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A Filter Paper Assay for Low Cellulase Activities and the Cultivation of Trichoderma Reesei on Acid Whev and Sweet Whev Permeate

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A Filter Paper Assay for Low Cellulase Activities and the Cultivation of Trichoderma Reesei on Acid Whev and Sweet Whev Permeate AN ABSTRACT OF THE THESIS OF Tor Soren Nordmark for the degree of Master of Science in Food Science and Technology presented on November 24. 1993 . Title: A Filter Paper Assay for Low Cellulase Activities and the Cultivation of Trichoderma reesei on Acid Whev and Sweet Whev Permeate Abstract approved: Michael H. Penner The traditional filter paper assay for saccharifying cellulase originally described by M. Mandels et al (1976) has been modified to make possible low activity determinations of Trichoderma cellulases. The enzymatic activity appears to decline during a prolonged incubation period if no precautions have been taken. By means of adding bovine serum albumin and potassium chloride as protein stabilizers and sodium azide as an antimicrobial agent filter paper activities in the range from 0.02 to 0.37 (IUPAC assay, 1987) can be estimated by extending the incubation time up to 20 hours. Filter paper activity values obtained by this method may be compared to those obtained by the IUPAC assay by using a conversion factor from 1.4 to 1.7. Acid whey and sweet whey permeate have been investigated as media for growth and metabolite production by Trichoderma reesei QM 9414 using shake flask cultures and spore inocula. In the case of acid whey the mycelial growth after 2 weeks is 13 mg dry weight /ml substrate. The specific growth rate is 0.29 / day. The fungus appears to metabolize the whey protein the first 2 weeks. The alkalinity of acid whey rises continuously over a three week period up to a pH of 8.5. In the case of whey permeate the maximal mycelial weight gain is 4.4 mg/ml which appears after 8 days. A rise in net soluble protein level comes after 3-5 days and reaches a maximum value of 0.23 mg/ml after 2 weeks. The pH of whey permeate rises continuously to 7.5 after 3 days and then slowly declines. The net production of cellulases is low on both media. Dilution 1:6 of the acid whey, supplementation with ammonium sulfate and pH- adjustments did not enhance the production of cellulases. Acid whey supports a significant growth and sweet whey permeate shows potential for extracellular protein production. A literature review surveys the composition and uses of acid whey, environmental aspects of whey wastes, the fungus Trichoderma reesei, the mode of action of the Trichoderma reesei cellulase system and the structure of cellulose in cotton and wood. A Filter Paper Assay for Low Cellulase Activities and the Cultivation of Trichoderma reesei on Acid Whey and Sweet Whey Permeate by Tor Soren Nordmark A THESIS submitted to Oregon State University in partial fulfillment of the requirements for the degree of Master of Science Completed November 24, 1993 Commencement June 1994 APPROVED: Professor of Food Science and Technology in charge of major y" 1r \ >lead of the Department of Food Science and Technology ,ry in v — | i »ii \y t . i Dean of GrackJaie Schoo} \ Date thesis is presented November 24. 1993 Typed by Tor Soren Nordmark Acknowledgement I wish to thank Dr. Michael Penner for encouragement and supportive ideas throughout the experimental work as well as assistance during the preparation of this thesis. His ability to give me freedom in determining research approaches and topics also stimulated the progress. Dr. Alan Bakalinsky offered me laboratory space and added valuable views to my microbiological perspectives. This work has been supported by funds from the Western Dairy Foods Research Center and the Oregon State University Agricultural Experiment Station to which I wish to express my gratitude. Finally I would like to express my gratefulness to Ean-Tun Liaw and other colleagues in the department who generously made arrangements in their own work in order to accommodate for my experiments and with whom I had many helpful discussions. I I Table of Contents 1. General Introduction and Literature Review 1.1 Project goals 1 1.2 Background 1.2.1 Environmental aspects on whey wastes 2 1.2.2 Whey and whey permeate 4 1.2.3 Industrial processing of whey media 9 1.2.4 The microfungus Trichoderma reesei 11 1.2.5 The Trichoderma reesei cellulase system 14 1.2.6 Cellulose structure and cellulase action 16 1.2.7 The measurement of cellulolytic enzyme 2 2 activity 1.3 Research outline 24 2. The Application of the Filter Paper Assay for Cellulase at Low Activity Levels 2.1 Introduction 26 2.2 Materials and methods 2.2.1 Materials 30 2.2.2 Enzyme preparation 3 0 11 I 2.2.3 Enzymatic hydrolysis 31 2.3 Results and discussion 2.3.1 Principles of an improved assay 3 3 2.3.2 Activity development studies 3 8 2.3.3 Stabilization studies 44 2.4 Conclusion 50 3. Cultivation of Trichoderma reesei on Acid Whey and Sweet Whey Permeate 3.1 Introduction 51 3.2 Materials and methods 3.2.1 Substrates 54 3.2.2 Microbe 55 3.2.3 Incubation experiments 55 3.2.4 Reagents and analytical procedures 56 3.3 Results and discussion 3.3.1 The development of mycelial biomass 58 3.3.2 Metabolite production and effects on 61 substrate composition. 3.4 Final comments 80 4. Bibliography 82 Appendices A. A protocol for the maintenance and handling of 9 7 Trichoderma reesei cultures on potato dextrose agar B. Spore counting on PDA plates by using Rose Bengal 102 C. A protocol for the modified IUPAC filter paper 103 assay D. An experiment demonstrating the modified IUPAC 109 filter paper assay E The pigment production by Trichoderma reesei 113 on liquid media F. A protocol for preincubation and postincubation 114 processing when recovering proteins on whey media List of Figures Fig. 1. The taxonomy of T. longibrachiatum. 1 3 Fig. 2. The Trichoderma reesei cellulase system. 15 Fig. 3. Cellulose structure. 17 Fig. 4. Attachment of cellulases to the cellulose. 19 Fig. 5. Modes of cellulase action. 21 Fig. 6. The relation between the amount of glucose 36 released in a filter paper assay and the logarithm of (A) the enzyme concentration and (B) the incubation time. Fig. 7. The level of reducing sugar as glucose produced by 4 0 Trichoderma reesei QM 9414 (A) and viride (B) cellulase during a time extended filter paper assay in the absence of stabilizers plotted against the product of enzyme concentration (E as dilution level) and time (t in hours). Fig. 8. The amount of reducing sugar as glucose released 41 by Trichoderma viride cellulase at different concentrations of non-macerated Whatman #1 filter paper and cellulase activities. Fig. 9. Incubation of Trichoderma reesei QM 9414 cellulase 4 3 when the substrate (Whatman #1 filter paper) is either (A) always present or (B) present only during an assay hour ending by the times shown. In case A four groups of different dilutions each cover the points where E x t is equal to the original value, whereas in case B all enzyme is undiluted. V I Fig. 10. The level of reducing sugar as glucose produced by 4 7 Trichoderma reesei (A) and viride (B) cellulase during time extended filter paper assays when stabilizers are used plotted against the product of enzyme concentration (E as dilution factor) and time (t in hours). All standard deviation bars are covered by the symbols. Fig. 11. The result of filter paper assays (triplicate 48 samples) using stabilized (S) and non-stabilized (NS) Trichoderma reesei (A) and viride (B) cellulase. The conversion factors (f) between the apparent activities are 1.4 and 1.7 respectively. Fig. 12. The growth of T. reesei on filter-sterilized acid 59 whey. Fig. 13. The growth of T. reesei on filter-sterilized whey 6 2 permeate. Fig. 14. The development of pH when T. reesei is cultivated 6 3 on thermally sterilized acid whey. Fig. 15. The development of soluble protein as BSA when T. 6 5 reesei is grown on thermally sterilized acid whey. Fig. 16. The development of soluble protein as BSA when T. 6 6 reesei is grown on filter-sterilized acid whey. Fig. 17. The amount of glucose released in a filter paper 6 8 assay (IUPAC) when T. reesei is cultivated on filter-sterilized acid whey. Fig. 18. The time course of pH when T. reesei is grown on 6 9 filter-sterilized acid whey that has been diluted 1:6. VI I Fig. 19. The development of pH when T. reesei is grown on 71 filter-sterilized sweet whey permeate. Fig. 20. The development of soluble protein as BSA when T. 7 2 reesei is grown on filter-sterilized sweet whey permeate. Fig. 21. The amount of glucose released in a filter paper 74 assay (IUPAC) when T. reesei is cultivated on filter-sterilized sweet whey permeate. Fig. 22. The effect of 0.1 % (w/v) Solkafloc addition on the 76 level of soluble protein as BSA when T. reesei is grown on filter-sterilized sweet whey permeate Fig. 23. The change in lactose level with and without added 78 0.1 % (w/v) Solkafloc when T. reesei is grown on filter-sterilized sweet whey permeate. Fig. 24. If 0.001 % Rose Bengal is added to the PDA then the 102 7". reesei colonies will appear more distinct and limited as can be seen from the photos above. Higher concentrations can kill the fungus. Fig. 25. The amount of reducing sugar as glucose released 111 in a modified filter paper assay for different incubation times and for 2 dilutions. T. viride cellulase is used. VIII List of Tables Table 1.
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