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Aspects of Nitrogen Metabolism in Polyporus

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Aspects of Nitrogen Metabolism in Polyporus ASPECTS OF NITROGEN METABOLISM IN POLYPORUS TUMULOSUS A THESIS SUBMITTED FOR THE DEGREE OF MASTER OF SCIENCE IN THE UNIVERSITY OF NEW SOUTH WALES BY JOHN FRANCIS WILLIAMS 8I0MEOICAL roH ^^UBRARIE^^y MIN JO * SCHOOL OF BIOLOGICAL SCIENCES UNIVERSITY OF NEW SOUTH WALES JANUARY 1957 TO DECEMBER I960 This work was carried out as a part-time study between January 1957 and December I960, in the School of Biological Sciences of the University of New South ./ales. The material incorporated in this thesis has not been submitted towards a degree in any other University. With the exception of the data in Table 19 and the phenolic acid analyses reported in Figure 1$, the results submitted are my own unaided work. Williams. TABLE OF CONTENTS Acknowledgements i Summary ii 1. INTRODUCTION 1 SURVEY OF LITERATURE - NITROGEN METABOLISM IN FUNGI Introduction 3 t 2. INORGANIC NITROGEN METABOLISM 4 Nitrate Assimilation 4 Nitrite Nitrogen 10 Hyponitrite Reduction 12 Hydroxylamine Reduction 14 Oxime Pathway 15 Ammonia Nitrogen 16 Nitrogen IS 3. THE UTILIZATION OF AMINO ACIDS BY FUNGI 19 The Catabolism of Amino Acids 20 Synthesis and Interconversion of Amino Acids in Fungi 24 (1) The Glutamic Acid Group 27 (2) The Aspartic Acid Group 36 (3) Lysine 42 (4) The Pyruvic Acid Group of Amino Acids 43 (5) Histidine 46 (6) The Serine Group 51 (7) The Aromatic Amino Acids 56 4. URSA AND UREIDES 62 The Occurrence of Urea and its Precursors in Fungi 62 5. THE METABOLISM OF THE NUCLEIC ACIDS AND THEIR CONSTITUENTS 70 The Degradation of Nucleic Acid Derivatives by Fungi 70 The Uptake, Interconversion and Synthesis of Purines and Pyrimidines by Fungi 75 6. PROTEINS AND PEPTIDES 55 Peptides of Fungi and Actinomycetes 91 7. AMINES 94 EXPERIMENTAL SECTION 5. PART A - MATERIALS AND METHODS 95 Section (1) The Test Organism 95 (2) The Culture Medium 99 (3) . Cultivation and Harvesting 100 (4) The Mycelium 101 (5) The Culture Liquor 102 (6) The Cell Wall Fraction 102 (7) The Mycelial Extract 103 (5) Acid Hydrolysis of Mycelium and Cell Wall Fraction 103 (9) Alkaline Hydrolysis of Mycelium and Cell Wall Fraction 104 (10) Analytical Methods other than Chromatography 104 (13.) Paper Chromatography 105 (a) The Qualitative Identification of Amino Acids 105 (b) The Sulphur-Containing Amino Acids 115 (c) Urea, Ureides and Guanidine Bases 116 (d) Amino Sugars 116 (e) Visualising Agents for the Guanidino Group of the ’’Unknown Amino Acid” 117 (12) The Quantitative Analysis of Amino Acids 117 The Estimation of Error and the Statistical Treatment of Results 124 9. PART B - METABOLIC STUDIES ON POLYPORUS TUMULOSUS 131 Physiological Aspects 132 Measurement of Growth 132 Growth of P. tumulosus on WD5 134 The Effect of Environmental Conditions 146 The Effect of Aeration 147 The Effect of Temperature 143 pH and Growth 146 The Optimal Concentration of Nutrients 149 The Amino Acid Composition of P, tumulosus Fractions 153 The Amino Acid Composition of the Culture Medium 171 The Cell Nall Protein 172 Urea and Ureides 175 The Unknown Amino Acid 173 REFERENCES 131 i ACKNOWLEDGEMENTS I wish to express thanks to my Supervisor, Professor B. J. Ralph, for his guidance and assistance during the course of this work, and to Mr. R. Crowden, formerly of this Department, for many instructive and helpful discussions and for his collaboration in the physiological studies of P. tumulosus. Miss E. Perry, Mr. F. Wickenden, Mr. P. Brady and Mr. R. Simson of the Research Laboratory, William Arnott Pty. Limited prepared and photographed the figures in this thesis and have my gratitude• Dr. S. Ratner (Public Health Research Institute, New York, U.S.A.) and Dr. F. Morrison, Australian National University kindly donated samples of Argininosuccinic acid and Lombricine. My thanks are due to Miss Pam Keyes, who typed the final draft of this thesis, and above all, thanks to my wife who, for four years, has shared the life of a part-time student. ii SUMMARY Principal aspects of the nutrition and metabolism of nitrogen in fungi are discussed. Physiological studies of the growth of P. tumulosus by still culture on modifications of William Saunders glucose-salts medium were made and have shown that in some circumstances limitation of optimal growth in the organism is the result of unfavourable carbon to nitrogen ratios. The accumulation of phenolic and certain amino acids in culture media and extracts of the organism, as shown by sequential growth studies, is consistent with such C/N imbalance. The amino acid composition of proteins of the mycelium and cell wall, together with that of the culture medium and mycelial extract, is described. It is established that an unidentified amino acid exists in the mycelial extract of this organism. The sequential appearance of this acid during growth and its chemical and possible biochemical relations to other amino acids are discussed. Ureidosuccinic acid was found on chromatograms of P. tumulosus mycelial extracts; this ureide has not previously been reported in fungi. Allantoin, Allantoic Acid and an unidentified ureide (possibly Allantoxanic or Uroxanic Acid) were found in extracts of autolysing cells. 1 INTRODUCTION This investigation was primarily undertaken as part of a comprehensive survey, by this Department, on the biochemistry of the wood-rotting Basidiomycete fungus, Polyporus tumulosus Cooke, and secondarily to form a supplementary study to a parallel investigation on the phenolic acid metabolism of this organism. Crowden (1) has identified aromatic compounds in P. tumulosus derived from the Shikimic acid path of biosynthesis and has established that many of these compounds bear a pronounced resemblance to tyrosine. With the provision of this background, research was directed to those aspects of nitrogen metabolism in this organism, which would relate to the nature and incidence of aromatic amino acids. The complementary nature of the metabolic studies on phenolic acids and nitrogen metabolism established the need to standardize culture conditions for the organism. The definition of the scope of this thesis limited the investigation to the identification of nitrogenous metabolites in the fractions; culture media, mycelia, and mycelial extracts, with a directive to attend in particular to amino acids. The metabolic studies of the organism cultured on WD5- glycine and WD5-alanine were dictated by an investigation of the pattern of phenolic acid side chain synthesis. Principally these studies were to serve as an introduction to 2 the role of nitrogen in these transformations and the establishment of the necessary techniques for the more comprehensive investigation of the organism when grov/n with a simple ammonium salts medium (WD5). A survey of some aspects of the literature on the metabolism and nutrition of nitrogen in fungi has been made. The recent comprehensive review of Cochrane (2) has provided the chief background against which this survey has been planned, but most of the material reviewed has been obtained from the sections dealing with nitrogen metabolism in the annual review series of Biochemistry, Microbiology and Plant Physiology. In addition, material not available from these periodicals has been obtained diirectly from various journals, Proceedings of Symposia, Chemical and Biological Abstracts. The review deals principally with those aspects of the nitrogen metabolism of fungi which are directly related to the experimental work. While the magnitude of certain aspects of the reported results may not appear to justify such a detailed literature survey, it was deemed necessary for adequate appraisal of these results. ASPECTS OF NITROGEN METABOLISM IN POLYPORUS TUMUL05US SURVEY OF LITERATURE Nitrogen Metabolism in Fungi - 3 - NITROGEN METABOLISM IN FUNGI INTRODUCTION Irrespective of the organism the continuance of life and the synthesis of cytoplasm are dependent on the availability of the same basic materials, namely sources of carbon, nitrogen, water, and mineral salts, together with some mechanism for providing energy in a form that can be utilized in biological systems. All autotrophs derive their nitrogen from an inorganic source and, depending on the organism, use molecular nitrogen, ammonium ion, nitrate or nitrite. The heterotrophs on the other hand have a specific requirement for one or more substances and are unable to grow in their absence from the culture medium. The need of a culture for nitrogen is not optimal. It is governed primarily by the supply of an energy yielding carbon compound. Steinberg and Bowling (3) in their studies with Aspergillus niger demonstrated this and established a strict proportionality between carbohydrate supply and nitrogen demand. Generally nitrogen is taken up from an organic or inorganic medium during the growth phase while amino acids and water soluble nitrogenous compounds, enzymes etc. are found during the latter stages of grov/th and autolysis. It has been found in studies on Scopularopsis brevicaulis that the organism utilizes nitrogen from autolysed cells to - 4 - synthesise new tissue (4). INORGANIC NITROGEN METABOLISM Under this heading will be reviewed the role of nitrates, nitrites and ammonium ion as nutrients and metabolites. Inorganic Nitrate Assimilation The assimilation of nitrate nitrogen by fungi involves the reduction of nitrate to ammonia, resulting in an oxidation-reduction change of the nitrogen atom from + 5 to -3, involving a net change of 8 electroins, as shown in Table No. 1 TABLE No. 1. Oxidation-Reduction States off Nitrogen and its Compounds Oxidation-Reduc tion Nitrogen State of N atom Compounds +6 n°3, n206 hn03 +4 no2, n204 +3 N203 hno2 +2 NO +1 N2°- H2N2°2 0 W2 -1 nh2oh -2 nh2 - NH2 -3 NH3 A number of enzymes, which catalyze the reduction nitrate to ammonia via nitrite and hydroxylamine, have 5 characterised in bacteria and fungi. Reduced pyridine nucleotides act as electron donors and all steps seem to involve metal-dependent flavo-proteins. The assumption is that each enzymic step involves a two electron or proton change.
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