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WILLIAM ZEV HASSID O cto b er 1,1 899-A pri! 2S, 1974 BY CLINTON BALLOU AND HORACE A. BARKER ZEV (ZE'EV) HASSID was born in Jaffa, Palestine, probably on October 1, 1899, although he seemed uncertain about the date of his birth and sometimes gave the year as 1897 or 1901. The name William was added after he came to the United States. His parents, Mordecai and Esperanza Hassid (Chassid), were born in Poland, but were Russian citizens at the time of his birth. His father was a lumber merchant who brought Jum- ber from Russia to Palestine. When Zev was four years old, the family moved to a farm in the vicinity of Kremenetz in the Russian Ukraine, and his childhood was spent in this rural en- vironment. He often wandered in the adjacent woods and fields, hunted for birds' nests, and associated with shepherds, so much so that his father scolded him and said that if he did not spend more time on his studies he would qualify for nothing more than a herdsman. Russian and Yiddish were Zev's native languages. Little is known about his early education except that he was required to study traditional Jewish religious material. He did not ac- cept this instruction readily and in later life rebelled by dis- sociating himself from most formal religious activities. Never- theless, his early training was evidenced by a familiarity with and an occasional quotation from scriptures. In 1912, Hassid was sent back to Palestine with a group of 197
198 BIOGRAPHICAL MEMOIRS Jewish children to continue his education in a Hebrew lan- guage school. His parents hoped that he would be admitted to the "Gymnasia Hertzlia" in Tel Aviv, but instead he was sent to the recently founded Agricultural High School in the Jewish settlement of Petah-Tikva. The curriculum included Hebrew, French, and Arabic languages, Hebrew religious studies, his- tory, geography, and considerable science, plus professional studies in soil, plant nutrition, subtropical horticulture, animal husbandry, laboratory practice in analytical methods, and field experience in agricultural techniques. He graduated in 1916. With the outbreak of WorIct War I, Hassid was cut off from communication with his parents, who tract remained in Russia. Consequently, his only income was what he could earn himself. While still in school and after graduation, he worked as a laborer in the orange groves and other farms near Petah-Tikva, Nikva- Israel, and Ben Shemen. His income was small, his food and clothing correspondingly poor. For a considerable period he lived on bread, watery soup, pumpkin porridge, and oranges; he was often hungry. His clothing was worn and ragged. Once his shoes were stolen, and he had to go barefoot for a considerable time. During this period Hassid was frequently ill. He sufferer! repeated attacks of malaria and dysentery; he also contracted typhoid fever, which he barely survived. Palestine was controlled by the Turks until the British army invaded the country in 1917. Following the Balfour DecIa- ration that same year, many young Jews joined the British army to help liberate the country from the Turks and create conditions favorable for the establishment of a Jewish home- land. Hassid volunteered for army service early in 1918, partly for patriotic reasons and partly to improve his standard of liv- ing. Initially, he was rejected by the army because of his poor health, but finally, through the intervention of James de Roths- child, an officer in the British army, he was accepted into the 38th brigade of the Royal Fusiliers (1st Judeans), later referred
WILLIAM ZEV HASSID 199 to as the Jewish Legion. At this time he became a British citi- zen and served for two years in the army, mainly as a clerk at supply depots and at General Headquarters, 3rd Echelon. He never engaged in combat; however, on at least one occasion he was close enough to the front to be within artillery range, and a large shell landed close to him but failed to explode. He also guarded prisoners and supplies in transit. The latter duty re- quired him to travel as far as Beyreuth in Lebanon and Alex- andria in Egypt. At the railway station in Beyreuth he once witnessed the ceremonial arrival of Lawrence of Arabia. In Alexandria he first heard of the University of California from a fellow soldier, Assaf Gur (Grazovsky), who had studied there. When Hassid completed his military service with the rank of corporal in August 1920, he was awarded the British War Medal and the Victory Medal, bearing the inscription, "The great war for civilization." His superior officer provided him with the following recommendation: "Corporal Hassid has been employed as a clerk in the Battalion Orderly Room and at Rec- ords, 3rd Echelon. He is a conscientious, painstaking worker, reliable and capable, and as a soldier has borne an exemplary character." After leaving the army, Hassict decided to use the accumu- lated savings from his pay to go to California to study agronomy at the University, with the intention of returning ultimately to Palestine to assist in the development of scientific agriculture. He traveled by way of Paris, New York, and Chicago, staying briefly in each city, and finally arrived in Berkeley in late 1920. His funds were almost exhausted, so he supported himself by doing odd jobs in stores and restaurants in Berkeley in the winter and by working as a farm hand in the vicinity of Fresno in the San Joaquin Valley in the spring and summer. He also taught Hebrew in the local synagogue. Hassid first registered at the University of California in Au- gust 1921. However, his knowledge of English was so limited
200 BIOGRAPHICAL MEMOIRS that he could! not follow lectures well enough to take notes, and even reading textbooks required more time than he could spare between jobs. So after a week of frustration and mounting ten- sion, he took a leave of absence from the University and moved to Fresno where he spent two years as a student in Fresno State College, majoring in Letters and Science with an emphasis on Chemistry, French language, and Mathematics. The following year he enrolled at the Southern Branch of the University of California at Los Angeles. His course grades were about aver- age; he even failed one course in quantitative analysis and had to repeat it later. His undistinguished record during this period probably resulted from his inadequate command of English and the fact that he had to earn a living while attending school. In August 1924, Hassid returned to the Berkeley campus of the University. He majored first in chemistry but later changed to general literature, the field in which he obtained a Bachelor of Arts degree in December 1925. Following graduation, he im- mediately started graduate studies in the School of Education, and in December 1926 he received a Certificate of Completion with majors in chemistry and general literature and minors in mathematics and physics. In the same month Hassid obtained a General Secondary School Credential from the State Board of Education, but he never taught in public schools. Instead, he worked for some months as a chemical analyst in a commercial company. By September 1927, and possibly earlier, Hassid took a position as research assistant under Professor D. R. Hoagland in the Division of Plant Nutrition of the Agricultural Experi- ment Station. His main duties were the routine analysis of plant materials and soils for a variety of inorganic constituents, but he also obtained experience in growing plants in culture solution and studying the absorption of various nutrients. This work renewed Hassid's interest in plant research, and in August 1928 he again enrolled in the University, this time as a graduate stu-
WILLIAM ZEV HASSID 201 dent in Plant Nutrition. During the following two years, while still working half time or more, he took courses in botany, plant physiology, and plant nutrition and prepared a Master of Science thesis dealing with the structure of the four isomers of penta-O- acetyl-D-galactose. Professor Hoagland formally supervised his thesis research, but the nature of the problem clearly indicates that it was inspired and guided mainly by Professor Walter H. Dore, who was interested in the structure of carbohydrates and had applied X-ray diffraction methods to their elucidation. After receiving the master's degree in August 1930, Hassid started to prepare for a doctorate in Plant Physiology. While visiting the beaches south of San Francisco, he observed the abundant fleshy marine algae, which appeared to consist largely of polysaccharides, and he decided to investigate the structure of the major component. This was the subject of his Ph.D. thesis, which was completed and accepted in December 1934, with Professor Dore as the chairman of the committee. Hassid had worked almost independently, however, using the methods developed by Haworth for carbohydrate structure determina- tion, which were unfamiliar to Dore. Professor T. D. Stewart of the chemistry department, a member of the thesis committee, was particularly impressed by the clear results and logical pre- sentation of Hassid's thesis, and his enthusiastic reaction helped Hassid to obtain an appointment the following year as a Junior Chemist in the Division of Plant Nutrition of the Agricultural Experiment Station. The circumstances of Hassid's appointment as a Junior Chemist are amusing and illustrative of the method of making appointments in 1935. Professor Hoagland lectured and taught laboratory courses in plant biochemistry during the Fall term. Hassid had served for several years as a teaching assistant in the laboratory course, and Hoagland, who directed an active re- search program and served as Chairman of the Division, had come to depend on him for the preparation of reagents, setting
202 BIOGRAPHICAL MEMOIRS up of equipment, and much of the instruction. But after re- ceiving his Ph.D. in 1934, Hassid began to look for a better position than was provided by his assistantship. In the early summer of 1935, he received an offer of a position ant! told Pro- fessor Hoagland that he was planning to leave before the be- ginning of the Fall term. Hoagland was upset at this news because he was preparing to attend a botanical congress in Amsterdam and would not return to Berkeley until the begin- ning of the term and then would have no experienced assistant for the laboratory course. He finally asked Hassid whether he would stay on if he received an Experiment Station appoint- ment. Hassid agreed. Hoagland consulted with Dean Hutchison, who found that the available funds were insufficient to provide the usual starting salary of $2,000 per year, but he could offer $1,800. Hassid accepted this although it was considerably be- low what he was offered outside the University. He never re- gretted his decision. In 1939 he received the additional title of Instructor, and so was launched upon his academic career. Hassid's independent scientific research began with an in- vestigation of the ethanol-extractable carbohydrates in the ma- rine alga Iridea laminarioides, and this work led step-by-step to an interest in the biochemistry of carbohydrates that he sus- tained in one form or another for almost thirty-five years. In the initial study, he identified dulcitol (galactitol) as a major component of this plant extract and observed that reducing sugars were notably absent from acid hydrolysates. He then purified and characterized an abundant polysaccharide from the same organism and showed it to be a sulfated polygalactan. From methylation studies, he was able to conclude that the galac- tosyl units were probably joined by 1~4 glycosidic linkages ant} that the sulfate was probably esterified with the hydroxyl group on carbon 6. These results led him to speculate that the metab^ olism in this alga might involve a relationship between galac-
WILLIAM ZEV HASSID 203 titoT and galactan analogous to that between glucose and starch in higher plants, but he never followed up this idea. Hassid's study of the structure of the algal galactan was the first of a long series of investigations of polysaccharides. Many of the preparations were provided by colleagues with whom he was always happy to collaborate. C. B. Lipman called his atten- tion to a very viscous substance produced from mannitol by an unidentified bacterium that had been isolated "from a mud brick taken from a wall in an old Roman village which was built about 400 AD in the western desert of Egypt." In a paper with W. L. Chandler (1937), Hassid characterized the viscous material as a polysaccharide containing about ten anhydroglu- cose units. In the following years he published papers dealing with the molecular structure of canna starch (with W. H. Dore), dog liver glycogen (with I. L. Chaikoff), the dextran formed from sucrose by Betacoccus arabinosaceus (with H. A. Barker), an insoluble polysaccharide derived from Saccharomyces cerevisiae (with M. A. Joslyn and R. M. McCready), and glycogen and starch derived from sweet corn (Zea mays) (with R. M. Mc- Cready). The existence of the enzyme phosphorylase that converts glycogen and inorganic phosphate to a-D-glucose 1-phosphate had been demonstrated by C. F. Cori and G. Cori in 1936, and the reversal of this reaction was reported in 1939 by W. Kiessling. Hassid and R. M. McCready (1941) undertook the structural analysis of the biosynthetic product and showed that it had starchlike properties but that the molecules were unbranched in contrast to the highly branched natural polymer. In a short review for Chronica Botanica published in 1942, Hassid related the prevalent view that "the enzyme phosphorylase, and not amylase as had been previously assumed, is chiefly responsible for the synthesis and breakdown of starch in the plant." Mc- Cready and Hassid developed a convenient method for prepar-
204 BIOGRAPHICAL MEMOIRS ing pure a-D-glucose 1-phosphate on a relatively large scale, thus making this important compound readily available for bio- chemical studies. They also developed a procedure for deter- mining the relative amounts of amylose and amylopectin in starch based upon the large difference in the absorption coe~- cients of iodine complexes of the two components. The absorp- tion coefficients of different starch samples were found to correlate well with the degree of hydrolysis of the components by,8-amylase. Their results supported the conclusion of K. H. Meyer that amylose consists of long unbranched chains of glycosyl units. This experience with carbohydrates prepared Hassid for an important collaborative effort with S. Ruben and M. D. Kamen (1939) representing the first application of radioactive carbon to the study of photosynthesis. The short-lived C-isotope (20.5 minutes half-life) had become available through Kamen's asso- ciation with the Radiation Laboratory at the University in Berkeley, and a study was carried out to determine the distribu- tion of the label from C-carbon dioxide when fed to barley leaves in the light or after they had been kept in the dark for various periods of time. Although some of the label was incorpo- rated into carbohydrate, the results indicated that most of it was present in the plant in a water-soluble noncarbohydrate form. In an extension of this work, the green alga Chlorella pyrenoidosa was utilized in place of barley leaves, allowing a much more efficient incorporation of the radiocarbon label. Studies of the kinetics of incorporation in the light and dark, and on the reversibility of the reaction, were carried out with Ruben and Kamen (1940~. Although it was concluded that "the greater fraction if not all the COOS has been reduced to POOH," no specific identification of the initial product of photosynthesis was made other than that there were "at least one alcoholic hydroxyl and one carboxyl group in the active
WILLIAM ZEV HASSID 205 molecules." The product was later identified as phosphoglyceric acid by M. Calvin and his associates. Other than a brief collaboration with S. Aronoff, A. Benson, and M. Calvin (1947) on the distribution of label from SCOW in photosynthesizing plant tissue, Hassid's research on photo- synthesis was not continued and his involvement appears to have been based more on an interest in carbohydrate structural analysis than in the fundamentals of carbon fixation in plants. However, it is apparent from the later turn of events that this introduction to the utility of radioactive tracer techniques for elucidation of biochemical processes had a strong influence on his development. Hassid's initial appointment as a Junior Chemist in the Agricultural Experiment Station was followed by promotion to the academic staff as an Instructor in Plant Nutrition (1939) and as Assistant Professor in 1941. About this time, he devel- oped an association with fellow colleagues H. A. Barker and M. Doudoroff that grew into a close scientific collaboration and a lifelong friendship. His first joint study with Barker concerned the structure of an extracellular (1~6) dextran produced by the bacterium Leuconostoc mesenterioides when grown on sucrose. Less than three years later, he was involved with Doudoroff and N. Kaplan in the beginning of an important study on the bio- synthesis of sucrose. Doudoroff had been interested in the bac- terium Pseudomonas saccharophila because it oxidized sucrose faster than the component monosaccharides glucose and fructose. This was demonstrated to result from the presence of an enzyme that converts sucrose to a-D-glucose 1-phosphate and D-fructose. Doudoroff, Kaplan, and Hassid found that this reac- tion can be reversed, leading to the formation of sucrose as de- tected by the production of a nonreducing substance that yielded reducing sugar on acid hydrolysis. That it was indeed sucrose was proved by Hassid, Doudoroff, and Barker (1944)
206 BIOGRAPHICAL MEMOIRS when they prepared 2.5 grams of the crystalline disaccharide by the action of sucrose phosphorylase on a mixture of 15 grams each of a-D-glucose 1-phosphate and D-fructose and showed that its properties were identical with those of commercial sucrose. The enzymatic synthesis of sucrose resulted in some pub- licity that came to the attention of officials of the Coca-Cola company, who were having difficulty obtaining sucrose because of wartime rationing. The company sent a representative to Berkeley to ascertain whether commercial quantities of sucrose could be made by the enzymatic method. Hassid and his asso- ciates were away on vacation at the time, so the Coca-Cola emissary discussed the problem with Professor Hoagland and reported that his company was prepared to provide $500,000 for research on this enzyme if a commercial process of sucrose synthesis seemed feasible. Unfortunately, Professor Hoagland was pessimistic about the possibility of sweetening Coca-Cola by this method, and so further support of research of sucrose phosphorylase was left to the University and the U.S. Public Health Service. Later studies on sucrose phosphorylase showed that it could transfer D-glucose to ~-sorbose and D-threo-pentulose to form nonreducing disaccharide analogs of sucrose and to ~-arabinose to yield 3-O-a-D-glucopyranosyl-~-arabinose. The mechanism of the phosphorylase reaction was investigated by isotope exchange reactions with 32P-orthophosphate, which was shown to ex- change into nonradioactive a-D-glucose 1-phosphate. With alternative monosaccharide acceptors such as ~-sorbose, the enzyme was shown to transfer D-glucose from sucrose in the ab- sence of orthophosphate. Clearly, the reaction involved an inter- mediate D-glucosyl-enzyme that could be formed either from a-D-glucose 1-phosphate or from an a-D-glucosyl derivative such as sucrose. Near the end of the 1940s, Hassid's research began to take
WILLIAM ZEV HASSID 207 a new direction as he concentrated on the synthesis of radio- carbon-labeled sugars and on the chemical preparation of sugar phosphates. E. W. Putman was a major collaborator in develop- ing methods for preparing labeled sugars from plant tissue after their biosynthesis by the photosynthetic fixation of i4C-carbon dioxide. They applied the new methods of paper chromatog- raphy to the preparation of uniformly i4C-labeled carbohydrates of high specific activity, including glucose, fructose, galactose, sucrose, and starch. Hassid generously supplied radioactive sugars both to his colleagues and to many scientists throughout the country before they became commercially available. These studies on the preparation of labeled sugars were extended to investigations of the route by which the carbon dioxide, once fixed as glyceric acid phosphate, was converted into various carbohydrates, as well as the processes by which labeled hexose was taken up and utilized for synthesis of sucrose and cellulose. About this time, Hassid was joined by two graduate stu- dents, V. Ginsburg and E. F. Neufeld, in an active program dealing with the role of sugar nucleotides in the interconversion of carbohydrates in higher plants. In the initial studies they were joined by P. K. Stumpf, who had been appointed to the Department of Plant Nutrition in 1948, and who, as an under- graduate student at Harvard University, had worked on puri- fication of potato phosphorylase with D. E. Green. Although Stumpf is best known for his investigations on lipid metabolism in plants, his earlier studies in sugar metabolism were influen- tial in the development of Hassid's interests in this field. The discovery by L. F. Leloir (1951) of uridine diphosphate D-glucose and the demonstration that this substance served as a glucosyl donor for synthesis of disaccharides focused attention on the sugar nucleotides as intermediates in the interconversion of carbohydrates, and Hassid directed his concern to these sub- stances in higher plants. With Ginsburg and Stumpf (1956), he
208 BIOGRAPHICAL MEMOIRS investigated the occurrence of uridine diphosphate derivatives of D-glucose, D-galactose, D-xylose, and ~-arabinose in mung bean (Phaseolus aureus), the latter two derivatives being found for the first time in nature. The same source yielded the uridine diphosphate derivatives of N-acetyl-D-glucosamine and D-glucu- ronic acid (with J. Solms and D. S. Feingold), whereas the guanosine diphosphate derivatives of ~-galactose and D-mannose later were identified in the red alga Porphyra perforate (with I. C. Su), and guanosine diphosphate D-mannuronic acid was isolated from the brown alga Fucus gardneri (with T. I. Lin). These studies on the natural occurrence of sugar nucleotides in plants were paralleled by investigations of their biosynthesis by the pyrophosphorylase reaction. A series of papers with Neu- felcI, Putman, Feingold, Ginsburg, and others delineated the presence in higher plants of pyrophosphorylases that formed the respective uridine diphosphate hexoses from reaction of uridine triphosphate with the 1-phosphate esters of a-D-galactose' a-D- xylose, ,8-~-arabinose, a-D-glucuronic acid, a-D-galacturonic acid, and N-acetyl-a-D-glucosamine. Because these studies required the sugar 1-phosphates as substrates and such substances were not generally available at the time, considerable effort was de- voted to the improvement of published syntheses and the de- velopment of new ones for the preparation of glycosyl phos- phates. Later studies dealt with the enzymic phosphorylation of several sugars, including D-galactose, ~-arabinose, D-glucuronic acid, and D-galacturonic acid (with Neufeld, Feingold, and others). Hassid's international reputation attracted senior scientists from around the world to work in his Berkeley laboratory. One of these was Winifred N. Watkins from the Lister Institute of Preventive Medicine in London, who is noted for her studies on the structure and biosynthesis of the blood group antigens. In 1961 Watkins and Hassid undertook a study of the biosyn- thesis of lactose in mammary tissue. It had been claimed by
WILLIAM ZEV HASSID 209 i. G. Gander, W. E. Petersen, and P. D. Boyer (1957) that bovine mammary tissue contained enzymes that converted uridine di- phosphate D-galactose and a-D-glucose 1-phosphate to lactose 1-phosphate, and that the latter was hydrolyzed to free lactose. However, since uridine diphosphate D-galactose and a-D-glucose 1-phosphate are in ready equilibrium by way of uridine diphos- phate D-glucose, one would expect i4C-D-glucose to be incorpo- rated equally into the two parts of lactose by this pathway, an expectation that was contrary to recorded observations by other workers. Watkins and Hassid reinvestigated this matter and established that uridine diphosphate D-galactose and free D-glu- cose were the precursors of lactose in lactating guinea pig and bovine mammary tissue and that the product was lactose rather than lactose 1-phosphate. In addition to these results on lactose synthesis, Watkins and Hassid (1962) also made the important observation that mam- mary tissue contained an enzyme activity that transfers D-galac- tose to N-acetyl-D-glucosamine. Noting the occurrence in milk of oligosaccharides that contained this lactosamine unit, and finding that different mammary gland preparations gave dif- ferent relative amounts of ~4C-lactose and N-acetyl-~4C-lactos- amine when incubated with uridine diphosphate i4C-D-galactose, they concluded that "different enzymes are responsible for the synthesis of the two compounds." This conclusion, and a later one by Helene Babad and Hassid (1966) that the soluble, puri- fied lactose synthetase from milk was "very labile to further purification," proved to be incorrect, for it was found subse- quently by K. E. Ebner and others (1967) that the mammary gland,8-D-galactosyltransferase is under the control of a specifi- city-altering protein, a-lactalbumin. In the presence of a-lactal- bumin, the preferred acceptor is D-glucose, and lactose is the product, whereas in absence of the protein, N-acetyl-D-glucos- amine acts as the acceptor to yield N-acetyl-lactosamine. The observations of Watkins and Hassid can be explained by the
210 BIOGRAPHICAL MEMOIRS presence of variable amounts of a-lactalbumin in their lactose synthetase preparations, and the results of Babad and Hassid are explained as reflecting the first successful fractionation of these two proteins. The existence of enzymes that interconverted sugar nucleo- tides without degrading the molecule had become recognized by the latter part of the 1950s, and Hassid turned to the study of some of these reactions in plants. With Neufeld and Fein- gold, he demonstrated the enzymic conversion of uridine di- phosphate D-glucuronic acid to the nucleotide derivatives of galacturonic acid, xylose, and arabinose. In a later study, the enzyme activities that catalyzed these reactions were separated so that it was possible to show the 4-epimerization of uridine diphosphate D-glucuronic acid to the galacturonic acid deriva- tive as an isolated step and to demonstrate the decarboxylation of uridine diphosphate D-glucuronic acid to uridine diphosphate xylose. An enzyme activity was also found that epimerized the xylose derivative to the arabinose derivative. Throughout his career, Hassid was concerned with the fundamental question of how the sugar that is formed in a photo- synthesizing plant is converted to disaccharides such as sucrose and into polysaccharides such as starch and cellulose. His earliest experiments dealt with the infiltration of radiocarbon- labeled sugars into plant leaves, but later they became more sophisticated with the utilization of well-defined sugar nucleo- tides as specific donors in cell-free enzyme systems. It was known that uridine diphosphate D-glucose was a precursor of a ,B-1~4- glucan in ~cetobacter xylinum (L. Glaser, 1957) and of the a-1~4-glucan, glycogen, in liver (Leloir, 1957~. Feingold, Neu- feld, and Hassid (1958) reported the synthesis of a ,3-1~3-glucan from this sugar nucleotide by a digitonin-treatec] particulate transferase from mung bean, the product apparently being iden- tical to laminarin, whereas R. A. Dedonder and Hassid (1964) observed formation of a ,8-1 - 2-glucan in Rhizobium japonicum.
WILLIAM ZEV HASSID 211 In a similar fashion, uridine diphosphate xylose was shown to be the precursor of a,8-1 - 4-xylan in asparagus (~Asparagus of~ici- nalis). Hassid also investigated the synthesis of alginic acid (with T.-Y. Lin), the synthesis of pectin (with C. L. Villemez), and the methylation of the latter polysaccharicle to form the ether and ester derivatives (with H. Kauss). This interest in polysaccharide formation in plants led Has- sid naturally to an investigation of cellulose biosynthesis and perhaps to the culminating point of his scientific career. Even today the study of cellulose biosynthesis in higher plants is fraught with technical difficulties that have hindered the defi- nition of this system in the same detail that has been possible with other polysaccharide-forming reactions. In 1964, A. D. Elbein, G. A. Barber, and Hassid reported the formation of cellulose from guanosine diphosphate D-glucose and a particu- late enzyme preparation from mung bean seedlings. Contrary to the finding of Glaser with a bacterial system, the plant syn- thetase had no activity with uridine diphosphate D-glucose. The polysaccharide product was characterized primarily on the basis of its alkali insolubility, which was similar to that of cellulose, and on the formation of radioactive cellobiose and cellodextrins by acid hydrolysis and by acetolysis. The study was complicated by the concomitant formation of a glucomannan from endo- genous guanosine diphosphate mannose. However, a soluble enzyme system that made cellulose was eventually obtained (with H. M. Flowers, K. K. Batra, and l. Kemp, 1969), and this prod- uct was only slightly contaminated by mannose. Some controversy concerning cellulose biosynthesis arose when D. O. Brummond and A. P. Gibbons (1964) reported that uridine diphosphate D-glucose was a precursor of a cellulose-like polymer in Lupinus albus, and L. Ordin and M. A. Hall (1967) observed a similar reaction in Avena saliva. These reports stim- ulated Hassid to restudy the roles of both uridine and guanosine diphosphate D-glucose in the synthesis of alkali-insoluble poly-
212 BIOGRAPHICAL MEMOIRS saccharides in Phaseolus aureus. He eventually established, with H. M. Flowers, K. K. Batra, and J. Kemp (1968), that the poly- mer formed with uridine diphosphate D-glucose in L. albus was an alkali-insoluble,8-1~3-glucan rather than cellulose, whereas that produced by A. saliva was a mixed ,8-1~3- and ,8-1~4-glu- can. C. M. Tsai and Hassid (1971) succeeded in separating the two enzymic activities of A. saliva that make the two polysac- charides, and they found that the type of glucan made was de- pendent on the concentration of the sugar nucleotide donor. From this brief survey, we can see that there were few fea- tures of carbohydrate metabolism in plants that escaped Hassid's touch, and much that we know about the role of sugar nucleo- tides in the interconversion of carbohydrates in plants is a direct result of his persistent effort. From the incorporation of labeled precursors into monosaccharides, to the conversion of the mono- saccharides to their glycosyl 1-phosphates, to the action of the pyrophosphorylases in the synthesis of the nucleoside diphos- phate sugars, to the interconversion of the resulting sugar nucleotides, to the polymerization of the activated monosac- charides yielding disaccharides and the homopolysaccharides, and finally to the modification of the polysaccharides by methyl- ation in summary, to almost every aspect of carbohydrate metabolism Hassid contributed his full and devoted attention. Nor did his efforts slacken with age, for he continued working and writing and thinking science as though it was among the most important things in life, and to him it was. He also believed in the importance of communication as a force in scientific progress, for he was a prolific writer of reviews and contributed heavily to books and serials dealing with carbo- hydrates, the total of such articles being almost fifty. Hassid's many contributions on the structure and synthesis of plant carbohydrates were recognized by a number of honors and awards. He received the first Sugar Research Award (1945) of the National Academy of Sciences (jointly with M. Doudoroff
WILLIAM ZEV HASSID 213 and H. A. Barker), the Charles Reid Barnes Honorary Life Membership Award of the American Society of Plant Physiolo- gists (1964), and the C. S. Hudson Award of The American Chemical Society (1967~. He was elected to membership in the National Academy of Sciences (1958) and the American Acad- emy of Arts and Sciences (1969), and he was honored at the 6th International Symposium on Carbohydrate Chemistry (1972) as one of three outstanding senior American carbohydrate chemists. He was elected Chairman of the Division of Carbo- hydrate Chemistry ~ 1949-1950), American Chemical Society, and he served as a member of numerous editorial boards, in- cluding those of the Journal of Biological Chemistry, Annual Review of Biochemistry, Carbohydrate Research, Phytochem- tstry, and A nalytical Biochemistry. Hassid married Lila Berlin Fenigston in 1936. They had no children, but any void this may have created in their lives was filled by the many friends who shared the warm hospitality of their home in the Berkeley hills. Lila was a gracious and viva- cious hostess and an accomplished violinist who made their home a center for friendly social gatherings and for the per- formance of chamber music. She also had a talent for making fine English translations of Yiddish poetry, which were either published in Jewish periodicals or presented orally in a series of radio programs. Lila's appreciation and contributions to the arts provided a happy counterpoint to the scientific life of her husband. Perhaps as a result of his severe childhood illnesses and early deprivations, Hassid never enjoyed robust health. For many years, he suffered from the effects of high blood pressure, and he had debilitating attacks of hyperthyroidism and hepatitis. In his early sixties, he suffered a severe coronary occlusion from which he never fully recovered. As he grew older he developed increasing coronary complications that finally resulted in his death on April 2S, 1974. He left many friends who will remem-
214 BIOGRAPHICAL MEMOIRS her him as a friendly, gentle, and soft-spoken person, but one who on rare occasions, when sufficiently annoyed, could display a strong temper. His personal warmth and generosity, coupled with his sincerity and modesty, attracted many friends, whom he treasured and often regarded as somewhat larger than life. He took pride in the accomplishments of his colleagues and students and spent much time and effort in helping to further their careers and in nominating them for promotions, awards, and honors of various sorts. He never tired nor stinted in help- ing those who he felt were deserving. THE INFORMATION in this memoir relating to Hassid's early life in Russia and Palestine was obtained from my (H.A.B.) conversations with him, from letters (dated June 2, 1974 and June 25, 1974>, and from an unpublished account of Hassid's early life written by Dr. Rivka Ashbel (prepared for the celebration of Petah-Tikva's seven- tieth anniversary and based on conversations between Hassid and Ashbel in 1949~. Additionally, we had available a number of docu- ments including Hassid's passport application, naturalization papers, and records of his service in the British army and service medals. Most of this material and some of Hassid's manuscripts and letters are kept in a file in the Bancroft Library at the University of California, Berkeley.
WILLIAM ZEV HASSID BIBLIOGRAPHY 1933 215 Occurrence of dulcitol in Irideae laminarioides (Rhodophyceae). Plant Physiol., 8:480-82. The isolation of a sodium sulfuric acid ester of galactan from Irideae laminarioides (Rhodophyceae). i. Am. Chem. Soc., 55:4163-67. 1935 The structure of sodium sulfuric acid ester of galactan from Irideae laminarioides (Rhodophyceae)..~. Am. Chem. Soc., 57:2046-450. 1936 Determination of reducing sugars and sucrose in plant materials. Ind. Eng. Chem. Anal. Ed., 8: 138-40. Carbohydrates in Irideae laminarioides (Rhodophyceae). Plant Physiol., 11:461-63. Comparison of the total nitrogen in wheat seeds by the Gunning (modified Kjeldahl) and the Dumas combustion methods. l. Am. Chem. Soc., 58:2075. 1937 With W. L. Chandler. The isolation of a new polysaccharide synthe- sized by a soil microorganism. J. Biol. Chem., 117: 203-7. Determination of sugars in plants. Ind. Eng. Chem. Anal. Ed., 9:228-29. With W. H. Dore. The molecular structure of canna starch. J. Am. Chem. Soc., 59:1503-8. 1938 With I. L. Chaikoff. Phosphorylation of glycogen in vitro. Science, 88: 15-16. With I. L. Chaikoff. The molecular structure of liver glycogen of the dog. J. Biol. Chem., 123: 755-59. Isolation of hexosemonophosphate from pea leaves. Plant Physiol., 13:641-47.
216 BIOGRAPHICAL MEMOIRS 1939 With S. Ruben and M. D. Kamen. Radioactive carbon in the study of photosynthesis. l. Am. Chem. Soc., 61:661-63. A water-soluble glucosan from barley roots. A. Am. Chem. Soc., 61:1223-25. With S. Ruben, M. D. Kamen, and D. C. deVault. Photosynthesis with radio-carbon. Science, 90:570-71. 1940 With R. M. McCready. Determination of starch in plants. Ind. Eng. Chem., 12:142~4. With M. D. Kamen and S. Ruben. Radioactive nitrogen in the study of N2 fixation by non-leguminous plants. Science, 91:578-79. With H. A. Barker. The structure of dextran synthesized from su- crose by Betacoccus arabinosaceus, Orla-lensen. l. Biol. Chem., 1 34: 163-70. With S. Ruben and M. D. Kamen. Photosynthesis with radioactive carbon. II. Chemical properties of the intermediates. T. Am. Chem. Soc., 62:3443-50. 1941 With M. A. Joslyn and R. M. McCready. The molecular constitution of an insoluble polysaccharide from yeast, Saccharomyces cere- visiae. J. Am. Chem. Soc., 63: 295-98. With R. M. McCready. The molecular constitution of glycogen and starch from the seed of sweet corn (`Zea mays). J. Am. Chem. Soc., 63: 1632-35. With R. M. McCready. Transformation of sugars in excised barley shoots. Plant Physiol., 16:599-610. With R. M. McCready. The molecular constitution of enzymatically synthesized starch. I. Am. Chem. Soc., 63:2171-73. 1942 Recent work on the structure of plant polysaccharides. Chronica Botanica, 7:135-37. With R. M. McCready. Semimicrodetermination of carbon. Ind. Eng. Chem. Anal. Ed., 14:525-26.
WILLIAM ZEV HASSID 217 With R. M. McCready. Identification of sugars by microscopic appearance of crystalline osazones. Ind. Eng. Chem., 14:683-86. 1943 With M. Doudoroff and N. Kaplan. Phosphorolysis and synthesis of sucrose with a bacterial preparation. I. Biol. Chem., 148:67-75. With G. T. Cori and R. M. McCready. Constitution of the poly- saccharide synthesized by the action of crystalline muscle phos- phorylase. J. Biol. Chem., 148:89-96. With R. M. McCready. The separation and quantitative estimation of amylose and amylopectin in potato starch. I. Am. Chem. Soc., 65:115~57. With R. M. McCready. The molecular constitution of amylose and amylopectin of potato starch. I. Am. Chem. Soc., 65:1157-61. With E. E. Baker and R. M. McCready. An immunologically active polysaccharide produced by Coccidioides i~r~mitis, Rixford and Gilcl~rist. J. Biol. Chem., 149:303-11. The molecular constitution of starch and the mechanism of its for- mation. Quarterly Review of Biology, 18:311-30. 1944 With R. M. McCready. The preparation and purification of glucose 1-phosphate by the aid of ion exchange absorbents. l. Am. Chem. Soc., 66:560-63. With H. A. Barker and M. Doudoroff. Enzymatic synthesis of crystal- line sucrose. Science, 100:51. Chemistry of carbohydrates. Annual Review of Biochemistry, 13: 59-92. With M. Doudoroff and H. A. Barker. Enzymatically synthesized crystalline sucrose. l. Am. Chem. Soc., 66:1416-19. With M. Doudoroff and H. A. Barker. Synthesis of two new carbo- hydrates with bacterial phosphorylase. Science, 100:315-16. With W. L. McRary, W. H. Dore, and R. M. McCready. Inulin in guayule. Parthenium argentatum Gray. J. Am. Chem. Soc., 66: 1970-72. 1945 Review of the Annual Review of Biochemistry, Vols. 12 and 13. J. Am. Chem. Soc., 67:505.
218 BIOGRAPHICAL MEMOIRS The molecular constitution of starch. Wallerstein Lab. Commun., 8:34 45. With M. Doudoroff and H. A. Barker. Isolation and structure of an enzymatically synthesized crystalline disaccharide, d-glucosido-L- sorboside. l. Am. Chem. Soc., 67:139~97. Recent advances in the molecular constitution of starch and gly- cogen. Fed. Proc., 4:227-34. Review of the Annual Review of Biochemistry, Vol. 14. Ann. Revs. Inc., of Stanford. J. Am. Chem. Soc., 67:2277-78. 1946 The mechanism of breakdown and formation of starch and glycogen. Wallerstein Lab. Commun., 9: 135~4. With M. Doudoroff, H. A. Barker, and W. H. Dore. Isolation and structure of an enzymatically synthesized crystalline disaccha- ride, D-glucosido-D-ketoxyloside. J. Am. Chem. Soc., 68:1465-67. With W. R. Meagher. Synthesis of maltose 1-phosphate and D-xylose 1-phosphate. J. Am. Chem. Soc., 68:2135-37. 1947 With M. Doudoroff and H. A. Barker. Studies with bacterial sucrose phosphorylase. I. The mechanism of action of sucrose phos- phorylase as a glucose-transferring enzyme (trans-glucosidase). I. Biol. Chem., 168: 725-32. With M. Doudoroff and H. A. Barker. Studies with bacterial sucrose phosphorylase. II. Enzymatic synthesis of a new reducing and of a new non-reducing disaccharide. I. Biol. Chem., 168:733-46. With S. Arnoff, A. Benson, and M. Calvin. Distribution of C~4 in photosynthesizing barley seedlings. Science, 105:664-65. With M. Doudoroff. EnzymaticalIy synthesized disaccharides. Arch. Biochem., 14:29-37. With M. Doudoroff and H. A. Barker. Studies with bacterial sucrose phosphorylase. III. Arsenolytic decomposition of sucrose and of glucose 1-phosphate. J. Biol. Chem., 170: 147-50. 1948 With l. Katz and M. Doudoroff. Arsenolysis and phosphorolysis of the amylose and amylopectin fractions of starch. Nature, 161:96-97.
WILLIAM ZEV HASSID 219 With M. Doudoroff, A. L. Potter, and H. A. Barker. The structure of an enzymatically synthesized reducing disaccharide, D-glucosido- L-arabinose. i. Am. Chem. Soc., 70:305-10. With E. W. Putman, G. Krotkov, and H. A. Barker. Preparation of radioactive carbon-labeled sugars by photosynthesis. J. Biol. Chem., 1 73:785. With A. L. Potter, I. C. Sowden, and M. Doudoroff. Alpha-L glucose-l-phosphate. i. Am. Chem. Soc., 70: 1751-52. With A. L. Potter. Starch. I. End-group determination of amylose and amylopectin by periodate oxidation. J. Am. Chem. Soc., 70:3488-90. With A. L. Potter. Starch. II. Molecular weights of amyloses and amylopectins from starches of various plant origins. I. Am. Chem. Soc., 70:3774-77. With M. Doudoroff. Enzymatically synthesized polysaccharides and disaccharides. Fortschr. Chem. Organ. Naturstoffe, 5:101-27. 1949 With M. Doudoroff, E. W. Putman, A. L. Potter, and l. Lederberg. Direct utilization of maltose by Escherichia coli. J. Biol. Chem., 179:921-24. With H. WoIochow, E. W. Putman, M. Doudoroff, and H. A. Barker. Preparation of sucrose labeled with Ci4 in the glucose or fructose component. J. Biol. Chem., 180: 1237. With A. L. Potter and M. A. Joslyn. Starch. III. Structure of apple starch. J. Am. Chem. Soc., 71:4075. 1950 With E. W. Putnam. Transformation of sugars in plants. Annul Rev. Plant Physiol. 1: 109-24. With M. Doudoroff. Synthesis of disaccharides with bacterial en- zymes. Advances in Enzymology, 10:123-43. With E. W. Putman, A. L. Potter and R. Hodgson. The structure of crown-gall polysaccharide. I. l. Am. Chem. Soc., 72:5024. Enzymatic synthesis of sucrose and other disaccharides. Advances in Carbohydrate Chemistry, 5:29~8.
220 BIOGRAPHICAL MEMOIRS 1951 With A. L. Potter. Starch. IV. The molecular constitution of amylose subfractions. l. Am. Chem. Soc., 73:593. With A. L. Potter. Starch. V. The uniformity of the degree of branching in amylopectin. J. Am. Chem. Soc., 73:997. With J. Katz. Arsenolysis of amylose and amylopectin. Arch Bio- chem. 30:272-81. With D. A. Rappoport and H. A. Barker. Fermentation of L- arabinose-l-C~4 by Lactobacillus pentoaceticus. Arch. Biochem. Biophys., 31:326. With S. Nussenbaum. Enzymatic synthesis of amylopectin. I. BioI. Chem. 190:673-83. With D. A. Rappoport. Preparation of L-arabinose-l-Ci4. J. Am. Chem. Soc., 73:5524. Metabolism of polysaccharides and disaccharides. In: Phosphorus Metabolism, ed. W. D. McElroy and B. Glass, vol. 1, pp. 11~2. Baltimore: Johns Hopkins Press. With H. A. Barker. Degradation and synthesis of complex carbo- hydrates. In: Bacterial Physiology, ed. C. H. Werkman and P. W. Wilson, pp. ~548-65. N.Y.: Academic Press. With M. Doudoroff and H. A. Barker. Phosphorylases phosphor- olysis and synthesis of saccharides. In: The Enzymes, ed. l. B. Sumner and K. Myrback, vol. 1, part 2, 101~39. N.Y.: Academic Press. 1952 With S. Nussenbaum. Estimation of molecular weight of starch poly- saccharides. Determination of their reducing end groups. Analyt- ical Chemistry, 24:501. With S. Abraham and I. L. Chaikoff. Conversion of Ci4-palmitic acid to glucose. II. Specific glucose carbons labeled. I. Biol. Chem., 195:567-81. With E. W. Putman. Isolation and purification of radioactive sugars by means of paper chromatography. J. Biol. Chem., 196:749-52. With S. Nussenbaum. Mechanism of amylopectin formation by the action of Q-enzyme. I. Biol. Chem., 196:785-92. With S. Abraham and E. W. Putman. Distribution of radioactivity
WILLIAM ZEV HASSID 221 in photosynthetically prepared Ci4-labeled glucose. Arch. Bio- chem. Biophys., 41: 61-63. 1953 Starch. In: Organic Chemistry, An Advanced Treatise, ed. H. Gil- man, vol. 4, pp. 901-50. N.Y.: John Wiley and Sons. With R. C. Bean, E. W. Putman, and R. E. Trucco. Preparation of i4C-labeled D-galactose and glycerol. J. Biol. Chem., 204:169. Mechanisms of complex saccharide formation. Biologisch iaarboek, 20: 50-55. 1954 With E. W. Putman. Structure of galactosylglycerol from Irideae laminarioides. J. Am. Chem Soc., 76:2221. With M. Heidelberger and M. D. Aisenberg. Glycogen, an immuno- logically specific polysaccharide. journal of Experimental Medi- cine, 99:343-53. With W. H. Wadman and A. B. Anderson. The structure of an arabogalactan from Jeffrey Pine (Pinus [eJ5Freyi). J. Am. Chem. Soc., 76:4097-100. Biosynthesis of complex saccharides. In: Chemical Pathways in Metabolism, ed. D. M. Greenberg, pp. 235-75. N.Y.: Academic Press. With E. W. Putman. Sugar transformation in leaves of Canna indica. I. Synthesis and inversion of sucrose. l. BioI. Chem., 207: 885-902. 1955 With R. C. Bean. Assimilation of C14O2 by a photosynthesizing red alga, Iridophycus flaccidum. J. Biol. Chem., 212:411-25. With ~. Edelman and V. Ginsburg. Conversion of monosaccharides to sucrose and cellulose in wheat seedlings. l. Biol. Chem., 213: 843-54. With R. C. Bean. Glucose oxidase in Iridophycus flaccidum. Fed. Proc. 14:179. With E. W. Putman and C. F. Litt. The structure of D-glucosyI-D- xylose synthesized by maltose phosphorylase. l. Am. Chem. Soc., 77:4351.
222 BIOGRAPHICAL MEMOIRS With R. C. Bean. Synthesis of disaccharides with pea preparations. J. Am. Chem. Soc., 77:5737-38. With R. C. Bean. Assimilation of C~4-labeled carbon dioxide by a photosynthesizing red alga, 1. Iaminarioid ies. Fed. Proc., 13: 179. With R. McCready. a-D-Glucose 1-phosphate. In: Biochemical Preparations, ed. W. W. Westerfield et al., vol. 4. pp. 63-70. N.Y.: John Wiley and Sons, Inc. 1956 With E. F. Neufeld. Hydrolysis of amylose by p-amylase and Z- enzyme. Arch. Biochem. Biophys., 59:405-519. With R. C. Bean. Carbohydrate oxidase from a red alga, Iridophycus f accid um. J. Biol. Chem., 218:425-36. With V. Ginsburg and P. K. Stumpf. Uridine diphosphate pentoses in mung bean seedlings. Fed. Proc., 15:262. With E. W. Putman and V. Ginsburg. Metabolism of galactose in Canna leaves and wheat seedlings. Biochem. Biophys. Acta, 20: 17-22. With V. Ginsburg and E. F. Neufeld. Enzymatic synthesis of uridine diphosphate xylose and uridine diphosphate arabinose. Proc. Natl. Acad. Sci. U.S.A., 42:333-35. With R. C. Bean. Enzymatic oxidation of glucose to gIucosone in a red alga.Science, 124:171-72. With V. Ginsburg. Pentose metabolism in wheat seedlings. J. Biol. Chem., 223:277-84. With V. Ginsburg and P. K. Stumpf. The isolation of uridine di- phosphate derivatives of D-glucose, D-galactose, D-xylose and L-arabinose from mung bean seedlings. J. Biol. Chem., 223: 977-83. 1957 With i. Solms and D. S. Feingold. Uridine diphosphate N-acetyl- glucosamine and uridine diphosphate glucuronic acid in mung bean seedlings. J. Am. Cl~em. Soc., 79:2342-43. With S. Abraham. Chemical procedures for analysis of polysaccha- rides. In: Methods in Enzymology, ed. S. P. Colowick and N. O. Kaplan, vol. 3, pp. 3~50. N.Y.: Academic Press.
WILLIAM ZEV HASSID 223 With R. M. McCready. Preparation of a-D-glucose 1-phosphate by means of potato phosphorylase. In: Methods of Enzymology, ed. S. P. Colowick and N. O. Kaplan, vol. 3, pp. 137~3. N.Y.: Aca- demic Press. With E. F. Neufeld, V. Ginsburg, E. W. Putman, and D. Fanshier. Formation and interconversion of sugar nucleotides by plant extracts. Arch. Biochem. Biophys., 69:603-16. With E. W. Putman. Anomeric 1-dicyclohexylammonium phosphate esters of D-glucopyranose, D-galactopyranose, D-xylopyranose, and L-arabinopyranose. I. Am. Chem. Soc., 79:5057. With l. Solms. Isolation of uridine diphosphate N-acetylglucosamine and uridine diphosphate glucuronic acid from mung bean seedlings. l. Biol. Chem., 228:357-64. With C. E. Ballou. Phosphate esters. In: The Carbohydrates, ed. W. Pigman, pp. 172-87. N.Y.: Academic Press. With C. E. Ballou: Oligosaccharides. In: The Carbohydrates, ed. W. Pigman, pp. 473-533. N.Y.: Academic Press. 1958 Enzymatic synthesis (including brief review of formation in photo- synthesis) and interconversion of the monosaccharides. In: Hand buch der Pflanzenthysio logic, ed. W. RuhI and, vol. . 6, pp. 47-62. Berlin: Springer Verlag. The synthesis and transformation of the oligosaccharides in plants, including their hydrolysis. In: Handbuch der PQanzenphysi- ologie, ed. W. Ruhland, vol. 6, pp. 125-36. Berlin: Springer Verlag. With E. F. Neufeld and D. S. Feingold. Enzymatic conversion of uridine diphosphate D-glucuronic acid to uridine diphosphate gal acturonic acid, uridine di phosphate xylose, and uridine di- phosphate arabinose. J. Am. Chem. Soc., 80:4430. With D. S. Feingold and E. F. Neufeld. Synthesis of a ,8-1,3-linked glucan by extracts of Phaseolus aureus seedlings. [. Biol. Chem., 233:783-88. With D. S. Feingold and E. F. Neufeld. Enzymic synthesis of uridine diphosphate glucuronic acid and uridine diphosphate galac- turonic acid with extracts from Phaseolus auresus seedlings. Arch. Biochem. Biophys., 78:401-6.
224 BIOGRAPHICAL MEMOIRS 1959 With D. S. Feingold and E. F. Neufeld. Xylosyl transfer catalyzed by an asparagus extract. l. Biol. Chem., 234:488-89. With E. F. Neufeld and D. S. Feingold. Sugar nucleotides in the interconversion of carbohydrates in higher plants. Proc. Natl. Acad. Sci. U.S.A., 45:905-15. With E. F. Neufeld and D. S. Feingold. Enzymic phosphorylation of D-glucuronic acid by extracts from seedlings of Phaseolus aureus. Arch. Biochem. Biophys., 83:96-100. 1960 With E. F. Neufeld and D. S. Feingold. Phosphorylation of D- galactose and L-arabinose by extracts from Phaseolus aureus seedlings. l. Biol. Chem., 235:906-9. With D. S. Feingold and E. F. Neufeld. The 4-epimerization and decarboxylation of uridine diphosphate D-glucuronic acid by extracts from Phaseolus aureus seedlings. l. Biol. Chem., 235: 910-13. With G. Kessler and D. S. Feingold. Utilization of exogenous sugars for biosynthesis of carbohydrates in germinating pollen. Plant Physiol., 35:505-9. With J. C. Su. Identification of guanosine diphosphate 1-galactose, guanosine diphosphate D-mannose, and adenosine 3',5'-pyro- phosphate from red alga, Porphyra perforate, J. G. Agardh. {. Biol. Chem., 235:36-7. Biosynthesis of complex saccharides. In: Metabolic Pathways, ed. D. M. Greenberg, vol. 1, pp. 251-300. N.Y.: Academic Press. Carbohydrate. In: Encyclopedia of Science and Technology, pp. 451- 58. N.Y.: McGraw-Hill. Monosaccharide. In: Encyclopedia of Science and Technology, pp. 578-83. N.Y.: McGraw-Hill. 1961 With G. Kessler, E. F. Neufeld, and D. S. Feingold. Metabolism of D-glucuronic acid by Phaseolus aureus seedlings. l. Biol. Chem., 236: 308-12. With W. M. Watkins. Enzymatic synthesis of lactose from uridine
WILLIAM ZEV HASSID 225 diphosphate D-galactose and D-glucose. Biochem. Biophys. Res. Commun., 5:260-64 With E. F. Neufeld, D. S. Feingold, S. M. Ilves and G. Kessler. Phosphorylation of D-galacturonic acid by extracts from germi- nating seeds of Phaseolus aureus. J. Biol. Chem., 236:3102-5. 1962 The biosynthesis of poIysaccharides from nuceloside disphosphate sugars. In: Biochemical Society Symposium No. 21, pp. 63-79. Cambridge, Eng.: Cambridge University Press. With Jong-Ching Su. Carbohydrates and nucleotides in the red alga Porphyra perforate. I. Isolation and identification of carbo- hydrates. Biochemistry, 1: 468-74. With l. C. Su. Carbohydrates and nucleotides in the red alga Porphyra perforate. II. Separation and identification of nucleo- tides. Biochemistry, 1:47~80. With W. Watkins. The synthesis of lactose by particulate enzyme preparations from guinea pig and bovine mammary glands. I. Biol. Chem., 237:1432~0. With E. F. Neufeld. Glycosidic bond exchange (survey). In: The Enzymes, ed. P. D. Boyer, H. Lardy, and K. Myrback, vol. 6, pp. 277-315. N.Y.: Academic Press. 1963 With G. A. Barber. In vivo oxidation of L-rhamnose and L-glucose by higher plants. Bulletin of the Research Council of Israel, 1 lA4:249-52. With R. B. Frydman. Biosynthesis of sucrose with sugar cane leaf preparations. Nature, 199:382-83. With R. B. Frydman and E. F. Neuleld. Thymidine diphosphate D-galactose pyrophoshorylase of Phaseolus aureus. Biochim. Biophys. Acta, 77:332-34. With E. F. Neufeld. Biosynthesis of saccharides from glycopyranosyl esters of nucleotides ("sugar nucleotides"~. In: Advances in Car- bohydrate Chemistry, ed. M. L. Wolfrom, vol. 18, pp. 309-56. N.Y.: Academic Press. With D. S. Feingold and E. F. NeufeId. Preparation of UDP-D- xylose and UDP-L-arabinose. In: Methods in Enzymology, ed.
226 BIOGRAPHICAL MEMOIRS S. P. Colowick and N. O. Kaplan, vol. 6, pp. 782-87. N.Y.: Academic Press. 1964 With D. S. Feingold and E. F. Neufeld. Enzymes of carbohydrate synthesis. In: Modern Methods of Plant Analysis, ed. H. F. Linskens, B. D. Sanwal, and M. N. Tracy, vol. 7, pp. 47~519. Berlin: Springer-Verlag. With D. B. E. Stroud. Synthesis of 4-keto sugar phosphates. Biochem. Biophys. Res. Commun., 15:65-69. With A. D. Elbein and G. A. Barber. The synthesis of cellulose by an enzyme system from a highe, plant. l. Am. Chem. Soc., 86: 309-10. With E. F. Neufeld. Quantitative determination of starch in plant tissues. In: Methods of Carbohydrate Chemistry, ed. R. L. Whistler, vol. 4, pp. 33-35. N.Y.: Academic Press. Carbohydrates. In: McGraw-Hill Yearbook of Science and Tech- nology, pp. 169-73. McGraw-Hill Book Co. With T. Y. Lin. Isolation of guanosine diphosphate D-mannuronic acid from the marine brown alga, Fucus gardneri Silva. l. Biol. Chem., 239:945~6. With Helene Babad. A soluble lactose-synthesizing enzyme from bovine milk. J. Biol. Chem., 239: PC946~8. With G. A. Barber. The formation of guanosine diphosphate D- glucose by enzymes of higher plants. Biochim. Biophys. Acta, 86:399~02. With R. A. Dedonder. The enzymatic synthesis of a (,B-1,2-~-linked glucan by an extract of Rhizobium japonicum. Biochim. Bio- phys. Acta, 90:239. With G. A. Barber and A. D. Elbein. The synthesis of cellulose by erlzyme system from higher plants. J. Biol. Chem., 239:4056. With G. A. Barber and A. D. Elbein. Synthesis of cellulose by an enzyme system from a higher plant. Sixth International Congress of Biochemistry, New York. (Abstract) 1965 Cellulose, synthesis of. In: McGraw-Hill Yearbook of Science and Technology,p. 137.N.Y.:McGraw-HillBook Co.
WILLIAM ZEV HASSID 227 With E. G. Castanera. Properties of uridine diphosphate D-glu- curonic acid decarboxyIase from wheat germ. Arch. Biochem. Biophys., 110:462-74. With S. Haq. Biosynthesis of sucrose phosphate with sugar cane leaf chloroplasts. Plant. Physiol., 40:591-94. With G. A. Barber. Synthesis of cellulose by enzyme preparations from the developing cotton boll. Nature, 207:295-96. Biosynthese van cellulose. Umschau in Wissenschaft und Technik. No. 21, p. 681. With l. B. Pridham. Biosynthesis of raffinose. Plant Physiol. 40: 98~86. 1966 With C. L. Villemez and T. Y. Lin. Biosynthesis of the polygalac- turonic acid chain of pectin by a particulate enzyme preparation from Phaseolus aureus seedlings. Proc. Natl. Acad. Sci. U.S.A., 54: 1626-32. With T. Y. Lin. Isolation of guanosine diphosphate uranic acids from a marine brown alga, Fucus gardneri Silva. J. Biol. Chem., 241:3282-83. With T. Y. Lin, A. D. Elbein, and l. C. Su. Substrate specificity in pectin synthesis. Biochem. Biophys. Res. Commun., 22:650-57. With A. D. Elbein. The enzymatic synthesis of a glucomannan. Biochem. Biophys. Res. Commun., 23:311-18. With Helene Babad. Soluble uridine diphosphate D-galactose: D- glucose ,6-4-D-galactosyltransferase from bovine milk. l. Biol. Chem., 241 :2672-78. With J. B. Pridham. A preliminary study on the biosynthesis of hemicelluloses. Proceedings of the Biochemical Society. Biochem- ical journal, 100:21 P. With C. L. Villemez and A. L. Swanson. Properties of a polygalac- turonic acid-synthesizing enzyme system from Phaseolus aureus seedlings. Arch. Biochem. Biophys., 116:446-52. With R. W. Bailey. Xylan synthesis from uridine-diphosphate-D- xylose by particulate preparations from immature corn cobs. Proc. Natl. Acad. Sci. U.S.A., 56: 1586-93. With T. Y. Lin. Pathway of alginic acid synthesis in the marine brown alga, Fucus gardneri Silva. l. Biol. Chem., 241:5284-97. Some aspects of sugar nucleotide metabolism. In: Current Aspects of
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