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MICHEGAN STATE UNIVERSITY Evelyn Anne Horenstein 1965 THESIS LIBRARY Michigan Statfl University

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MICHEGAN STATE UNIVERSITY Evelyn Anne Horenstein 1965 THESIS LIBRARY Michigan Statfl University PHOTOMORPHGGENI’ESIS EN A NEW AQUATEC E’UNGUS BLASTQCLQMELLA BRHANNECA That. {01’ Hm Degree a? DB. D. MICHEGAN STATE UNIVERSITY Evelyn Anne Horenstein 1965 THESIS LIBRARY Michigan Statfl University This is to certify that the ' thesis entitled Photomorphogenesis in a new aquatic fungus, Blastocladiella britannica presented by Evelyn Anne Horenstein has been accepted towards fulfillment of the requirements for Eh . D. degree in _an:an.y_ WOW Major professor Date May 6, 1955 0-169 '73 K: $4 b4 w ‘p (1) Hi (I It (r L.' ' ' (1' m (1‘ 1. t- .1: :1 i) (1’ if '(1 ,1.) '(3 L). ‘1 ;< (U (i) L) ,I 7 » (I) ABSTRACT PHOTOMORPHOGENESIS IN A NEW AQUATIC FUNGUS BLASTOCLADIELLA BRITANNICA by Evelyn Anne Horenstein The developmental history in pure culture of a single— spore isolate of a Blastocladiella was investigated in some detail. No other species in the Blastocladiaceae which has been subjected to rigorous examination displays such a high degree of morphological variability--a variability which is probably phenotypically controlled. This characteristic led to the isolation of several distinct substrains, one of which was incapable of producing resistant sporangia and another (strain B 101) produced them in abundance. As a consequence of these and other observations, this single- celled fungus was designated a new species, B. britannica° In strain B 101, formation of resistant Sporangia was affected by several environmental factors, but most strik- ingly by white light. When grown on agar media, it produces: (a) in the dark, about 90-100% brown, thick-walled resistant sporangia (RS) with a generation time of about 65 hr.; (b) in white light, nearly colorless, thin-walled (TW) Sporangia with a generation time of about 30 hr. However, neither the absence nor the presence of light is required continuously t r «v- 6:- . v..- l-.__,‘rb... - l .4..- (ll Evelyn Anne Horenstein throughout the entire growth period for the genesis of one or the other of these morphological forms° The organism's early stages of development are quite plastic; cells which start their growth in the light, and which are, therefore, on the TW pathway, can be induced to revert to RS types by eliminating the light. Conversely, cells which start growth in the dark (i.e. along the RS pathway) can be transformed into TW types by exposure to light. In both instances, however, a stage in development is reached beyond which addition or withdrawal of illumination can no longer effect morphogenetic reversal. At this point of no return, the cells have become committed to one pattern or the other. To study photomorphogenesis at a chemical level, it was necessary to grow the organism in large quantities under conditions which induced reproducible morphological uniform- ity and which could be controlled as precisely as possible. Consequently, a method was devised for growing synchronized, single generations of B. britannica (a million or more cells at a time) uniformly suspended in agitated media, wherein the effects of light and dark on morphogenesis were demon- strable as well as the reversal of morphogenesis by altera- tion of the light and dark regime before a point of no return. Up to a stage just preceding the end of the generation time of a TW sporangium, no morphological differences are discernible under the light microscope between it and a (I! V [u {3 t ll) a. ,‘ () cu-.. _. l 9‘ H ‘0 h I ”\V‘c.rl ssh-l who» . At.” ,t \t (3" -..~..- . V . U ‘ ‘4 (-u W \— ‘ o M 9 s v- K x ‘ t. \A s y ,— 9.. Evelyn Anne Horenstein dark-grown developing RS at a corresponding chronological age. Yet, within the next 1-2 hr., the entire protoplast of a TW is cleaved into hundreds of uninucleate, uniflagellate motile spores, and this new generation of cells is then dis- charged. On the other hand, the thalli growing in the dark have reached only the half-way point in their ontogeny; they continue to enlarge for several hours thereafter and then gradually differentiate into mature RS. In synchronous cultures, dry weight/cell increases exponentially at the same rate in light and dark. On the other hand, the capacity for uptake of glucose by cells of various ages grown in the dark exceeds that of light-grown cells. Furthermore, just as the course of development can be reversed by excluding or supplying light before their respective points of no return, so, too, their capacities for glucose uptake can be similarly reversed. However, the point of no return for glucose uptake precedes the point of no return for morphogenesis by several hours. The light- sensitive glucose uptake by B. britannica may be a factor in the determination of the ultimate morphology of this organism. PHOTOMORPHOGENESIS IN A NEW AQUATIC FUNGUS BLASTOCLADIELLA BRITANNICA BY Evelyn Anne Horenstein A THESIS Submitted to Michigan State University in partial fulfillment of the requirements for the degree of DOCTOR OF PHILOSOPHY Department of Botany and Plant Pathology 1965 ACKNOWLEDGMENTS I am very grateful to Dr. L. G. Willoughby who graciously sent us the culture of Blastocladiella britannica, thus affording me the opportunity to study this organism. Thanks are due to Professor William M. Seaman, who kindly provided the Latin translation for the diag- nosis of Blastocladiella britannica. For his capable guidance, his never-failing en- couragement and expressions of confidence, and his enthusiastic support, I should like to express my very deepest gratitude to Professor Edward C. Cantino who, more than anyone, is responsible for the successful completion of this thesis. *************** ii To Mother, Dad, and Don iii TABLE OF CONTENTS Page INTRODUCTION . 1 LITERATURE REVIEW. 4 Description of the Blastocladiaceae with Emphasis on Unique Characteristics of the Family. 4 The motile spore . 6 The resistant sporangium . 10 Life Cycles in the Blastocladiales . 12 Brachyallomyces type . 12 Cystogenes type. 12 Euallomyces type . 15 Differentiation of the Resistant Sporangium in Blastocladiella emersonii. 15 Induction of resistant sporangium formation. 15 Formulation of a hypothesis. 15 Biochemistry of morphogenesis. 19 Period of exponential growth . 22 Period of differentiation. 27 MATERIALS AND METHODS. 32 Culture Procedures . 52 Analytical Procedures. 37 OBSERVATIONS AND EXPERIMENTAL. 41 Conditions for Optimal Growth. 41 Morphological Variability. 42 Pure Strains of TW Colonies. 47 The RS Colonial Strain . 50 Life history . 50 Effect of temperature on RS formation. 51 Effect of bicarbonate on RS formation. 51 Effect of glucose on RS formation. 52 Effect of light on RS formation. 54 Isolation of RS strains yielding 100% indi- vidual RS plants . 54 Development of Synchronous Cultures of Resistant Sporangia. 60 Growth and Morphological Characteristics of Syn- chronized Single Generation Cultures . 68 iv TABLE OF CONTENTS - Continued Page Growth pattern in the dark 70 Growth pattern in the light. 71 Point of no return in development. 76 Glucose Uptake Capacity. 79 Reversal of GUC. 81 The point of no return for GUC 82 DISCUSSION . o . The Taxonomic Position of Blastocladiella britan- nica O O O O O O I O O O O 85 The Aquatic Phycomycetes and Biological Research Morphogenesis in Blastocladiella britannica. 95 Morphological variability. 95 Photomorphogenesis . 97 SUMMARY. 101 LIST OF REFERENCES . 102 APPENDICES . 108 LIST OF TABLES TABLE Page Lactic acid production by B. britannica. 41 II Morphological types produced by B. britannica. 46 III Relationships among volume per cell, weight per cell, and weight per unit volume of TW cells at maturity. 76 IV GUC and intracellular pools. 117 Effect of phosphate on GUC . 119 Effect on GUC of different sugars during growth . 120 VII Effect on GUC of the presence of other sugars. 122 vi LIST OF FIGURES FIGURE Page 1. Life cycles in the Blastocladiales . 14 2. Enzyme reversal in B. emersonii. 24 5. Comparative activities of isocitritase in syn— chronously developing OC and RS plants of B. emersonii during ontogeny . 26 Transformations in total RNA during morpho- genesis of RS. 28 Transformations in composition of RNA during . sol morphogenes1s of RS. 28 Changes in protein nitrogen, plant volume, and dry wt. during RS morphogenesis in B. emersonii. 50 Increase in chitin and melanin content during RS differentiation in B. emersonii . 50 Changes in lactate pool, glucose-6-ph05phate dehydrogenase, glucose~consumption, and poly— saccharide pool during RS differentiation in B. emersonii . 50 Developmental potentialities in B. britannica. 45 10. Size of spores from the TW and RS strains. 48 11. Effect of different concentrations of PYG (2% agar) on the composition of populations derived from spores of TW colonies . 12. Effect of different concentrations of PYG (2% agar) on the composition of populations derived from spores of RS colonies . 55 15. Effect of different concentrations of glucose (in PYG) on RS formation . 55 14. Effects of semi-anaerobic conditions on the composition of populations derived from spores of RS colonies . 55 vii .,4 ~I) L!) IL. '4') r\l «\1 1‘\’ U) ('\] [14 LIST OF FIGURES - Continued FIGURE Page 15. Effect of light on RS plant development. 55 16. The relation between duration of light and RS formation on medium PYG. 56 17. Results of attempts to select populations of vigorous RS producers. 59 18. Growth curves of individual RS plants. 59 19. Incidence of RS after transfer from liquid to solid media. 64 20. Growth of RS cells in the dark . 72 21. Effect of pH upon the growth rate of TW cells in the light. 72 22. Effect of population density upon the growth of TW cells . 74 25. Effect of population density upon the volume of TW cells at generation time. 74 24. Effect of population density upon the generation time of TW cells . 75 25. Relationship between the volume of the TW cell at generation time and its dry wt.
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