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Beschreibung

Biosurfactants and Sustainability A timely and authoritative collection of resources on the sustainable production of biosurfactants In Biosurfactants and Sustainability, a team of distinguished researchers presents emerging themes in the rapidly evolving field of biosurfactants. The editors have chosen work that focuses on biosurfactants as eco-friendly and versatile compounds of interest in societies seeking sustainable forms of development. The book examines biosurfactants in the context of biorefineries and in the exploration of extremophilic microorganisms for biosurfactant production. The included works discuss biosurfactant production from different lignocellulosic and amylaceous raw materials, as well as oilseeds and other agro-industrial byproducts. Readers will also find: * A thorough introduction to microorganisms producing biosurfactants, as well as sustainable biosurfactant production in biorefineries * Comprehensive explorations of the challenges of biosurfactant production in fermentation processes * Practical discussions of bioreactors and metabolic engineering used in biosurfactant production * Fulsome treatments of biosurfactant production using enzyme and novel biosurfactant applications in nanotechnology, health, agriculture, and environmental cleanup Perfect for researchers and professionals with an interest in biosurfactant application and biotechnology processes, Biosurfactants and Sustainability will also benefit academic researchers, industry scientists, and engineers in biotechnology, microbiology, biomass conversion, environmental science and engineering.

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Biosurfactants and Sustainability

From Biorefineries Production to Versatile Applications

Edited by

Paulo Ricardo Franco Marcelino Lorena School of Engineering University of São Paulo Brazil

Silvio Silverio da Silva Lorena School of Engineering University of São Paulo Brazil

Antonio Ortiz Lopez Department of Biochemistry and Molecular Biology University of Murcia Spain

 

 

 

This edition first published 2023

© 2023 John Wiley & Sons Ltd

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions.

The right of Paulo Ricardo Franco Marcelino, Silvio Silverio da Silva and Antonio Ortiz Lopez to be identified as the authors of the editorial material in this work has been asserted in accordance with law.

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In view of ongoing research, equipment modifications, changes in governmental regulations, and the constant flow of information relating to the use of experimental reagents, equipment, and devices, the reader is urged to review and evaluate the information provided in the package insert or instructions for each chemical, piece of equipment, reagent, or device for, among other things, any changes in the instructions or indication of usage and for added warnings and precautions. While the publisher and authors have used their best efforts in preparing this work, they make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives, written sales materials or promotional statements for this work. The fact that an organization, website, or product is referred to in this work as a citation and/or potential source of further information does not mean that the publisher and authors endorse the information or services the organization, website, or product may provide or recommendations it may make. This work is sold with the understanding that the publisher is not engaged in rendering professional services. The advice and strategies contained herein may not be suitable for your situation. You should consult with a specialist where appropriate. Further, readers should be aware that websites listed in this work may have changed or disappeared between when this work was written and when it is read. Neither the publisher nor authors shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages.

Library of Congress Cataloging-in-Publication Data

Names: Marcelino, Paulo Ricardo Franco, editor. | Silva, Silvio Silvério da, editor. | Ortiz Lopez, Antonio, editor.

Title: Biosurfactants and sustainability : from biorefineries production to versatile applications / edited by Paulo Ricardo Franco Marcelino (University of São Paulo, Lorena School of Engineering, Lorena, São Paulo, Brazil), Silvio Silverio da Silva (University of São Paulo, Biotechnology Department, Lorena, São Paulo, Brazil), Antonio Ortiz Lopez (University of Murcia, Departamento de Bioquímica y, Murcia, Spain).

Description: Hoboken, NJ : John Wiley & Sons Ltd., 2023. | ECIP galley contains incomplete author affiliation for Antonio Ortiz Lopez. | Includes bibliographical references and index.

Identifiers: LCCN 2023000550 (print) | LCCN 2023000551 (ebook) | ISBN 9781119854364 (hardback) | ISBN 9781119854371 (adobe pdf) | ISBN 9781119854388 (epub) | ISBN 9781119854395 (ebook)

Subjects: LCSH: Biosurfactants.

Classification: LCC TP248.B57 B563 2023 (print) | LCC TP248.B57 (ebook) | DDC 668/.1--dc23/eng/20230213

LC record available at https://lccn.loc.gov/2023000550

LC ebook record available at https://lccn.loc.gov/2023000551

Cover Image: © süleyman Ibişov/EyeEm/Adobe Stock Photos

Cover Design: Wiley

Set in 9.5/12.5pt STIXTwoText by Integra Software Services Pvt. Ltd, Pondicherry, India

Contents

Cover

Title page

Copyright

List of Contributors

Foreword

Introduction

Biosurfactants: Concept, Biological Functions, Classification, General Properties and Applications

1 Microorganisms Producing Biosurfactants in the Current Scenario

1.1 Introduction

1.2 Microbial Biosurfactants

1.2.1 Structure and Classification of Biosurfactants

1.2.2 Biosurfactants Producing Yeasts

1.2.3 Biosurfactants Produced by Extremophile Microorganisms

1.3 Industrial Applications of Biosurfactants

References

2 Selection of Biosurfactant-Producing Microorganisms

2.1 Introduction

2.2 Traditional Methods of Detection

2.2.1 Direct Measure of Surface/interfacial Activity

2.2.2 Indirect Measure of Surface/interfacial Activity

2.2.3 Effects of Culture Media Based on Agro-industrial By-products on Properties of BS

2.3 High-throughput Analysis Method for the Screening of Potential Biosurfactants Producers

2.4 Screening of Microorganisms Biosurfactants and Lipases Producers

2.5 Conclusion and Future Perspectives

References

3 Metabolic Engineering as a Tool for Biosurfactant Production by Microorganisms

3.1 Metabolic Engineering and Biosurfactants

3.2 Regulation and Heterologous Production of Biosurfactants

3.3 Extension of Substrate Range for Biosurfactant Production

3.4 Improvement of Overall Cellular Physiology

3.5 Elimination or Reduction of By-product

3.6 Future Perspectives

3.7 Conclusions

References

4 Biosurfactant Production in the Context of Biorefineries

4.1 Biorefineries in Contemporary Society

4.2 Biomass and Biorefineries: Industrial By-products as Raw Materials for Biorefineries

4.3 Biosurfactant Production in the Context of Lignocellulosic Biorefineries

4.4 Biosurfactant Production in the Context of Oleaginous Biorefineries

4.5 Biosurfactant Production in the Context of Starchy and Biodiesel Biorefineries

4.6 Conclusion

References

5 Biosurfactant Production by Solid-state Fermentation in Biorefineries

5.1 Introduction

5.2 Advantages of Biosurfactant Production by Solid-State Fermentation

5.3 Suitable Biomasses for Biosurfactant Production in Biorefineries

5.4 Microorganisms Used in Biosurfactant Production by Solid-state Fermentation

5.5 Raw Materials Used in Solid-state Fermentation for Biosurfactant Production

5.6 Pretreatment of Raw Materials for the Production of Biosurfactants in Solid-state Fermentation

5.7 Physicochemical Factors of Solid-state Fermentation

5.8 Strategies for Scaling-up of Solid-state Fermentation for Biosurfactant Production

5.9 Conclusion

References

6 An Overview of Developments and Challenges in the Production of Biosurfactant by Fermentation Processes

6.1 Introduction

6.2 Current Market and Potential Applications of Biosurfactants

6.3 Biosurfactant as a Sustainable Alternative: Factors Influencing its Production

6.3.1 Factors Involved in the Biosurfactant Production

6.4 Strategies and Main Challenges for Biosurfactant Production

6.4.1 Process Configurations as Strategies for Biosurfactant Production

6.4.2 Bioreactors Used in the Biosurfactants Production: Types, Advantages, and Disadvantages

6.4.3 Biosurfactant Separation Processes

6.5 Future Perspectives and Conclusion

References

7 Enzymatic Production of Biosurfactants

7.1 Introduction

7.2 What are the Biosurfactants Produced Enzymatically? Esterification Reactions of Sugars and Fatty Acids Catalyzed by Enzymes

7.2.1 Esterification Reactions of Sugars and Fatty Acids Catalyzed by Enzymes

7.3 Enzymes and Methods for Biosurfactant Production: Bioreactors and Ways of Conducting Enzymatic Processes

7.4 Advantages and Disadvantages of Enzymatic Biosurfactant Production

7.5 Potential Use of Enzymes for the Production of Biosurfactants

7.6 Production of Biosurfactants by the Enzymatic Route in Biorefineries: Demand for More Modern Production Processes

7.7 Conclusion

References

8 Co-production of Biosurfactants and Other Bioproducts in Biorefineries

8.1 Introduction

8.2 Microbial Surfactant Production

8.3 Co-production of Biosurfactants in a Biorefinery

8.3.1 Co-production of Biosurfactants and Polyhydroxyalkanoates

8.3.2 Co-production of Biosurfactants and Enzymes

8.3.3 Co-production of Biosurfactants and Lipids

8.3.4 Co-production of Biosurfactants and Ethanol

8.4 Conclusions

References

9 Biosurfactants in Nanotechnology: Recent Advances and Applications

9.1 Introduction

9.2 Biosurfactants and their Types

9.2.1 Glycolipid Biosurfactants

9.2.2 Rhamnolipids

9.2.3 Trehalolipids

9.2.4 Sophorolipids

9.2.5 Mannosylerythritol Lipids

9.2.6 Lipopeptide Biosurfactants

9.2.7 Phospholipid Biosurfactants

9.2.8 Polymeric Biosurfactants

9.3 Properties of Biosurfactants

9.3.1 Surface and Interface Activity

9.3.2 Efficiency

9.3.3 Foaming Capacity

9.3.4 Emulsification/Emulsion Forming and Emulsion Breaking

9.3.5 Tolerance for Temperature and pH Tolerance

9.3.6 Low Toxicity

9.3.7 Biodegradability

9.4 Conventional Methods for Biosurfactant Production

9.5 Commercial Applications of Biosurfactants

9.5.1 Application of Biosurfactants in Agriculture

9.5.2 Application of Biosurfactants in Nanotechnology

9.5.3 Applications of Biosurfactants in Commercial Laundry Detergents

9.5.4 Application of Biosurfactants in Medicine

9.5.5 Application of Biosurfactants in the Food Processing Industry

9.5.6 Application of Biosurfactants in the Cosmetic Industry

9.5.7 Application of Biosurfactants in Petroleum

9.5.8 Application of Biosurfactant in Microbial-enhanced Oil Recovery

9.6 Biosurfactants in Nanotechnology (Biosurfactant Mediated Synthesis of Nanoparticles)

9.6.1 Glycolipids Biosurfactants Produced Nanoparticles

9.6.2 Lipopeptides Biosurfactants Produced Nanoparticles

9.7 Conclusions

References

10 Interaction of Glycolipid Biosurfactants with Model Membranes and Proteins

10.1 Introduction

10.2 Interaction of Glycolipid Biosurfactants with Model Membranes

10.2.1 Rhamnolipids

10.2.2 Trehalose Lipids

10.2.3 Other Glycolipids

10.3 Interaction of Glycolipid Biosurfactants with Proteins

10.3.1 Rhamnolipids

10.3.2 Trehalose Lipids

10.3.3 Mannosylerythritol Lipids

10.4 Conclusions

References

11 Biosurfactants: Properties and Current Therapeutic Applications

11.1 Production of Microbial Biosurfactants

11.2 Anti-tumoral Activity of Biosurfactants

11.3 Anti-inflammatory Activity of Biosurfactants

11.4 Anti-microbial Activity of Biosurfactant

11.4.1 Biosurfactants as Anti-bacterial Agents

11.4.2 Biosurfactants as Anti-viral Agents

11.4.3 Biosurfactants as Anti-fungal Agents

11.5 Other Therapeutic Applications of Biosurfactants

11.6 Concluding Remarks

References

12 Fungal Biosurfactants: Applications in Agriculture and Environmental Bioremediation Processes

12.1 Biosurfactants as Agrochemicals

12.1.1 Biosurfactants as Herbicide Adjuvants

12.1.2 Biosurfactants and Antifungal Activity

12.1.3 Biosurfactants as Insecticidal Adjuvants

12.2 Insecticidal Biosurfactants for Use against Disease Vector Insects

12.3 Fungal Biosurfactants in Bioremediation Processes

References

13 New Formulations Based on Biosurfactants and Their Potential Applications

13.1 Introduction

13.2 General Chemical and Biochemical Aspects

13.3 Downstream Processing

13.4 Biosurfactants in Cosmetics and Personal Care

13.5 Biosurfactants in Medicine and Pharmaceutics

13.6 Biosurfactants in Food and Feed

13.7 Biosurfactants in Pesticides, Insecticides, and Herbicide Formulations

13.8 Biosurfactants in Civil Engineering

13.9 Miscellaneous

13.9.1 Detergent Formulations

13.9.2 Bioremediation Purposes

13.9.3 Nanoparticle Synthesis

13.9.4 Polymer Synthesis

13.10 Overview of the Biosurfactant Market

13.11 Conclusions and Future Perspectives

References

14 Techno-economic-environmental Analysis of the Production of Biosurfactants in the Context of Biorefineries

14.1 Introduction

14.1.1 Background

14.1.2 Surfactant Versus Biosurfactant

14.1.3 Biosurfactant Market, Producers, and Patents

14.1.4 Biosurfactant Production Routes

14.2 Economic Aspects of the BS Production

14.3 Environmental Aspects

14.4 Biosurfactant Production Synergies in the Brazilian Biorefineries Context

14.5 Conclusion

References

Index

End User License Agreement

List of Tables

CHAPTER 01

Table 1.1 Classification of biosurfactants...

Table 1.2 Microorganisms and substrates...

CHAPTER 02

Table 2.1 Microbial strains...

Table 2.2 Microbial strains...

CHAPTER 03

Table 3.1 Compilation of strains...

CHAPTER 04

Table 4.1 Chemical composition...

Table 4.2 Main types of pre-treatments...

Table 4.3 Some studies on the...

CHAPTER 05

Table 5.1 Microorganisms, culture medium...

Table 5.2 Common methods...

CHAPTER 06

Table 6.1 Examples of biosurfactant...

Table 6.2 Advantages and disadvantages...

CHAPTER 07

Table 7.1 Main advantages and...

Table 7.2 Enzymes applied...

Table 7.3 Advantages and disadvantages...

CHAPTER 08

Table 8.1 Feedstock biorefinery...

Table 8.2 Co-production...

Table 8.3 Enzymes with detergent...

CHAPTER 09

Table 9.1 Different kinds of...

CHAPTER 11

Table 11.1 Biosurfactants...

Table 11.2 Anti-tumor activity...

Table 11.3 Mechanism of action...

Table 11.4 Anti-bacterial activity...

CHAPTER 12

Table 12.1 Biosurfactants produced...

CHAPTER 13

Table 13.1 Most relevant...

Table 13.2 Different purification...

Table 13.3 Main biosurfactant...

CHAPTER 14

Table 14.1 BS production around...

Table 14.2 Economic indicators...

CHAPTER c00

Table I.1 Types of biosurfactants...

List of Illustrations

CHAPTER 01

Figure 1.1 Fatty acid synthesis...

Figure 1.2 Industrial applications...

CHAPTER 02

Figure 2.1 Modified figure...

CHAPTER 03

Figure 3.1 Schema of the relationship...

Figure 3.2 Metabolic pathways...

CHAPTER 04

Figure 4.1 Structure and...

Figure 4.2 Fermentable sugars...

Figure 4.3 Main inhibitor compounds...

CHAPTER 05

Figure 5.1 Microbial groups...

Figure 5.2 Main types of bioreactors...

CHAPTER 06

Figure 6.1 The main applications...

Figure 6.2 Types of...

CHAPTER 07

Figure 7.1 Main types of reactors...

Figure 7.2 Usual stages in industrial...

Figure 7.3 Industrial process steps...

CHAPTER 08

Figure 8.1 PHA and rhamnolipid...

CHAPTER 09

Figure 9.1 Classification and...

Figure 9.2 Different properties...

Figure 9.3 Conventional method...

Figure 9.4 Schematic representation...

CHAPTER 10

Figure 10.1 General structure...

CHAPTER 11

Figure 11.1 Production and biochemical...

CHAPTER 12

Figure 12.1 Example structural...

CHAPTER 13

Figure 13.1 Chemical structures...

CHAPTER 14

Figure 14.1 Schematic representation ...

Figure 14.2 Integrated production...

Figure 14.3 Integrated production...

CHAPTER c00

Figure I.1 Types of biosurfactant...

Figure I.2 Chemical structures of...

Guide

Cover

Title page

Copyright

Table of Contents

Foreword

List of Contributors

Begin Reading

Index

End User License Agreement

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List of Contributors

Elisa Amorim Amâncio TeixeiraDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Francisco J. ArandaDepartment of Biochemistry and Molecular Biology-A University of Murcia Murcia, Spain

Nagamani BalagurusamyLaboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila Torreón, México

Thércia R. BalbinoBiotechnology Department University of São Paulo São Paulo, Brazil

Cristiani BaldoDepartment of Biochemistry and Biotechnology Londrina State University Londrina, Brazil

F.G. BarbosaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Fernanda G. BarbosaBiotechnology Department University of São Paulo São Paulo, Brazil

Heitor B. S. BentoDepartment of Bioprocess Engineering and Biotechnology São Paulo State University São Paulo, Brazil

Charles Lowell CantrellNatural Products Utilization Research Unit United States Department of Agriculture Oxford MS, USA

Felipe R.CarlosPostgraduate Program in Biotechnology Federal University of Alfenas Alfenas, Brazil

M.J. Castro-AlonsoDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Maria Jose Castro-AlonsoBiotechnology Department University of São Paulo São Paulo, Brazil

Jonas ContieroUniversidade Estadual Paulista “Julio de Mesquita Filho” Instituto de Biociências São Paulo, Brazil

Débora Luiza Costa BarretoDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Lívia da Costa CoelhoDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Patrícia C. M. Da RósDepartment of Chemical Engineering, University of São Paulo, São Paulo, Brazil.

Mayanne Karla da SilvaDepartamento de Microbiologia Universidade Federal de da Silva Minas Gerais Minas Gerais, Brazil

Silvio S. Da SilvaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Vinícius Luiz da SilvaMicroGreen – Soluções Biotecnológicas Piracicaba São Paulo, Brazil

G. L. de ArrudaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Ana Karine F. de CarvalhoPostgraduate Program in Biotechnology Federal University of Alfenas Alfenas, Brazil

María de Lourdes Froto MadariagaLaboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila Torreón, México

Stephanie Evelin de Oliveira AlvesDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Yasmim Senden, dos SantosDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Láuren Machado Drumond de SouzaDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Stephen Oscar DukeNational Center for Natural Products Research School of Pharmacy Oxford MS, USA

Embrapa Andrew Milli, EliasInstrumentação São Paulo, Brazil

Carlos Antonio Espinosa-LavenantLaboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila, Torreón, México

Juan Gerardo Flores-IgaLaboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila Torreón, México

Felipe Fernando FurlanDepartment of Chemical Engineering Federal University of São Carlos São Paulo, Brazil

Bruno C. GambaratoDepartment of Material Science University Center of Volta Redonda Volta Redonda, Brazil

Roberto de Campos GiordanoDepartment of Chemical Engineering Federal University of São Carlos São Paulo, Brazil

Vívian Nicolau GonçalvesDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Teran HernándezFernando, Laboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila Torreón, México

Vitor B. HidalgoPostgraduate Program in Biotechnology Federal University of Alfenas Alfenas, Brazil

Avinash P. IngleDepartment of Agricultural Botany Dr. Panjabrao Deshmukh Krishi Akola, India

Julio Bonilla JaimeEscuela Superior Politécnica del Litoral ESPOL, Centro de Investigaciones Biotecnológicas del Ecuador (CIBE) Guayaquil, Ecuador

Talita M. LacerdaBiotechnology Department University of São Paulo São Paulo, Brazil

Jonathan Coronel LeónEscuela Superior Politécnica del Litoral ESPOL, Centro de Investigaciones Biotecnológicas del Ecuador (CIBE) Guayaquil, Ecuador

Andreza Aparecida LongatiDepartment of Materials and Bioprocess Engineering State University of Campinas São Paulo, Brazil

Roberta Barros LovaglioUniversidade Federal de São Carlos São Carlos, Brazil

P.R.F. MarcelinoDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Nathália Sá Alencar do Amaral MarquesCatholic University of Pernambuco (UNICAP) Boa Vista, Brazil

Rogger Alessandro Mata da CostaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Laura Beatriz Miranda MartinsDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Everson Alves MirandaDepartment of Materials and Bioprocess Engineering, State University of Campinas São Paulo, Brazil

Mangesh Moharil Department of Agricultural Botany Dr. Panjabrao Deshmukh Krishi Akola, India

Dayana Montero RodríguezCatholic University of Pernambuco (UNICAP) Boa Vista, Brazil

Fabiana Guillen Moreira GasparinDepartment of Biochemistry and Biotechnology Londrina State University Londrina, Brazil

Salvador S. MuñozBiotechnology Department University of São Paulo São Paulo, Brazil

Júlia de Paula Muzetti RibeiroDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Antonio OrtizDepartment of Biochemistry and Molecular Biology-A University of Murcia Murcia, Spain

Fernanda PalladinoInstitute of Biological Sciences Federal University of Minas Gerais Belo Horizonte-MG, Brazil

C.A. PradoDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Natana Gontijo RabeloDepartamento de Microbiologia Universidade Federal de Minas Gerais Minas Gerais, Brazil

Mahendra RaiDepartment of Biotechnology Sant Gadge Baba Amravati University Maharashtra, India

Maria Ines RezendeDepartment of Biochemistry and Biotechnology Londrina State University Londrina, Brazil

T.M. RochaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Rita C.L.B. RodriguesLorena Engineering School University of São Paulo Lorena-SP, Brazil

Cintia M. RomeroPlanta Piloto de Procesos Industriales Microbiológicos, Consejo Nacional de Investigaciones Científicas y Técnicas Tucuman, Argentina

Luis Galarza RomeroEscuela Superior Politécnica del Litoral ESPOL, Centro de Investigaciones Biotecnológicas del Ecuador (CIBE) Guayaquil, Ecuador

Carlos A. RosaMicrobiology Department Federal University of Minas Gerais Belo Horizonte-MG, Brazil

Luiz Henrique RosaDepartamento de Microbiologia Universidade Federal de Minas Gerais Belo Horizonte, Brazil

Daylin Rubio-RibeauxDepartment of Biotechnology University of São Paulo, São Paulo, Brazi

Diana A. SanchezBiotechnology Department University of São Paulo São Paulo, Brazil

S. Sánchez-MuñozDepartment of Biotechnology University of São Paulo São Paulo, Brazil

J.C. SantosDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Monica C. SantosBiotechnology Department University of São Paulo São Paulo, Brazil

Shreshtha SaxenaDepartment of Agricultural Botany Dr. Panjabrao Deshmukh Krishi Akola, India

Gilda Mariano SilvaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

José A. TeruelDepartment of Biochemistry and Molecular Biology-A University of Murcia Murcia, Spain

Martha Inés Vélez-MercadoLaboratorio de Biorremediación Ciudad Universitaria de la Universidad Autónoma de Coahuila Torreón, México

M.C.A. VianaDepartment of Biotechnology University of São Paulo São Paulo, Brazil

Thiago A. VieiraBiotechnology Department University of São Paulo São Paulo, Brazil

Foreword

Several famous universities and research institutes have come together inthis book to offer the principles and fundamentals of biosurfactants, ecologically versatile compounds, their diverse sources and production mechanisms, as well as a vast field of applications in industry, agriculture, health, and in different environmental areas. Gathering knowledge on the basis of a solid theoretical background and bold applications is the biggest challenge facing Academia today, whose mission is to educate new generations and produce knowledge based on research, and integrate it into innovation for industrial partners and society. This book makes it accessible to undergraduate and graduate students, researchers, and professionals in the field, and demonstrates how the science of biosurfactants and the technology developed for their production and use can be applied to mission-oriented research to improve processes, improve materials and products, to meet the well-being of the population and promote sustainable development. May new disruptive ideas be born from this book, so that the frontier of knowledge is always extended for the benefit of society, with respect for the environment and the preservation of the planet. I wish you good reading, good study, and a good source of ideas!

Liedi Bernucci

Professor at Escola Politecnica of the University of São Paulo

President of Technological Research Institute of the State of São Paulo

Introduction

Paulo Ricardo Franco Marcelino1,*, Carlos Augusto Ramos1, Guilherme de Oliveira Silva1, Ramiro Reyes Guzman1, Silvio Silverio da Silva1, and Antonio Ortiz Lopez2

1 Laboratório de Bioprocessos e Produtos Sustentáveis (LBios), Escola de Engenharia de Lorena (EEL), Universidade de São Paulo (USP), Brazil2 Departamento de Bioquímica y Biología Molecular-A, Facultad de Veterinaria, Universidad de Murcia, Spain* Corresponding author

Biosurfactants: Concept, Biological Functions, Classification, General Properties and Applications

Biosurfactants are compounds with amphipathic structures obtained from natural sources such as plants (saponins), animals (bile salts) and microorganisms (glycolipids, lipopeptides, lipoproteins, polymerics and others) (Figure I.1) [1]. They present highlighted physicochemical and biological properties, such as surfactant and/or emulsifier, antimicrobial, antitumor, larvicide, mosquitocide, anti-inflammatory, immunomodulatory, which give them numerous applications [2–7]. For a long time the concept of biosurfactants was restricted only to microbial surfactants, but more recently Wim Soetaert, in a lecture at the Workshop on biosurfactants – Berlin, 2014, began to divide biosurfactants into first- and second-generation compounds, according to the origin of these substances.

Figure I.1 Types of biosurfactant: (A) plant biosurfactants: saponins, (B) animal biosurfactants: bile salts, (C) microbial biosurfactants: glycolipids, lipopeptides, lipoproteins and polymerics.

First-generation biosurfactants or “green surfactants” are conceptualized as compounds extracted and purified from vegetable and animal raw materials or produced entirely from renewable resources through chemical syntheses. The main examples include saponins, sugar esters, alkyl polyglycosides, and alkanolamides. Second-generation biosurfactants are compounds entirely produced from renewable resources and through a biological process (biocatalysis or fermentation), being exemplified mainly by microbial surfactants of the glycolipid and lipopeptide type [8, 9].

Microbial surfactants have several physiological functions such as emulsification, solubilization and intracellular transport of insoluble compounds in aqueous media, cell release in biofilms, antimicrobial activity, and quorum sensing [1, 4].

The need for some microorganisms to emulsify, solubilize, and transport compounds insoluble in aqueous medium to intracellular compartments, facilitating the consumption of these substrates for energy generation, has already been studied in depth. In addition to energy production, this biological function is of fundamental importance for technologies for the bioremediation of insoluble organic compounds in aqueous media [10–16].

The production of biosurfactants by microorganisms can also be considered a regulatory mechanism for cell adherence and release on surfaces, during a signaling/communication process dependent on population density, the so-called quorum sensing [17–19]. Adherence and cell release may occur due to surface properties (surface tension, surface enthalpy per unit area, surface composition, among others), the interfaces of supports, and microbial membranes. In addition, the microbiological properties and characteristics of the surrounding medium, such as temperature, pH, ionic strength, and availability of nutrients, are also determinant in the reported process [20–22]. Biosurfactants are molecules of fundamental importance in cell adherence and release processes. Microorganisms can use cell wall surfactant substances to regulate surface properties, aiming to adhere to or detach from a given location according to their need to find new habitats with greater availability of nutrients or to get rid of environments with unfavorable conditions [23, 24].

Biosurfactant compounds are also important for microorganisms because they have antimicrobial activity, becoming a defense mechanism in a competitive environment. Lipopeptides and glycolipids are the main biosurfactants with antimicrobial properties [25–28]. The antimicrobial action of biosurfactants is due to the interaction of these compounds with short-chain phospholipids found in the plasma membrane, causing an increase in membrane permeability through its solubilization, rupture, and disruption [29]. Among the biosurfactants with antimicrobial action, the class of lipopeptides is the most reported and studied [28]. With the recent pandemic caused by the SARS-COV2 coronavirus, some works have also highlighted the antiviral potential of biosurfactants [30–32].

The physical-chemical properties and classifications of biosurfactants are due to their structural characteristics. According to Bognolo [33] and Banat et al. [34], in biosurfactants, hydrophobic portions consisting of hydrocarbon chains with one or more fatty acids are observed, which can be saturated, unsaturated, hydroxylated or branched, linked to a hydrophilic portion, which it can be an ester, a hydroxyl group, a phosphate, a carboxylate, a carbohydrate, an amino acid, or a peptide. Most biosurfactants are neutral or anionic (present negatively charged polar groups), ranging from small fatty acids to large polymers. Based on the nature of their hydrophilic groups, biosurfactants are also commonly classified according to their chemical composition, as can be seen in Table I.1 [35, 36].

Table I.1 Types of biosurfactants and their main representatives.

Type of biosurfactant

Main representatives

Saponins or saponosides

Steroidal saponins

Triterpene saponins

Glicolipids

Rhamnolipids

Sophorolipids

Mannosylerythritol lipids (MEL)

Cellobiose lipids/ustilagic acid

Xylolipids

Trealolipids

Mycolates/trealomycolates

Oligosaccharide lipids

Lipid polyols

Fatty acids, neutral lipids and phospholipids

Fatty acids

Neutral lipids

Phospholipids

Polymeric surfactants

Emulsan

Biodispersant

Liposan

Carbohydrate-lipid-protein complex

Lipo heteropolysaccharides

Mannoproteins

Mannan-lipid

Lipopeptides and lipoproteins

Surfactin

Iturin

Serrawettin

Subtilisin

Cerelipin

Gramicidin

Viscosine

Amphisine

Tolaasin

Syringomycin

Peptide-lipids (lysine lipid and ornithine lipid)

Particulates

Membrane vesicles, fimbriae, and whole cells

Although the majority of biosurfactants reported in the literature are microbial metabolites associated, partially or not associated with cell growth, Kappeli and Finnerty [37] and Santos et al. [38] reported that some microbial cells, due to their high surface hydrophobicity, can also be considered surfactants by themselves. As an example, we have some hydrocarbonclastic microorganisms and species of Cyanobacteria, Staphylococcus, Serratia, Acinetobacter calcoaceticus, and Pseudomonas marginalis. There are also reports of some microbial extracellular vesicles that have high surfactant activity being classified as particulate biosurfactants.

In addition to natural and synthetic biosurfactants, others can be obtained through enzymatic/chemical transformations in a pre-existing structure, the so-called semi-synthetic or modified biosurfactants [39, 40]. Such modification mechanisms allow not only the structural change of the molecule but also the physical-chemical and biological properties. Zinjarde and Ghosh [41] reported the use of enzymatic systems, such as lipases and glycosidases, in the structural modification of precursor molecules of biosurfactants, such as carbohydrates and lipids. Delbeke and collaborators [42] synthesized a quaternary ammonium sophorolipid. From a sophorolactone produced by Starmerella bombicola, organic reactions were carried out in order to insert an amine group in the biosurfactant molecule. The modified sophorolipid obtained was tested against pathogenic bacteria, proving to be an excellent antimicrobial in the treatment of gram-positive strains, with greater potential than gentamicin sulfates already used in standard antibiotic-therapies, proving to have great promise in the clinical area. Recently, rhamnolipids were also functionalized with the amino acids arginine and lysine, obtaining cationic rhamnolipids (RLs) derivatives (Figure I.2) with outstanding physicochemical properties, in addition to presenting notable DNA binding affinity and good antimicrobial activity against gram-positive bacteria, including methicillin-resistant Staphylococcus aureus [40].

Figure I.2 Chemical structures of new semi-synthetic or modified surfactants based on rhamnolipids and amino acids: dirhamnolipid-arginine derivative and dirhamnolipid-lysine derivative.

The classification of biosurfactants can also be given according to their molecular weights. Some authors commonly classify them as high molar mass compounds (polysaccharides, lipopolysaccharides, lipoproteins, or complex mixtures of these biopolymers) or low molar mass (glycolipids and lipopeptides) [43]. This classification, despite still being widely used, has received criticism, since low molar mass molecules tend to have more surfactant characteristics, while high molecular mass molecules have emulsifying characteristics. Because of this, these molecules have different physicochemical and biological characteristics, and it is not correct to classify them all as biological surfactants. According to some authors, the correct way to classify these molecules would be to name molecules with a high molar mass of bioemulsifiers, while those with low molar mass would be called biosurfactants [44].

Biosurfactants are of fundamental importance in the current scenario because they are considered ecologically correct products, due to their low-/non-toxicity and high biodegradability. In addition, when compared to synthetic surfactants, they are more efficient, as they exhibit reduced surface and interfacial activities; are tolerant of high temperatures and extremes in pH and ionic strength; are specific and biocompatible [3, 33, 38, 45–48].

Biosurfactants are considered versatile molecules due to their wide possibilities in industrial applications. They can be used in the oil, chemical, food, and pharmaceutical industries and also in environmental and agricultural applications [4, 6, 38, 49–55]. However, these biomolecules are still not widely used due to high production costs, associated with inefficient product recovery methods and the use of expensive substrates [4]. In addition, several research groups have been seeking a better understanding of the biochemical production pathways for these products and thus developing new strategies to increase production.

The production of biosurfactants in the context of biorefineries can be a way to reduce the production costs of these molecules. In recent years, studies have begun on the production of biosurfactants in these biofactories and the technical and economic viability of these processes and products.

This book intends to present an overview of trends in the production of biosurfactants in biorefineries using fermentative and enzymatic processes and the versatility of applications of these bioproducts in the contemporary world that seeks sustainable development.

In Chapter 1, the main microorganisms produced by biosurfactants will be shown and how they can have their potential explored in different biorefineries. In addition, it will be briefly presented how extremophile microorganisms can be used in the biosurfactant production.

In Chapter 2, topics related to the selection of microorganisms that produce biosurfactants and lipase enzymes will be addressed, from conventional microorganisms to extremophiles.

In Chapter 3 some concepts related to the use of metabolic engineering tools in the microbial production of biosurfactants will be presented.

In Chapter 4, the production of biosurfactants in the context of biorefineries will be discussed. The possibility of biosurfactant production as a value-added product in biorefineries will be shown. In addition, the main biomasses used and their pre-treatments and important aspects of fermentation processes in this context will also be presented.

In Chapter 5, the production of biosurfactants by solid-state fermentation will be discussed, which is of fundamental importance in lignocellulosic biorefineries, but still little explored in the literature.

In Chapter 6, the development and challenges in the production of biosurfactants will be discussed. Some of the main production problems and how they can be overcome will be highlighted. It is noteworthy that this chapter will address some important bioprocess engineering concepts for the production of biosurfactants, mainly in submerged fermentations.

In Chapter 7, important concepts in the enzymatic production of biosurfactants will be discussed. The enzymes and conditions used in the enzymatic synthesis of biosurfactants, the bioreactors and the advantages and disadvantages of this form of sustainable production of biosurfactants will be discussed.

In Chapter 8, the production of biosurfactants will be discussed concomitantly with other important bioproducts for biorefineries, such as microbial polyesters, enzymes, lipids, and ethanol.

In Chapter 9, the use of biosurfactants in nanotechnological processes will be discussed, a topic of relevance in the current scenario.

In Chapter 10, the interaction of biosurfactants with models of membranes and proteins will be discussed. This theme is extremely important to understand future applications of biosurfactants, mainly in the pharmaceutical area, as antimicrobials.

In Chapter 11, the properties and therapeutic applications of biosurfactants will be discussed. It will be shown briefly from the purification processes necessary for the application of these metabolites in the pharmaceutical area to their antimicrobial, antitumor and anti-inflammatory potential.

In Chapter 12, the production of biosurfactants by fungi and the application of these metabolites in agriculture and bioremediation processes will be discussed.

In Chapter 13, the application of biosurfactants in the development of new products will be discussed. It will be shown how important biosurfactants are for the pharmaceutical, food and even civil engineering industries.

In Chapter 14, a brief discussion of the aspects necessary for techno-economic-environmental analysis of the production of biosurfactants in the context of biorefineries will be presented.

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1 Microorganisms Producing Biosurfactants in the Current Scenario

Fernanda Palladino1, Rita C.L.B. Rodrigues2, Yasmim Senden dos Santos1, and Carlos A. Rosa1,*

1 Microbiology Department, Institute of Biological Sciences, Federal University of Minas Gerais, 31270–901, Belo Horizonte-MG, Brazil2 Biotechnology Department, Lorena Engineering School, University of São Paulo, 12602–810, Lorena-SP, Brazil* Corresponding author

1.1 Introduction

Surfactants are compounds used in different industry segments (oil, detergent soap, food, and beverages) [1, 2]. They are amphipathic molecules that reduce the surface tension at the oil–water interface, increasing the solubility of immiscible substances in water. The global demand for surfactants is overgrowing, and according to market surveys, the outlook is for more than 520 billion tonnes of surfactants to be on the global market by 2022 [3]. The production of synthetic surfactants began in the first half of the 20th century. However, with a more expressive development of the petrochemical industry after World War II, the production of these compounds became more significant since the petrochemical industry provides raw materials to produce synthetic surfactants [4]. In the last decades, surfactants have attracted attention for their wide range of applications in modern society and their harmful effects on the environment. They have low biodegradability because they are derived from petroleum ecotoxicity and bioaccumulation [5].

In recent years, there has been an increase in the search for biodegradable compounds of natural origin that can be a more sustainable alternative to synthetic surfactants, corresponding to environmentally friendly and versatile products with extensive industrial applications [6, 7]. Natural surfactants are synthesized from living organisms, such as microorganisms (biosurfactants), plants (saponins), and vertebrates (bile salts) [8]. However, the best alternative for the production of surfactants that are less harmful to the environment is those produced by microorganisms, as they offer advantages over synthetic surfactants, such as lower toxicity, more significant foaming potential, biodegradability, thermal stability, selectivity and specificity, resistance to extreme values of pH and ionic strength, and solubility in alkaline medium [9, 10]. Although several studies involve bacteria in producing biosurfactants, yeasts are being studied because some bacteria are not GRAS (generally regarded as safe). Thus, this chapter will address microorganisms classified as the best biosurfactant producers and their applications.

1.2 Microbial Biosurfactants

Biosurfactants are a structurally diverse group of surface-active molecules produced by microorganisms [11]. These compounds can be synthesized by bacteria, yeasts, and filamentous fungi from various substrates, such as oils, alkanes, sugars, and industrial by-products [12]. Its production occurs when microorganisms are growing aerobically. During growth, they are secreted into the culture medium and help translocate and transport hydrophobic substrates across cell membranes, making them more available for metabolism and uptake and allowing microorganisms to develop on immiscible substrates [13]. Due to their characteristics, biosurfactant molecules have several properties, such as emulsification, detergency, wetting, foaming, dispersion, solubilization, and antimicrobial activity [12].

Several microorganisms can produce molecules with interfacial activity. There has been increasing interest in identifying and isolating new microorganisms that produce surfactant molecules with good characteristics, such as low critical micelle concentration (CMC), low toxicity, reduced surface tension, high emulsification activity,and ion complexation metallic [14, 15]. Studies approach biosurfactants of bacterial origin, produced by Pseudomonas, Acinetobacter, Bacillus, and Arthrobacter; however, the yeast species capable of producing these compounds are gaining more and more attention from the world scientific community probably due to these bacterial genera having a pathogenic nature, making their application in the food industry restricted. On the other hand, yeast has the advantage that some species have GRAS status, which means that microorganisms with this status are not pathogenic or toxic and can be applied in medicinal and food products [8]. Some yeasts of the genus Candida, Saccharomyces, Starmerella, Rhodotorula, Pseudozyma, Yarrowia, Pichia, Ustilago, Schizonella, Kluyveromyces, Wickerhamiella, Kurtzmanomyces, Debaryomyces, Cutaneotrichosporon, Spathaspora, and Scheffersomyces e Meyerozyma have been studied and reported in the literature as good biosurfactants producers [13, 16–18].

1.2.1 Structure and Classification of Biosurfactants

Biosurfactants are surface-active compounds produced by microorganisms, which have different structures obtained from renewable resources, possessing both hydrophilic and hydrophobic moieties [19]. Its amphiphilic structure modifies the surface properties or interfaces of complex systems. Biosurfactants are being studied as alternatives to cationic surfactants (quaternary ammonium), anionic surfactants (sodium lauryl sulfate, alkylbenzenesulfonate, alkyl ether phosphate), and amphoteric surfactants (phosphatidylcholine), mainly in the pharmaceutical industry [20].

Biosurfactants can be divided into two categories according to molecular weight [15]. One of these categories comprises low molecular weight, effective surface, and interfacial tension reducers. The other category is high molecular weight, called bioemulsifiers, which can form and stabilize emulsions but not necessarily reduce surface tension [21, 2, 22, 23]. Table 1.1 presents a classification of biosurfactants and producing microorganisms.

Table 1.1 Classification of biosurfactants and producing microorganisms.

Molecular weight

Biosurfactant

Microorganism

References

Low molecular weight

Rhamnolipids

Pseudomonas

sp.

1

Phulpoto et al. [

24

]

Aspergillus

sp.

3

Kiran et al. [

25

]

Planococcus

spp.

1

Gaur et al. [

26

]

Trehaloselipids

Rhodococcus

sp.

1

Bages-Estopa et al. [

27

]

Sophorolipids

Starmerella bombicola

2

Ceresa et al. [

28

]

Mannosylerythritol lipids (MELs)

Pseudozyma tsukubaensis

2

Andrade et al. [

29

]

Ustilago maydis

3

Becker et al. [

30

]

Glucose lipids

Alcanivorax borkurnensis

1

Yakimov et al. [

31

]

Cellobiose lipids

Pseudozyma aphidis

2

,

P. hubeiensis

2

Morita et al. [

32

]

Fatty acids

Corynebacterium lepus

1

Cooper et al. [

33

]

High molecular weight

Phospholipid

Sphingobacterium

sp.

1

Burgos-Díaz et al. [

34

]

Surfactin/iturin/ fengycin

Bacillus subtilis

1

Arima et al. [

35

]

Surfactin/fengycin

Alcaligenes aquatilis

1

Yalaoui-Guellal et al.[

36

]

Polymyxines

Paenibacillus polymyxa

1

Deng et al. [

37

]

Viscosin

Pseudomonas fluorescens

1

Bonnichsen et al. [

38

]

Serrawettin

Serratia marcescens

1

Clements et al. [

39

]

Emulsan

Acinetobacter venetianus

1

Castro et al. [

40

]

Liposan

Candida lipolytica

2

Cirigliano et al. [

41

]

Vesicle

Acinetobacter

sp.

1

Kappeli et al. [

42

]

1Bacteria

2Yeast

3Filamentous fungus. Vieira et al., 2021 / with permission of Elsevier

The main parameter used to identify the presence of biosurfactants in a system is the reduction of the surface tension of the medium. This substance in a system forms micelles (amphipathic molecules), which aggregate into hydrophilic and hydrophobic portions. More micelles lead to lower surface tension and are directly influenced by the increased concentration of biosurfactants [43, 44]. However, screening methods to identify these compounds based solely on surface tension reduction may eliminate bioemulsifiers as they will not always significantly reduce surface tension. Therefore, different tests must be performed, not just surface tension measurements, which are often used as preliminary tests [45].

There are many types of biosurfactants based on their chemical nature, such as glycolipids, lipopolysaccharides, oligosaccharides, and lipopeptides reported as being produced by various bacterial genera [46, 47]. Among the most studied is the genus Pseudomonasand Bacillus. Silva et al. [48] evaluated the commercial production of a biosurfactant from Pseudomonas cepacia CCT6659 grown in industrial by-products in a 50 L semi-industrial bioreactor for use in removing hydrocarbons from oily effluents. A concentration of 40.5 gL-1 was achieved on scale-up, and the surface tension was reduced to 29 mN m-1. Huang [49] isolated Serratia marcescens ZCF25 from the sludge of an oil tanker. This author carried out tests for biosurfactants production and observed that the biosurfactants produced was classified as a lipopeptide that managed to reduce the water surface tension from 72–29.50 mN m-1, showed high tolerance in a wide range of pH (2–12), high temperatures (50–100 oC) and salinity (10–100 gL-1). This microorganism was considered promising to be used in industrial applications and bioremediation. Hu et al. [50] evaluated the biosurfactant production by Bacillus subtilis ATCC 21332 in a 7 L and 100 L bioreactor using fish peptone generated by enzymatic hydrolysis of crushed tuna (Katsuwonus pelamis). Results showed that Bacillus subtilis ATCC 21332 could effectively use fish waste peptones for surfactin production. The highest surfactin productivity achieved in pilot-scale experiments was 274 mg L-1. Phulpoto et al. [24] evaluated several microorganisms isolated from freshwater lakes to produce biosurfactants. Among them, bacteria of the Pseudomonas and Bacillus species stood out, presenting an emulsification index of 32.70% and 55% and surface tension of 33.15 mN m-1 and 34.15 mN m-1, respectively. Liu et al. [51] evaluated the biosurfactant production by Bacillus licheniformis. The highest emulsification index was obtained using lactose-based mineral salt solution. The samples were characterized, and the biosurfactant found was a series of lipopeptides containing C13-, C14-, C15-, C16-surfactin, and other types of lipopeptides. They also observed that the biosurfactant altered the wettability of the hydrophobic surface of core slices with different permeability. This biosurfactant can maintain emulsification activity at pH 4–11, 85 oC, 25% NaCl solution, or 17.5% CaCl2 solution. Although there are several studies involving bacteria in the production of biosurfactants, yeasts are being studied.

1.2.2 Biosurfactants Producing Yeasts

The attention to the use of yeasts as biofactories in the production of biosurfactants is because some species are GRAS, making their application in the food, pharmaceutical, and cosmetic industries safer. For this reason, research involving the production of biosurfactants by yeasts has increased, and several strains of non-pathogenic species have been reported as good producers [52]. The genus Candida is a well-recognized producer of biosurfactants [53]. Some examples of Candida species able to produce biosurfactants are lipopeptide produced by Candida lipolytica [54], sophorolipids produced by Candida bombicola [55] or by Candida apicola [56] and mannosylerythritol lipid produced by Candida antarctica [57].

Some microorganisms can produce biosurfactants when they grow on different substrates, ranging from carbohydrates to hydrocarbons. Different carbon sources alter the structure of the biosurfactants produced and, consequently, their emulsifying properties. The biosurfactants production by yeasts is influenced by carbon and nitrogen sources and the presence or absence of phosphorus, iron, magnesium, and manganese in the substrate. Carbon sources can be alone or combined with organic or inorganic nitrogen sources such as yeast extract, ammonium salts, and nitrate salts [58]. Other parameters can change the yield of the process, such as temperature, pH, speed of agitation, aeration of the medium, and fermentation time. Table 1.2 shows microorganisms, substrates, and parameters used to produce biosurfactants [58, 8].

Table 1.2 Microorganisms and substrates for the production of biosurfactants.

Microorganisms

Carbon source

Surface tension (mNm

−1

)

E24 (%)

Cultivation

Method

References

Yarrowia lipolytica

Glucose (10 g L

-1

)

34.7

68

Flasks

Yalçin et al. [

18

]

Apiotrichum loubieri

35.3

67

Flasks

Pseudomonas aphidis

Glucose (40 g L

-1

)

32.83

ns

*

Flasks

Niu et al. [

59

]

Candida glaebosa

Glycerol (40 g L

-1

)

ns

*

30

Flasks

Bueno et al. [

53

]

Bacillus Subitilis

Glucose (4% w v

− 1

)

25.9

ns

*

Flasks

Queiroga et al. [

60

]

Bacillus licheniformis

Lactose (10 g L

-1

)

ns

*

96

Flasks

Liu et al. [

51

]

Naganishia adellienses

Sugarcane straw hemicellulosic hydrolysate (40 g L

-1

Xylose)

ns

*

52

Flasks

Chaves et al. [

9

]

Cutaneotrichosporon mucoides

Sugarcane bagasse hemicellulosic hydrolysate (40 g L

-1

Xylose)

ns

*

70

Flasks

Marcelino, et al. [

17

]

Meyerozyma guilliermondii

ns

*

70

Flasks

Trichosporon montevideense

Sunflower oil (20 g L

-1

)

44.9

75.80

Flasks

Monteiro et al. [

61

]

Aspergillus niger

Banana stalks powder

ns

*

57

Solid State

Asgher et al. [

62

]

Candida sphaerica

Refinery residue of soybean oil (90 g L

-1

) and corn steep liquor (90 g L

-1

)

25.00

ns

*

Flasks

Luna et al. [

63

]

27.48

ns

*

Bioreactor

Candida tropicalis

Sugarcane molasses (25 g L

-1

), Corn steep liquor (25 g L

-1

), Waste frying oil (25 g L

-1

),

29.98

ns

*

Flasks

Almeida et al. [

64

]

34.12

ns

*

Bioreactor

Pseudomonas aeruginosa

Corn steep liquor (10% v v

−1

), sugarcane molasses (10% v v

−1

)

31.4

59.0

Flasks

Gudiña et al. [

65

]

*ns (Not specified)

Sophorolipids are among the best-known and most studied biosurfactants, offering several advantages over synthetic surfactants [66]. Its production has been reported in the literature by the yeasts Starmerella bombicola, S. batistae, S. apicola, Rhodotorula bogoriensis, Wickerhamiella domercqiae, S. riodocensis, S. stellata, e Candida sp.NRR Y-27208 [67, 68].

Most biosurfactants are generally produced when cultures reach the stationary phase of growth. However, some species may show small production during the exponential growth phase. Sophorolipid biosynthesis is stimulated under nitrogen-limiting conditions since the enzymatic activity is high under these conditions. Furthermore, it occurs at the end of the exponential phase and the beginning of the microorganism’s stationary growth phase [69]. The proportion, type, and production of acidic or lactone forms of sophorolipids depend on some variables, such as the composition of the medium (carbon, nitrogen, and salt sources), production lineage, environmental conditions (pH, temperature, agitation, time, and aeration) and the type of cultivation process (batch or continuous) [70, 66]. The sophorolipid production is highly stimulated when two carbon sources of lipophilic origin (alkanes, saturated and unsaturated fatty acids, alcohols, oils, and fats) and hydrophilic (glucose) are present in the culture medium. Production can be lower when there are no hydrophobic sources in the middle.

The amphiphilic nature of biosurfactant creates surface tension at the interphase of two mediums and thus, helps microorganisms utilize hydrophobic carbon sources as nutrients and protects microorganisms from adverse environmental conditions [71]. Furthermore, the type of hydrophobic source will influence the sophorolipid final composition [14]. evaluated the production of sophorolipid by Starmerella bombicola using biodiesel as a hydrophobic substrate. They used a fed-batch fermentation system and obtained higher yields than flask cultivation (58.1–224.2 gL-1). It showed good surfactant properties, such as surface tension of 34.2 nM m-1 and a critical micellar concentration of 25.1 mg L-1.

Two basic approaches are adopted globally for cost-efficient production of biosurfactants: (1) utilization of abundant, inexpensive, and waste biomass as a substrate for the production media resulting in low initial raw material costs required for the process, and (2) development and optimization of bioprocesses for maximizing biosurfactant production and recovery, leading to reduced operating costs [72, 73]. By-products from different industrial segments have been used for the biotechnological production of biosurfactants. Among them, we can have glycerol [74]; animal fat [75]; soy oil refinery by-product [76]; a by-product of a peanut oil refinery [77]; residual frying oil [78]; clarified cashew apple juice [79]; corn steep liquor [80], cassava wastewater [81], vinasse [79]; molasses [82]; sugarcane bagasse hemicellulosic hydrolysate [83]. A single carbon source is used in biosurfactants produced via microbial fermentation (carbohydrate or a combination of two substrates – sugars and lipids) [22].