

chemist. At that time, the chemical institute was housed in an old 

hospital near the main buildings of Cologne University. Meanwhile, 

the area has been used to build new facilities, the new 

Institute of Genetics being one of the latest. During their diploma 

or even doctoral thesis work, the trainees had still to fully pay for

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any equipment and material. During the practical courses, bench 

space was limited to about 1.5 meter in width, and normally 40–50 

people had to share a huge laboratory. Despite this high density, 

there was only limited contact or communication among the 

fellows. Actually, everyone buried himself in his own subject, 

mostly not realizing what his next neighbors were engaged in. 

The only common activity consisted in (rather boring) two-weekly 

seminars, where every fellow had to report news he found in 

periodicals he had been assigned to. The situation improved 

during my thesis work: we obtained more space, communication 

grew more intense, and resulted in a good co-operation between 

the lab fellows. 

I had already experienced that chemistry in some respect is a 

dangerous subject (not to mention the sometimes stinky air in the 

lab), and nearly every one of my colleagues had to pass through 

one or the other bad experience. I remember three personal 

accidents from this time, which, however, ended without too 

serious consequences. During our practical courses I was obliged 

to synthesize a mustard gas-like compound, which handled even 

under extreme precautions, caused a cauterization of the corneas 

of both my eyes; fortunately it was cured by special treatment and 

staying in complete darkness for 1 week. Two compounds I 

prepared for my thesis work invoked explosions upon gentle 

distillation, which ruined all glassware in the lab. 

My thesis work started in 1960 under the guidance of Professor 

Leonhard Birkofer, who at that time was the only organic chemist 

in Cologne since Professor Kurt Alder had deceased in 1958. His 

main interests concentrated on natural compounds, such as dyes of 

flowering plants like Petunia which he obtained from a cooperation 

with the institute of botany, as well as silicon-organic 

compounds. But the problem I had to investigate was the 

possibility of synthesizing asymmetric diamino acids which had 

never become known before. This turned out to be very tricky, 

because these compounds were extremely unstable. However, some 

of the precursors I had to prepare yielded novel N-heterocyclic 

compounds upon further reaction, and in the end I had collected 

and characterized some 20 of such derivatives [1,2]. InJune1962, 

I finished my PhD after having passed oral exams in three subjects, 

organic chemistry, physics, and physical chemistry, with the best 

marks, for which I was awarded a prize by our university. During 

this period of time, I gathered a first teaching experience as an


assistant in chemical courses for medical students and candidates 

for a teaching post. Usually, in those years young chemists had no 

problem to find a suitable job, but after a number of interviews in 

leading German chemical companies, I decided not to take a 

position in industry, rather I was attracted by the possibility to do 

basic research. As no free post for a research assistant was open at 

the chemical institute, my boss recommended to apply as a post-doc 

at the newly founded Institute of Genetics in Cologne. In 

September 1962, I was engaged as a post-doctoral fellow by 

Hans-Georg Zachau, who headed the department for nucleic acid 

research, after I had convinced Max Delbrück, appointed the first 

director of the institute, in an interview that I had an honest 

interest and a good qualification to do research in molecular 

biology. Undoubtedly, this was the decisive switch in my career. 

One has to recollect that around that time – and even for so 

many years to follow – no training in biochemistry, never mind 

molecular biology, was offered at German universities. During my 

studies of organic chemistry, protein chemistry was touched only 

peripherally, nucleic acid chemistry simply did not exist. So my 

first notion of this field stems from a fascinating lecture by late 

Fritz Cramer talking about methods of oligonucleotide synthesis, 

when applying for the vacant chair in Cologne. Of course it took 

some effort for a ‘‘beginner’’ to grasp the essentials of biochemistry, 

genetics, and molecular biology. But excellent books helped 

open this new world, for example, Biochemistry by Peter Karlson, 

Classical and Molecular Genetics by Carsten Bresch and Rudolf 

Hausmann (both from the Institute of Genetics); books on nucleic 

acids such as by A. Michelson or Lord Todd. 

Institute of Genetics and tRNA 

How different became life in the Institute of Genetics! While in 

organic chemistry there were only two departments, each employing 

some three assistants and the same number of technicians, the 

Institute of Genetics accommodated five departments, each – on 

average – endowed with six scientific co-workers and roughly the 

same number of technicians. Conceived as an inter-disciplinary 

research institute, Genetics was open to collaborators from 

different fields, such as physics, chemistry, biology, or medicine, 

and most remarkably, to foreign co-workers. A completely new

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H. FELDMANN 

experience for me was to find a perfect ‘‘infrastructure’’: 

secretariat, workshop, library, cleaning kitchen, etc. 

A completely new experience for me was team-work, the 

superior maxim in every department of the Institute of Genetics. 

People collaborated and talked to each other very openly. 

Contacts between people from the single departments were 

guaranteed by weekly seminars and colloquia. In the seminars, 

all researchers and doctoral fellows had to elaborate on a given 

topic. One of the outstanding issues in discussions at that time 

(1962/1963) was the Genetic Code: triple, quadruple, comma-free 

or not? [3]. The colloquia were covered by invited speakers. For 

1963 only, I have counted 68 renowned scientists from all over the 

world visiting the institute. When Max was around, he used to sit 

in the first row in order to ‘‘control’’ the speaker. If he felt 

something mysterious in this presentation, he turned to the 

audience: ‘‘Everybody got it?’’ If there was no clear-cut answer, 

the baffled speaker was urged: ‘‘Better say it again!’’ 

The relaxed atmosphere of the institute became manifest 

through the many parties, for which Max had a foible. A most 

spectacular venue was the farewell party for Max on July 19, 

1963, when he had decided to return to CalTech at Pasadena. I 

have tried to document this venue by editing the original 

manuscript we used for the performances. In our sketch, Max 

was damned to be tied and boiled by the ‘‘wild Zachaus’’’ in a 

huge container, which we used for our large scale tRNA 

preparations. To mimic the boiling water, we had put some 

dry ice together with a little water onto the bottom. After a while 

Max sighed in great pain: ‘‘Can’t you at least remove the dry ice; 

my back is already burning?’’ Even after Max had left the 

institute this atmosphere was not lost, even during the ‘‘stormy 

time’’ the crew had to pass. Figure 1 shows Carnival decorations 

Rainer Thiebe and I fabricated to illustrate this phase of 

depression. It would, of course, be really tempting to tell 

more anecdotes about people and life in the institute during the 

first years. 

Clearly, the unconventional setup of the Institute of Genetics 

made it the birthplace for Molecular Biology in Germany. In 1962, 

I could engage myself in this field, experiencing and applying new 

techniques devoted to the analysis of transfer ribonucleic acids 

(tRNA), the subject Hans Zachau’s group (Figure 2) had decided 

to work on. My first task became what was called the ‘‘2u/3u


Fig. 1. Carnival at the Institute of Genetics in Cologne, 1964. From left to right: 

Commander Max Delbrück, Lieutenant Walter Harm, Vice-Commander Peter 

Starlinger, and Able-bodied Seaman Hans Zachau. 

problem’’: it had been established that for protein synthesis the 

amino acids are hooked onto specific tRNAs via an amino ester 

bond to the ribose of the A residue of the ‘‘3u-CCA-end’’ and from 

there are transferred to the growing peptide chain, but it 

remained open whether the 2u- or the 3u-OH group of the ribose 

was involved. In a series of experiments, using chemical and NMR 

(nuclear magnetic resonance) spectroscopic methods, we compared 

a number of synthetic amino adenosyl esters with 

aminoacyl adenosine isolated from total tRNA that had been 

charged with amino acids by amnioacyl tRNA synthetases. While 

all synthetic compounds exhibited a ratio 30:70 for the 2u- versus 

the 3u-esters (probably due to acyl migration under the reaction 

conditions), at least 95% of the ‘‘natural’’ aminoacyl adenosine 

consisted of the 3u-ester. With this finding, we arrived at the 

conclusion that the 3u-linkage was the one active in aminoacyl 

tRNA [4–8]. From theoretical considerations, it had been argued 

that the 2u-hydroxyl of the terminal ribose should be the more 

reactive in the amino acylation reaction. Much later, this 

contradiction was solved by showing that two structurally 

different classes of aminoacyl tRNA synthetases do exist, class I

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H. FELDMANN 

Fig. 2. Zachau’s group (‘‘3rd Floor’’) at the Institute of Genetics in Cologne, 

1962. From top to bottom and left to right: Susanne Notz, Hans Zachau, Fritz 

Melchers, Dieter Dütting, Horst Feldmann, Anita Mosch, Hugo Gottschling, 

Gisela Schultz, Paula Prüfert, Rainer Thiebe, Wolfgang Karau, Gudrun Patzelt, 

and Heidi Heusinger. 

enzymes transferring the amino acid to the terminal 3u-hydroxyl, 

class II enzymes to the terminal 2u-hydroxyl prior to a rearrangement 

yielding the 3u-derivative, which in all cases is the active 

form of the charged tRNAs in protein synthesis. 

In 1963/1964, I became integrated into Hans Zachau’s main 

project, deciphering the primary structure of the major serine 

specific tRNAs from yeast. This subject again challenged the 

analytical skills of a chemist, but at the same time met my interest 

in molecular architecture. In fact, it was pure and hard chemistry 

at the beginning. We had to isolate the starting material from 

large quantities of brewer’s yeast. I guess during this period 

I myself worked up about a ton of yeast slurry I had to supply from


‘‘SESTER Kölsch’’ brewery in Cologne. Each batch (about 100 kg 

yeast slurry) had to be stirred with 200 liters of a phenol/buffer 

mixture for several days, then the aqueous solution was decanted, 

the soluble RNA precipitated by adding 250 liters ethanol, and 

harvested by sedimentation in a huge centrifuge. The raw tRNA, 

resembling a brownish shoe polish, was purified on a DEAE 

cellulose column and yielded some 30 grams of white material [9]. 

Later, C.F. Boehringer and Soehne, Mannheim, set out to produce 

yeast tRNA on an industrial scale following this protocol, and we 

were lucky that they supplied enough of this material to us free of 

charge. The ensuing steps, namely the purification of the serine 

specific tRNA, consisted of a series of counter-current distributions 

in different solvent systems, whereby every fraction had to 

be tested for amino acid acceptor activity (i.e. in this case, 

enzymatic charging with radioactively labeled serine for measurement). 

We were fortunate that Zachau’s group received sufficient 

support to buy the most modern equipment available on the 

market: an automatic counter-current machine composed of 

300 tubes with 20 ml capacity each manufactured by E.C. 

Apparatus Co., Swarthmore, Pa., and an automatic liquid 

scintillation counter developed and sold by Hewlett-Packard 

Company. 

In principle, these devices worked reliably, but any operator’s 

error could have fatal consequences. Once a tube of the countercurrent 

machine had been broken, a whole battery of 10 tubes 

had to be replaced, because no glassblower around was able to 

repair it, so we had to wait for an original set supplied by the 

company. A nasty but advisable procedure was to clean the 

apparatus after each use by washing it with sulfochromic acid to 

avoid potential contamination by ‘‘finger’’ ribonuclease. 

At the end of 1965, we finished sequencing of yeast serine tRNA 

[10–12], shortly after Robert W. Holley’s group in Ithaca had 

published the sequence of the first tRNA from yeast [13]. The 

analytical procedures involved complete and partial digestions 

with T1 and pancreatic ribonucleases, resolved on extremely thin 

DEAE (diethylaminoethyl) cellulose columns in 7 M urea, 

subsequent digestions with snake venom phosphodiesterase or 

micrococcal nuclease, followed by paper chromatography and 

spectrometric identification of the single constituents generated 

from the oligonucleotides by alkaline hydrolysis. In all, we 

collected several thousand UV-spectra, all recorded by hand.

290 

H. FELDMANN 

In fact, we had solved the primary structures of two very closely 

related molecules not completely separated by counter-current 

distribution, termed Ser I and Ser II. But when I analyzed each of 

the side fractions, it became clear that Ser I and Ser II, both 

present in about equal amounts in our preparations, differed in 

only three nucleotides (one in the TCC loop and two in the extra 

loop) [14]. Later, when the complete sequence of the genome had 

been determined (see below), no gene for Ser I could be identified 

in lab strain aS288C but 11 copies for Ser II. The ‘‘secret’’ of Ser I 

has never been solved. Probably, commercial brewer’s yeast we 

used (C836) consisted of two related species. Along with the 

sequence determination, I became involved in characterizing the 

structures of two ‘‘odd’’ nucleotides, N 6 -acetylcytidin and 

isopentenyl-adenosin (iPA), the latter of which we had found in 

serine tRNA at the 3u side of the anticodon [15]. 

A report on the serine tRNAs [16] was included in the Cold 

Spring Harbor Symposium on Quantitative Biology devoted to the 

‘‘Genetic Code,’’ 1966. This offered me the splendid opportunity 

not only to follow the current achievements reported at this venue 

but to visit a number of laboratories in the US, at the University 

of Albany, at the Roswell Park Memorial Institute in Buffalo, at 

the University of Illinois in Urbana, at Oak Ridge Natl. Lab. in 

Tennessee, and at the University of Chicago. I also visited old 

friends from Genetics, Fritz Melchers at the Salk Intitute in 

La Jolla, San Diego, Max Delbrück with his wife Manny and 

Charles David at CalTech in Pasadena, and Thomas Trautner at 

the UCSF, Berkeley. In order to take the cheapest way of flying to 

New York and back, I choose a prop-jet of Icelandic Airlines 

starting from Luxembourg, which offered a 24 hour stop-over in 

Reykjavik. There I met Johann Gudmundsson, who had obtained 

his diploma in chemistry during my time in Cologne and had 

taken a position in a fish cannery in Reykjavik. On the spot, he 

managed a sight-seeing tour in his veteran Ford car and showed 

me around a lot of the impressing country. This was ‘‘Iceland in 

24 hours,’’ but the impressions from this intense trip should stay 

for the rest of my life. 

With so many contacts and after having done some work in my 

‘‘new field,’’ it would have been easy for me to find a post-doctoral 

position in the United States, but just during my time in Genetics 

our two daughters had been born (Barbara in 1963 and Miriam in 

1966), and so we preferred to stay in old Germany.


Institute of Physiological Chemistry in Munich 

tRNA Biogenesis 

In early 1967, we had to say goodbye to Cologne and Genetics: 

Hans Zachau had been offered a chair at the Institute of 

Physiological Chemistry in Munich, and I was happy to get a 

tenure position there and to start my own laboratory. My first 

contact with yeast extended to a general topic of my research 

during the years to follow. What I had learned in Molecular 

Biology from the ‘‘early days’’ on, I tried to pass on to our 

students, and I am happy that even many of the medical students 

realized the importance of molecular biology for modern medicine. 

At first, I continued to work on the multiplicity of serine tRNAs, 

trying to isolate isoacceptors (e.g. the one specific for the AGU/ 

AGC codons), but this failed because of the minimal quantities of 

this species occurring in yeast [17]. (Later my lab at least 

succeeded in characterizing the three genes encoding this tRNA.) 

So I decided to work on the methionine specific yeast tRNAs, as 

we could be sure that there were two discernible activities, one for 

the initiation and one for the elongation of peptide chains [18]. 

After initial experiments, we decided to completely change our 

former sequencing strategy, now employing [ 32 P]-labeled material 

and adopting the elegant ‘‘Sanger technique’’ [19]. I got 

accustomed to grow 24 liters batches of yeast cells fed with 

200 mCi [ 32 P] phosphate each time we needed fresh tRNA. All 

manipulations for harvesting the cells could be done during night 

time leaving no traces of radioactivity in the lab or elsewhere. 

Supernatants were put to a device in the basement and stored in 

big tanks for decontamination, solid waste was kept in big iron 

casks for more than 10 half-life times of [ 32 P] so that virtually no 

radioactivity remained. A more serious issue was to convince our 

colleagues that, with appropriate precautions such as special 

safety hoods and a CO2 extinguisher, there was no problem 

performing high voltage electrophoresis in tanks filled with 

20 liters kerosin for cooling. Fortunately, I could pay a short visit 

to the Sanger lab at MRC (Medical Research Council), Cambridge, 

where Bart Barrell showed me several tricks how to run the 

analyses and how to interpret the radioautographs obtained after 

2D-electrophoresis. We published the primary structure of the 

yeast non-initiator methionine tRNA (tRNA Met 

3 ) [20,21] after Tom 

RajBhandary’s lab had finished that of tRNA Met 

i [22].

292 

H. FELDMANN 

Besides this structural work I became interested in the 

characterization of precursors to tRNA in yeast as a model system. 

tRNA precursors had already been studied in E. coli and phage T4 

[23,24] and Sidney Altman and collaborators had detected that 

endonuclease P was necessary to produce mature tRNA from its 

precursors [25], but no details were known in eukaryotes [26]. We 

used gel electrophoresis of ‘‘soluble’’ RNA preparations of yeast 

cells pulse-labeled with high doses of [ 32 P] phosphate and found 

that some distinct bands migrating slower than mature tRNA 

appeared. The smaller of these products contained minor amounts 

of the tetranucleotide TCCGp characteristic for tRNA but only 

traces of other minor nucleotides, while the larger of these 

products contained the tetranucleotide UUCGp but no minor 

nucleotides. Thus we concluded that these bands represented 

precursors to tRNA. This was confirmed by incubating material 

isolated from several of these gel bands in vitro in a yeast cell 

lysate, which yielded mature tRNA in kind of a two-step process 

[27]. Soon after, Goodman, Olson, and Hall reported on the first 

yeast tRNA gene to have an intervening sequence [28]. 

In conjunction with our efforts to identify the precursors to 

tRNA, we developed a two-dimensional gel electrophoretic system 

that enabled us to reproducibly map 40–50 individual tRNA 

species including isoacceptors from yeast, as well as individual 

precursors [29]. tRNAs were identified by (i) co-electrophoresis 

of purified [ 32 P] tRNAs with non-labeled bulk tRNA; 

(ii) comparison of patterns derived from pure tRNAs with bulk 

tRNA; (iii) fingerprinting of spots from pure [ 32 P] tRNA species; 

(iv) electrophoresis of bulk tRNA charged with one [ 3 H]- or [ 14 C]amino 

acid, whereby the aminoacyl tRNA was stabilized prior to 

electrophoresis by transforming the amino group into a 100-fold 

more stable OH-group. We realized that this technique of high 

resolution capacity – among other applications – was not only 

helpful to islolate specific yeast tRNAs but to identify the amino 

acid accepted by them, likewise also for determining the specific 

tRNAs contained in a tRNA population from any organism. One 

example is kindly mentioned by Guy Dirheimer, whom I met for 

the first time in the 60’s, in volume 44 of these Personal 

Recollections [30]. He sent Jean Weissenbach to my lab to learn 

the details of the procedure. They applied it to separate and 

isolate the mitochondrial tRNAs from yeast and started sequencing 

of several of those (e.g. [31]), a subject the Strasbourg lab


very successfully continued until 1986, also paying attention to 

the non-canonical codon recognition and the biogenesis of minor 

nucleotides in yeast mitochondrial tRNAs [32,33]. 

The collaboration between Strasbourg and Munich was intensified 

by mutual visits of our whole groups during these years. 

Excellent opportunities for contacts with the many colleagues 

working in the tRNA field and exchange of new results and ideas 

were offered by the yearly tRNA workshops each one organized by 

a particular laboratory at most attractive places world-wide. Of 

those I keep a deep memory (and the Abstract booklets, too) are 

the ones in Cambridge (1970), Göttingen (1971), Nof Ginossar 

(Kibbuz in Israel), organized by Uriel Littauer in 1975, Sandbjerg 

(Alsen, Denmark; 1976 – because it took me back to the area 

where I had spent 2 years as a boy), Aarhus (1978), Strasbourg 

(1980), Tokyo (1983), Taos (New Mexico; 1985), and Umea˚ (1987). 

Most impressive was the workshop in Japan held in Hakone, a 

resort area by the foot of Fujiyama at a hotel built in 1899 for 

a visit of the German Emperor. I extended the trip with my 

friend Wolfgang Wintermeyer to visit Tokyo, Kyoto and Nara, 

Singapore, Hong Kong, Bangkok, and Taipei, which was easy to 

arrange as we flew by Singapore Airlines who offered multiple 

stop-overs at low cost. 

A very attractive and scenic place was Taos, where the meeting 

in 1985 was held in a motel, not far from the Taos skiing area, 

which some of the participants enjoyed for one free day. The 70 

fantastic slopes had been built by an Austrian fellow and given 

fairy tale’s names like Schneewittchen (Snow White), Rumpelstilzchen, 

etc. To reach the meeting, I had to rent an Ugly Duck in 

Albuquerque, but afterwards this car was good enough to carry 

me about 2,000 miles around Four Corner’s Monument. I enjoyed 

the picturesque landscape with its big miracles of nature among 

others Mesa Verde, Monument Valley, Painted Desert, Petrified 

Forest, and Grand Canyon. 

The two-dimensional gel electrophoresis system prompted 

Walter Kleinow from the Zoological Institute in Cologne, who 

worked on mitochondria of Locusta migratoria, to start collaboration 

with us. From the minute amounts of RNA he was able to 

prepare, we succeeded to first detect the unusually low (GþC) 

content for mitochondrial rRNA and tRNA and than to resolve 

B27 tRNA spots, which migrated faster than the cytosolic tRNAs 

indicating smaller sizes of the mitochondrial tRNAs [34]. That the

294 

H. FELDMANN 

population of 27 tRNAs was far below the minimal number of 

tRNAs to translate all sense codons according to the Wobble 

hypothesis (32 in cytoslic tRNA) had just been reported by 

Dirheimer and colleagues (e.g. [35]) from their investigations on 

yeast mitochondrial tRNAs. However, the smaller size and the 

low (GþC) of the locust mitochondrial tRNAs were unexpected 

and pointed to further peculiarities of these molecules. 

The low amounts of material available to do further studies on 

locust tRNAs tempted me in 1976 to ‘‘bit to the side’’ and to 

devote part of our lab activity (Figure 3) to the isolation and 

analysis of mtDNA and tRNA from rat liver mitochondria. 

Around this time, some data on these issues had been published 

but they were rather inconsistent. So I interested a new doctoral 

student, Rüdiger GroXkopf, to take up this work in 1977/1978. In 

the beginning we had to isolate the starting material each time 

from sacrificed rats as our laboratory had no permission for 

cloning as yet. Also home-made restriction enzymes had to be 

employed and I am still grateful to the colleagues from Zachau’s 

group for their generous gifts of several enzymes. So, as a basis for 

Fig. 3. My collaborators in 1977: Antonin Eigel, Christa BleifuX, Maria 

Wagner, Petra Müller, Gabriele Goertz, and Rüdiger GroXkopf.


further experiments we established a reliable restriction map of 

the mtDNA [36], and finally could take advantage of cloning. 

Using the Maxam-Gilbert technique [37], large portions of the 

mtDNA were sequenced and analyzed [38–40]. These publications 

document that we did not give up our efforts, though I became 

aware on a trip to Spetses in 1979 that Fred Sanger’s group in 

Cambridge was about to publish the full sequence of human 

mtDNA [41,42]. But we consoled ourselves that we were two 

people that had started in ignorance, and that they were by 14 

people. We were satisfied, however, that the work of Dirheimer 

and his colleagues on yeast mt tRNA or ours on locust mt tRNA 

had been extended to human mitochondria. This knowledge on 

the human mitochondria later formed a basis to recognize the 

many diseases caused by mtDNA mutations. 

Yeast tRNA Genes and Ty Elements 

Once we had learned that even eukaryotic tRNA genes occupy 

more genomic space than prescribed by the structural part, we 

became interested in the problem, how many genes do code for a 

particular tRNA species in yeast and how are these genetic 

entities arranged within the genome? Two organisms had been 

studied thus far in some detail: E. coli [43] and Xenopus laevis 

[44]. For E. coli, it had been established that tRNA genes 

sometimes are encountered as singular transcriptional units but 

that the majority of them were found to be arranged as 

multimeric transcriptional units, either in polycistronic entities, 

or interspersed into ribosomal RNA transcriptional units. For 

eukaryotes it was known that tRNA genes occur at a higher 

redundancy than in prokaryotes. Particularly in Xenopus oocytes 

gene redundancy can amount to several hundred copies per tRNA 

species. Moreover, it had been demonstrated that here the tRNA 

genes together with non-transcribed spacers form clusters of 

serially repeated sequences. Our initial experiments using tRNA- 

DNA hybridization indicated the occurrence of B10 gene copies 

for some of the major tRNA species from yeast tested [45], but I 

was mislead to interpret our results as several (isogenic or nonisogenic) 

copies being clustered like in Xenopus [44] or in 

Drosophila [46]. A more reliable statement was obtained by 

measuring the lengths of transcriptional units of tRNA genes by

296 

H. FELDMANN 

the UV-light technique [47]. Clearly, the transcriptional units 

were no longer than 200 bp for almost all of the 13 tRNA species 

tested and we concluded that these tRNA genes are scattered 

throughout the genome in singular entities. In 1978, we began 

to investigate the genomic organization of yeast tRNA genes at 

the molecular level. This became feasible only when we had 

adopted the genetic engineering techniques developed from the 

mid-1970s on. 

The beginning of genetic engineering undoubtedly was the 

successful approach of Paul Berg and his collaborators to show 

that recombinant DNA could be maintained in a host cell [48]. 

I vividly remember a long night session with full moon in the 

courtyard of a monastery at the Summer School 1971 held in 

Erice, where Berg, Sanger, and Tomkins chaired a discussion on 

the three paradigm shifts initiating a revolution in Molecular 

Biology: the discovery and use of restriction enzymes [49–51]; the 

utilization of recombinant DNA; and the necessity for developing 

methods allowing the determination of long DNA sequences, 

which had to follow principles different from the ones applied 

to the sequencing of RNA and became reality in the years to 

follow [52–54]. 

Though since 1972, several methods had been developed for 

cloning and characterization of recombinant DNA molecules, it 

was only after the Asilomar Conference on Recombinant DNA 

[55] that safe and simple procedures and bacterial vehicles could 

be propagated for extensive use of cloning recombinant DNA 

molecules. A big advantage of cloning vehicles based on phage 

lambda was the larger size of DNA sequences that could be 

accommodated. Those latter properties were shared by the 

cosmids, plasmid gene-cloning vectors packageable in phage 

lambda heads. These methods were applied to yeast genes, too. 

The transformation of yeast cells by replicating hybrid plasmids, 

however, was the first successful transformation of a eukaryotic 

cell and marked a break-through for yeast molecular biology 

[56,57]. 

Our first approach to clone various tRNA genes from yeast was 

only semi-selective and largely followed conventional cloning 

techniques, but our aim was clear: we wanted to study the 

structure and genomic environment of these genes and learn how 

they are expressed. Judith Olah coming as a post-doc was to 

initiate this work [58]. Unfortunately, in October 1979 an


unforeseen incision happened: I suffered a stroke and became 

paralyzed on my left body side. This put me back for a very long 

time. Luckily for me, I did not loose speech or mental capacity. 

Though I recovered after some years of physical restriction, at the 

age of 47 I had no longer a chance to look for a position abroad. So, 

I modestly continued the work on tRNA genes with the invaluable 

help of a very nice lab crew [59–61]. 

Once we had collected data for several tRNA genes, we noticed 

that their flanking sequences revealed extended blocks of 

homology, including those for isoacceptors as well as those for 

non-related genes [61]. These similarities were largely detected 

by eye-inspection, as computer-assisted searches at that time 

were not yet available. The occurrence of repetitive elements in 

this context was a novel observation, since all sequences of yeast 

tRNA genes determined by our colleagues did not reach very far 

into the flanking regions. We speculated that the sequence 

elements might be of functional significance for the expression 

(like the conserved internal A and B boxes or the 3u-termination 

signal) or for the process of dispersion of the multiple tRNA gene 

copies over the yeast genome. In 1982, with more detailed 

sequence information, we obtained evidence [62] that most of the 

repetitive sequences represented specimen of the newly detected 

class of yeast Ty1 elements or their dispersed LTR remnants (solo 

delta’s) [63]. Shortly after their discovery, Ty1 elements had been 

shown to mediate DNA rearrangements (e.g. [64,65]), and, in 

accord with their capability of transposition, they could be moved 

to new chromosomal loci into pre-existing Ty1 elements by a gene 

conversion mechanism or be excised from a given chromosomal 

locus leaving behind only one of their deltas. 

In consecutive studies we were able to confirm that the 

5u-flanking sequences of tRNA genes constituted preferred 

integration sites for Ty transposition, and that these were 

localized in region-specific distances upstream of the genes; in 

many cases, multiple integration and excision events were 

documented genome-wide [66–68]. The phenomenon of preferred 

integration of Ty elements upstream of tRNA genes was later 

inforced by other groups investigating Ty-host interactions (e.g. 

[69,70]). Although target site selection is still not well understood 

for this general class of elements, it is becoming clear that Ty 

elements target their integration to very specific regions of their 

host genomes, probably in order to prevent disturbances of cell

298 

H. FELDMANN 

integrity. Targets containing genes transcribed by RNA polymerase 

III (Pol III) were found up to several 100-fold more active 

as integration targets for Ty1 than ‘‘cold’’ sequences lacking such 

genes. High-frequency targeting was dependent on Pol III 

transcription, and the region specific integration was found to 

occur in an upstream window of B700 bp [71]. The pattern of 

insertion was non-random and not distributed equally throughout 

the genome, but periodic, with peaks separated by B80 bp. 

Recently, it has been demonstrated that ATP-dependent chromatin 

remodeling by Isw2p upstream of tRNA genes leads to 

changes in chromatin structure and Ty1 integration site selection, 

and that Bdp1p, a component of the RNA polymerase III 

transcription factor TFIIIB, is required for targeting of Isw2 

complex to tRNA genes [72]. 

Over the years, details on the structural and functional 

organization of the yeast transposons were worked out by several 

groups. The Ty elements became useful models for the epitome of 

retrotransposition, and manifold aspects pertinent to this theme 

are still under study to present. First indications that Ty 

elements represent autonomous genetic entities which direct 

expression of endogenous genes was obtained from experiments 

in Kingsman’s laboratory [73]. Soon it was established that Ty1 

followed a retrovirus-like strategy for the expression of a large 

fusion protein [74]. Concomitantly, a second class of variant Ty 

elements, Ty2, was shown to obey a similar sequence organization 

and expression strategy as the Ty1 class elements [75]. Other 

experiments confirmed that Ty elements transpose through an 

RNA intermediate [76]. The retrovirus-like gene organization in 

Ty1 became also evident from its complete nucleotide sequence 

[77,78]. 

In the Ty1/2 elements, two open reading frames, TYA and TYB, 

comprise sequences encoding the retrovirus-like gag and pol 

proteins, whereby a translational frameshift (in a þ1 mode) can 

occur in the region overlapping TYA and TYB [79] thus producing 

a gag-pol polyprotein. Finally, the minimal site for ribosomal 

frameshifting in Ty1/2 was determined to be a seven nucleotide 

sequence which induces tRNA slippage involving a minor tRNA 

species [80]. This finding rendered an explanation at the 

molecular level as to why the gag versus pol protein precursors 

were produced in a ratio of 20 to 1: translation of TYA was 

stopped at a usual stop codon in this minimal site, while


read-through by frameshifting was limited by the availability of a 

rare tRNA. 

Further evidence for the similarity between the Ty elements 

and retroviruses was provided by the finding that the RNA 

transcript of Ty together with the smaller proteins processed out 

of the precursor proteins by the Ty protease [81] are associated 

with virus-like particles in yeast [82]. In contrast to retroviruses, 

however, these Ty virus-like particles (Ty-VLP) turned out not to 

be infectious and hence are trapped within its host. 

The detailed characterization of Ty3 (a gipsy type retrotransposon) 

revealed [83] that this element also transposes 

via VLPs as transposition-competent particles and exhibits 

translational frameshifting in a þ1 mode. Previous studies by 

the group of Susan Sandmeyer had shown that Ty3 (or its Long 

Terminal Direct Repeats (LTRs), sigma) insertions had occurred 

consistently in a 16–19 bp distance upstream of several tRNA 

genes. 

In this concert, Rolf Stucka identified Ty4 as a new type of yeast 

elements occurring in low-copy number, belonging to the class of 

copia elements and possessing a gene organization and expression 

strategy similar to Ty1/2; Ty4 also integrates into tRNA upstream 

regions [84,85]. However, we found that transposition capability 

was rather low [86]. From a collaboration with Cécile Neuvéglise 

and her colleagues on the genomic evolution of retrotransposons 

in Hemiascomycetous yeasts it appeared that Ty4 is an evolutionary 

recent element [87]. 

The last retrotransposon found in yeast, Ty5, revealed a 

number of features deviant from those of the other Ty elements: 

its preferred target sites were identified to be silent chromatin 

regions, such as origins of replication at the telomeres and silent 

mating type loci [88]. Targeting was found to be mediated by 

interactions between Ty5 integrase and silencing proteins, and it 

was argued that recognition of specific chromatin domains may be 

a general mechanism by which retrotransposons and retroviruses 

determine integration sites [89]. 

The second question, whether there was a transcriptional 

interference between Ty insertions and tRNA genes was 

answered positively by our first experiments using micro-injection 

of various such constructs into Xenopus oocytes [90]: particular 

segments revealed a stimulatory effect on tRNA gene transcription. 

As this constituted a heterologous system, we sought to

300 

H. FELDMANN 

prove our findings in the homologous system. In order to be able 

to identify and quantitate transcripts from an individual gene, 

Robert Krieg and Rolf Stucka synthesized a unique ‘‘artificial 

tRNA gene’’ (SYN2): it was tagged by an intron-like sequence 

that could not be spliced out from its long precursor but otherwise 

behaved like resident tRNA genes [91]. This gene combined with 

various Ty constructs and integrated as a single copy each into 

the yeast genome was used to monitor the transcriptional 

interference between Ty (and segments thereof) and a flanking 

tRNA gene as well as the chromatin conformation of the 

stable transcription complex and its flanking regions [91,92]. 

Figure 4 shows a photograph of my collaborators involved in this 

work at a birthday trip in 1988. 

We observed that there is a modest stimulatory effect (like in 

the majority of regulatory systems in yeast) of Ty or LTR 

insertions upstream of a tRNA gene on its expression in vivo. 

Transcriptional interference between Ty1 insertions and two 

POL III-transcribed genes was later also shown in the cases of 

Fig. 4. People of my lab crew around 1988: Gertrud Mannhaupt, Hans 

Lochmüller, Susanne Mitzel, Robert Krieg, Rolf Stucka, Christa Schwarzlose, 

and Uschi Obermeier.


tagged SNR6 and SUP2 [93]; vice versa, RNA analysis indicated a 

modest tRNA position effect on Ty1 transcription at native 

chromosomal loci. Further, this study revealed that tRNA genes 

exert a modest inhibitory effect on adjacent pol II promoters, a 

result that was confirmed in other experiments [94]. 

The problem of correlating tRNA gene expression and chromatin 

structure is more complex. The data we and others (e.g. Refs 

[91–93], and references cited therein) obtained supported the 

following model: (i) tRNA genes counteract the formation of a 

canonical chromatin structure over a window reaching from 

B30 bp each upstream and downstream. In other words, actively 

transcribed tRNA genes have to be kept free of nucleosomes. 

(ii) The general pattern tRNA genes exhibit in DNaseI digestion 

experiments is a triplet of hypersensitive sites resulting from 

protection of sequences at the A and B box elements and 

accessibility upstream and downstream from the structural gene 

and between the A and B boxes, reflecting the binding of TFIIIC to 

the intragenic promoter and the tight binding of TFIIIB to 

the upstream transcription initiation site (B30 bp in length). 

(iii) Accessibility of this site by TFIIIB is crucial for active tRNA 

gene transcription, so that this region has to be kept in a 

nucleosome-free configuration. (iv) In DNaseI experiments the 

adjacent hypersensitive site(s) indicating canonical nucleosome 

spacing are located B170 bp and B340 bp upstream from the 

initiation start site of actively transcribed tRNA genes. The first 

upstream nucleosome in these instances is found positioned in a 

way as to form a boundary induced by the transcription complex. 

(v) A prerequisite for the induction of such a constellation is that 

the formation of the transcriptional complex outweighs the 

formation of nucleosomes, a situation that preveiled in competiton 

experiments. (vi) Whenever the sequences upstream of a tRNA 

gene are ‘‘favorable’’ to assist this positioning effect, transcription 

is enabled at a normal or even slightly elevated level. In 

‘‘unfavorable’’ cases, however, nucleosomes can be formed over 

these sequences thus exerting a constraint for transcriptional 

initiation. We documented this correlation in a number of 

experiments using appropriate constructs and mapping hypersensitive 

sites and transcriptional efficacy concomitantly [91,92]. The 

highest transcriptional rates were always found in constructs, in 

which Ty elements, delta or tau sequences had been placed into 

‘‘native’’ distances upstream of a tRNA gene.

302 

The Yeast Genome Project 

H. FELDMANN 

In the years between 1985 and 1988, our laboratory work slowly 

developed into a new phase. During their thesis work, Joachim 

Hauber and Peter Nelböck-Hochstetter had used high-molecular 

weight yeast DNA to clone fragments 35–40 kb in length into a 

cosmid vector (pY3030) that W. Piepersberg from the Munich 

Institute of Genetics had kindly provided to us. Both of them felt 

that it was timely to turn to a genomic scale, dissecting the 

organization of genomic entities along whole chromosomes. With 

the help of a technician Peter managed to establish a cosmid 

bank of 3,000 distinct clones, to prepare DNA from these by 

‘‘mini-preps’’ and to fix it onto filters in a slot-blot apparatus 

within 4 weeks. This procedure was repeated a second time and 

we ended up with two ordered cosmid libraries in all representing 

the 12.8 Mb yeast genome at P ¼ 99.99%, that is 12 times the 

genome equivalent. Together with Rolf Stucka, Peter Nelböck 

established a physical map of chromosome II by means of 

‘‘chromosomal walking’’ and succeeded in localizing a variety of 

tRNA genes, Ty elements, and genes for diverse functions. 

Though these results were only published in form of doctoral 

theses, somehow we had sort of a favorable undercover press, 

which caused André Goffeau from Louvain-la-Neuve to ring me 

up in the lab 1 day in summer 1988. He asked me whether we 

would like to contribute to an assessment on ‘‘Sequencing of the 

Yeast Genome’’ he was preparing for the BRIDGE Programme of 

the European Communities [95]. We were enthusiastic about this 

initiative and, together with Yde Steensma from Leyden, 

produced an overview on how to take advantage of ordered 

cosmid libraries for such an ambitious project. It is a pity that this 

assessment to which a number of renowned colleagues participated, 

has never been made publicly available. It contains some 

ideas or predictions which never became reality, but many others 

that did. André tells more details about the difficulties he 

encountered when considering the launching of chromosome III 

sequencing [96], which for a number of reasons had been chosen 

to be the first chromosome to be tackled in 1989. After he had 

solved the bureaucratic obstacles, the enterprise run rather 

smoothly thanks to his untiring effort as well as that of Steve 

Oliver from Manchester as the ‘‘DNA coordinator’’ and the 35 

European colleagues who participated in this project. Gertrud


Mannhaupt, whom I could hire as a senior collaborator, together 

with Irene Vetter, took care of our share in chromosome III 

sequencing but Gertrud finally became the ‘‘good spirit’’ of all our 

further scientific enterprises. We were happy that we received EC 

support for our sequencing contributions, about 5 ECU per final 

base pair assembled as André had promised and accomplished. 

Moreover, the payment was never delayed as soon as the 

sequences had been proven and annotated. The collection and 

assembly of the sequence data had been put in the hands of MIPS 

(the Martinsried Institute for Protein Sequences), that had been 

developed by Werner Mewes at the Martinsried Max-Planck- 

Institute for Biochemistry. He had set up the necessary 

informatic infrastructure which he – sometimes under considerable 

bureaucratic difficulties – was able to offer for the whole time 

of the yeast genome project. As I myself had fun in computing, we 

were in close and most friendly contact all these years, and still 

are after he moved to another university institution near Munich. 

Before the sequencing of chromosome III was started in 1989, 

André had initiated regular meetings of a ‘‘Steering Committee’’ 

that should supervise current and future activities. As soon as the 

first data had been collected, the sequencers and the committee 

met at regular intervals for progress reports and to exchange 

know-how and expertise. We followed the proposal of André and 

Steve to publish coherent regions from chromosome III, whenever 

the analyses had yielded useful information on new genes [97–99]. 

Before chromosome III had been finalized in 1992 [100], André 

and the ‘‘network’’ had to decide how to continue. 

As our initial complete chromosome II cosmid library [101] was 

not found suitable to serve as DNA material to start the project 

(the DNA was from an industrial strain), Rolf Stucka took the 

initiative to develop a new library from the lab strain aS228C, 

which had been given preference in the discussions of the 

Steering Committee. Thus we were happy to be awarded the 

contract for sequencing chromosome II (B820 kb) this time. 

Concomitantly, Bernard Dujon’s laboratory which had prepared 

an ordered yeast cosmid library by means of another vector [102], 

started sequencing chromosome XI (B670 Kb) and finished in 

1994 [103]. Though we were paid as ‘‘DNA coordinator,’’ EC had 

reduced the subsidiary amount allocated for each final bp to 2 

ECU. Luckily for us, the helper in the hour of pecuniary need was 

the German Ministry for Science and Technology (BMFT) which

304 

H. FELDMANN 

raised our income per bp by another 2 ECU. I might insert here 

that our lab largely had to rely on funds (for staff, equipment, and 

consumables) we had to invite from grant-giving institutions. 

While, for example, the Deutsche Forschungsgemeinschaft (DFG) 

in those years was not prepared to subsidize any sequencing 

project, the BMFT was open for this type of funding, because they 

could expect to gain from novel technical developments and to 

draw new insights from initiatives such as the yeast genome 

project. Indeed, it was obvious for anyone that the final goal of 

this project was much beyond establishing the complete sequence 

of a small eukaryote, namely to use this information for 

concurrent or subsequent functional analyses. As a fact, the 

wealth of information obtained in the yeast genome project 

turned out to be extremely useful as a reference against which 

sequences of human, animal or plant genes, and those of a 

multitude of unicellular organisms under study could be 

compared. André has given credit [96] to BMFT’s ‘‘outspoken 

support y (without which) y the EC would not have been 

engaged in sequencing the yeast genome.’’ 

While Gertrud Mannhaupt with three technicians took care of 

the sequencing and the analysis of particular regions of chromosome 

II [104–107] and later those from chromosome XV [108], 

Rolf Stucka was providing cosmid clones to the 18 European 

sequencing laboratories. We used to travel to the internal 

meetings with an 8 meter long detailed map of chromosome II 

to co-ordinate the program and to avoid too much overlapping 

during sequencing. It is remarkable that Rolf found time to follow 

his own projects, one of which was aimed at aspects of sugar 

metabolism – in a nice collaboration with Carlos Gancedo from 

Madrid [109–112]. Before the sequence of chromosome II had 

been solved [113], our lab concentrated on a new theme (see 

below). None the less, we eagerly followed the rapid developments 

in sequencing the many chromosomes that were left and 

brilliantly managed and brought to success by André Goffeau 

within the next 2 years [114–116]. As André Goffeau mentioned 

[96] he succeeded ‘‘to keep a collaborative spirit in the international 

yeast-sequencing community’’ by arranging the full 

participation of some American groups (particularly with the 

help of Mark Johnston) as well as that of Howard Bussey’s, Bart 

Barrell’s, and Peter Philippsen’s groups to the project. The initial 

aversion of the American scientific community to engage in an


international network for sequencing the yeast genome was made 

clear to me during the 1990 Cold Spring Harbor Meeting of 

‘‘Genome Mapping and Sequencing’’ to which I participated. 

André had asked me to present our intentions in a short ‘‘free’’ 

communication, but the organizers curtly refused. Only Dr Ito, a 

Japanese who worked at the Max-Planck-Institute for Genetics in 

Berlin and who chaired this session, was kind enough to allot 

5 minutes for my presentation. 

The sequencing project was followed by several collaborative 

initiatives for functional analyses of the wealth of novel genes 

that had been detected. We participated in a German network 

chaired by Karl-Dieter Entian from Frankfort [117], and I was 

able to document our continuous affiliation with the yeast 

projects in separate chapter of three books [118–120]. 

Yeast 26S Proteasome and Triple A Proteins 

In connection with our work on Ty elements and the yeast 

genome sequencing project we became interested in yeast 

transcription factors. By looking into the TATA box binding 

factor, Rolf Stucka found that the protein sequence had a 

bipartite structural symmetry – probably the consequence of an 

ancestral gene duplication – that helped explain its saddle-like 

structure [121]. The theme ‘‘transcription factors’’ became a 

major issue when I got involved in setting up a special DFG 

program devoted to ‘‘Factors and Mechanisms of Gene Activation’’ 

(see below). In this context, we wanted to search for new 

factors involved in Ty expression. Peter Nelböck had just 

published a paper [122] in which he reported on a putative novel 

protein interacting with the human immunodeficiency virus tat 

transactivator, called TBP-1 [122]. As HIV in many respects 

resembled Ty, we asked for a probe to search for similar factors in 

our yeast cosmid library. Already the first experiments Rolf 

Stucka undertook in 1992 were successful: a dozen different 

clones could be isolated and sequenced by our lab crew. To our 

surprise, the most prominent feature of the encoded proteins was 

a highly conserved domain containing nearly 200 amino acids, in 

one or two copies, encompassing two sequence elements characteristic 

of a novel type of putative ATPases. The first 

observation of this new category of proteins and its designation

306 

H. FELDMANN 

‘‘AAA proteins’’ stems from a paper of the laboratory of Wolf 

Kunau [123]. They noticed that the novel motif occurred in three 

groups of proteins fulfilling different cellular functions, such as 

Sec18p and NSF; Cdc48p, p97, and VCP; and TBP-1 (quotations, 

see Ref. [124]) and argued that ‘‘these proteins are members of a 

novel family of putative ATPases and may be descendants of one 

common ancestor.’’ Indeed, within a short time this family was 

found to be more widespread than initially thought and 

comprised a large variety of members in both eukaryotic and 

prokaryotic organisms. The first international conference on AAA 

proteins was held in Gif-sur Yvette in 1995 and the name for this 

family accepted by the community. 

When we published our results, further members of the AAA 

family had been characterized ([124]; and references cited 

herein). From comparisons we inferred that our collection 

specified four members of the yeast 26S proteasome, Yta1p, 

Yta2p, Yta3p, and Yta5p, whereby Sug1/Cim3 (as well as Cim5/ 

Yta3) had been discovered [125] during our work on this subject; 

the sixth yeast proteasomal AAA protein was recognized as Sug2 

[126]. In order to keep a unified nomenclature, these AAA 

ATPases, meanwhile identified as subunits of the regulatory 

particle of the 26S proteasome, were called Rpt1 through Rpt6, 

whereas the non-AAA proteins were designated Rpn1–14 [127]. 

Likewise interesting became the Yta10, Yta11, and Yta12 

proteins, which were highly similar in structure and also revealed 

the presence of the novel ATPase domain but bearing additional 

modules. Homologies with ftsH/hflB from E. coli were obvious as 

well as sequences typical for mitochondrial ‘‘matrix-targeting 

domains.’’ These and further notions led us to propose [124] ‘‘that 

Yta10, Yta11, and Yta12 may represent subunits of a proteasomelike 

complex in yeast mitochondria. This complex might be 

similar and evolutionarily related to the cytosolic 26S protease 

and thus constitute a mitochondrial proteolytic system in addition 

to the one that bears similarity to the Lon proteases y’’ Reimund 

Tauer engaged in this subject [128] and – in a fruitful cooperation 

with Thomas Langer and his group [129] working in Walter 

Neupert’s department – it was established that these three AAA 

proteins formed a novel ATP-dependent complex in the inner 

membrane of mitochondria with proteolytic and chaperone-like 

activities [130], whereby the proteolytic activity was found to be 

exerted by a Zn-dependent metallo-protease module linked to the


AAA domain. Thomas Langer and his collaborators took care of 

the further characterization of this ‘‘mitochondrial quality 

control system’’ [131] and since worked out a wealth of important 

aspects in the biogenesis of mitochondria [132]. Together with 

Thomas, we organized the second international workshop on 

Triple A proteins in 1997 at Tutzing (upper Bavaria) supported by 

EMBO. 

The S. cerevisiae genes encoding the proteasomal entities are 

single copy throughout, and the majority of them are essential for 

cell viability. As I was convinced that the yeast ubiquitinproteasome 

system (similar to the ribosome) fulfils the requirements 

of a regulatory network, in which expression of the single 

genes is co-ordinated, we followed this issue. Again, it turned out 

a justified speculation. First, we detected a unique upstream 

nonamer box (GGTGGCAAA) which we called PACE (proteasome 

associated control element) to occur in the promoter regions of 28 

out of the 31 proteasomal genes, including the RPT genes and 

most of the RPN (non-AAA) genes, and in B50 further yeast 

genes involved in the ubiquitin-proteasome pathway as well as in 

genes mediating diverse regulatory functions in yeast. We set out 

to identify the cognate DNA binding factor and were surprised to 

come across Rpn4, a subunit of the 19S regulatory cap. The 

structure and the role of Rpn4 to act as a transcriptional activator 

was substantialized in a number of experiments [133], for which 

we took advantage of a reporter system we had developed many 

years before [134]. Unfortunately, after this seminal contribution 

I had to quit my lab due to age. 

In the following years, it was demonstrated that Rpn4 links 

base excision repair with proteasomes [135]. Further ingenious 

studies by Alex Varshavsky’s lab revealed that Rpn4p is a ligand, 

substrate, and transcriptional regulator of the 26S proteasome 

and exerts a negative feedback control (e.g. [136–139]). RPN4 

appears to be under control of several stress factors, such as 

Yap1p and Pdr1/3p [140]. On the other hand, Rpn4p strongly 

mediates the cell’s adaptation to arsenic-induced stress as 

revealed by expression profiling [141]. Filamentous-form growth 

is controlled by many modules in an integrated network, in which 

the proteasome system is probably integrated through Rpn4 

[142]. Also in mammals and Drosophila (for review [143]), a 

similar system appears to be operative, but no molecular details 

are available as yet.

308 

H. FELDMANN 

Administration and Teaching Medical Students 

In 1967, when we had moved to Munich, for some period of time I 

was to take care of all kinds of administrative business: purchase 

and maintenance of equipment, accounting, etc. One task that 

met my interests was to keep liaison with the university building 

department, which had begun to raise an eight-storey new 

institute in our courtyard that should offer more space to the 

two new chairs of biochemistry (see Ref. [144]) and physiology. 

Theodor Bücher, who initially was the only head of the institute 

of physiological chemistry, in his negotiations with the Bavarian 

Ministry of Science had accomplished that three new departments 

were created. Though we could improve some internal facilities 

during the construction of the new building (such as lab 

equipment, isotope labs) we were not able to prevent serious 

mistakes that are still persistent despite many necessary and 

costly repair over the years. For my part I was happy to move into 

a new lab and a well-equipped isotope lab in 1971. 

All the years I worked at the Institute of Physiological 

Chemistry in Munich, I had to look after the training of medical 

students at several levels. These teaching obligations and the 

administrative loads connected with them were greatly put on the 

shoulders of the ‘‘younger’’ staff, because most of the professors 

preferred to concentrate on giving the general lectures. 

One of my first activities was to organize a serious practical 

course in biochemistry, setting up reasonable experiments. The 

most intriguing problem was to develop a fixed schedule to hurry 

so many students through these venues within the minimal time 

of four terms: logistics had to consider subject and number of 

students as well as time, space, and staff available, in other words 

the time-table had to be strictly non-overlapping for each 

individual. Whoever has encountered such a demand, knows 

quite well that it can hardly be solved by computer programs. 

With my colleague Joachim Otto from the neighboring department 

we managed to develop a ‘‘master-plan’’ which was followed 

through 25 years. In 1974, the faculty entrusted me with the coordination 

of pre-clinical education and I had to chair the 

meetings of a committee consisting of representatives from every 

discipline, university administration, and examination board. In 

the beginning this task took much of my time, but in the end the 

meetings had to take place only twice a year.


Analytica 

In 1977, I was elected member of the Scientific Committee for the 

bi-annual Conference of Bioanalytics held in Munich, connected 

to a world-wide respected trade fair in biochemical, biomedical, 

and bioanalytical instrumentation, the Analytica. I was then 

nominated to take care of the formalities connected to the award 

of a special prize Biochemical Analytics (later: Molecular 

Bioanalytics), and to act as its Secretary. The prize was 

inaugurated by a donation from Boehringer Mannheim GmbH 

and later continued by Roche Diagnostics GmbH. The statutes 

specify that the prize should be awarded for outstanding work in 

the field of molecular bioanalysis, that is for the development of 

novel methods and for new scientific contributions to the 

advancement of molecular and biochemical analysis in diagnostics 

and therapy. The awardees are nominated by a prize committee 

consisting of personalities representing the Gesellschaft für 

Biochemie und Molekularbiologie, the company, and the biochemical, 

biophysical and medical research, and the prize is 

handed over every 2 years at the Analytica Conference. For many 

years, when the German Society for Clinical Chemistry was the 

body scientifically responsible for all enterprise connected to the 

Analytica, the prize winners and some hundred guests were 

invited to Bayerischer Hof in Munich to celebrate the event 

(Figure 5). 

I am delighted that the list of the Prize winners (Table 1) names 

eminent researchers, who indeed made outstanding contributions 

to the fields specified above. A satisfaction for all those colleagues 

that participated in the selection of these persons was that in 

several cases this Prize was given to the awardees well in advance 

of the Nobel Prize. At the same time, it documents some of the 

highlights in the development of molecular biology. 

FEBS 

I became aware of FEBS (Federation of European Biochemical 

Societies) through their early meetings held in Warsaw (1966), 

Oslo (1967), and Prague (1968). These meetings provided 

excellent opportunities for a ‘‘beginner’’ to follow novel

310 

Fig. 5. Analytica 1982: With my wife at the Analytica reception. 

H. FELDMANN 

developments in biochemistry and molecular biology and to 

present his own results in short talks. 

In 1983, I was fortunate to be nominated by the Gesellschaft für 

Biologische Chemie (GBM) for membership in the Advanced 

Courses Committee (ACC) of FEBS. I took this post with pleasure, 

because I felt in absolute agreement with the main objective of 

FEBS that since its foundation in 1964 has been ‘‘to advance basic 

research and education in biochemistry, molecular and cellular 

biology, and molecular biophysics on a European level.’’ My 

predecessor as chairman of the ACC was Giorgio Bernardi, who 

succeeded in continuously raising the number of courses held per 

year from only a few in the beginning to more than a dozen, 

before he had to retire from this duty in 1986. I remember my 

first participation to an ACC meeting that was held on the 

occasion of a lecture course in Maria Alm (Austria) on the 

Biochemistry of Ageing organized by Fritz Cramer and Brian 

Clark in 1984. In 1986, FEBS Council appointed me to become 

Giorgio’s successor. I took up this office in 1987 and was 

reappointed for two further 3-year periods in 1990 and 1993. 

I was lucky to work with a Committee the members of which 

were enthusiastic in contacting colleagues from all over Europe,


TABLE 1 

Awardees of the Prize Biochemical Analytics/Molecular Bioanalytics 

1980 Fred Sanger and Alain 

Coulson, Walter Gilbert 

and Alan Maxam 

1982 César Milstein and George 

Köhler 

Development of modern 

techniques for sequencing 

DNA 

Development of monoclonal 

antibodies 

1984 Ed Southern Development of the DNA 

hybridization technique: 

1986 Brigitte Wittmann-Liebold, 

Leroy Hood and 

M.W. Hunkapiller 

‘‘Southern blot’’ 

Development of methods and 

instrumentation for 

sequencing proteins at a micro 

scale 

1988 Charles Cantor and 

Development of pulsed field 

David Schwartz 

electrophoresis 

Sir Alec Jeffreys DNA fingerprinting 

1990 Karin Mullis and H.A. Erlich Development of the PCR 

1992 Jean Lawrence and 

David Ward 

technique 

Development and applications of 

non-radioactive highly 

sensitive in situ hybridization 

techniques 

1995 Gregory Winter Isolation of high affinity human 

antibodies directly from large 

1996 Mario Capecchi and 

Rudolf Jaenisch 

1998 Arthur B. Pardee (Dana- 

Farber Cancer Institute, 

Boston) and Peng Liang 

(Vanderbilt Cancer 

Center, Nashville 

Tennessee) 

2000 Franz Hillenkamp (Institut 

für Medizinische Physik 

und Biophysik, Münster) 

and Michael Karas 

(Institut für Chemie, 

Frankfurt/Main) 

synthetic repertoires 

In recognition of their 

pioneering work on the specific 

integration of DNA in 

mammalian cells and for 

establishing transgenes as a 

basic tool for research in 

molecular biology and 

medicine 

Development and applications of 

Differential Display 

Development of Matrix-Assisted 

Laser Desorption/Ionization 

Mass Spectrometry of 

Biopolymers, one of the 

essential technologies in 

genomics and proteomics

312 

H. FELDMANN 

who would be willing to run a FEBS Course. Though the funds for 

FEBS Courses were raised to 1 Mio Deutsche Mark per annum, we 

had to set certain limits for the amount of money given as a 

support to each course. So it was highly appreciated if organizers 

were able to invite co-sponsorship from other grant giving 

institutions. One particular advantage of running a FEBS Course, 

however, was that Youth Travel Grants were provided to assist 

attendance at these by younger scientists. As half of the FEBS 

Courses budget was designed for this purpose, up to 25% of the 

participants in a lecture course and all of the participants in a 

practical course could profit from this type of support. In 

accordance with FEBS’ general policy, fellowships were preferably 

awarded to young scientists from Eastern European countries, who 

otherwise would have had little chance to receive funds from their 

national institutions. Another aspect connected to this issue was 

that the ACC sought to invite colleagues from these countries to 

organize FEBS Courses at their home institutions, an encouragement 

that in fact paid out successfully. During my time as 

chairman, the ACC consisted of 10 members: eight colleagues from 

different Constituent Societies as well as the FEBS Secretary 

General and the FEBS Treasurer. This arrangement has been 

kept, but fortunately more colleagues from former Eastern 

countries became members of the ACC since. In all these years, 

the ACC received enough applications to sort out inappropriate 

ones. Priority was given to practical courses, because the 

committee felt that this type of venue would be of greatest benefit 

to young researchers who had no other opportunities to experience 

novel laboratory techniques or to learn techniques, which they 

wanted to apply in new projects. Thus the practical courses 

complete the intentions of the FEBS fellowships’ program. Indeed, 

some of the practical courses were so successful that the organizers 

and the ACC decided to repeat them, sometimes in subsequent 

years or in a series. I gratefully recollect that for one particular 

course the main organizer [Wilhelm Ansorge from EMBL 

(European Molecular Biology Laboratory)] repeatedly undertook 

to transfer all special equipment and instruments needed for this 

course to a place that had no supplies of this kind. The significance 

of the Advanced Courses Programme is also documented by the 

fact that students themselves, the Young Scientists movement, 

took the initiative to organize a successful series of courses entitled 

‘‘Young Scientists’ view of molecular biology and biotechnology.’’


Extraordinary venues I remember were two FEBS courses 

organized by Giorgio Bernardi, one in Cairo and one in Harare, 

Zimbabwe. He felt that students from African countries should 

profit from recent developments in Europe. In conjunction with 

these courses Giorgio arranged for private tours with some 

colleagues (I think he was fond of traveling). As I shared his view 

to spend a few extra days on ones own expenses to see a country 

whenever one had an opportunity to visit it, I undertook an 

unforgettable tour from Cairo to Upper Egypt with Marianne 

Grunberg-Manago. In Harare, Marianne, Brian Clark and his 

wife, the wife of Francois Gros, and I rented an old Peugot and 

went cross-country for 10 days. 

Personally, I am most grateful to FEBS that they have 

supported the Spetses Summer Schools on Molecular Biology 

(see below) from 1983 to present in co-sponsorship with NATO 

(North Atlantic Treaty Organization) and EMBO, and finally 

have decided to give full financial aid to these well-known venues 

together with EMBO. The years in FEBS were always exciting 

and enjoyable. I was glad to meet and to work with so many nice 

and enthusiastic colleagues from so many different countries, 

above all the members of the Executive (Figure 6) and the ACCs 

(Figure 7), but not to forget, the organizers of the FEBS Courses 

and the numerous student participants at courses which I had a 

chance to attend. I vividly remember the splendid and jovial 

atmosphere at the Committee meetings governed by hospitality 

and friendship and many exhilarating episodes that occurred at 

these occasions. At least I cannot repress one that happened when 

the ACC met in Amsterdam (Figure 7) and tell it with our host’s, 

Karel Wirtz from the University of Utrecht, own words [145]: ‘‘It 

also gave me a chance to make the committee members familiar 

with the capricious nature of Dutch wind and water. Having been 

asked to organize the meeting in Amsterdam a 60-feet sailing 

barge of the early 1900s was chartered (the whole enterprise 

turned out to be less expensive than a hotel). This ship offered a 

bunk for each member and a spacious room under deck where we 

could discuss and review the applications. Arrived on Friday late 

afternoon we sailed from Amsterdam harbour the next morning 

to cross the Ijsselmeer. During an 8-hour sailing trip we finished 

the agenda while a two-men crew made sure we reached the port 

of Hoorn at the north side of this large body of water. On Sunday 

morning we had to hoist the sails again to return to Amsterdam.

314 

H. FELDMANN 

Fig. 6. FEBS Executive Committee Meeting, Budapest 1990. Sitting: Prakash 

Datta and Doria Cavallini. Standing: Horst Kleinkauf, John Mowbray, Karl 

Decker, Guy Dirheimer, Vito Turk, Horst Feldmann, and Carlos Gancedo. 

On our way back, the wind had picked up to force 7 while heavy 

showers tested our endurance, Horst being in the kitchen to 

prepare lunch, lost his balance to become encased knee deep in 

macaroni. Somehow he still managed to produce a tasty meal.’’ 

He forgot to mention that the ship just went about without a 

warning. It was a pleasant feeling to know that the ACC got in 

best hands in 1996 with Karel as my successor. 

In a way, I miss all these activities, but I am grateful that despite 

my retirement I still have an opportunity to keep contact with 

many friends from my time at FEBS. In 1995, I was awarded the 

FEBS Ferdinand Springer Lecture Tour that offered the splendid 

opportunity to visit a number of Institutes around Europe and to 

talk about my current scientific work. Since 1995 I am acting as an 

Editor to FEBS Letters, the journal founded in 1968 by Prakash 

Datta for rapid publication of relevant novel reports and reviews 

in biochemistry, biophysics, molecular biology, and related topics; 

Giorgio Semenza, then Managing Editor of FEBS Letters, brought 

me in. In 2003, FEBS gave me an opportunity to finish a book 

Forty Years of FEBS – 1964 to 2003 – a Memoir [145], which was


Fig. 7. ACC Meeting, Amsterdam 1992. On a sailing tour on the IJsselmeer. 

From left to right: Julio Celis, Horst Feldmann, John Mowbray, Slobodan 

Barbaric, Vito Turk, Paulette Vignais, and Thanos Evangelopoulos. 

presented at the 40th Anniversary of FEBS celebrated at the 

FEBS Congress 2004 in Warsaw. 

Spetses Summer Schools 

The Greek island of Spetses is the place I had a chance to visit 

more often than any other one in beautiful Greece; in fact, I can 

call it ‘‘my second home.’’ I immediately fell in love with the 

island when I was admitted as a participant to the 3rd NATO 

Advanced Study Institute held there in 1969. 

Actually, Marianne Grunberg-Manago had started this series of 

International Summer Schools on Molecular Biology in 1966 and 

through her untiring initiative [146] the Schools have been kept 

as a series of well-known annual lecture courses until to date. 

There was a college (Anargyrios and Korgialenios School) large 

enough to accommodate students and a hotel at a short distance 

from it both of them adjacent to a good beach. The island was 

small enough to facilitate contacts between students and 

professors and it was large enough to provide peace and quiet.

316 

H. FELDMANN 

Fig. 8. Participants to the Spetses Summer School on Molecular Biology in 

1992. 

My active involvement in the Spetses Summer Schools dates 

back to the year 1970/1971. Hans Zachau was asked to organize 

the third School in order to bring in the Germans as a third party 

and to make the School an annual event, the directorship being 

rotated regularly between France, England, and Germany. (For 

some interval, the directorship has included Tom Caskey 

(Houston) and John Hershey (Davis) from the US to reach a 

4-yearly rotation.) From then on I helped Hans Zachau as one of 

the German co-organizers, and after 1988 I took over as a German 

organizer (Figure 8). 

Initially, the Spetses Summer Schools were sponsored exclusively 

by NATO. But their strict rules soon caused the organizers 

to apply for further grants from EMBO (sponsor since 1972), and 

also from FEBS (sponsor since 1983), as to allow to invite and 

support lecturers and students from non-NATO countries. In 

recent years, the School relied on financial support from EMBO 

and FEBS only. It may well be that some colleagues were 

skeptical that Spetses had been established as sort of a ‘‘club’’ and 

should not be funded. But the facts show that over the years 

nearly 500 (different!) renowned lecturers came to the island to


teach some 5,000 young pre- and post-doctoral researchers. 

Indeed, one can realize until to date that Spetses participants 

form a community, and I am still in contact with some of the 

students (particularly Russians) who attended the School several 

years back. To give an account on the Spetses’ activities, I 

compiled an overview at a website [147]. The topics dealt with in 

these venues reflect important moments in the development of 

molecular biology over the last 40 years. Thus, the intentions of 

these courses of familiarizing young researchers with novel 

insights and recent advances in this field completely met those 

of the grant-giving institutions. 

In 1996, I arranged for a 3-day Workshop to celebrate the 30th 

Anniversary of the Summer Schools at Spetses and was able to 

invite former organizers and lecturers of outstanding merit. As 

the authorities and the inhabitants of Spetses were proud and 

most grateful that they were selected to host an International 

Course of this calibre for so many years, all previous organizers 

were presented with a brass plaque by the Spetses Mayor to 

affirm their honorary citizenship. A similar venue was repeated 

successfully to celebrate the 40th Anniversary in 2006, organized 

by Brian Clark and myself. It was again a wonderful occasion that 

so many old companions were to meet many of whom had not 

seen each other for nearly 40 years. As I promised to Marianne, 

when she fell seriously sick, as long as I could to have an eye on 

Spetses and to engage in keeping the tradition of the School. 

Gene Technology 

I have already briefly recapitulated the story of recombinant 

DNA. In 1971 Cetus and in 1976 Genentech companies were 

founded. The first pharmaceutical products based on recombinant 

DNA were somatostatin (1977); insulin (1978); growth hormone 

(1979); and interferon (1980) (e.g. [148]). Also Suisse scientists 

and industries engaged early in gene technology. In Germany, 

however, the application of gene technology for industrial 

purposes lacked behind considerably, though we had little 

problems to get permission for cloning in the lab after the 

German guidelines had been worked out and published. For 

industry, there were two reasons for reluctance: (i) at the 

beginning, the bosses leading big German chemical or

318 

H. FELDMANN 

pharmaceutical firms publicly stated ‘‘y there is no need to 

engage in this doubtful enterprise, if it turns out promising, we 

will buy the know-how y’’; (ii) later, the first attempts to set up 

an industrial production (e.g. for human insulin in recombinant 

bacteria) failed as German authorities voted down a bill. In all 

honesty I have to say that fortunately the situation in 

biotechnology has changed thanks to the engagement from 

politics, research and industry, so that these days biotechnology 

has a good and respected standing world-wide. 

As I felt (fortunately not being the only one) that it was timely 

around the early 1980s to familiarize at least those people 

interested in the ‘‘chances and risks’’ of gene technology with the 

new developments, I accepted several invitations to discuss the 

relevant items with chemists, geneticist, pharmacists, 

medical doctors or even ‘‘laymen,’’ at congresses or privately 

(e.g. [149–151]). In 1981, German industry no longer could deny 

that gene technology was attractive. But except a few smaller 

companies, who showed a growing interest in adapting novel 

techniques, there was no sincere attempt from ‘‘in-house’’ to 

train their employees. Rather came an impetus from the 

Gesellschaft Deutscher Chemiker, who asked me in 1982 to 

organize an advanced vocational training course on ‘‘Methods 

and Results of Gene Technology’’ for some 20 participants at our 

institute. I could solicit the help of some of my junior colleagues 

(Fritz Fittler, Urs Hänggi, Peter Philippsen, Rolf Streeck, and 

Wolfgang Wintermeyer), but generous funding allowed me to also 

invite foreign lecturers. This course was offered and successfully 

repeated 3 times in the years after, until 1985. A remarkable 

feature was that there was a growing interest of patent attorneys 

in these courses. A number of large and well respected offices had 

been established in Munich, whose clientele recruited from 

renowned biotechnical companies world-wide, the reason being 

that the European Patent Office had been installed in Munich in 

1977. The contacts brought about by the courses stimulated 

several of our young doctoral students to start a career as patent 

attorneys as well (in all about 10). 

The courses on gene technology in Munich had raised the 

particular interest of Boehringer GmbH, who wanted to set up a 

similar advanced vocational training course for their staff, which 

we called ‘‘Novel Methods in Gene Technology.’’ With my good 

colleagues and friends Wolfram Hörz and Gustav Klobeck both


from our institute, we managed to familiarize the participants 

with the latest practical and theoretical developments in 

molecular biology and genetic engineering in 3-day courses. I 

remember we started this series in 1985 and repeated these 

annual venues at least 10 times, until 1995. These venues had 

arisen from the good relations we had with some influential 

colleagues from Boehringer Company. The enterprise was 

abruptly stopped when the firm was taken over by Roche to 

become ‘‘Roche Diagnostics’’ in 1998. 

A very pleasant cooperation with the German Society for 

Animal Husbandry was launched by Horst KräuXlich, the director 

of the Munich Institute of Animal Breeding. As the members of 

Deutsche Gesellschaft für Züchtungskunde (German Society for 

Animal Breeding) had a growing interest in learning the 

capabilities of recombinant DNA technology and had heard about 

our courses, they invited us to run a lecture course for them in 

1986. It was really satisfying to see that our seeding message was 

to bear fruit in the years to follow. The animal breeders 

successfully combined micro-injection of recombinant genes with 

in vitro fertilization to pigs, cattle, sheep, and other domestic 

animals [152]. The DFG supported these activities by generous 

grants in a special program, whereby the applications of the single 

members had to be evaluated bi-annually and the forthcoming 

results to be presented in common sessions. It was always a 

pleasure for me to be accepted by these colleagues as an 

‘‘honorary animal breeder.’’ 

Sonderforschungsbereich 190 

The DFG supported the research of our group during my entire 

time. In 1989, members of the Munich scientific community 

interested in mechanisms of gene activation sought a closer contact 

with each other. We wanted to build a forum that should foster a 

practical cooperation and a direct exchange of results and ideas and 

should overcome the disadvantage that the respective institutes 

were scattered throughout Munich. This was exactly the concept of 

the Sonderforschungsbereiche (SFBs), quite a number of which 

existed all over Germany. Prerequisites for funding were to present 

a timely and attractive theme and the willingness of up to 20 groups 

to cooperate for better or worse. The application presented to DFG

320 

by Herbert Jaeckle, then the new head of the Munich Institute of 

Genetics, Wolfram Hörz from our institute and myself was accepted 

and financing started in 1990. The responsibility for smooth rolling 

was laid in my hands as chairman of SFB190 in consecutive 3-year 

periods (from 1990 through 1998), and Wolfram Hörz was elected 

my successor for the fourth period (the maximal life-time for an 

SFB, until the end of 2001). An ongoing problem was to catch the 

attention of new groups whenever current members dropped out. 

Many of the group leaders left Munich as they had been offered 

chairs in other universities or directorships at foreign Max-Planck- 

Institutes, and in order to keep a ‘‘critical mass’’ these groups had 

to be replaced by newcomers. But we succeeded to solve this 

problem as Munich was an attractive place which helped recruitment 

of capable colleagues. 

Other Encounters 

H. FELDMANN 

Before I engaged in committees, boards, or in the organization of 

international meetings myself, I was attending several venues 

Hans Zachau had initiated and organized as chairman [144]. 

Together with members from the Russian Academy of Sciences he 

started the series of German–Russian Symposia, with the first 

bilateral conference held in Munich in 1976. The continuation of 

these venues for over 20 years opened the extravagant chance for 

me to visit several interesting places in Russia and the former 

Soviet Republic. On the Russian side, the symposia were 

organized by scientists from the Engelhardt Institute of Molecular 

Biology in Moscow. The meetings always started with a 

short stay in Moscow and from there the participants were taken 

by air plane to the various locations the organizers had chosen. 

Every time Tatiana Venkstern, a scientist from the Engelhardt 

Institute, was responsible to take care of us and to act as our 

motherly travel marshal. All these visits (Zagorsk, Erevan, and 

Armenia, 1981; Irkutsk and Lake Baikal, 1989; Suzdal, Vladimir, 

and Riga, 1993), but also the friendship with Tatiana, left 

unforgettable moments in my life. Through these tours, I realized 

how difficult life for the majority of the population was those 

years and what I could personally do to alleviate the frustrating 

situation our Russian colleagues had to struggle with. One late 

compensation consisted in the possibility to raise money from the


German Volkswagen-Foundation to start a fruitful cooperation 

with Vadim Karpov’s group from the Engelhardt Institute in 

Moscow [133]. 

My engagements in immediate duties, in faculty or university 

affairs have been mentioned above. Commitments at the national 

level included services to the DFG (as a member of many panels of 

experts), the Alexander v. Humboldt-Foundation and the German 

Studienstiftung. I always kept good relations with EMBO after I 

had been elected EMBO member in 1979. My activity for the 

German Society for Biochemistry and Molecular Biology (GBM) 

largely concentrated on collaboration with them during my time 

as Secretary for the Analytica. But I am proud that the GBM logo 

I designed for them 30 years back is still in use. 

What concerns my international commitments, not mentioned 

above, I acted for some years (1993–1997) as an advisor for 

the German-Israeli Cooperation Programme in Biotechnology 

(DISNAT) maintained on the German side by the Ministry of 

Research and Technology. Very pleasant memories are connected 

to many events I was invited to by my French colleagues, better to 

say friends. They had noticed that I was able to follow lectures 

and discussions in their own language. This permitted me to 

serve as examiner for several thèse de troisième cycle and 

qualification pour l’enseignement supérieur at University Louis 

Pasteur in Strasbourg, or in evaluations of CNRS programs at 

Gif-sur-Yvette. When the French yeast community started 

their Génolevures program, a project aimed at the ‘‘Genomic 

exploration of the Hemiascomycetous Yeasts,’’ I was happy to help 

Jean-Luc Souciet from Strasbourg who is acting as a coordinator 

of the Génolevures project [153]. Since then I have been 

invited to their meetings in Paris or Strasbourg, which I consider 

a very selfless gesture supporting our long-lasting friendly 

relationships. 

Epilogue 

Contrary to many colleagues whose personal recollections I read 

with great interest, I have to confess that I never enjoyed the close 

guidance by teachers belonging to particular academic schools. 

This does not exclude that being trained as a chemist I never gave 

up to look at the world in a sense of chemistry – at least with one

322 

H. FELDMANN 

eye. This perception was intensified by several chances to meet 

eminent scientists at the Lindau Nobel Prize Meetings during my 

time at university. My wife and I still keep good memories of the 

4th Meeting of Awardees in chemistry in 1961. A treasure for me 

is the reprint of a remarkable essay by J.H. van’t Hoff explaining 

the value of ‘‘Imagination in Science’’ which my scientific 

‘‘grandfather’’, Richard Kuhn, had edited and handed out to the 

students [154]. 

From my childhood on I was used to grasp and accept what I felt 

was interesting or suitable for me, be it knowledge, reflections, 

experiences, skills, or obligations. So I am deeply thankful to all 

people who have contributed to my development in one or the 

other way, in more or less intense connections. This does not 

imply that I was always prepared to adopt their opinions or way of 

action, particularly of those who in my eyes displayed attitudes 

unacceptable for me. 

Of course, my gratitude includes all my gifted and motivated 

co-workers, some of whom I cited by names above, but also those 

who have not been mentioned explicitly. Our lab always was an 

assembly of chemists, biologists, medical students, and technicians, 

individualists originating from regions all over Germany. 

More splashes of color were contributed by people coming from 

various nationalities for different but often overlapping periods of 

time. In this respect I remember: Hermann Falter, a Bavarian- 

Canadian postdoc; Erwin Meixner, an Austrian PhD student; 

Giuseppe Pirro (Pino), an Italian post-doc from Modena; Antonin 

Eigel, a Czech immigrant; Anni Fradin, a student from Bretagne 

who lost her supervisor in Gif-sur-Yvette and worked for her 

dissertation in my lab on tRNAMet and methylation in yeast; Noel 

Deering, an Irish technician; Rick Baker, an American post-doc 

coming with a Humboldt stipend; Pamela Smith, a British 

secretary fluent in German, Italian, and Spanish; and Vadim 

Karpov with Olga Preobrashinskaia and two students, all coming 

from the Moscow Engelhardt Institute to collaborate with us 

on Rpn4. 

The same gratitude applies to all colleagues with whom I shared 

my daily life during my time at the Munich institute, which was 

like a safe haven for me. They helped me in scientific as well as in 

personal affairs. I do not repeat that I enjoyed to get acquainted 

with so many people outside the institute, some of them becoming 

collaborators or real friends. Even if there will be little chance to


meet one or the other of them again, they are alive in my memory. 

It is them who have to decide on the quality of my endeavor. 

Privately, my family was my best support and place of refuge, 

although they may not have realized this during the time I was 

busy or away from home. As said my wife gave birth to two 

daughters, and initially I was disappointed not to have a son. But 

I soon comprehended that daughters are much more devoted to a 

father than sons can be. Fortunately my wife took care of the 

children in any situation in life, which granted me a feeling of 

independence. Otherwise I feel lucky that they share one or the 

other of my interests: both of them loved and performed 

(classical) music and participated as supernumeraries in opera 

or ballett. Barbara before she took up and finished medicine was 

educated and got a diploma as a ballett dancer, Miriam is fond of 

fine arts, languages, and gardening. 

As a zoon politicon I never became a member of a political party, 

because such an interest had been spoiled for me radically by my 

experiences as a youngster. Not even could I accept to be 

monopolized by an interest group whose policy I did not agree 

with or to be sqeezed into a corset hierarchy; though in German 

university one has still to deal with this habitude, the most 

miserable outcome being a condescending attitude of some staff 

people towards their ‘‘subordinates.’’ 

Like many other things doing science is fun. Whenever 

somebody has developed a liking for science, I feel he will stick 

to it for ever. I found this notion expressed in all recollections of 

scientists I read thus far: pursuing scientific research gives 

satisfaction and pleasure. Indeed, this must be due to the fact that 

a scientist enjoys many privileges. As an occupation it bears its 

challenges, joys but also disappointments. I hope my recollections 

preferably mirror the pleasant aspects, because these are best to 

remember and to forget about the rest. 

Die Erinnerung ist das einzige Paradies aus dem wir nicht vertrieben 

werden können. 

ACKNOWLEDGMENT 

Jean Paul 

I wish to thank Giorgio Semenza for critical reading of the 

manuscript.

324 

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[98] Mannhaupt, G., van der Linden, G., Vetter, I., Maurer, K., Pilz, U., 

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[102] Thierry, A., Gaillon, L., Galibert, F. and Dujon, B. (1995) Construction of 

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[103] Dujon, B. et al. (1994) Complete DNA sequence of yeast chromosome XI. 

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[104] Mannhaupt, G., Stucka, R., Ehnle, S., Vetter, I. and Feldmann, H. (1992) 

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The excision repair gene RAD16 located in this region belongs to a novel 

group of double-finger proteins. Yeast 8, 397–408. 

[105] Schaaf-Gerstenschläger, I., Mannhaupt, G., Vetter, I., Zimmermann, 

F.K. and Feldmann, H. (1993) TKL2, a second transketolase gene of 

Saccharomyces cerevisiae. Cloning, sequence and deletion analysis of the 

gene. Eur. J. Biochem. 217, 487–492. 

[106] Mannhaupt, G., Stucka, Ehnle, S., Vetter, I. and Feldmann, H. (1994) 

Analysis of a 70 kb region on the right arm of yeast chromosome II. Yeast 

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[107] Johnson, D.R., Cook, S.J., Feldmann, H. and Gordon, J.I. (1994) 

Suppressors of nmt1-181, a conditional lethal allele of the Saccharomyces 

cerevisiae myristoyl-CoA: Protein N-myristoyltransferase gene, reveal 

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[108] Mannhaupt, G., Vetter, I., Schwarzlose, C., Mitzel, S. and Feldmann, H. 

(1996) Analysis of a 26 kb region on the left arm of yeast chromosome XV. 

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[109] Gonzalez, M.I., Stucka, R., Blazquez, M.A., Feldmann, H. and Gancedo, 

C. (1992) Molecular cloning and characteristics of CIF1, a yeast gene 

necessary for growth on glucose. Yeast 8, 183–192. 

[110] Pammer, M., Briza, P., Ellinger, A., Schuster, T., Stucka, R., Feldmann, 

H. and Breitenbach, M. (1992) DIT101 (CDS2, CAL1), a cell cycleregulated 

yeast gene required for synthesis of chitin in cell walls and 

chitosan in spore walls. Yeast 8, 1089–1099. 

[111] Stucka, R., Blázquez, M.A. and Feldmann, H. (1993) The fdp1 and cif1 

mutations are caused by different single nucleotide changes in the yeast 

CIF1 gene. FEMS Microbiol. Lett. 107, 251–254. 

[112] Blázquez, M.A., Stucka, R., Feldmann, H. and Gancedo, C. (1994) 

Trehalose-6-P synthase is dispensable for growth on glucose but not for 

spore germination in Schizosaccharomyces pombe. J. Bacteriol. 176, 

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[113] Feldmann, H. et al. (1994) Complete DNA sequence of yeast chromosome 

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[114] Vassarotti, A., Dujon, B., Mordant, P., Feldmann, H., Mewes, W. and 

Goffeau, A. (1995) Structure and organization of the European yeast 

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[115] Goffeau, A., Barrell, B.G., Bussey, H., Davis, R.W., Dujon, B., Feldmann, 

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[117] Entian, K.D., Schuster, T., Hegemann, J.H., Becher, D., Feldmann, H., et al. 

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[122] Nelböck, P., Dillon, P.J., Perkins, A. and Rosen, C.A. (1990) A cDNA for a 

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peroxisome biogenesis, encodes a member of a novel family of putative 

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[124] Schnall, R., Mannhaupt, G., Stucka, R., Tauer, R., Ehnle, S., 

Schwarzlose, C., Vetter, I. and Feldmann, H. (1994) Identification of a 

set of yeast genes coding for a novel family of putative ATPases with high 

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[125] Ghislain, M., Udvardy, A. and Mann, C. (1993) S. cerevisiae 26S protease 

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[127] Finley, D., Tanaka, K., Mann, C., Feldmann, H., Hochstrasser, M., 

Vierstra, R., Johnston, S., Hampton, R., Haber, J., McCusker, J., Silver, 

P., Frontali, L., Thorsness, P., Varshavsky, A., Byers, B., Madura, K., 

Reed, S.I., Wolf, D., Jentsch, S., Sommer, T., Baumeister, W., Goldberg, 

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[128] Tauer, R., Mannhaupt, G., Schnall, R., Pajic, A., Langer, T. and 

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[130] Arlt, H., Tauer, R., Feldmann, H., Neupert, W. and Langer, T. (1996) 

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nonamer box found upstream of 26S proteasomal and other genes in 

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[134] Mannhaupt, G., Pilz, U. and Feldmann, H. (1988) A series of shuttle 

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[135] Jelinsky, S.A., Estep, P., Church, G.M. and Samson, L.D. (2000) 

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A Life with Yeast Molecular Biology - Prof. Dr. Horst Feldmann

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