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MICROBIAL FERMENTATIONS IN NATURE AND AS DESIGNED PROCESSES Fermentation is one of the most important metabolic tools that biology has developed and microorganisms in many ways seem to have become the true masters of fermentative metabolism. Each of the fermentative microbial functions evolved to fit an energetic opportunity, and each function has ecological value. This book provides its readers with: * Understanding regarding the commonalities and distinctions between aerobic and anaerobic fermentations as performed by microorganisms. * A summary of knowledge regarding the ways in which animals and plants depend upon symbiotic interactions with their fermenting microbial partners including the deconstruction of complex polysaccharides. Information is also included about how those natural technologies constitute adaptation into designed processes for anaerobic degradation of lignocellulosic materials. * The important role of rhizosphere microbes that facilitate availability of inorganic and organic phosphates for plants. These phosphates get stored in the plant's seeds. After ruminant animals ingest the seeds, enzymes produced by gastrointestinal microbial fermentation allow the animals to utilize their dietary phosphates. * History of how microbial fermentation has been harnessed from prehistoric times to the present for processing and preserving food products for humans and fodder for our domesticated animals. * Insight into the ways that microbial fermentations are used as an engineering tool for producing chemicals, including enzymes and pharmaceuticals, which improve the health of ourselves and our domesticated animals. * Perspectives on possible future research directions for the field of applied microbial fermentation that will help to advance agriculture and industry.

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Table of Contents

COVER

TITLE PAGE

COPYRIGHT PAGE

DEDICATION

LIST OF FIGURES

LIST OF CONTRIBUTORS

PREFACE

SECTION I: AN INTRODUCTION TO MICROBIAL FERMENTATION

CHAPTER 1: MICROBIAL FERMENTATION

1.1 INTRODUCTION TO FERMENTATION PROCESSES

1.2 EXAMPLES OF MICROBIAL FERMENTATIONS IN NATURE

1.3 DESIGNING FERMENTATION PROCESSES

1.4 THE DISPOSAL OF WASTE PRODUCTS FROM FERMENTATIONS

1.5 FERMENTATIONS THAT PRODUCE AND PRESERVE FOOD FOR HUMANS

1.6 SILAGE FERMENTATIONS THAT PRODUCE AND PRESERVE FOOD FOR LIVESTOCK

1.7 TOBACCO FERMENTATIONS

1.8 COMPOST FERMENTATIONS AND THE ANAEROBIC DIGESTION OF FOOD WASTES FOR GENERATING BIOGAS

1.9 FERMENTATIONS THAT PRODUCE CHEMICALS, ENZYMES, AND PHARMACEUTICALS

1.10 SUMMARY

REFERENCES

CHAPTER 2:

RHODOTORULA TORULOIDES

AS A BIOFACTORY OF CAROTENOIDS, LIPIDS, AND ENZYMES

2.1 INTRODUCTION

2.2

RHODOTORULA TORULOIDES

2.3 MECHANISM OF LIPID BIOSYNTHESIS

2.4 FACTORS AFFECTING INTRACELLULAR LIPID BIOSYNTHESIS

2.5 CAROTENOID BIOSYNTHESIS BY

RHODOTORULA TORULOIDES

2.6 MECHANISM OF CAROTENOID BIOSYNTHESIS

2.7 STRATEGIES TO INCREASE CAROTENOID SYNTHESIS

2.8 BIOSYNTHESIS OF PHENYLALANINE AMMONIA‐LYASE BY

R. TORULOIDES

2.9 CONCLUSIONS AND FUTURE TRENDS

REFERENCES

SECTION II: THE ROLE OF MICROBIAL FERMENTATION IN DIGESTION PROCESSES

CHAPTER 3: GUT MICROBIAL ECOLOGY

3.1 INTRODUCTION

3.2 VARIATION ALONG THE GUT

3.3 PHYLOGENY, GUT MORPHOLOGY, AND FEEDING STRATEGY SHAPE MICROBIAL COMMUNITIES

3.4 A PREDICTIVE ECOLOGICAL FRAMEWORK FOR THE GUT

3.5 APPLICATIONS TO HOST MANAGEMENT

3.6 CONCLUSION

REFERENCES

CHAPTER 4: FERMENTATION IN THE RUMEN

4.1 INTRODUCTION

4.2 NUTRITIONAL PHYSIOLOGY OF RUMINANTS

4.3 THE RUMEN MICROBIAL COMMUNITY

4.4 BIOCHEMISTRY OF RUMEN FERMENTATION

4.5 MANIPULATION OF RUMEN FERMENTATION

4.6 CONCLUSIONS

REFERENCES

CHAPTER 5: MICROBIAL DEGRADATION OF LIGNOCELLULOSE IN NATURAL AND ENGINEERED SYSTEMS – FROM THE SMALLEST TO THE BIGGEST BIOREACTOR

5.1 INTRODUCTION TO LIGNOCELLULOSE AND ITS DEGRADATION IN VARIOUS SYSTEMS

5.2 ENZYMATIC MECHANISMS OF LIGNOCELLULOSE DEGRADATION AND CARBOHYDRATE‐ACTIVE ENZYMES

5.3 ANAEROBIC DIGESTION OF PLANT BIOMASS

5.4 OVERVIEW OF THE MOST EFFICIENT LIGNOCELLULOSE‐DEGRADING SYSTEMS DEVELOPED BY NATURE

5.5 FROM COMPARISON TO BIOPROSPECTING: WHAT CAN WE LEARN FROM THE TERMITE GUT SYSTEM TO IMPROVE INDUSTRIAL LIGNOCELLULOSE BIOPROCESSING

5.6 BEYOND THE STANDARD LIGNOCELLULOSE DEGRADATION: NEW APPLICATIONS AND FUTURE CHALLENGES

5.7 CONCLUSIONS

ACKNOWLEDGMENTS

REFERENCES

SECTION III: USING MICROBIAL FERMENTATION TO PRODUCE BEER AND WINE

CHAPTER 6: ARCHEOLOGICAL EVIDENCE FOR FERMENTED ALCOHOLIC BEVERAGES IN RITUAL FEASTS OF NEOLITHIC CHINA

6.1 INTRODUCTION

6.2 METHODS: IDENTIFYING BREWING METHODS IN NEOLITHIC CHINA

6.3 BREWING BEER AND NEOLITHIC REVOLUTION IN THE EARLY NEOLITHIC (ca. 9000 TO 7000 cal. BP)

6.4 DEVELOPMENT OF DIVERSE BREWING METHODS IN THE MIDDLE NEOLITHIC (ca. 7000 TO 4600 cal. BP)

6.5 INDIVIDUALIZED DRINKING TRADITION AND DEVELOPMENT OF SOCIAL COMPLEXITY IN THE LATE NEOLITHIC (4600 TO 3800 BP)

6.6 DISCUSSION AND CONCLUSIONS

ACKNOWLEDGEMENTS

REFERENCES

CHAPTER 7: BREWS OF THE PAST: BIOARCHAEOLOGY OF MICROBIAL FERMENTATION

7.1 INTRODUCTION

7.2 HISTORIC ALE AND BEER

7.3 GOLDEN AGES OF BREWING

7.4 EARLY BEERS. EGYPTIAN AND MESOPOTAMIAN

7.5 GRUIT ALES

7.6 MONASTIC BREWING AND EARLY‐HOPPED ALES

7.7 VICTORIAN AND EDWARDIAN HIGH‐HOPPED BEERS

7.8 CONCLUSIONS

REFERENCES

CHAPTER 8: AN ACCIDENTAL ART FORM: SPONTANEOUS FERMENTATION OF BEER

8.1 INTRODUCTION

8.2 FERMENTATION AND ITS SPONTANEITY

8.3 MUCH A BREW ABOUT NOTHING

8.4 CURTAIN CALL

REFERENCES

CHAPTER 9: JAPANESE TRADITIONAL FERMENTATION USES SOLID‐STATE FUNGAL CULTIVATION TO PRODUCE SAKE

9.1 INTRODUCTION TO TRADITIONAL JAPANESE FERMENTATIONS PERFORMED USING

ASPERGILLUS ORYZAE

9.2 PREPARATION OF RICE KOJI BY CONTROLLED FUNGAL GROWTH

9.3 FERMENTATION OF THE RICE AS A MASH TO PRODUCE ETHANOL

9.4 SUMMARY

REFERENCES

SECTION IV: USING MICROBIAL FERMENTATION TO PRODUCE FOOD AND FODDER

CHAPTER 10: SAFETY DEMONSTRATION OF FOOD AND FEED CULTURES

10.1 INTRODUCTION

10.2 FERMENTED FOOD PRODUCTS AND INTERNATIONAL TRADE

10.3 FOOD FERMENTATION. FOOD AND FEED CULTURES

10.4 HISTORY OF (SAFE) USE

10.5 SAFETY CONSIDERATIONS

10.6 OPPORTUNISTIC INFECTIONS BY LACTIC ACID BACTERIA

10.7 ANTIBIOTIC RESISTANCE

10.8 FOOD FERMENTATION AS END USE OF MICROBIAL FOOD CULTURES: HISTORY OF SAFE USE

10.9 BENEFIT – RISK ASSESSMENT OF FOOD AND FEED CULTURES

10.10 UNDERLYING FACTORS AND SPECIFIC HEALTH CONDITIONS OF CONCERN VS. OPPORTUNISTIC INFECTIONS

10.11 OPPORTUNISTIC INFECTIONS AND MICROBIAL FOOD CULTURES: GUIDELINES TO HEATH CARE PRACTITIONERS

10.12 FOOD FERMENTATION AS END USE OF FOOD AND FEED CULTURES

10.13 CONCLUSION

REFERENCES

CHAPTER 11: STARTER CULTURES AND THEIR ROLE IN FERMENTED FOODS

11.1 INTRODUCTION

11.2 FERMENTATION

11.3 FERMENTED FOODS

11.4 STARTERS

11.5 THE ROLE OF STARTERS IN FOOD SAFETY

11.6 THE ROLE OF STARTERS IN FOOD SENSORY PROPERTIES

11.7 STARTERS PROMOTE INNOVATION IN FERMENTED FOODS

11.8 THE HEALTH BENEFITS OF FERMENTED FOODS

11.9 CONCLUSIONS

ACKNOWLEDGMENTS

REFERENCES

CHAPTER 12: AFRICAN FERMENTED FOODS AND BEVERAGES. POTENTIAL IMPACT ON HEALTH

12.1 INTRODUCTION

12.2 SOME STAPLE FERMENTABLE FOODS IN AFRICA

12.3 THE DIVERSITY OF GUT MICROBIOTA

12.4 TRADITIONAL FERMENTATION STRATEGIES

12.5 FERMENTED NON‐ALCOHOLIC STARCHY FOODS

12.6 COMMON AFRICAN FERMENTED ALCOHOLIC BEVERAGES

12.7 SOME AFRICAN FERMENTED ANIMAL PRODUCTS

12.8 POPULAR AFRICAN TRADITIONAL FERMENTED FOODS AND BEVERAGES

12.9 EXAMPLES OF UNDESIRABLE EFFECTS OF FERMENTED PRODUCTS

12.10 CONCLUSIONS

FURTHER READING

CHAPTER 13: FERMENTED FOODS OF SOUTH ASIA

13.1 INTRODUCTION

13.2 ORIGIN, HISTORY, AND ROLE OF FERMENTATION

13.3 FERMENTATION OF FOODS

13.4 SOUTH ASIAN COUNTRIES AND INDIGENOUS FERMENTED FOODS – INDIA

13.5 INDIGENOUS FERMENTED FOODS OF SRI LANKA

13.6 INDIGENOUS FERMENTED FOODS OF BHUTAN

13.7 FERMENTATION – AN ECONOMIC APPROACH

13.8 FOOD SAFETY AND FOOD PRESERVATION

13.9 FOOD SAFETY RISKS IN TRADITIONAL FOODS

13.10 SUMMARY AND FUTURE PROSPECTS

13.11 CONCLUSION

REFERENCES

CHAPTER 14: THE ENSILING FERMENTATION OF FORAGE CROPS

14.1 INTRODUCTION

14.2 THE ENSILING FERMENTATION

14.3 THE FOUR PHASES OF ENSILING

14.4 STEPS OF ON‐FARM ENSILING

14.5 ENVIRONMENTAL ASPECTS OF ENSILING

14.6 UNCONVENTIONAL ENSILING

14.7 DEVELOPMENT OF SILAGE RESEARCH TOOLS

14.8 CONCLUSIONS

ACKNOWLEDGMENT

REFERENCES

SECTION V: A CLOSING PERSPECTIVE OF MICROBIAL FERMENTATION

CHAPTER 15: THE INTERSECTION OF MICROBIAL FERMENTATION AND EVOLUTIONARY ECOLOGY: WE PROVIDE THE HABITAT, YOU PROVIDE THE FERMENTATION

15.1 INTRODUCTION TO THE COMPETITIVE ECOLOGY OF MICROBIAL FERMENTATION

15.2 EXAMPLES OF SYMBIOTIC NICHE COOPERATIONS BETWEEN MICROBES AND THEIR ANIMAL HOSTS THAT DEPEND UPON MICROBIAL FERMENTATION

15.3 EXAMPLES OF USING SYMBIOTIC NICHE COMPETITIONS THAT RELY UPON MICROBIAL FERMENTATION TO PRODUCE FOOD FOR HUMANS AND LIVESTOCK

15.4 EXAMPLES OF USING MICROBIAL FERMENTATION TO PRODUCE INDUSTRIAL CHEMICALS

15.5 SUMMARY

REFERENCES

INDEX

END USER LICENSE AGREEMENT

List of Tables

Chapter 1

TABLE 1.1 Examples of bacteria that are present in food fermentations.

TABLE 1.2 Examples of fungi that are present in food fermentations.

TABLE 1.3 Examples of bacteria that are present in silage fermentations.

TABLE 1.4 Examples of fungi and oomycetes that are present in silage fermen...

TABLE 1.5 Examples of bacteria that are present in tobacco fermentations.

TABLE 1.6 Examples of fungi that are present in tobacco fermentations.

TABLE 1.7 Examples of bacteria that are present in compost fermentations.

TABLE 1.8 Examples of fungi that are present in compost fermentations.

TABLE 1.9 Examples of Archaea used in the fermentation production of chemic...

TABLE 1.10 Examples of bacteria used in the fermentation production of chem...

TABLE 1.11 Examples of fungi used in the fermentation production of chemica...

TABLE 1.12 A master list of viral families that affect prokaryotes.

TABLE 1.13 A master list of viral families and unassigned (floating) genera...

TABLE 1.14 Examples of how purified products from microbial fermentations a...

Chapter 2

TABLE 2.1 Lipid synthesis and growth of various Rhodotorula toruloides stra...

TABLE 2.2 Carotenoid biosynthesis by different Rhodotorula toruloides strai...

Chapter 3

TABLE 3.1 Predicted responses to major drivers of gut microbial diversity....

Chapter 5

TABLE 5.1 Main industrial processes relying on the microbial degradation of...

TABLE 5.2 Composition of the most common lignocellulosic biomasses of indus...

TABLE 5.3 Characteristics of the most efficient insect gut lignocellulose d...

Chapter 6

TABLE 6.1 Neolithic sites revealed alcohol remains in China.

Chapter 7

TABLE 7.1 Major components of gruit and their characteristics.

TABLE 7.2 Metagenomic profile of bacteria in three bottles retrieved from t...

Chapter 8

TABLE 8.1 Microbial cast list: in order of major appearance.

a

Chapter 10

TABLE 10.1 Microbiological cut‐off values (mg/l).

Chapter 11

TABLE 11.1 Classification of fermented foods based on their main fermentati...

TABLE 11.2 Categories of fermented foods (FF): examples and recent literatu...

TABLE 11.3 Main microbial groups of starter cultures.

Chapter 12

TABLE 12.1 The most at‐risk of food insecurity among the 54 African countri...

Chapter 13

TABLE 13.1 History of fermented foods.

TABLE 13.2 Indigenous fermented foods of Gujarat and Rajasthan.

TABLE 13.3 Indigenous fermented foods of Uttarakhand, Uttar Pradesh, Haryan...

TABLE 13.4 Indigenous fermented foods of Himachal Pradesh.

TABLE 13.5 Indigenous fermented foods of Madhya Pradesh and Goa.

Chapter 14

TABLE 14.1 Typical fermentation end‐products of various silage.

TABLE 14.2 The effect of maturity stage on yields and wheat plants composit...

TABLE 14.3 Results of microbiome analysis of 90–100 days laboratory silages...

List of Illustrations

Chapter 2

FIGURE 2.1 Metabolic pathways of

Rhodosporidium toruloides

. The metabolites ...

FIGURE 2.2

Rhodotorula toruloides

carotenoid biosynthetic pathway.

Chapter 3

FIGURE 3.1 (a) Grime's ecological model describes three primary strategies i...

Chapter 4

FIGURE 4.1 Overall scheme of rumen digestion of polysaccharides (inside of s...

FIGURE 4.2 Metabolism of pyruvate to acetate and butyrate. Main routes of ca...

FIGURE 4.3 Propionate formation via randomizing (succinate) and non‐randomiz...

FIGURE 4.4 Main pathways of metabolic hydrogen transactions in rumen ferment...

FIGURE 4.5 Simplified scheme of rumen, small intestine, and liver nitrogen m...

Chapter 5

FIGURE 5.1 Schematic overview of the anaerobic digestion of plant biomass an...

FIGURE 5.2 Simplified structure of the main lignocellulosic polysaccharides ...

FIGURE 5.3 Characterization of the termite gut system and proposed exploitat...

FIGURE 5.4 Comparison of microbial communities and their polysaccharide degr...

FIGURE 5.5 Glucoside hydrolases in microbial genomes from the anaerobic dige...

FIGURE 5.6 Comparison of the main glycoside hydrolase (GH) families targetin...

FIGURE 5.7 Anaerobic fungi in full‐scale anaerobic digestion systems (a) and...

Chapter 6

FIGURE 6.1 Examples of ancient fungal elements found on Neolithic pottery ve...

FIGURE 6.2 Globular jars unearthed from major sites in early Neolithic China...

FIGURE 6.3 Regional archeological cultural development and alcohol‐related p...

FIGURE 6.4 Examples of serving and drinking vessels during the late Neolithi...

Chapter 7

FIGURE 7.1 Embossed and screw top Victorian beer bottles.

FIGURE 7.2 Profiles of major features of brewing through periods of developm...

FIGURE 7.3 Example ingredients of gruit. (a) Bob myrtle/sweet gale; (b) Juni...

FIGURE 7.4 (a) Bottle #2 from the Wallachia shipwreck in preparation for ana...

Chapter 8

FIGURE 8.1 Allagash Brewing Company coolship room. Windows are propped open ...

FIGURE 8.2 Allagash Brewing Company aging barrels. “CS” refers to “Coolship....

Chapter 9

FIGURE 9.1 An electron microscopy image of an

A. oryzae

conidiophore and a p...

FIGURE 9.2 Haze komi and invasive growth of

A. oryzae

(green) into steamed r...

FIGURE 9.3 A tank to create the fermentation mash and fresh sale.

Chapter 11

FIGURE 11.1 Major mycotoxins and fungal species that produce them.

Chapter 12

FIGURE 12.1 Cassava roots and leaves.

FIGURE 12.2 Moringa leaves and trees.

FIGURE 12.3

Uapaka kirkiana

or wild loquat (

mushuku/muzhanje

);

Ziziphus maur

...

FIGURE 12.4 A diet high in fermented food or beverage decreases inflammatory...

FIGURE 12.5 Mitochondrial yeast metabolism, maltose being the best sugar sou...

FIGURE 12.6

Brassica tournefortii

on the left. Coffee beans in the middle. K...

FIGURE 12.7 Typical Sudanese

kissra

making and a full meal with several ferm...

FIGURE 12.8 Ripe bananas being prepared for fermentation in Burundi.

FIGURE 12.9 Preparation of wild water mellon wine.

FIGURE 12.10 Cashew apple fruit on the left with the unique external seed an...

FIGURE 12.11 Traditional preparation of amarula drink (“ocanhu”) in Mozambiq...

FIGURE 12.12 Images of masau (

Ziziphus mauritiana

), tambarind (

Tamarindus in

...

FIGURE 12.13 Milk and butter

smen

.

FIGURE 12.14 The multipurpose dawadawa tree, known as the African locust bea...

FIGURE 12.15 Commercially cassava beer is produced in Mozambique, brand name...

FIGURE 12.16 Palm wine created from the sap of various species of palm trees...

FIGURE 12.17 Aspartame, a well‐known artificial sweetener of sodas, candy, c...

Chapter 13

FIGURE 13.1 Types of food fermentation.

FIGURE 13.2 Fermented foods of Tamil Nadu.

FIGURE 13.3 Fermented foods of Kerala.

FIGURE 13.4 Fermented foods of North East–Indo‐Nepal Himalayan region.

FIGURE 13.5 Fermented foods of Sri Lanka.

Chapter 14

FIGURE 14.1 Some forage crops for ensiling.

FIGURE 14.2 The major steps of silage making.

FIGURE 14.3 Assessing corn grain maturity by milk line. The Milk Line (the b...

FIGURE 14.4 The effect of applying

L. plantarum

at ensiling on rate of pH de...

FIGURE 14.5 Ideal (left) vs. poor sealing (right).

FIGURE 14.6 Silage silo types.

FIGURE 14.7 Anaerobic mini‐silos (Weck™, Wehr‐Offlingen, Germany). The sprin...

Guide

Cover Page

Title Page

Copyright Page

Dedication

LIST OF FIGURES

LIST OF CONTRIBUTORS

PREFACE

Table of Contents

Begin Reading

Index

WILEY END USER LICENSE AGREEMENT

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MICROBIAL FERMENTATIONS IN NATURE AND AS DESIGNED PROCESSES

Edited by

CHRISTON J. HURST

Cincinnati, OH, USA

Universidad del Valle, Santiago de Cali, Valle, Colombia

This edition first published 2023© 2023 John Wiley & Sons Ltd

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DEDICATION

Daniel Yee‐Chak Fung was born in Hong Kong on May 15, 1942 and loving support from his family enabled Daniel to emerge from the Pacific War with a tremendous sense of optimism about life. Daniel eventually began his university training with a scholarship at the International Christian University in Tokyo, Japan, where he received a Bachelor of Arts Degree during 1965. He next earned a Master of Science degree in Public Health at the University of North Carolina, Chapel Hill during 1967. Soon afterward, during 1969 he was awarded a Doctor of Philosophy degree from Iowa State University. Daniel then found a place that truly felt like home when he became a professor of food science at Kansas State University in Manhattan, Kansas. He also found humor in explaining to his relatives that yes, he was at a university in Manhattan, but it was in the middle of the Kansas prairie and not in New York City. Daniel’s professional interest was to understand and explain the microbiology of food science. He earned tremendous international recognition for his efforts on the rapid detection of microbial contamination in food. Although, perhaps Daniel had the most fun when teaching a course on authentic Chinese cooking in what he described as small‐town USA.

I had the extremely good fortune to meet Daniel around 1990 during the annual meetings of the American Society for Microbiology. From then onward, I hoped to see him each time that I attended a microbiology conference. Daniel was one of the most enthusiastic people who ever could be imagined. His positive attitude constantly was encouraging to everyone around him. His smile and kindness seemed to brighten any room.

Eventually, many years ago I came to guess that something was wrong with Daniel’s health when he did not return a telephone call. I then spoke with the chairman of his department and learned that Daniel was in a group home for Alzheimer’s patients. I sent to Daniel a greeting card and included a photograph of myself with my name on the back of the photograph. I guessed that Daniel might recognize my face as being familiar even if remembering my name and the circumstances of our acquaintance might be challenging. Unfortunately, my good friend Daniel passed away on December 1, 2019.

Knowing Daniel was a great honor. The memory of his presence remains with me as a source of inspiration and I do very proudly dedicate my efforts on this book to him.

Daniel Yee‐Chak Fung 1942–2019

LIST OF FIGURES

2.1

Metabolic pathways of

Rhodosporidium toruloides

. The metabolites connected by arrows (a single arrow represents a step) and involved enzymes (named in the center of the arrow) are represented. Abbreviations are shown for the commonly used names of metabolites and enzymes. The endoplasmic reticulum, mitochondria, lipid body, and peroxisome are also present as cellular compartments. The enzymes with names in green coincide with genes that have been previously defined. FA is an abbreviation for fatty acid; FAME is an abbreviation for fatty acid methyl ester; FFA is an abbreviation for free fatty acid; FAEE is an abbreviation for fatty acid ethyl ester; TAG is an abbreviation for triacylglycerol, and VLFA is an abbreviation for very long chain fatty acid.

2.2

Rhodotorula toruloides

carotenoid biosynthetic pathway.

3.1

(a) Grime’s ecological model describes three primary strategies in plants (ruderal, competitive, and stress‐tolerant) in response to the environment. Grime originally defined “competitive species” as competing for resources and/or overlapping niche space. (b) Here, I adapt that framework to predict which types of microbial taxa are favored in species with different gut lengths and associated transit time; feeding strategies, specifically with regard to dietary fiber; and oxygen levels present in the gut. In my microbial model, “opportunistic species” include potential pathogens, and “competitive species” include microbial taxa that compete amongst themselves and with the host for easily digestible nutrients (e.g. sugar, starch, proteins, and lipids).

4.1

Overall scheme of rumen digestion of polysaccharides (inside of sky blue squircle), fermentation of hexoses and pentoses to pyruvate via glycolysis (inside of red squircle), pyruvate metabolism to volatile fatty acids (inside of purple squircle), pentose cycle and phosphoketolase cleavage (inside of green squircle), and hydrogenotrophic methanogenesis (inside of orange squircle).

4.2

Metabolism of pyruvate to acetate and butyrate. Main routes of carbon compounds are marked with sky blue arrows. Alternative pathway involving butyryl‐CoA: acetate CoA transferase is shown in gray arrows. Link with propionate randomizing pathway involving succinyl‐CoA: acetate CoA transferase is shown in purple. Generation of ATP by substrate‐level phosphorylation is shown with light green arrows. Reactions of cofactor reduction are shown in red arrows. Abbreviations: ADP, adenosine diphosphate; ATP, adenosine triphosphate; CO

2

, carbon dioxide; Fe

ox

, oxidized ferredoxin; Fe

red

2−

, reduced ferredoxin; H+, proton; HS‐CoA, coenzyme A; NAD+, oxidized nicotinamide adenine dinucleotide; NADH, reduced nicotinamide adenine dinucleotide; P

i

, phosphate group.

4.3

Propionate formation via randomizing (succinate) and non‐randomizing (acrylate) pathways. Main routes of carbon compounds are marked with sky blue arrows. Reactions involving transfer of coenzyme A are shown with gray arrows. Generation of ATP by substrate level and electron transport‐linked phosphorylation is shown with light green arrows. Reactions of cofactor re‐oxidation are shown in red arrows. Abbreviations: ADP, adenosine diphosphate; ATP, adenosine triphosphate; B

7

, biotin; B

12

, cobalamin; CO

2

, carbon dioxide; H

+

, proton; HS‐CoA, coenzyme A; NAD

+

, oxidized nicotinamide adenine dinucleotide; NADH, reduced nicotinamide adenine dinucleotide; P

i

, phosphate group.

4.4

Main pathways of metabolic hydrogen transactions in rumen fermentation. Reduction of NAD

+

to NADH in glycolysis and of ferredoxin in pyruvate oxidative decarboxylation is depicted in sky blue arrows. Formate formation in pyruvate oxidative decarboxylation and dissociation into carbon dioxide (CO

2

) and dihydrogen (H

2

) previously to its incorporation into methane (CH

4

) and other electron sinks is shown in gray arrows. Under low dihydrogen concentration NADH is re‐oxidized by [FeFe] A3 type hydrogenases in confurcation with reduced ferredoxin (Fd

red

2−

; yellow arrows). Under relatively high dihydrogen concentration NADH donates electrons in intracellular reactions and reduced ferredoxin is re‐oxidized to its oxidized species (Fd

ox

) by fermentative, dihydrogen‐evolving [FeFe] A1 type hydrogenases (purple arrows). Hydrogenotrophic methanogenesis incorporates dihydrogen and carbon dioxide (sky blue arrows). Incorporation of metabolic hydrogen into methyl‐reducing and fermentative methylotrophic methanogenesis is depicted in light green arrows.

4.5

Simplified scheme of rumen, small intestine, and liver nitrogen metabolism. Endogenous protein is not depicted for simplicity. Liver synthesized urea is recycled to the rumen through the rumen wall and in saliva (not depicted for simplicity). Abbreviations: AA, amino acids; N, nitrogen; NH

4

+

, ammonium.

5.1

Schematic overview of the anaerobic digestion of plant biomass and its position within the lignocellulose biorefinery concept.

5.2

Simplified structure of the main lignocellulosic polysaccharides and their enzymatic degradation mechanisms (a). Lignin degradation pathway is omitted here. Classification of main enzymes involved in lignocellulosic polysaccharides degradation (b). EC, enzyme commission number; CAZyme, carbohydrate‐active enzyme; GH, glycoside hydrolase; CE, carbohydrate esterase.

5.3

Characterization of the termite gut system and proposed exploitation of its potential toward improved lignocellulose utilization in engineered systems. Figure inspired by the work from Godon et al. (2013) and further modified. The highly compartmented gut model is characteristic of soil feeding termites, e.g.

Labiotermes

sp. The gut compartments involved in the degradation of the main lignocellulose fractions were identified after Marynowska et al. (2022). PFR, plug flow reactor; CSTR, continuously stirred tank reactor.

5.4

Comparison of microbial communities and their polysaccharide degrading potential in the anaerobic digestion (AD) reactors and the termite gut (Termite) system. Multidimensional scaling (MDS) of the calculated Jaccard similarity coefficient comparing the genomic content of the carbohydrate active enzymes (CAZymes) encoded in microbial genomes from the two systems (a). This subfigure was compiled based on microbial genomes generated in previous studies (Campanaro et al. 2020; Hervé et al. 2020). Main bacterial phyla present in the different types of AD reactors (i.e. alimented with different feedstocks), and higher termites feeding on distinct substrates (b). This subfigure was compiled based on (Calusinska et al. 2018b; Marynowska et al. 2020). WWTP, wastewater treatment plant, OFMSW, organic fraction of municipal solid waste.

5.5

Glucoside hydrolases in microbial genomes from the anaerobic digestion reactors (AD) and the termite gut system (Termite). (a) Shared diversity of glucoside hydrolase (GH) families in bacterial genomes from AD and the termite gut systems. (b) Average GH family diversity per microbial genome (only genomes with >70% completeness and <10% contamination were used), displayed for the main bacterial phyla. All pair‐wise comparisons are statistically significant (t.test, p < 0.005). (c) Linear discriminant analysis (LDA) of GH gene families differentially enriched in microbial genomes (community‐wise) in the two systems. This figure was compiled based on microbial genomes generated in previously published studies (Campanaro et al. 2020; Hervé et al. 2020).

5.6

Comparison of the main glycoside hydrolase (GH) families targeting the different lignocellulose fractions and other non‐lignocellulosic polysaccharides and their phylogenetic distribution among major bacterial phyla, for the anaerobic digestion reactors (a) and the termite gut system (b). This figure was compiled based on microbial genomes generated in previously published studies (Campanaro et al. 2020; Hervé et al. 2020).

5.7

Anaerobic fungi in full‐scale anaerobic digestion systems (a) and their lignocellulolytic potential (b and c). OFMSW, organic fraction of the municipal solid waste; WWTP, wastewater treatment plant; PL, polysaccharide lyase; GT, glycosyl transferase; GH, glycoside hydrolase; CE, carbohydrate esterase; CBM, carbohydrate‐binding module; AA, auxiliary enzyme. This figure was compiled based on own, unpublished data (a), and relying on the analysis of Neocallimastigomycota genomes available in public databases (b and c).

6.1

Examples of ancient fungal elements found on Neolithic pottery vessels in China. 1, 2:

Monascus

cleistothecia; 3, 4:

Aspergillus

vesicles; 5: cf.

28 pt

hyphae; 6:

Rhizopus

sporangium, showing structure of collarette and columella; 7:

Mucor

sporangium connecting to hypha; 8:

Rhizopus

or

Mucor

sporangia; 9: yeast cells in budding position; 10: a cluster of yeast cells.

6.2

Globular jars unearthed from major sites in early Neolithic China, ca. 9000 to 7000 cal. BP. 1: Dadiwan; 2: Guantaoyuan; 3: Baijia‐Lingkou; 4: Peiligang; 5: Jiahu; 6: Pengtoushan; 7: Kuahuqiao; 8: Shangshan; 9: Qiaotou; 10: Xiaohuangshan; 11: Cishan; 12: Houli.

6.3

Regional archeological cultural development and alcohol‐related pottery vessels in the Yellow River valley, the middle Neolithic period. (a) Distribution of the Beixin‐Dawenkou and Yangshao cultures. (b) Alcohol‐related pottery vessels in the Yangshao culture; 1: stove‐cauldron; 2: funnel; 3: amphora; 4, 5:

dakougang

vats (from Xipo). (c). Alcohol‐related pottery vessels in the Beixin‐Dawenkou cultures; 1: globular jar; 2:

gui

pitcher; 3, 4: goblets; 5–8:

dakougang

vats (from Yuchisi).

6.4

Examples of serving and drinking vessels during the late Neolithic Longshan culture. (a) Distribution of the Longshan culture in the Yellow River region. (b) Examples of alcohol‐related vessels from the Shandong Longshan culture; 1: cup; 2, 3: goblets; 4: pitcher. (c). Examples of alcohol‐related vessels from Shimao; 5: cups; 6, 7: pitchers.

7.1

Embossed and screw top Victorian beer bottles.

7.2

Profiles of major features of brewing through periods of development. Wheat – proportion used as a major ingredient: barley — proportion used as a major ingredient; fruit & herbs — prevalence in recipes, particularly as gruit; ABV — general level of alcohol achieved: shelf life — likely drinkable period (days to months); yeast management — awareness of value of yeast repatching; dispense — sophistication of draught and packaged product.

7.3

Example ingredients of gruit. (a) Bob myrtle/sweet gale; (b) Juniper berries; (c) Yarrow.

7.4

(a) Bottle #2 from the Wallachia shipwreck in preparation for analysis. (b) Yeast colonies isolated from Wallachia shipwreck bottle.

8.1

Allagash Brewing Company coolship room. Windows are propped open at all times and the wooden ceiling is important for harboring organisms. The fans help circulate airflow. The beer sits overnight and is then drained back into the main brewing building to be placed into barrels.

8.2

Allagash Brewing Company aging barrels. “CS” refers to “Coolship.” Apple refers to the fresh pomades added while lavender refers to the lavender‐smoked malt used. Beer was poured into barrels to begin aging on March 27th, 2019, November 20th, 2020, or January 4th, 2022.

9.1

An electron microscopy image of an

A. oryzae

conidiophore and a photo of rice

koji

.

9.2

Haze komi and invasive growth of

A. oryzae

(green) into steamed rice (red).

9.3

A tank to create the fermentation mash and fresh sale.

11.1

Major mycotoxins and fungal species that produce them.

12.1

Cassava roots and leaves.

12.2

Moringa leaves and trees.

12.3

Uapaka kirkiana

or wild loquat (

mushuku/muzhanje

);

Ziziphus mauritiana

in an open informal market.

12.4

A diet high in fermented food or beverage decreases inflammatory proteins, reduces molecular signs of inflammation, and improves immune response. Therefore, improving digestive health with a high‐quality fermented diet may improve immunity against infections and diseases.

12.5

Mitochondrial yeast metabolism, maltose being the best sugar source. Secondary metabolites and metabolism intermediates are shown in ethanol formation.

12.6

Brassica tournefortii

on the left. Coffee beans in the middle. Kombucha on the right.

12.7

Typical Sudanese

kissra

making and a full meal with several fermented foods.

12.8

Ripe bananas being prepared for fermentation in Burundi.

12.9

Preparation of wild water mellon wine.

12.10

Cashew apple fruit on the left with the unique external seed and women preparing cashew fruit wine in Guiné‐Bissau.

12.11

Traditional preparation of amarula drink (“ocanhu”) in Mozambique.

12.12

Images of masau (

Ziziphus mauritiana

), tambarind (

Tamarindus indica

), millet (

Cenchrus americanus

), bulrush (

Scirpoides holoschoenus

), monkey orange msala (

Strychnos spinosa

), and baobab (

Adansonia digitata

) fruit.

12.13

Milk and butter

smen

.

12.14

The multipurpose dawadawa tree, known as the African locust bean tree (

Parkia biglobosa

and

Parkia filicoidea

).

12.15

Commercially cassava beer is produced in Mozambique, brand name

Impala

.

Chibuku

sorghum beer is made in several countries.

Burukutu

preparation in Nigeria.

12.16

Palm wine created from the sap of various species of palm trees; hibiscus wine.

12.17

Aspartame, a well‐known artificial sweetener of sodas, candy, chewing gum, and energy drinks, is a synthetic chemical composed of the amino acids phenylalanine and aspartic acid, with a methyl ester. The methanol produced by the metabolism of aspartame is absorbed and quickly converted into formaldehyde and then completely oxidized to formic acid. The acceptable daily intake (ADI) for aspartame is 40 mg/kg of body weight. Ethanol, drinking alcohol, is mainly produced from maize fermentation. Ethylene glycol, used as antifreeze, if ingested is rapidly absorbed from the gastrointestinal tract and slowly absorbed through the skin or lungs.

13.1

Types of food fermentation.

13.2

Fermented foods of Tamil Nadu.

13.3

Fermented foods of Kerala.

13.4

Fermented foods of North East–Indo‐Nepal Himalayan region.

13.5

Fermented foods of Sri Lanka.

14.1

Some forage crops for ensiling.

14.2

The major steps of silage making.

14.3

Assessing corn grain maturity by milk line. The Milk Line (the border between the deep yellow and the milky parts) proceeds from the external perimeter toward the center of the cobs as the plant matures (A and B, respectively).

14.4

The effect of applying

L. plantarum

at ensiling on rate of pH decrease in wheat silage.

14.5

Ideal (left) vs. poor sealing (right).

14.6

Silage silo types.

14.7

Anaerobic mini‐silos (Weck™, Wehr‐Offlingen, Germany). The springs which secure the lids enable gas release from inside only but not air penetration.

LIST OF CONTRIBUTORS

Lourdes Georgina Michelena Álvarez, ICIDCA‐Instituto Cubano de Investigaciones de los Derivados de la caña de azúcar. Ciudad de La Habana, Cuba

Roberto Arredondo Valdés, Faculty of Chemical Sciences of the Autonomous University of Coahuila, Coahuila, México

Victoria Bell, Faculty of Pharmacy, University of Coimbra; Pólo das Ciências da Saúde, Azinhaga de Santa Comba, Coimbra, Portugal

HD Bhimani, Division of Microbial & Environmental Biotechnology, Navsari Agricultural University, Gujarat, India

Simana Bora, Department of Physics, Silapathar College, Silapathar, Department of Botany, Bahona College, Assam, India

François Bourdichon, Facoltà di Scienze Agrarie, Alimentarie Ambientali, Università Cattolica Del Sacro Cuore, Piacenza, Italy

Magdalena Calusinska, Environmental Research and Innovation Department, Luxembourg Institute of Science and Technology, Belvaux, Luxembourg

Nathaly Cancino‐Padilla, Centro Regional de Investigación Carillanca; Instituto de Investigaciones Agropecuarias INIA, Vilcún, La Araucanía, Chile

Sagarika Ekanayake, Department of Biochemistry, Faculty of Medical Sciences, University of Sri Jayewardenepura, Nugegoda, Sri Lanka

Dania Alonso Estrada, Faculty of Chemical Sciences of the Autonomous University of Coahuila, Coahuila, México

Evelyn Faife Pérez, ICIDCA‐Instituto Cubano de Investigaciones de los Derivados de la caña de azúcar. Ciudad de La Habana, Cuba

Tito H. Fernandes, CIISA ‐ Centre for Interdisciplinary Research in Animal Health, Faculty of Veterinary Medicine, University of Lisbon, Lisboa, Portugal. Former CEIL, Lúrio University, Nampula, Mozambique

Alessandra Fontana, Facoltà di Scienze Agrarie, Alimentarie Ambientali, Università Cattolica Del Sacro Cuore, Piacenza, Italy

Gomathy M, Department of Agricultural Microbiology, Tamil Nadu Agricultural University, Tamil Nadu, India

Xavier Goux, Environmental Research and Innovation Department, Luxembourg Institute of Science and Technology, Belvaux, Luxembourg

José Guina, Instituto Superior de Estudos Universitários, Polytechnic University, Nampula, Mozambique

Christon J. Hurst, Cincinnati, OH, USA; Universidad del Valle, Santiago de Cali, Valle del Cauca, Colombia

Anna Iliná, Faculty of Chemical Sciences of the Autonomous University of Coahuila, Coahuila, México.

Priya John, Department of Plant Pathology, Navsari Agricultural University, Gujarat, India

Marta Laranjo, MED‐Mediterranean Institute for Agriculture, Environment and Development & CHANGE‐Global Change and Sustainability Institute, IIFA‐Institute for Advanced Studies and Research, Universidade de Évora, Évora, Portugal

Li Liu, Department of East Asian Languages and Cultures, Stanford University Stanford, CA, USA

Tong Liu, Department of Forest Ecology and Management, Swedish University of Agricultural Sciences, Umeå, Sweden

Uma Maheswari, Department of Agricultural Microbiology, Tamil Nadu Agricultural University, Tamil Nadu, India

Manoj M, Department of Microbiology, Navsari Agricultural University, Gujarat, India

José Luis Martínez‐Hernández, Faculty of Chemical Sciences of the Autonomous University of Coahuila, Coahuila, México

Erin A. McKenney, Department of Applied Ecology, North Carolina State University, Raleigh, NC, USA

Lorenzo Morelli, Facoltà di Scienze Agrarie, Alimentarie Ambientali, Università Cattolica Del Sacro Cuore, Piacenza, Italy

Arianna Núñez Caraballo, Autonomous University of Coahuila, Coahuila, México

Nayra Ochoa Viñals, Faculty of Chemical Sciences of the Autonomous University of Coahuila, Coahuila, México

Ken Oda, National Research Institute of Brewing, Higashi‐hiroshima, Hiroshima, Japan

Vania Patrone, Facoltà di Scienze Agrarie, Alimentarie Ambientali, Università Cattolica Del Sacro Cuore, Piacenza, Italy

Rajakumar D, Department of Agronomy, Tamil Nadu Agricultural University, Tamil Nadu, India

Anu S. Rajan, Department of Agricultural Microbiology, Kerala Agricultural University, Kerala, India

Ramya P, Department of Agricultural Microbiology, Tamil Nadu Agricultural University, Tamil Nadu, India

Sabarinathan KG, Department of Agricultural Microbiology, Tamil Nadu Agricultural University, Tamil Nadu, India

Dilip Saikia, Department of Physics, Silapathar College, Silapathar, Department of Botany, Bahona College, Assam, India

Cassandra Suther, Department of Food Science, University of Massachusetts Amherst, Amherst, MA, USA

Norio Takeshita, Microbiology Research Center for Sustainability (MiCS), Faculty of Life and Environmental Sciences, University of Tsukuba, Tsukuba, Japan

Dilip Tamang, Kakajan Govt. M.E. School, Assam, India

Theradimani M, Department of Plant Pathology, Tamil Nadu Agricultural University, Tamil Nadu, India

Keith Thomas, Brewlab Ltd., University of Sunderland, England

Emilio M. Ungerfeld, Centro Regional de Investigación Carillanca; Instituto de Investigaciones Agropecuarias INIA, Vilcún, La Araucanía, Chile

Nelson Vera‐Aguilera, Centro Regional de Investigación Carillanca; Instituto de Investigaciones Agropecuarias INIA, Vilcún, La Araucanía, Chile

Zwi G. Weinberg, Forage Preservation and By‐Products Research Unit, Agricultural Research Organization, The Volcani Institute, Rishon Le Zion, Israel (Retired)

Maria Westerholm, Department of Molecular Sciences, Swedish University of Agricultural Sciences, Uppsala, Sweden

Tharanee Wijayaratne, Department of Biochemistry, Faculty of Medical Sciences, University of Sri Jayewardenepura, Nugegoda, Sri Lanka

PREFACE

Microbial Fermentations in Nature and as Designed Processes

Christon J. Hurst, Editor

Fermentation is one of the most important metabolic tools that biology has developed and microorganisms do in many ways seem to have become the true masters of fermentative metabolism. There are times when microbial fermentation processes seem to occupy nature’s center stage, although microbial fermentation processes often function less obviously by serving as part of biology’s stabilizing foundation and by providing the backstage support for more noticeable activities.

Animals and plants very importantly depend upon their own fermentative capabilities at times and in places where neither aerobic nor anaerobic respiration are energetically suitable. Those capabilites are overshadowed by the extensive and reliably successful symbiotic interactions which the same animals and plants have developed with their fermenting microbial partners.

Each of the fermentative microbial functions evolved to fit an energetic opportunity and each function has ecological value. It is important for us to acknowledge those values and understand the balance that exists between beneficial versus detrimental perspectives of microbial fermentation. For example, fermentative recycling processes which are so crucial in soil and aquatic sediments, and which also have a necessary role in the gastrointestinal tract of animals, can seem detrimental when they contribute to disease in animals and plants.

This book provides a summary of our understanding with regard to the commonalities and distinctions of aerobic versus anaerobic fermentations as performed by microorganisms. The authors explain how animals, including ruminants and termites, have evolved to use their symbiont fermentative microbes as tools for deconstruction of complex polysaccharides, and how we have adopted those natural technologies into our designed processes for anaerobic degradation of lignocellulosic materials. The important role of rhizosphere microbes that facilitate availability of inorganic and organic phosphates for plants also is explained. Those phosphates get stored in the plant’s seeds. And, after animals ingest the seeds, enzymes produced by symbiotic microbial fermentation allow the animals to utilize their dietary phosphates.

The authors provide understanding of the ways that fermentation has been harnessed from prehistoric times to the present for food processing and food preservation, and that technology valuably helps to sustain our species by allowing seasonal harvests to provision us throughout the year. The authors also explain how natural microbial activities have been similarly harnessed for creation of fermented fodder that sustains many of our domesticated agricultural animals.

During my efforts to understand the microbiology of food fermentations, I have asked myself several times if using microbial fermentation to produce and preserve food items may represent a collaborative genetic selection process. The microbes which are prevalent in spontaneous fermentations of food would have selected for a human population which could survive ingesting those microbes. Similarly, those microbes which are spontaneously present in the silage materials that we produce as fodder for our ruminant livestock would have exerted a selection process on our livestock. In cooperative response, we have learned to design food and silage fermentation processes in ways that would favor growth of the most easily survivable microbial populations. Those microbial populations have become our symbionts. That collaborative selection process has not been perfect as evidenced by the opportunistic pathogenicity associated with some of the microbes that exist in fermented food products and fermented fodder.

This book also offers insight into the ways that we have learned to use microbial fermentations as an engineering tool for producing chemicals, including enzymes and pharmaceuticals, which improve the health of ourselves and our domesticated animals. Additionally, the book offers some insight into possible future research directions for the field of applied microbial fermentation that will help to advance agriculture and industry.

I wish to thank Scott Robbins and Andrea Siefring‐Robbins who own Urban Stead Cheese, which is an artisanal cheese company in Cincinnati, Ohio, for very kindly having allowed me to watch their crew perform the almost magical process by which microbial fermentation produces cheddar cheese.

This book is the twenty‐second and final microbiology volume that I will have edited during a period of thirty four and a half years, beginning in February of 1989 when I started organizing the first volume. Those volumes have contained in total five hundred and thirty eight chapters of which I wrote fifty two. I have enjoyed the honor of representing the six hundred and eighty seven colleagues who joined me on this long interesting adventure as authors and coauthors, especially those who also joined me as coeditors, and I offer my thanks to each of them. Edited books are collaborative efforts similar to good fermentations, intended to share collective knowledge with an appreciative audience. Many of those authors and coauthors kindly have contributed to more than a single book project. In particular, I thank Professor Gideon Wolfaardt who graciously always agreed to participate each of the five times he was asked to write a chapter that would help us present the unfolding story of science. Gideon, collaborative projects have success because of the helpful and reliable efforts made by people such as you. A dream of mine would be to invite all of those colleagues to meet together in a large pub, I would buy a pint of nicely fermented beer for each, and then I would raise my glass to them and say “Together, we have accomplished something very good”. That could not happen, and many contributors to the earlier books have since passed away. Instead, I will think of them and say these lines from an old song,

“So fill to me the parting glass … Good night and joy be to you all”.

Christon J. Hurst in 2022

Christon J. Hurst

Cincinnati, Ohio

SECTION IAN INTRODUCTION TO MICROBIAL FERMENTATION

CHAPTER 1MICROBIAL FERMENTATION: UNDERSTANDING AND USING ONE OF NATURE'S TOOLS

CHRISTON J. HURST1,2

1Cincinnati, OH, USA

2Universidad del Valle, Santiago de Cali, Valle del Cauca, Colombia

CONTENTS

1.1 Introduction to Fermentation Processes

1.2 Examples of Microbial Fermentations in Nature

1.2.1 Fermentation Contributes to the Nutrient Recycling Capabilities Within Extreme Environments

1.2.2 Fermentative Microbes Support the Biological Availability of Phosphates in Soil

1.2.3 Fermentative Microbes Are Important for the Biological Availability of Dietary Phosphate

1.2.4 Natural Ethanol Production in Sap, Its Ingestion by Animals, and Human Intestinal Production of Ethanol

1.3 Designing Fermentation Processes

1.3.1 The Levels of Liquid and Oxygen Are Perhaps the Two Most Defining Characteristics of Fermentations

1.3.2 Microbial Ecology Largely Determines the Chemical Profile of Designed Fermentation Products

1.3.3 Spontaneous Fermentations and the Use of Back Slopping

1.3.4 The Importance of Using Defined Starter Cultures and Monitoring Fermentations for Product Quality Control

1.4 The Disposal of Waste Products from Fermentations

1.5 Fermentations That Produce and Preserve Food for Humans

1.5.1 Food Fermentation Is Largely About Lactic Acid and Ethanol

1.5.2 Fermentations That Preserve Fruits and Vegetables

1.5.3 Fermentations of Grains and Other Seeds Including Beans

1.5.4 Fermentations That Produce Alcoholic Beverages

1.5.5 Fermentations That Produce Vinegar

1.5.6 Fermentations That Create Dairy Products

1.5.7 Fermentations That Preserve Meat Products

1.5.8 The Microbial Safety of Fermented Food

1.6 Silage Fermentations That Produce and Preserve Food for Livestock

1.6.1 The Effect of Nitrate in Fertilizers

1.6.2 The Microbial Safety of Fermented Fodder

1.7 Tobacco Fermentations

1.8 Compost Fermentations and the Anaerobic Digestion of Food Wastes for Generating Biogas

1.9 Fermentations That Produce Chemicals, Enzymes, and Pharmaceuticals

1.9.1 Basic Reactor Designs

Submerged Fermentations

Solid State Fermentations

1.9.2 The Types of Microbes That Are Used for These Fermentations

1.9.3 The Types of Products and Their Applications

1.10 Summary

References

1.1 INTRODUCTION TO FERMENTATION PROCESSES

When we consider the ways in which living beings gain energy by degrading organic compounds, most of us first think of the aerobic respiration process which relies upon oxygen as its terminal electron acceptor and generates water molecules. Nearly all of the eukaryotic species that possess mitochondria, including many groups of microorganisms, use aerobic respiration with oxygen as their electron acceptor. Humans gain almost all of our energy by that process.

There are groups of microbes that survive and thrive under conditions where the presence of oxygen is limited. Those microbes often will degrade organic compounds by using anaerobic respiration processes which rely upon either nitrate, sulfate, or elemental sulfur as terminal electron acceptors (Lecomte et al. 2018). Microbes additionally may use metals and metalloids as electron acceptors, and that can occur under aerobic as well as anaerobic conditions (Hurst 2021).

Metabolic fermentation processes degrade organic compounds in ways that use either the same organic molecule, or a different organic molecule, as the terminal electron acceptor. Metabolic fermentation generates a wide range of products including organic acids and their conjugate bases, alcohols, aldehydes, ketones, carbon dioxide, and hydrogen. The amount of energy that can be gained by using metabolic fermentation to catabolize carbohydrates, lipids, and proteins is low in comparison with the amount of energy that could be gained if the organism were able to process those same compounds by either aerobic or anaerobic respiration. The energetic inefficiency of using fermentation to degrade organic materials is associated with the fact that potential energy remains in the molecular bonds of the organic products. Among the particularly well‐known examples of metabolic fermentation processes are those which rely upon pyruvic acid as final electron acceptor to produce lactic acid, and the use of acetaldehyde as final electron acceptor which produces ethanol.

Macroorganisms do at times use metabolic fermentation processes as a supplemental means for obtaining energy. Animals, including humans, rely upon fermentation to produce lactic acid within our muscle tissues when those tissues are stressed by a shortage of intracellular oxygen. Plants growing in soil use fermentation processes in their root tissues when the surrounding soil is saturated with water and the level of oxygen supplied from photosynthesis performed in aerial parts of the plants is insufficient to supply the metabolic needs of those root tissues. Rice plants very notably can produce acetaldehyde and ethanol as a short‐term emergency survival mechanism during darkness and oxygen deprivation, as might occur when the soil is completely flooded with turbid water (Boamfa et al. 2003).

Microorganisms living freely in the environment will use metabolic fermentation processes to satisfy their energy needs when the conditions of their surroundings are unfavorable for supporting either aerobic or anaerobic respiration. There additionally are symbiotic partnerships by which macroorganisms serve as physical hosts for many of the microbial fermenters and these are interdependencies that enable the macroorganisms to gain benefits from their fermenting microbial partners. Examples of such beneficial partnerships are the microbial fermentations that naturally occur in the gastrointestinal tracts of animals (Mississippi State University 2022) and humans (Oliphant and Allen‐Vercoe 2019). Coevolution has developed those gastrointestinal processes as a way of helping both the animals and microbes to optimally benefit from the ingested food. Our developments in food technology include using Propionibacterium and Anaerotignum propionicum (previously Clostridium propionicum) for the propionic acid fermentations of cheese, and we understand that the natural ecology of those microbes includes residence in the rumen and intestinal tract, as well as on the skin of mammals (Ciani et al. 2010). Plants also have many types of partnerships with fermenting microbes, with an example being the fact that bacteria in the rhizosphere use metabolic fermentation processes to generate organic acids which assist with the availability of phosphates, as will be described a bit later in this chapter.

Many of the natural microbial fermentation processes that are so important to putrefaction are proteolytic and they serve by liberating amino acids from proteins. Our designed microbial fermentation processes that create miso and soy sauce from soybeans, and those processes that produce meat products like ham and salami, are examples of similarly using proteolytic microbial fermentation to liberate amino acids. And yet, there are unfortunate times when gastrointestinal microbes that naturally represent those same nutrient recycling processes cause disease (Diether and Willing 2019) including associations with intestinal infections, peritoneal infections, and gangrene (Ciani et al. 2010). Fermentations which produce amino acids are termed amino acid fermentations.

We additionally have applied our skills at technological design processes to harness some of the other microbial metabolic fermentation mechanisms. The numerous applications of these processes prominently include the preservation of food that is intended for us to consume (Tables 1.1 and 1.2).

The microbial ecology of food preservation greatly depends upon fermentations which accomplish anaerobic conversions of sugars into acids and ethanol. As you already may have guessed, fermentations, and the microbes which perform those metabolic activities, often are named descriptively based on the desired product. Acetic acid, butyric acid, lactic acid, and propionic acid, respectively, represent products from acetic acid fermentations, butyric acid fermentations, lactic acid fermentations, and propionic acid fermentations. The fermentations which typically produce ethanol are termed either alcoholic or ethanolic fermentations. All of these acids and the ethanol are chemical weapons that fermenting microbes use in an effort to control the ecology of their environment and these chemical products are important to the microbial successions that occur during fermentation processes.

Anaerobic lactic acid fermentations are performed by lactic acid bacteria and also by some yeasts. These fermentation processes are described as either homolactic or heterolactic. Homolactic fermentation converts one molecule of glucose initially into two molecules of pyruvate, then converts those into two molecules of lactic acid. Heterolactic fermentation performs that same conversion of glucose into lactic acid, but then further converts one of the lactic acid molecules into amolecule of carbon dioxide plus a molecule of ethanol. The products of heterolactic fermentation thus are a combination of carbon dioxide, ethanol, and lactic acid. Generation of the three additional organic acids that have earned prominence in the creation and preservation of food, those being propionic, butyric, and acetic, all follow lactic acid production. Fermentation by propionic acid bacteria anaerobically converts lactic acid to acetic and propionic acids, carbon dioxide, and water. Fermentation by butyric acid bacteria anaerobically converts lactic acid to acetic and butyric acids, carbon dioxide, and hydrogen. Butyric acid production occurs when the pH of either food or silage fermentations is not sufficiently low to exclusively allow control by microbes that produce lactic acid fermentation.

TABLE 1.1Examples of bacteria that are present in food fermentations.

Genus

Species

Primary Material Being Fermented

Product

Geographical Origin

Acetobacter

aceti

Milk

Fermented beverage (Kefir)

North Caucasus

Acetobacter

aceti

Palm wine

Palm wine vinegar

Philippines

Acetobacter

aceti

Previously alcoholic fermented coconut water (liquid from coconuts plus sugar)

Coconut water vinegar

Philippines

Acetobacter

aceti

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

aceti

Previously alcoholic fermented molasses (Molasses beer)

Molasses vinegar

Global

Acetobacter

cerevisiae

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

fabarum

Cacao seeds

Cocoa beans

Ghana

Acetobacter

fabarum

Milk

Fermented beverage (Kefir)

North Caucasus

Acetobacter

lovaniensis

Milk

Fermented beverage (Kefir)

North Caucasus

Acetobacter

lovaniensis

Sugar

Fermented beverage (Tibicos, also called Water Kefir)

Global

Acetobacter

malorum

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

oeni

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

orientalis

Milk

Fermented beverage (Kefir)

North Caucasus

Acetobacter

orleanensis

Vanilla

pods

Vanilla

Mexico

Acetobacter

pasteurianus

Cacao seeds

Cocoa beans

Equatorial South America

Acetobacter

pasteurianus

Cassava

Beer (Cassava beer, Chicha)

Africa (Cassava beer), Brazil (Chicha)

Acetobacter

pasteurianus

Chañar fruit

Beer (Chicha)

Chile

Acetobacter

pasteurianus

Kañiwa

Beer (Chicha)

Andean South America

Acetobacter

pasteurianus

Maize

Beer (Chicha)

Andean South America and Costa Rica

Acetobacter

pasteurianus

Oca

Beer (Chicha)

Brazil

Acetobacter

pasteurianus

Palm fruit

Beer (Chicha, Palm beer)

Central and South America (Chicha), Fiji (Palm beer)

Acetobacter

pasteurianus

Peanut

Beer (Chicha)

Peru

Acetobacter

pasteurianus

Potato

Beer (Chicha, Potato beer)

Germany and Netherlands (Potato beer), Northern South America (Chicha)

Acetobacter

pasteurianus

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

pasteurianus

Previously alcoholic fermented pineapple peel and sugar

Pineapple peel vinegar

Asia, Latin America

Acetobacter

pasteurianus

Quinoa

Beer (Cusqueña)

Peru

Acetobacter

pasteurianus

Rice

Beer (Chicha, Rice beer)

Japan (Rice beer), South America (Chicha)

Acetobacter

pomorum

Previously alcoholic fermented fruit juices

Vinegar

Global

Acetobacter

tropicalis

Vanilla

pods

Vanilla

Mexico

Acetobacterium

tundrae

Barley

Beer (Historical ale)

United Kingdom

Acidipropionibacterium

acidipropionici

Milk

Cheese (Swiss type)

Europe

Acidipropionibacterium

jensenii

Milk

Cheese (Swiss type)

Europe

Acidipropionibacterium

thoenii

Milk

Cheese (Swiss type)

Europe

Actinotignum

urinale

Barley

Beer (Historical ale)

United Kingdom

Aerococcus

viridans

Fish with rice

Fermented seasoning (Pla ra)

Thailand

Anaerotignum

propionicum

Milk

Cheese

Global

Bacillus

amyloliquefaciens

Barley

Beer (Historical ale)

United Kingdom

Bacillus

amyloliquefaciens

Lentils

Griddle cake (Dosa)

India

Bacillus

amyloliquefaciens

Rice

Steamed cake (Idli)

India and Sri Lanka

Bacillus

amyloliquefaciens

Soybean

Fermented soybean (Hawaijar, Peruyaan)

India

Bacillus

atrophaeus

Vanilla

pods

Vanilla

Mexico

Bacillus

cereus

Rice

Steamed cake (Idli)

India and Sri Lanka

Bacillus

cereus

Soybean

Fermented soybean (Hawaijar), Curry (Kinema)

India

Bacillus

licheniformis

Barley

Beer (Historical ale)

United Kingdom

Bacillus

licheniformis

Locust bean

Dry condiment (Eware, Irú)

Africa

Bacillus

licheniformis

Soybean

Fermented paste or curry (Bekang), Curry (Kinema)

India

Bacillus

pumilus

Locust bean

Dry condiment (Eware, Irú)

Africa

Bacillus

pumilus

Soybean

Fermented paste (Tungrymba), Paste or curry (Bekang)

India

Bacillus

pumilus

Vanilla

pods

Vanilla

Mexico

Bacillus

subtilis

Barley

Beer (Historical ale)

United Kingdom

Bacillus

subtilis

Cassava

Fermented cassava root subsequently granulated into a flour (Garri)

West Africa

Bacillus

subtilis

Locust bean

Dry condiment (Eware, Irú)

Africa

Bacillus

subtilis

Maize

Porridge (Ogi)

Nigeria

Bacillus

subtilis

Millet

Porridge (Ogi)

Nigeria

Bacillus

subtilis

Rice

Steamed cake (Idli)

India and Sri Lanka

Bacillus

subtilis

Sorghum

grain

Porridge (Ogi)

Nigeria

Bacillus

subtilis

Soybean

Coagulated soy milk (Tofu), Fermented bean served whole often as an alternative to meat (Hawaijar, Natto), Fermented paste (Miso, Tungrymbai)

China (Tofu), India (Hawaijar, Tungrymbai), Japan (Miso, Nattō)

Bacillus

subtilis

Tea leaves

Fermented tea leaves [To be brewed for beverage] (Qingzhuan brick tea)

China

Bacillus

subtilis

subsp.

subtilis

Vanilla

pods

Vanilla

Mexico

Bacillus

tequilensis

Rice

Steamed cake (Idli)

India and Sri Lanka

Bacillus

thuringiensis

Soybean

Curry (Kinema)

India

Bacillus

thuringiensis

Vanilla

pods

Vanilla

Mexico

Bifidobacterium

animalis subsp. lactis

Milk

Fermented beverage

Europe

Bifidobacterium

breve

Milk

Fermented beverage

Europe

Brachybacterium

alimentarium

Milk

Cheese (Gruyère type, including Beaufort)

France (Beaufort), Switzerland (Gruyère)

Brevibacillus

brevis

Soybean

Fermented paste or curry (Bekang)

India

Brevibacterium

jeotgali

Bivalve mollusks (intact)

Jeotgal

Korea

Brevibacterium

jeotgali

Cephalopods (intact)

Jeotgal

Korea

Brevibacterium

jeotgali

Crustaceans (intact)

Jeotgal

Korea

Brevibacterium

jeotgali

Fish (intact)

Jeotgal

Korea

Brevibacterium

jeotgali

Fish (internal organs)

Jeotgal

Korea

Brevibacterium

jeotgali