HANDBOOK OF FRUIT AND VEGETABLE FLAVORS -...

HANDBOOK OF FRUIT AND VEGETABLE FLAVORS

Feng ChenDepartment of Food Science and Human NutritionClemson University

L.M.L. NolletDepartment of EngineeringHogeschool Gent

A JOHN WILEY & SONS, INC., PUBLICATION

Edited by

Y.H. HuiScience Technology System

Technical Editors

Associate Editors

Raquel P.F. GuinéDepartamento de Indústrias Agro-AlimentaresEscola Superior Agrária de Viseu

M. Isabel Mínguez-MosqueraFood Biotechnology DepartmentInstituto de la Grasa (CSIC)

Fernando L.P. PessoaDepart de Engenharia Química, Escola de QuímicaUniv Federal do Rio de Janeiro

Jiwan S. SidhuDepartment of Family ScienceKuwait Women University

Peggy Stanfi eldConsultant

Olga Martín-BellosoDepartment of Food TechnologyUniversity of Lleida

Gopinadhan PaliyathDepartment of Plant AgricultureUniversity of Guelph

Jean-Luc Le QuéréInstitut National de la Recherche Agronomique (INRA)Unité Mixte de Recherche sur les Arômes (UMRA)

Nirmal SinhaResearch and DevelopmentGraceland Fruit, Inc.


Feng ChenDepartment of Food Science and Human NutritionClemson University

L.M.L. NolletDepartment of EngineeringHogeschool Gent

A JOHN WILEY & SONS, INC., PUBLICATION

Edited by

Y.H. HuiScience Technology System

Technical Editors

Associate Editors

Raquel P.F. GuinéDepartamento de Indústrias Agro-AlimentaresEscola Superior Agrária de Viseu

M. Isabel Mínguez-MosqueraFood Biotechnology DepartmentInstituto de la Grasa (CSIC)

Fernando L.P. PessoaDepart de Engenharia Química, Escola de QuímicaUniv Federal do Rio de Janeiro

Jiwan S. SidhuDepartment of Family ScienceKuwait Women University

Peggy Stanfi eldConsultant

Olga Martín-BellosoDepartment of Food TechnologyUniversity of Lleida

Gopinadhan PaliyathDepartment of Plant AgricultureUniversity of Guelph

Jean-Luc Le QuéréInstitut National de la Recherche Agronomique (INRA)Unité Mixte de Recherche sur les Arômes (UMRA)

Nirmal SinhaResearch and DevelopmentGraceland Fruit, Inc.

Copyright © 2010 by John Wiley & Sons, Inc. All rights reserved

Published by John Wiley & Sons, Inc., Hoboken, New JerseyPublished simultaneously in Canada

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Library of Congress Cataloging-in-Publication Data:

Handbook of fruit and vegetable fl avors / edited by Y.H. Hui. p. cm. Summary: “Acting as chemical messengers for olfactory cells, food fl avor materials are organic compounds that give off a strong, typically pleasant smells. Handbook of Fruit and Vegetable Flavors explores the fl avor science and technology of fruits and vegetables, spices, and oils by fi rst introducing specifi c fl avors and their commercialization, then detailing the technical aspects, including biology, biotechnology, chemistry, physiochemistry, processing, analysis, extraction, commodities, and requirements for application as food additives. With chapter authors representing more than ten different countries, this handy reference provides a comprehensive view of this evolving science.” – Provided by publisher. Summary: “This book provides a comprehensive reference on the fl avor science and technology of fruits and vegetables, spices, and oils. Beginning with an introduction on the specifi c fl avors and their commercialization, the book then details the technical aspects including biology, biotechnology, chemistry, physiochemistry, processing, analysis, extraction, commodities, and requirements for application as food additives. Regulatory considerations are discussed in relation to sanitation and safety in a fl avor manufacturing establishment” – Provided by publisher. ISBN 978-0-470-22721-3 (hardback) 1. Fruit–Flavor and odor–Handbooks, manuals, etc. 2. Vegetables–Flavor and odor–Handbooks, manuals, etc. 3. Oils and fats–Flavor and odor–Handbooks, manuals, etc. 4. Food–Sensory evaluation–Handbooks, manuals, etc. I. Hui, Y. H. (Yiu H.) TP440.H357 2010 664′.5–dc22 2010016633

Printed in the United States of America

10 9 8 7 6 5 4 3 2 1

www.copyright.comhttp://www.wiley.com/go/permissionhttp://www.wiley.com/go/permissionwww.wiley.com

v

PREFACE xi

CONTRIBUTORS xiii

LIST OF ABBREVIATIONS xix

SECTION A: FRUIT FLAVORS 1

PART I FRUIT FLAVORS: BIOLOGY, CHEMISTRY, AND PHYSIOCHEMISTRY 1

1. Fruits and Fruit Flavor: Classifi cation and Biological Characterization 3Yueming Jiang and Jun Song

2. Physiology and Biochemistry of Fruit Flavors 25Sonia Osorio, Cristina Muñoz, and Victoriano Valpuesta

3. Sensory Evaluation of Fruit and Vegetable Flavors 45Sara Bayarri and Elvira Costell

4. Fermentation and Fruit Flavor Production 59Gabriela Alves Macedo, Juliana Alves Macedo, and Luciana Francisco Fleuri

5. Environmental Effects on Flavor Changes 73Clara Pelayo-Zaldívar

PART II FRUIT FLAVORS: BIOTECHNOLOGY 93

6. Cell Culture for Flavor Production 95Adriane B.P. Medeiros, Suzan C. Rossi, and Carlos R. Soccol

7. Genetic Engineering of Fruit Flavors 101Asaph Aharoni and Efraim Lewinsohn

8. Bioconversion of Flavors 115P.F.F. Amaral, M.H.M. Rocha-Leão, and M.A.Z. Coelho

9. Plant Metabolic Pathways and Flavor Biosynthesis 129Carlos Sanz and Ana G. Pérez

CONTENTS

vi CONTENTS

PART III FRUIT FLAVORS: ANALYTIC METHODOLOGY AND CHEMICAL CHARACTERIZATIONS 157

10. History and Principles of Flavor Analysis 159F. Gutiérrez-Rosales

11. Advanced Analytic Methodology 177Jean-Luc Le Quéré

12. Extraction and Distillation 195Fernando L.P. Pessoa, Marisa F. Mendes, Eduardo M. Queiroz, and Silvio A.B. Vieira De Melo

13. Flavor Extraction: Headspace, SDE, or SFE 211Gerson L.V. Coelho, Marisa F. Mendes, and Fernando L.P. Pessoa

PART IV FLAVORS FOR FRUIT COMMODITIES 229

14. Annona Fruits 231Jorge A. Pino

15. Apple (Malus × domestica Borkh.) 247Maria Luisa López Fructuoso and Gemma Echeverría Cortada

16. Citrus Fruits and Oranges 265L.M.C. Cabral, A. Bravo, M. Freire JR., H.R. Bizzo, and V.M. Matta

17. Flavor in Grapes: Its Characterization and Commercial Applications 279N.R. Bhat, B.B. Desai, and M.K. Suleiman

18. The Aroma of Wine 303Juan Cacho and Vicente Ferreira

19. Mango Flavor 319O.P. Chauhan, P.S. Raju, and A.S. Bawa

20. Passion Fruit 345Narendra Narain, Nisha Nigam, and Mércia de Sousa Galvão

21. Pineapple (Ananas comosus [L.] Merril) Flavor 391Marta Montero-Calderón, M. Alejandra Rojas-Graü, and Olga Martín-Belloso

22. The Flavor of Plums 415E. Gómez-Plaza and C. Ledbetter

23. Strawberry Flavor 431Ana G. Pérez and Carlos Sanz

CONTENTS vii

24. Temperate Fruit Juice Flavors 451Terri D. Boylston

25. Fruits from Central and South America 463Jiwan S. Sidhu and Yearul Kabir

PART V FLAVORS OF SELECTED DRIED FRUITS 485

26. Overview of Flavors and Fruit Dehydration 487Regina Nabais

27. Flavors of Dried Apples 515Elena Venir

28. Flavors of Dried Apricots 531Mustafa Z. Özel and Fahrettin Göǧüş

29. Flavors of Dried Bananas 543Juan Wang, Bing Du, and Gong Ming Yang

30. Volatile Aroma/Flavor Components of Raisins (Dried Grapes) 549Ron G. Buttery

31. Flavors of Dried Pears 557Draženka Komes and Karin Kovačević Ganić

SECTION B: VEGETABLE FLAVORS 573

PART VI VEGETABLE FLAVORS: BIOLOGY, CHEMISTRY, PHYSIOCHEMISTRY, AND BIOTECHNOLOGY 573

32. Chemistry and Biochemistry of Some Vegetable Flavors 575Qayyum Husain

33. Traditional and New Analytical Methodology 627Emira Mehinagic and Jean-Luc Le Quéré

34. Vegetable Flavors from Cell Culture 663María Asunción Longo and María Angeles Sanromán

35. Flavor from Transgenic Vegetables 681Moustapha Oke and Gopinadhan Paliyath

36. Genetic Engineering of Fruit Flavors 693Jingyu Lin, Ningning Wang, Vincent P. Pantalone, and Feng Chen

37. Metabolic Factory for Flavors in Fruits and Vegetables 705Yoko Iijima

viii CONTENTS

PART VII VEGETABLE COMMODITIES 729

38. Avocado (Persea Americana Mill.) Oil 731Suely P. Freitas, Neusa P. Arruda, Clarice N. Gobbi, and Cláudia M. De Resende

39. Cabbage Flavor 741Theodore J.K. Radovich

40. Carrot Flavor 751Ahmad Sulaeman and Judy A. Driskell

41. Chili Flavor 775P.S. Raju, O.P. Chauhan, and A.S. Bawa

42. Corn Flavor 803Linda Pollak

43. Olive and Olive Oil 821Diego Luis García-González, María Teresa Morales, and Ramón Aparicio

44. Flavors in Onion: Characterization and Commercial Applications 849N.R. Bhat, B.B. Desai, and M.K. Suleiman

45. Onion: A Food, Spice, and Remedy in the Middle Eastern Gastronomy 873Pervin Basaran and Nese-Basaran-Akgul

46. Mushrooms in the Middle Eastern Diet 889Nese B. Akgul, Pervin Basaran, and Jiwan S. Sidhu

47. Flavoring Compounds in Red Pepper Fruits (Capsicum genus) and Processed Products 909Elisabet Fernández-García and Antonio Pérez-Gálvez

48. Potato Flavor 935Shelley H. Jansky

PART VIII OTHER VEGETABLES AND VEGETABLE PRODUCTS 947

49. Mexican Pickled Jalapeño Pepper 949María Eugenia Jaramillo-Flores, Lidia Dorantes-Alvarez, Raquel García-Barrientos, and Jorge Welti-Chanes

50. Dried Western Vegetable Products 963Leo Nollet

CONTENTS ix

51. Understanding Peanut Flavor: A Current Review 985E.R. Neta, T. Sanders, and M.A. Drake

52. Spices, Seasonings, and Essential Oils 1023Marisa F. Mendes and Fernando L.P. Pessoa

53. Flavor of Canola Oil 1033Silvana Martini

54. Flavors of Palm Oil 1051Ainie Kuntom and Abd. Azis Ariffi n

55. Sesame Oil 1071Neusa P. Arruda, Cláudia M. de Resende, and Suely P. Freitas

INDEX 1083

xi

For the last 50 years, our knowledge of the science and technology of food fl avors has increased tremendously. To distribute the information, publishers have released many professional reference books on the subject. In general, most of the books have some discussion on the fl avors of fruits and vegetables, though several of them specifi cally address these two important groups of food commodities. Information on the fl avors of fruits and vegetables is essential to the work of government, aca-demia, and industry. This book is an updated reference treatise on the fl avors of fruits and vegetables. It includes 55 chapters, with 31 on the fl avor of fruits and 24 on that of vegetables, covering the following topics:

• biology, chemistry, and biochemistry • biotechnology and genetic engineering • analytical methodology • processing technology • fresh and processed commodities • products derived from processed fruits and vegetables • regulatory consideration

There are several professional books on the subject matter and the preference for any particular one depends on the needs of the users. Although many topics are included in this volume, we do not claim that the coverage is comprehensive.

This work is the result of the combined efforts of more than 70 individuals from industry, government and academia worldwide. They represent the expertise of professionals from 18 countries including Belgium, Brazil, China, Canada, Croatia, Cuba, France, India, Indonesia, Israel, Italy, Japan, Kuwait, Malaysia, Mexico, Portugal, Spain, Turkey, and the United States. The editorial team consists of 12 established experts in the fl avors or processing of fruits and vegetables. Each con-tributor or editor was responsible for researching and reviewing subjects of immense depth, breadth, and complexity. Care and attention were paramount to ensure tech-nical accuracy for each topic. In sum, this volume is unique in many respects. It is our sincere hope and belief that it will serve as an essential reference on the fl avors of major plant foods.

We wish to thank all the contributors for sharing their expertise throughout our journey. We also thank the reviewers for giving their valuable comments leading to

PREFACE

xii PREFACE

improvements in the contents of each chapter. In addition, we thank members of the production team at John Wiley and Sons for their time, effort, advice, and exper-tise. All these professionals are the ones who made this book possible. You are the best judge of the quality of their work and we trust that you will benefi t from the fruits of their labor.

Y. H. Hui F. Chen

L. M. L. Nollet R. P. F. Guin é

J. L. Le Qu é r é O. Mart í n - Belloso

M. I. M í nguez - Mosquera G. Paliyath

F. L. P. Pessoa J. S. Sidhu

N. Sinha P. Stanfi eld

xiii

Nese B. Agkul, Department of Food Engineering, Faculty of Engineering, Hacettepe University, Ankara, Turkey

Asaph Aharoni, Department of Plant Sciences, Weizmann Institute of Science, Rehovot, Israel

P.F.F. Amaral, Departamento de Engenharia Bioqu í mica, Escola de Qu í mica/UFRJ, Rio de Janeiro, Brazil

Ram ó n Aparicio, Instituto de la Grasa, Padre Garc í a Tejero, Seville, Spain

Abd. Azis Ariffi n, Malaysian Palm Oil Board, Kuala Lumpur, Malaysia

Neusa P. Arruda, Instituto Federal do Rio de Janeiro, Rio de Janeiro, Brazil

Pervin Basaran, Department of Food Engineering, Suleyman Demirel University, Cunur, Isparta, Turkey

A.S. Bawa, Defence Food Research Laboratory, Siddarthanagar, Mysore, India

Sara Bayarri, Instituto de Agroqu í mica y Tecnolog í a de Alimentos, CSIC, Burjassot, Valencia, Spain

N.R. Bhat, Aridland Agriculture and Greenery Department, Food Resources and Marine Sciences Division, Kuwait Institute for Scientifi c Research, Safat, Kuwait

H.R. Bizzo, Embrapa Agroind ú stria de Alimentos, Av. das Am é ricas, Rio de Janeiro, Brazil

Terri D. Boylston, Department Food Science & Human Nutrition, Iowa State University, Ames, IA

A. Bravo, Curso de P ó s - gradua ç ã o em Ci ê ncia de Alimentos, Instituto de Qu í mica/UFRJ, Rio de Janeiro, Brazil

Ron G. Buttery, U.S. Department of Agriculture, Agricultural Research Service, Western Regional Research Center, Albany, CA

L.M.C. Cabral, Embrapa Agroind ú stria de Alimentos, Rio de Janeiro, Brazil

Juan Cacho, Laboratory for Flavor Analysis and Enology, Aragon Institute of Engineering Research, Analytical Chemistry Department, Faculty of Sciences, University of Zaragoza, Zaragoza, Spain

CONTRIBUTORS

xiv CONTRIBUTORS

O.P. Chauhan, Defence Food Research Laboratory, Siddarthanagar, Mysore, India

Feng Chen, Department of Plant Sciences, University of Tennessee, Knoxville, TN

Gerson L.V. Coelho, Departamento de Engenharia Qu í mica, Escola de Qu í mica, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil

M.A.Z. Coelho, Departamento de Engenharia Bioqu í mica, Escola de Qu í mica/UFRJ, Rio de Janeiro, Brazil

Gemma Echeverr í a Cortada, Postharvest Department, IRTA, Lleida, Spain

Elvira Costell, Instituto de Agroqu í mica y Tecnolog í a de Alimentos, CSIC, Burjassot, Valencia, Spain

Cl á udia M. de Resende, Universidade Federal do Rio de Janeiro, Instituto de Qu í mica, Rio de Janeiro, Brazil

B.B. Desai, Aridland Agriculture and Greenery Department, Food Resources and Marine Sciences Division, Kuwait Institute for Scientifi c Research, Safat, Kuwait

M é rcia de Sousa Galv ã o, Laborat ó rio de An á lise de Flavor, N ú cleo de P ó s - Gradua ç ã o em Ci ê ncia e Tecnologia de Alimentos, Universidade Federal do Sergipe, S ã o Crist ó v ã o - SE, Brazil

Lidia Dorantes - Alvarez, Escuela Nacional de Ciencias Biol ó gicas - IPN, Departa-mento de Graduados en Alimentos, Carpio y Plan de Ayala, Mexico, D.F.

M.A. Drake, Department of Food, Bioprocessing and Nutrition Sciences, North Carolina State University, Raleigh, NC

Judy A. Driskell, Department of Community Nutrition, Faculty of Human Ecology, Bogor Agricultural University (IPB), Indonesia; and Department of Nutrition and Health Sciences, University of Nebraska – Lincoln, Lincoln, NE

Bing Du, College of Food Science, South China Agricultural University, Guangzhou, China

Elisabet Fern á ndez - Garc í a, Grupo de Qu í mica y Bioqu í mica de Pigmentos, Departamento de Biotecnolog í a de Alimentos, Instituto de la Grasa (CSIC), Seville, Spain

Vicente Ferreira, Laboratory for Flavor Analysis and Enology, Aragon Institute of Engineering Research, Analytical Chemistry Department, Faculty of Sciences, University of Zaragoza, Zaragoza, Spain

Luciana Francisco Fleuri, Lab. de Bioquimica de Alimentos, DCA - FEA - UNICAMP, Campina SP, Brazil

M. Freire, Jr., Embrapa Agroind ú stria de Alimentos, Rio de Janeiro, Brazil

Suely P. Freitas, Universidade Federal do Rio de Janeiro, Escola de Qu í mica, Rio de Janeiro, Brazil

CONTRIBUTORS xv

Maria Luisa L ó pez Fructuoso, Department of Food Technology, University of Lleida, Lleida, Spain

Karin Kova č evi c ́ Gani ć , Faculty of Food Technology and Biotechnology, University of Zagreb, Zagreb, Croatia

Raquel Garc í a - Barrientos, Universidad Autónoma Metropolitana — Unidad Iztapalapa, Mexico, D.F.

Diego Luis Garc í a - Gonz á lez, Instituto de la Grasa, Padre Garc í a Tejero, Seville, Spain

Clarice N. Gobbi, Universidade Federal do Rio de Janeiro, Escola de Qu í mica, Rio de Janeiro, Brazil

Fahrettin G ö ǧ ü ş , The University of Pamukkale, Faculty of Science & Arts, Chemistry Department, Denizli, Turkey

E. G ó mez - Plaza, Food Science and Technology Department, University of Murcia, Murcia, Spain

F. Guti é rrez - Rosales, Instituto de la Grasa (CSIC), Seville, Spain

Qayyum Husain, Department of Biochemistry, Faculty of Life Sciences, A.M.U. Aligarh, India

Yoko Iijima, Kazusa DNA Research Institute, Kisarazu, Japan

Shelley H. Jansky, USDA - ARS and Department of Horticulture, University of Wisconsin - Madison, Madison, WI

Mar í a Eugenia Jaramillo - Flores, Escuela Nacional de Ciencias Biol ó gicas - IPN, Departamento de Graduados en Alimentos, Carpio y Plan de Ayala, Mexico, D.F.

Yueming Jiang, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou, China

Yearul Kabir, Department of Biochemistry and Molecular Biology, University of Dhaka, Dhaka, Bangladesh

Dra ž enka Komes, Faculty of Food Technology and Biotechnology, University of Zagreb, Zagreb, Croatia

Ainie Kuntom, Malaysian Palm Oil Board, Kuala Lumpur, Malaysia

C. Ledbetter, Crop Diseases, Pests & Genetics Research Unit, USDA, ARS, Parlier, CA

Jean - Luc Le Qu é r é , Institut National de la Recherche Agronomique (INRA), UMR 1129 Flavor, Vision and Consumer Behavior (FLAVIC), Dijon, France

Efraim Lewinsohn, Newe Yaar Research Center, Agricultural Research Organization, Ramat Yishay, Israel

Jingyu Lin, Department of Plant Sciences, University of Tennessee, Knoxville, TN

xvi CONTRIBUTORS

Mar í a Asunci ó n Longo, Department of Chemical Engineering, University of Vigo, Campus Universitario As Lagoas, Marcosende, Vigo, Spain

Gabriela Alves Macedo, Lab. de Bioquimica de Alimentos, DCA - FEA - UNICAMP, Campina SP, Brazil

Juliana Alves Macedo, Lab. de Bioquimica de Alimentos, DCA - FEA - UNICAMP, Campina SP, Brazil

Olga Mart í n - Belloso, Department of Food Technology, University of Lleida, Lleida, Spain

Silvana Martini, Department of Nutrition and Food Sciences, Utah State University, Logan, UT

V.M. Matta, Embrapa Agroind ú stria de Alimentos, Av. das Am é ricas, Rio de Janeiro, Brazil

Adriane B.P. Medeiros, Divis ã o de Engenharia de Bioprocessos and Biotecnologia, Departamento de Engenharia Qu í mica, Universidade Federal do Paran á , Curitiba, PR, Brazil

Emira Mehinagic, Groupe ESA, Laboratory GRAPPE, Angers, France

Marisa F. Mendes, Laborat ó rio de Termodin â mica Aplicada/Departamento de Engenharia Qu í mica, Universidade Federal Rural do Rio de Janeiro, Serop é dica, Rio de Janeiro, Brazil

Marta Montero - Calder ó n, Department of Food Technology, University of Lleida, Lleida, Spain

Mar í a Teresa Morales, Facultad de Farmacia, Universidad de Sevilla, Seville, Spain

Cristina Mu ñ oz, Departamento de Biolog í a Molecular y Bioqu í mica, Universidad de M á laga, M á laga, Spain

Regina Nabais, CERNAS — Centro de Recursos Naturais, Ambiente e Sociedade, Escola Superior Agr á ria de Coimbra, Instituto Polit é cnico de Coimbra, Bencanta, Coimbra, Portugal

Narendra Narain, Laborat ó rio de An á lise de Flavor, N ú cleo de P ó s - Gradua ç ã o em Ci ê ncia e Tecnologia de Alimentos, Universidade Federal do Sergipe, S ã o Crist ó v ã o - SE, Brazil

E.R.D. Neta, Department of Food, Bioprocessing and Nutrition Sciences, North Carolina State University, Raleigh, NC

Nisha Nigam, Department of Chemistry, R.D. National College, Bandra - West, Mumbai, India

Leo Nollet, University College Ghent, Member of Ghent University Association, Faculty of Applied Engineering Sciences, Ghent, Belgium

Moustapha Oke, Ontario Ministry of Agriculture, Food and Rural Affairs, Guelph, Ontario, Canada

CONTRIBUTORS xvii

Sonia Osorio, Departamento de Biolog í a Molecular y Bioqu í mica, Universidad de M á laga, M á laga, Spain

Mustafa Z. Ö zel, The University of Pamukkale, Faculty of Science & Arts, Chemistry Department, Denizli, Turkey

Gopinadhan Paliyath, Department of Plant Agriculture, University of Guelph, Guelph, Ontario, Canada

Vincent R. Pantalone, Department of Plant Sciences, University of Tennessee, Knoxville, TN

Clara Pelayo - Zald í var, Food Sciences, Department of Biotechnology, Universidad Aut ó noma Metropolitana - Iztapalapa, Mexico

Ana G. P é rez, Department of Physiology and Technology of Plant Products, Instituto de la Grasa, CSIC, Seville, Spain

Antonio P é rez - G á lvez, Grupo de Qu í mica y Bioqu í mica de Pigmentos, Departa-mento de Biotecnolog í a de Alimentos, Instituto de la Grasa (CSIC), Seville, Spain

Fernando L.P. Pessoa, GIPQ/DEQ/EQ/UFRJ, CT, Rio de Janeiro, Brazil

Jorge A. Pino, Instituto de Investigaciones para la Industria Alimentaria, Havana, Cuba

Linda Pollak, USDA - ARS Corn Insects and Crop Genetics Research Unit, Ames, IA

Eduardo M. Queiroz, Departamento de Engenharia Qu í mica, Escola de Qu í mica, Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil

Theodore J.K. Radovich, Department of Tropical Plant and Soil Sciences, University of Hawai ’ i at M ā noa, Honolulu, HI

P.S. Raju, Defence Food Research Laboratory, Siddarthanagar, Mysore, India

M.H.M. Rocha - Le ã o, Departamento de Engenharia Bioqu í mica, Escola de Qu í mica/UFRJ, Rio de Janeiro, Brazil

M. Alejandra Rojas - Gra ü , Department of Food Technology, University of Lleida, Lleida, Spain

Suzan C. Rossi, Divis ã o de Engenharia de Bioprocessos & Biotecnologia, Departamento de Engenharia Qu í mica, Universidade Federal do Paran á , Curitiba, PR, Brazil

T.H. Sanders, United States Department of Agriculture, Agricultural Research Service, Market Quality and Handling Research Unit, Raleigh, NC

Mar í a Angeles Sanrom á n, Department of Chemical Engineering, University of Vigo, Campus Universitario As Lagoas, Marcosende, Vigo, Spain

Carlos Sanz, Department of Physiology and Technology of Plant Products, Instituto de la Grasa, CSIC, Seville, Spain

xviii CONTRIBUTORS

Jiwan S. Sidhu, Department of Family Sciences, College for Women, Kuwait University, Safat, Kuwait

Carlos R. Soccol, Divis ã o de Engenharia de Bioprocessos & Biotecnologia, Departamento de Engenharia Qu í mica,Universidade Federal do Paran á , Curitiba, PR, Brazil

Jun Song, Agriculture and Agri - Food Canada, Atlantic Food and Horticulture Research Centre, Kentville, Nova Scotia, Canada

Ahmad Sulaeman, Department of Community Nutrition, Faculty of Human Ecology, Bogor Agricultural University (IPB), Indonesia; and Department of Nutrition and Health Sciences, University of Nebraska – Lincoln, Lincoln, NE

M.K. Suleiman, Aridland Agriculture and Greenery Department, Food Resources and Marine Sciences Division, Kuwait Institute for Scientifi c Research, Safat, Kuwait

Victoriano Valpuesta, Departamento de Biolog í a Molecular y Bioqu í mica, Universidad de M á laga, M á laga, Spain

Elena Venir, Department of Food Science, University of Udine, Udine, Italy

Silvio A.B. Vieira de Melo, Programa de Engenharia Industrial, Escola Polit é cnica, Universidade Federal da Bahia, Salvador - BA, Brazil

Juan Wang, College of Light Industry and Food Sciences, South China University of Technology, Guangzhou, China; and College of Food Science, South China Agricultural University, Guangzhou, China

Ningning Wang, College of Life Sciences, Nankai University, Tianjin, China

Jorge Welti - Chanes, Instituto Tecnol ó gico y de Estudios Superiores de Monterrey — Campus Monterrey, Mexico, D.F.

Gong Ming Yang, College of Food Science, South China Agricultural University, Guangzhou, China

xix

AECA aroma extract concentration analysis AEDA aroma extraction dilution analysis APCI - MS atmospheric pressure chemical ionization – mass spectrometry DAD diode array detection DSA descriptive sensory analysis GC gas chromatography GC - FTIR gas chromatography – Fourier transform infrared spectroscopy GC - MS gas chromatography – mass spectrometry GC - O gas chromatography – olfactometry HPLC high - performance liquid chromatography HPLC - DAD high - performance liquid chromatography diode array detection HPLC - DAD - MS/MS - ESI high - performance liquid chromatography – diode array detection – mass spectrometry/mass spectrometry – electrospray ionization HRGC high - resolution gas chromatography HRGC - MS high - resolution gas chromatography – mass spectrometry HS headspace HSE headspace extraction HSSE headspace sorptive extraction LC liquid chromatography LLE liquid - liquid extraction OPLC optimum performance laminar chromatography PTR - MS proton transfer reaction mass spectrometry SBSE stir bar sorptive extraction SDE simultaneous distillation - extraction SDEV simultaneous distillation - extraction under vacuum SFC supercritical fl uid chromatography SFE supercritical fl uid extraction SPE solid phase extraction SPME solid phase microextraction SPME - GC solid phase microextraction – gas chromatography

LIST OF ABBREVIATIONS

xx LIST OF ABBREVIATIONS

SPME - GC - MS solid phase microextraction – gas chromatography – mass spectrometry SSF solid - state fermentation TD - GC – MS thermal desorption – gas chromatography – mass spectrometry TLC thin - layer chromatography UAE ultrasound - assisted extraction VHS vacuum headspace

PART I

FRUIT FLAVORS: BIOLOGY, CHEMISTRY, AND PHYSIOCHEMISTRY

SECTION A: FRUIT FLAVORS

1. Fruits and Fruit Flavor: Classifi cation and Biological Characterization 3

2. Physiology and Biochemistry of Fruit Flavors 25

3. Sensory Evaluation of Fruit and Vegetable Flavors 45

4. Fermentation and Fruit Flavor Production 59

5. Environmental Effects on Flavor Changes 73

3

Handbook of Fruit and Vegetable Flavors, Edited by Y. H. HuiCopyright © 2010 John Wiley & Sons, Inc.

CHAPTER 1

Fruits and Fruit Flavor: Classifi cation and Biological Characterization

YUEMING JIANG 1 and JUN SONG 2

1 Chinese Academy of Sciences 2 Atlantic Food and Horticulture Research Centre

Fruit has always been a part of the human diet and is an important nutritional source, with high water content (70 − 85%) and a relatively high amount of carbo-hydrates but low contents of fat (less than 0.5%) and protein ( < 3.5%). It usually contains many useful vitamins as well as minerals, dietary fi ber, and antioxidants (Goff and Klee 2006 ; Knee 2002 ). From 2002 to 2007, there has been a steady increase in fruit production with 2.67% each year, partly in response to population growth and living standard improvement in most countries and effective encourage-ment by government health agencies of fruit consumption. In 2007, a total amount of 318.6 million tons of fruit was produced in the world, which is equivalent to 48.2 kg per capita of production and a fruit consumption of 12 kg per capita (Euromonitor 2008 ; http://faostat.fao.org ).

In this chapter, the botanical information, characterization, importance, and production of fruits are briefl y reviewed. The chapter provides general informa-tion about fruit and draws comparisons between fruit and fruit fl avor. Flavor characterization is also discussed in detail.

CLASSIFICATION OF FRUITS

There are different ways to classify fruit (Table 1.1 ). Generally speaking, the outer, often edible layer in fl eshy fruits is the pericarp, which develops from the ovary wall of the fl ower and surrounds the seeds. While the seeds are akin to the egg develop-ment in the ovary of a fowl, the pericarp may be assumed as the uterus. However, a small number of fruits do not fi t into this description. For example, in most nuts, the edible part is the seed but not the pericarp. In addition, many edible vegetables such as cucumber and squash are common pericarp and are botanically considered as fruits. In this chapter, the use of the term “ fruit ” will not refer to these vegetable

4 HANDBOOK OF FRUIT AND VEGETABLE FLAVORS

TABLE 1.1. Types of Fruit

True Berry Pepo Hesperidium False Berry

Aggregate Fruit

Multiple Fruit

Other Accessory

Fruit

Black currant Pumpkin Orange Banana Blackberry Pineapple Apple Red currant Cucumber Lemon Cranberry Raspberry Fig Peach Gooseberry Melon Lime Blueberry Boysenberry Mulberry Cherry Pomegranate Grapefruit Hedge apple Strawberry Avocado Kiwifruit Grape

fruits. In some fruits such as lychee and longan, the edible portion is actually an aril. From the botanical point, fruits can be classifi ed into simple fruits, aggregate fruits, and multiple fruits on the basis of anatomical attributes.

Simple Fruits

Simple fruits are formed from a single ovary and may contain one to many seeds, which have developed as part of the fruit. Simple fruits can be divided into two groups: fl eshy pericarp — berries, drupes, and pomes; and dry pericarp — nuts. Types of fl eshy and simple fruits are berry (red currant, gooseberry, and avocado), stone fruit or drupe (plum, cherry, peach, apricot, olive), false berry — epigynous accessory fruits such as banana and cranberry, and pome — accessory fruits such as apple and pear. In contrast to fl eshy and simple fruits, in nuts, it is the stony layer that surrounds the kernel of pecans and is removed when eating.

Aggregate Fruit

Aggregate fruits are formed from a single compound fl ower and contain many ovaries. Examples include strawberries, raspberries, and blackberries. An aggregate fruit or etaerio develops from a fl ower with numerous simple pistils. An example is the raspberry, whose simple fruits are termed as drupelets because each is like a small drupe attached to the receptacle. In some bramble fruits (such as blackberry), the receptacle is elongated and part of the ripe fruit, which makes the blackberry an aggregate - accessory fruit. The strawberry is also an aggregate fruit, in which the seeds are contained in achenes.

Multiple Fruit

Multiple fruits, such as pineapple, fi g, and mulberry, are formed from the fused ovaries of many separate but closely clustered fl owers. There are also many dry multiple fruits, for example, tulip tree (multiple of samaras), sweet gum (multiple of capsules), sycamore and teasel (multiple of achenes), and magnolia (multiple of follicles).

As described above, fruits can be summarized into eight types: (1) berry — simple fruit and seeds developed from a single ovary, (2) pepo — berries where the skin is

FRUITS AND FRUIT FLAVOR 5

hardened, (3) hesperidium — berries with a rind, (4) false berries — epigynous fruit made from a part of the plant other than a single ovary, (5) compound fruit — from several ovaries in either a single fl ower or multiple fl owers, (6) aggregate fruit — multiple fruits with seeds from different ovaries of a single fl ower, (7) multiple fruit — fruits of separate fl owers packed closely together, and (8) other accessory fruit — where the edible part is not generated by the ovary. Another common way to classify fruits is based on growing regions such as temperate zone fruits, sub-tropical fruits, and tropical fruits (Kader 2002 ).

SPECIES, VARIETIES, AND BIOLOGICAL CHARACTERISTIC OF MAJOR FRUITS

The major fruits, such as apple, pear, grape, strawberry, citrus, banana, and mango, currently contribute the most of the total world production. About two - thirds of the major fruits produced worldwide are consumed as fresh fruit.

As discussed above, fruits are classifi ed mainly on the basis of the ovary charac-teristic. In biology, fruit species can be classifi ed by their botanical origin. In this following section, species, varieties, biological characteristic, and production of major fruits are briefl y reviewed.

Apple

The genus Malus belongs to the Rosaceae family and forms with its closely related fruit ( Pyrus and Cydonia ) and ornamental ( Amelanchier , Aronia , Chaenomweles , Cotoneaster , Crateagus , Pyracantha , Sorbus ) genera, the subfamily Maloideae. Nowadays, Malus × domestica Borkh has been widely applied for apples.

World apple production reached 66 million tons in 2007 (Euromonitor 2008 ). Apple production is dominated by cultivars, such as “ Delicious, ” “ Gold Delicious, ” “ McIntosh, ” “ Jonathan, ” “ Cox ’ s Orange Pippin, ” “ Granny Smith, ” and “ Braeburn. ” In Asia, these varieties often replace the local varieties selected from the native species Malus prunifolia and its cultivated species Malus asiatica . China ’ s enormous growth in apple production is entirely due to the introduction of the “ Fuji ” cultivar.

Banana

Banana belongs to the genus Musa in the family Musaceae, order Zingiberales. The family Musaceae comprises two genera viz., Musa and Ensete . The genus Musa comprises all the edible bananas and plantains with over 50 species. Bananas are perennial monocotyledonous herbs that grow well in humid tropical and subtropical regions. The origin of banana is traced back to Southeast Asia in the jungles of Malaysia, Indonesia, or the Philippines. Banana originated from two wild diploid species namely, Musa acuminate Coll and Musa balbisiana Coll. M. acuminate is native of the Malay Peninsula and adjacent regions, while M. balbisiana is found in India eastward to the tropical Pacifi c.

Bananas are the fourth world ’ s most important food crop after rice, wheat, and maize, with production of 73 million tons in 2007 (Euromonitor 2008 ). The majority


of the banana crops are grown in the tropical and subtropical zones. From a consumer perspective, bananas are nutritious with a pleasant fl avor and widely consumed throughout the world. India is the world ’ s leading producer of banana and plantain, followed by Brazil and China.

Grape

The Vitis vinifera L. grape is one of the oldest cultivated plants and is thought to have originated in the region between the Mediterranean and the Caspian Sea. Cultivars of the vine slowly spread eastward across southern Asia and westward around the Mediterranean Sea. The Germplasm Resources Information Network ( http://www.ars - grin.gov ) of the United States Department of Agriculture describes the genera and 43 species, 5 natural hybrids, and 15 varieties of species in Vitis . V. vinifera is the most successfully used grape species with thousands of wine, table, and raisin grape cultivars grown throughout the world ’ s temperate zones.

Grapes are now grown in more than 90 countries of the world and become the world ’ s largest fruit crop with a total production of 69 million tons (Euromonitor 2008 ). The countries with the greatest acreage are Spain, France, Italy, Turkey, China, and the United States. The leading countries for the production of table grapes consumed as fresh fruit are China, Turkey, Italy, Chile, and the United States.

Citrus Fruit

Citrus, belonging to the family Rutaceae, is one of the world ’ s most important fruit. Citrus can be eaten as a fresh fruit, processed into juice, or added to dishes and beverages. The major types of edible citrus include citron ( Citrus medica L.); pomelo or shaddock ( Citrus grandis ); tangerine, mandarin, or satsuma ( Citrus reticulata Blanco); limes ( Citrus aurantifolia L.); sour orange ( Citrus aurantium L.); sweet oranges ( Citrus sinensis [L.] Osbeck); lemon ( Citrus limon L.); and grapefruit ( Citrus paradisi Macfad.). Brazil, the United States, and China are the three largest citrus producers in the world.

Strawberry

Strawberry belongs to the genus Fragaria . The genus is comprised of 32 species. Historically, several Fragaria species and novel hybrids have been brought into cultivation in different parts of the world, including Fragaria chiloensis in South America, and Fragaria moschata and Fragaria vesca in Europe. However, strawberry ( Fragaria × ananassa Duch) is one of the most widely grown small fruits in the world. The large modern fruit of today was developed in the early 18th century by the cross between the wild strawberry F. chiloensis and Fragaria virginiana .

Globally, a large part of the cultivated area is located in Europe, followed by Asia and North and Central America. In 2004, a total production of strawberry reached to 2.4 million tons in the world. The United States is the world ’ s leading strawberry producer with China, Spain, and Korea. Some countries like Turkey, Morocco, and Egypt have strongly increased their production.


Peach

Peach belongs to the Prunoideae subfamily of the family Rosaceae. In temperate regions, the family ranks third place in economic importance. The genus Prunus is characterized by species that produce drupes known as “ stone fruit. ” The edible portion of the fruit is a juicy mesocarp. There are three major groups of cultivars: nectarines, freestone peaches, and clingstone peaches. All commercial varieties of peach are Prunus persica (L.) Batsch, including nectarines differing from peach in the absence of pubescence ( “ fuzzless ” ) on the fruit surface. Peaches originated in China, with a cultivation history of over 4000 years. Peach is grown in all continents except Antarctica, and world peach production has increased steadily in recent year.

Pear

Pear species belong to the genus Pyrus , the subfamily Maloideae (Pomoideae) in the family Rosaceae. There are about 22 primary species in the genus , all of which originate in either Asia or Europe. The pear has been cultivated in China for at least 3000 years. There are two major species, European pear ( Pyrus communis L.) and Asian pear ( Pyrus pyrifolia L.), which are commercially cultivated. The fi rst species to be domesticated was P. pyrifolia (Burm.) Nakai because the wild type is edible but without selection. Later, the hardy northern Chinese type Pyrus ussuriensis Maxim probably became cultivated after selection from the wild type. Natural hybridization between these two wild species likely occurred in China to produce the modern “ Ussuri ” cultivars in northern China. In other parts of the world, cultivated pears have been derived from P. communis L., while P. communis var. pyraster and/or P. communis var. caucasica were probably the ancestors of the common pear of Europe, but “ French ” cultivars may be complex hybrids of these two.

Pear is the third important temperate fruit after grape and apple. Asia produces the most, followed by Europe, North and Central America, and South America. Among countries, China produced the most, followed by the United States, Italy, and Spain. Pears can be consumed as fresh fruit, fruit juice, cube for fruit salad, canned product, and dry fruit. About 80% of the total pear production is destined for fresh consumption.

Mango

The genus Mangifera , belonging to the dicotyledonous family “ Anacardiaceae, ” originates in the Indo - Burma region. Almost all the edible cultivars of mango are the single species Mangifera indica L., which originated in the Indian subcontinent. The few other species that contribute edible fruits are Mangifera caesia , Mangifera foetida , and Mangifera odorata , which are confi ned to the Malaysian region.

Mango is a very important tropical fruit and popularly known as the “ apple of the tropics. ” Mango is commercially grown in over 103 countries of the world. The major growing countries in the world are India, China, Mexico, Pakistan, Indonesia, Thailand, the Philippines, Brazil, Australia, Nigeria, and Egypt ( http://faostat.fao.org ). There are more than 1000 varieties of mango under cultivation, but only a few of them are grown on a commercial scale.


Papaya

Papaya ( Carica papaya L.) belongs to family Caricaceae, which consists of six genera including Carica a monotypic genus, Jacaratia (7 species), Jarilla (3 species), Cylicomorpha (2 species), Horovitzia (1 species), and Vasconcellea (21 species). Carica is the only genus of Caricaceae containing the domesticated species papaya , which is by far the most economically important and has a wide distribution through-out the tropics and subtropics of the world. Papaya probably originated in the lowland of Central America between southern Mexico and Nicaragua, and is now cultivated in many tropical and subtropical regions.

Papaya is a major tropical fruit grown commercially in India, Brazil, Mexico, Australia, Hawaii, Thailand, South Africa, the Philippines, Indonesia, and China. In recent years, intensive improvements and selections have given rise to a large number of papaya varieties, such as “ Kapoho Solo, ” “ Sun Rise, ” “ Sun Set, ” “ Waimanalo, ” “ Kamiya ” (United States), “ Pusa Delicious, ” “ Pusa Nanha, ” “ Pusa Dwarft, ” “ Surya ” (India), “ Cavite Special ” (the Philippines), “ Sainampueng, ” “ Kak Dum ” (Thailand), and improved “ Peterson, ” “ Guinea ” and “ Gold and Sunnybank ” (Australia).

Pineapple

Pineapple is a perennial monocot belonging to the family of Bromeliaceae, subfam-ily Bromelioideae. The Bromelioideae comprises 56 genera with more than 2000 species, which are classifi ed into three subfamilies: Pitcarnioideae, Tillandsioideae, and Bromelioideae. This last subfamily shows a tendency toward the fusion of fl oral parts, a trait most developed in Ananas . Many distinctions, particularly those related to fruit size and fertility, appear to be the direct result of human selection in the course of domestication.

Pineapple is the third most important tropical fruit after bananas and mangoes and has been cultivated in South America since the 15th century. Owing to its attrac-tive sweet fl avor, pineapple is widely consumed as fresh fruit, processed juice, and canned fruit, and is used as an ingredient in exotic foods. Five countries, Thailand, the Philippines, Brazil, China, and India, contribute to the major production in the world.

Plum

Plums belong to subfamily Prunoideae of the family of Rosaceae. Prunus species are divided into three major subgenera: Prunophora (plum and apricots), Amygdalus (peaches and almonds), and Cerasus (sweet and sour cherries). The subgenus Prunophora is divided into two main sections: Euprunus groups (plum species) and Armeniaca , which contains the apricot species. Plum has been domesticated inde-pendently in Europe, Asia, and America. In Europe, Prunus domestica L. is the most important source of fruit cultivars and has been grown for over 2000 years. In Asia, the Japanese plum Prunus salicina L. originates from China where it has been cul-tivated since ancient times. In north America, the third plum domestication source, a wide range of native species, such as Prunus americana Marsh., Prunus hortulana Bailey, Prunus angustifolia Marsh., and Prunus maritima Marsh., are present. The


major production of plum is located in Europe and Asia. In Europe, Germany is the leading producer.

FRUIT FLAVOR

The consumption of fresh fruit is dependent on the fruit quality (Baldwin et al. 2007 ; L ó pez et al. 2007 ). The quality of fresh fruit includes many aspects such as appear-ance, color, texture, fl avor, and nutritional value (Kader 2002 ; Song 2007 ). Among them, fl avor is one of the most important quality traits for fresh fruit (Dirinck et al. 1989 ; Dull and Hulme 1971 ; Maarse 1991 ; Reineccius 2006 ). Fruit fl avor is made up of sugars, acids, salts, bitter compounds such as alkaloids or fl avonoids, and aroma volatiles (Dirinck et al. 1989 ; Salunkhe and Do 1976 ; Song and Forney 2008 ). The fl avor of fresh fruit is determined by taste and aroma (odor - active compounds). The contribution of odor - active compounds to the fruit fl avors has gained increasing attention because these compounds are important for the characteristic fl avors of fruits (Baldwin 1993 , 2002b ; Br ü ckner 2008 ). The present chapter refers specifi cally the term “ fl avor ” to the volatile compounds. Volatile compounds in fruits are diverse, consisting of hundreds of different chemical compounds comprising only 10 − 7 – 10 − 4 of the fresh fruit weight (Berger 2007 ; Br ü ckner 2008 ). Although these volatile compounds are produced in trace amounts, they can be detected by human olfaction. The diversity is partially responsible for the unique fl avors found in different fruit species. The importance of volatile production in fruit related to its infl uencing factors has been intensively investigated and/or reviewed (Baldwin 2002 ; Dixon and Hewett 2000 ; Fellman et al. 2000 ; Forney et al. 2000 ; Song 2007 ; Song and Forney 2008 ).

Classifi cation of Volatile Compounds in Fruit Flavor

Chemical Structure Various types of fresh fruits produce distinct volatile profi les. Volatile compounds, which are produced by fresh fruits, are mainly comprised of diverse classes of chemicals, including esters, alcohols, aldehydes, ketones, lactones, and terpenoids (Table 1.2 ). However, some sulfur compounds, such as S - methyl thiobutanoate, 3 - (methylthio) propanal, ethyl 2 - (methylthio) acetate, ethyl 3 - (methylthio) propanoate, and 3 - (methylthio) propyl acetate, also contribute to the fl avor of fruit such as melons (Song and Forney 2008 ). Although an overwhelming number of chemical compounds have been identifi ed as volatile compounds in fresh fruit, only a fraction of these compounds have been identifi ed as impact compounds of fruit fl avor based on their quantitative abundance and olfactory thresholds (Cunningham and Barry 1986 ; Schieberle et al. 1990 ; Wyllie et al. 1995 ).

Biogenesis Volatile compounds forming the fruit fl avor are produced through many metabolic pathways during fruit ripening and postharvest storage, and depend on many factors related to the species, variety, climate, production, maturity, and pre - and postharvest handling. For most fruits, volatile production is closely related to fruit ripening. As direct products of a metabolic pathway or as a result of inte-ractions between pathways or end products, volatile compounds can be classifi ed by the biogenesis: fatty acids (FAs), amino acids, glucosinolates, terpenoid, phenol, and related compounds (Berger 2007 ). However, from the point of chemical

TABLE 1.2. Volatile Compounds Present in Fruit Flavor

Esters Alcohols Aldehydes Ketones Lactones Terpenoids

Butyl acetate Benzyl alcohol Acetaldehyde 2,3 - Butanedione γ - Butyrolactone β - Caryophyllene Butyl butanoate Butan - 1 - ol Benzaldehyde β - Damsenone

eucalyptol γ - Decalactone 1,8 - Cineole

Butyl hexanoate ( E ) - cinnamyl alcohol

( E ) - cinnamaldehyde

Eugenol δ - Decalactone Citral

Butyl - 2 - methyl butanoate 1 - Hexanol ( E , E ) - 2,4 - decadienal

2 - Heptanone γ - Dodecalactone β - Damascenone

Butyl propanoate ( E ) - 2 - hexenol Hexanal 4 - ( p - Hydroxyphenyl) - 2 - butanone

δ - Dodecalactone Dihydroedulan

Ethyl acetate ( Z ) - 3 - hexenol ( E ) - 2 - hexenal 3 - Hydroxy - 2 - butanone

γ - Jasmolactone Farnesyl acetate

Ethyl butanoate 1 - Octanol ( Z ) - 3 - hexenal β - Ionone γ - Octalactone Geraniol Ethyl 9 - decenoate ( Z ) - 6 - nonenol ( Z ) - 3 - hexenal Linalool δ - Octalactone Hotrienol Ethyl hexanoate Hexan - 1 - ol Nonanal 6 - Methyl - 5 - heptene -

2 - one α - Ionone

Ethyl 2 - methylbutanoate ( Z , Z ) - 3,6 - nonadienol

( Z ) - 6 - nonenal Nerolidol β - Ionone

Ethyl 3 - methylbutanoate 1 - Phenylethanol ( E , Z ) - 2,6 - nonadienal

1 - Octen - 3 - one Limonene

Ethyl 2 - methylpropanoate 2 - Phenylethanol ( E ) - 2 - nonenal 2 - Pentanone Linalool Ethyl 2 - methylbutanoate Phenylacetaldehyde ( Z ) - 1,5 - octadien - 3 -

one Myrtenol

Ethyl propanoate Terpenes Nerol Ethyl 2 - methylpropanoate α - Phellandrene Ethyl nonanoate α - Pinene ( E ) - 2 - hexenyl acetate β - Pinene

10

Esters Alcohols Aldehydes Ketones Lactones Terpenoids

( E ) - 3 - hexenyl acetate Terpinen - 4 - ol Hexyl acetate α - Terpineol Hexyl butanoate Terpinolene Hexyl propanoate α - Farensene Hexyl - 2 - methyl butanoate Methyl acetate Methyl cinnamate Methyl butanoate Methyl hexanoate Methyl nonanoate Methyl octanoate Methyl - 2 -

methylbutanoate

Methyl - 3 - methylbutanoate

2 - Methylbutyl acetate 3 - Methylbutyl acetate 2 - Methylpropyl acetate ( Z ) - 6 - nonenyl acetate ( Z , Z ) - 3,6 - nonadienyl

acetate

Pentyl acetate Benzyl acetate Propyl acetate Propyl - 2 - methyl

butanoate

11


characterization, volatiles can be classifi ed as esters, alcohols, aldehydes, ketones, lactones, and terpenoids (Table 1.2 ).

Volatile Compounds Formed from FAs FAs are precursors for a large number of volatile compounds. Many of them are important character - impacted aroma compounds that are responsible for fresh fruit fl avors. Those compounds are usually having straight - chain carbons ranged from C 1 to C 20 . Degradation of FAs occurs mainly by the three different oxidative routes: (1) α - and β - oxidation, (2) oxidation by the lipoxygenase pathway, and (3) autoxidation. The formation of fl avors via β - oxidation is exemplifi ed by considering fl avor formation in pears (Jennings 1967 ). The widest variety of fl avor compounds formed from lipids arises via lipoxygenase activity. Many of the aliphatic esters, alcohols, acids, and carbonyls found in fruits are derived from the oxidative degradation of linoleic and linolenic acids (Reineccius 2006 ). In addition, some of the volatile compounds derived from enzyme - catalyzed oxidative breakdown of unsaturated FAs may also be produced by autoxidation (Chan 1987 ). Autoxidation of linoleic acid produces the 9 - and 13 - hydroperoxides, whereas linolenic acid also produces 12 - and 16 - hydroperoxides (Berger 2007 ). Hexanal and 2,4 - decadienal are the primary oxidation products of linoleic acid, while autoxidation of linolenic acid produces 2,4 - heptadienal as the major product. Further autoxidation of these aldehydes leads to the formation of other volatile products (Chan 1987 ). As an alternative to the membrane catabolism, a hypothesis of low rate of de novo FA biosynthesis (free FA hypothesis) was proposed as the limiting factor for the aroma biosynthesis in fruit harvested too early (Song and Bangerth 2003 ). This hypothesis is also supported by the evidence that a close rela-tionship between low aroma volatile production, low free FA, and low ATP content in apple fruit (Song and Bangerth 2003 ; Tan and Bangerth 2001 ). Either oxidative degradation of FAs or newly biosynthesized free FAs are precursors responsible for the formation of straight - chain esters in many fruits, but their role in fl avor forma-tion needs to be clarifi ed.

Volatile Compounds Formed from Amino Acid Metabolism Amino acid metabolism generates aromatic, aliphatic, and branched - chain alcohols, acids, car-bonyls, and esters that are important to fruit fl avor (Reineccius 2006 ). Some volatile compounds can be produced by the action of enzymatic systems on amino acids. The major types of volatile compounds formed from the interaction of amino acids and sugars include aldehydes, alkyl pyrazines, alkyl thiazolines and thiazoles, and other heterocycles from the Strecker degradation (Maarse 1991 ). Amino acids are precursors for some branched aliphatic compounds such as 2 - methyl - 1 - butanol and 3 - methyl - 1 - butanol that are formed during the amino acid catabolism. These com-pounds can be further synthesized to form esters, which are important volatile compounds in many fruits with distinct “ fruity ” odor. As they share the same precur-sor pyruvate, which is generated from glycolysis, the interaction between FAs and branched amino acids is another important factor in the volatile biosynthesis of fruits. As apple fruits ripen, there is a great production of volatile compounds from branched amino acid pathway (Song 1994 ).

Volatile Compounds Formed from Carbohydrate Metabolism A large variety of volatile fl avors can be traced to carbohydrate metabolism (Berger 2007 ). As the


photosynthetic pathways involve turning CO 2 into sugars that are metabolized into other plant needs, for example, lipids and amino acids, nearly all plant fl avors come indirectly from carbohydrate metabolism. However, there are few fl avor constitu-ents that come directly from carbohydrate metabolism (Reineccius 2006 ).

Volatile Compounds Derived from Terpenoid Terpenoids are widely distrib-uted among fruits. There are two main types of terpenoids that may contribute signifi cantly to the fruit fl avor: (1) monoterpenes and sesquiterpenes and (2) irregu-lar terpenes mainly produced by catabolistic pathways and/or autoxidation (Berger 2007 ). The monoterpenes and sesquiterpenes are mainly formed by anabolic pro-cesses and, therefore, are present in intact plant tissue. However, the formation of some irregular terpenes cannot be explained by anabolic pathways in some fruits. These terpenoids are primarily oxidation - degraded products of the carotenoids.

Phenols and Related Compounds A large number of volatile phenols and related compounds occur in fruits, some of which are potent aroma compounds (Berger 2007 ). The majority of volatile phenols and related compounds are formed mainly through the shikimic acid pathway and are present either as free aglycones or bound glycosides that can be liberated by enzymatic hydrolysis. Generally, the volatile phenols and related compounds are benzene - substituted derivatives with methoxy and phenolic groups, often with an allyl, a vinyl, or an aldehyde group. Common fl avor compounds of this group are eugenol, vanillin, myristicin, apiole, elemicin, and benzaldehyde.

VOLATILE COMPOUNDS AND THEIR BIOLOGICAL CHARACTERISTIC OF MAJOR FRUITS

As described above, lipids, carbohydrates, proteins, and amino acids are enzymati-cally converted to volatile compounds. The characterization of fruit volatiles can be very complicated due to various infl uencing factors such as cultivars, fruit maturity, postharvest treatment, fruit sample (either intact fruit, slices, or homogenized samples), and analytic techniques (Berger 2007 ; Br ü ckner 2008 ; Cunningham and Barry 1986 ). Volatiles can be classifi ed as “ primary ” or “ secondary ” compounds, indicating whether they were present in intact fruit tissue or produced as a result of tissue disruption (Drawert et al. 1969 ). It should be pointed out that analysis of volatiles from either intact or disrupted fruit tissues will infl uence the aroma profi les and fi nal aroma interpretation. This following section reviews overall fl avor characterization of volatile compounds reported for some major fruits published in the past few years. The listed volatile compounds are those that are produced by fruit at a full ripe or close to consumption stage and summarized from different methodologies. In the following section, volatile compounds of major fruits are summarized in Table 1.3 .

Apple

More than 300 volatile compounds have been identifi ed in apple fruit (Dirinck et al. 1989 ). Only a few of these volatiles have been identifi ed as important active


TABLE 1.3. Volatile Compounds of Major Fruits

Fruit Name Volatile Compounds Reference

Apple Alcohols, aldehydes, 1 - butyl acetate, butyl 2 - methylbutanoate, β - damascenone, ethyl butanoate, ethyl butanoate, ethyl 2 - methylbutanoate, ethyl 2 - methylbutyl acetate, n - hexanal, 1 - hexanol, hexen - 1 - ol, hexyl acetate, hexyl butanoate, hexyl 2 - methylbutanoate, hexyl propanoate, ketone, 2 - methylbutanoate, methyl 2 - methylbutanoate, propyl 2 - methylbutanoate, pentyl acetate, 1 - propyl propionate, trans - 2 - hexenal, and trans - 2 - hexen - 1 - ol

Dixon and Hewett (2000) , Flath and others (1969) , Cunningham ( 1985 )

Banana Alcohols, amyl butanaote, butyl butanaote, esters, and isoamyl acetate

Jayanty and others (2002) , Maciel and others (1986)

Citrus fruit Acetaldehyde, acetoin, carvone, β - damascenone, ( E , E ) - 2,4 - decadienal, decanal, diacetyl, dodecanal, elinalool, ethanol, ethyl acetate, ethyl butanoate, ethyl propanoate, ethyl - 2 - methyl propanoate, ethyl - 2 - methyl butanoate, ethyl hexanoate, ethyl - 3 - hydroxy hexanoate, ethyl octanoate, ethyl decanoate, geranial citronellal, hexanal, ( E ) - 2 - hexenaln, ( E ) - 2 - hexen - 1 - ol, ( Z ) - 3 - hexenal, ( Z ) - 3 - hexen - 1 - ol, limonene, methyl butanoate, 3 - methyl butanol, neral, nonanal, ( E ) - 2 - nonenal, ( Z ) - 2 - nonenal, 1 - penten - 3 - one, 1 - octanol, octanal, 1 - octen - 3 - one, β - sinensal, α - terpincol, and terpinen - 4 - ol

Berger (2007) , Berry and others (1983)

Strawberry Butyrates, butyl acetate, 2,5 - dimethyl - 4 - hydroxy - 3(2H) - furanone, dimethyl - 4 - methoxy - 3(2H) - furanone, γ - decalactone, γ - dodecalactone, ethyl butanoate, ethyl cinnamates, ethyl hexanoate, ethyl 3 - methylbutanoate, ethyl propanoate, farnesyl acetate, furaneol, furaneol - β - glucoside, geraniol, 2 - heptanone, hexanal, ( E ) - 2 - hexenal, hexyl acetate, linalool, methyl cinnamates, methyl and ethyl acetates, methyl anthranilate, methyl butanoate, methyl 2 - methylbutanoate, methyl hexanoate, mesifurane, methional, propionates, and 1 - octen - 3 - one

Forney and others (2000) , Hakala and others (2002) , Sanz and others (1994) , Whitaker and Evans (1987)


Fruit Name Volatile Compounds Reference

Peach Benzaldehyde, benzyl alcohol, γ - caprolactone, cis - 3 - hexenyl acetate, β - damascenone, γ - decalactone, ( E , E ) - 2,4 - decadienal, δ - decalactone, γ - decalactone, dimethyl disulfi de, γ - dodecalactone, δ - dodecalactone, ethyl acetate, ethyl butanoate, ethyl octanoate, γ - decalactone, hexanal, ( Z ) - 3 - hexen - 1 - yl acetate, ( E ) - 2 - hexen - 1 - ol, ( Z ) - 3 - hexenal, γ - jasmolactone, linalool, methyl octanoate, γ - octalactone, δ - octalactone, 6 - pentyl α - pyrone, and terpinolene

Aubert and Milhet (2007) , Berger (2007) , Horvat and others (1992) , Narain and others (1990) , Visai and Vanoli (1997)

Pear Butyl acetate, butyl butanoate, hexyl acetate, ethyl hexanoate, ethyl octanoate, ethyl ( E ) - 2 - octenoate, ethyl ( E , Z ) - 2,4 - decadienoate, methyl ( E , Z ) - 2,4 - decadienoate, and pentyl acetate

Argenta and others (2003) , Kahle and others (2005) , Rapparini and Predieri (2003) , Rizzolo and others (1991)

Grape Methyl anthranilate Rosilllo and others (1999) , Whitaker and Evans ( 1987 )

Mango Camphene, butan - 1 - ol, car - 3 - ene, β - caryophyllene, p - cymene, cis - hex - 3 - en - 1 - ol, α - copanene, cyclohexane, dimethylcyclohexane, 1,1 - diethoxyethane, ethanol, ethylcyclohexane, ethyl butenoate, ethyl dodecanoate, ehtyl decanoate, ethyl octanoate, α - fenchene, 2 - furfural, hexanal, α - humulene, hydrocarbon, limonene, 1 - methylpropan - 1 - ol, methylcyclohexane, 3 - methylbutan - 1 - ol, myrcene, α - phellandrene, β - phellandrene, α - pinene, β - pinene, sabinene, sabinyl acetate, toluene, γ - terpinene, α - terpinolene, and xylene

MacLeod and Snyder (1985) , Macleod and Troconis (1982) , Malundo and others (2001) , Pino and Mesa (2006)

Papaya Linalool, ethyl acetate, phenylacetonitrile, benzyl isothiocynate, methyl butanoate, ethyl butanoate, 3 - methylbutanol, benzyl alcohol, α - terpineol, and butanol

Almora and others (2004) , Flath and others (1990) , Heidlas and others (1984) , Pino and others (2003)

Pineapple Acetoxyacetone, p - allyl phenol, γ - butyrolactone, β - hydroxyhexanoic acids, 4 - methoxy - 2,5 - dimethyl - 2(H) - furan - 3 - one, methyl esters of β - hydroxybutyric, γ - octalactone, 2 - propenyl hexanoate, sesquiterpene, 1 - ( E , Z ) - 3,5 - undecatriene, and 1 - ( E , Z , Z )3,5,8 - undecatetraene

Berger and others (1983, 1985) , Takeoka and others (1991) , Tokitomo and others (2005)

Plum Benzaldehyde, ( E , E ) - 2,4 - decadienal, δ - decalactone, γ - decalactone, ethyl nonanoate, ( Z ) - 3 - hexenal, linalool, methyl cinnamate, and δ - octalactone

Maarse (1991) , Horvat (1992)

TABLE 1.3. Continued


odor compounds being responsible for the characteristic aroma in most apple cultivars, such as β - damascenone, butyl, isoamyl, and hexyl hexanoate, along with ethyl, propyl, and hexyl butanoates (Cunningham 1985 ). The most abundant volatile components are esters, alcohols, aldehydes, ketones, and ethers, while esters are the principal compounds responsible for fruity odor (Fellman et al. 2000 ; Plotto et al. 2000 ). For example, ethyl 2 - methylbutanoate, 2 - methylbutyl acetate, and hexyl acetate contribute mostly to the characteristic aroma of “ Fuji ” apples, while ethyl butanoate and ethyl 2 - methylbutanoate are the active odor compounds in “ Elstar ” apples, and ethyl butanoate, acetaldehyde, 2 - methyl - 1 - butanol, and ethyl methylpro-panoate in “ Cox Orange ” (Acree et al. 1984 ; Berger 2007 ; Echeverria et al. 2004 ). Ethyl 2 - methylbutanoate also has a direct impact on “ Granny Smith ” apple fl avor (Lavilla et al. 1999 ).

Banana

The major volatile compounds in banana fruit are identifi ed as alcohols and esters, including amyl acetate, isoamyl acetate, butyl butyrate, and amyl butyrate. Esters predominate in the volatile fraction of banana fruit. Based on the combined analytic chemistry with sensory analysis, penten - 2 - one, 3 - methylbutyl, and 2 - methylpropyl esters of acetate and butyrate have been identifi ed as the most important banana fruit aroma (Berger et al. 1986 ). Isopentyl acetate and isobutyl acetate are also known as the most important impact compounds of banana aroma. The concentra-tions of acetates and butanoates increased during ripening of banana fruit (Jayanty et al. 2002 ). In addition, isoamyl alcohol, isoamyl acetate, butyl acetate, and elemi-cine were detected by olfactometric analyses as characteristics of banana odor (Boudhrioua et al. 2003 ).

Citrus

Citrus volatiles have been extensively examined over the past several decades. As the most foods of commercial interest, the volatile components of citrus juice have been known for some time. Table 1.2 lists the volatile compounds present in citrus juice, which were detected by gas chromatography (GC ) – olfactometry. Esters are important as they are responsible for the fl avor characteristic (Berger 2007 ), while the major esters are ethyl esters of C 3 to C 4 organic acids. Linalool is by far the most important alcohol. However, ketones, carvone, diacetyl, and acetoin are off - fl avors. Thus, the key fl avor compounds in fresh citrus fruit still need to be identifi ed.

Strawberry

Over 360 different volatile compounds have been identifi ed in strawberry fruit (Maarse 1991 ). Strawberry aroma is composed predominately of esters with alco-hols, ketones, lactones, and aldehydes being present in smaller quantities (Forney et al. 2000 ). Strawberries contain primarily straight esters, which comprise primarily of methyl, and ethyl acetates, butanoates, and hexanoates. Esters provide an aroma characteristic to the fruit (Gomes da Silav and Chavees das Neves 1999 ). Terpenoids and sulfur compounds may also have a signifi cant impact on the characteristic


aroma of strawberry fruit (Dirinck et al. 1981 ). The most important aroma compounds in strawberry fruit include ethyl cinnamates, methyl cinnamates, 2,5 - dimethyl - 4 - hydroxy - 3(2H) - furanone, furaneol, furaneol - beta - glucoside, dimethyl - 4 - methoxy - 3(2H) - furanone (mesifurane), methyl and ethyl acetates, propionates, and butyrates, which are responsible for fruity fl avor. A number of terpenes also contribute to the fl avor of strawberry fruit.

Peach

Approximately 100 volatile compounds have been identifi ed in peaches, including alcohols, aldehydes, alkanes, esters, ketones, lactones, and terpenes (Aubert et al. 2003 ; Visai and Vanoli 1997 ). The major volatile compounds are identifi ed as ethyl acetate, cis - 3 - hexenyl acetate, methyl octanoate, ethyl octanoate, γ - decalactone, benzyl alcohol, γ - caprolactone, and δ - decalactone. Among them, lactones, particu-larly γ - decalactone and δ - decalactone, have been reported as character - impacted compounds in peaches and are associated with C6 - aldehydes, aliphatic alcohols, and terpenes, which are responsible for fruity characteristics (Derail et al. 1999 ; Engel et al. 1988 ; Horvat et al. 1990 ; Narain et al. 1990 ). Nectarines produce less volatiles in total but more esters, linalol, and terpinolene and have more fruity and fl oral aroma notes than peaches (Visai and Vanoli 1997 ).

Pear

More than 300 volatile compounds have been identifi ed in pear, including aldehydes, alcohols, esters, ketones, and sulfur compounds (Rapparini and Predieri 2003 ). The most important character - impacted compounds of pears are listed in Table 1.3 . Methyl and hexyl esters of decadienoate are the character - impacted compounds of the European pear (Argenta et al. 2003 ; Kahle et al. 2005 ; Rapparini and Predieri 2003 ). Other volatile esters, for example, hexyl acetate, 2 - methylpropyl acetate, butyl acetate, butyl butanoate, pentyl acetate, and ethyl hexanoate possess strong pear - like aroma (Rapparini and Predieri 2003 ). Ethyl octanoate and ethyl ( E ) - 2 - octenoate contribute to sweet or fruity odors in pears, while a high concentration of 2,4 - decadienoates in fruit fl esh is accepted by consumers (Rizzolo et al. 1991 ). In addi-tion, hexanal, 2 - methylpropyl acetate, ethyl acetate, hexyl acetate, 3 - methylbutyl 2 - methylbutanoate, ethyl butanoate, and butanol are identifi ed as impact volatiles in “ Conference ” pears (Rizzolo et al. 2005 ).

Grape

The fl avor of grapes is made up of volatile alcohols, aldehydes, esters, acids, terpe-nols, and carbonyl compounds. Grape may be divided into aromatic and non-aromatic varieties. Free terpenols, for example, linalool and geraniol, have been identifi ed as major aroma compounds in both red and white grapes (Rosilllo et al. 1999 ). Octanoic acid and alcohols, particularly 2 - phenylethanol, are recognized after crushing (Rosilllo et al. 1999 ). In addition, esters and aldehydes were also reported in “ Aleatico ” grapes (Bellincontro et al. 2009 ). Fruit fl avor is highly correlated with consumer likings in table grapes.


Mango

Mango possesses a very attractive fl avor characteristic. About 270 volatile com-pounds from mango fruit were identifi ed. However, application of distillation extraction in combination with active odor value (aroma threshold) technologies exhibits that monoterpenes such as α - pinene, myrecene, α - phelladrene, σ - 3 - carene, p - cymene, limone and terpinolene, esters including ethyl - 2 - methylpropanaote, ethyle - butanoate, as well as ( E , Z ) - 2,6 - nonadienal, ( E ) - 2 - nonenal, methyl benzoate, ( E ) - β - ionone, decanal, and 2,5, - dimethyl - 4 - methoxy - 3(2H) - furanone are the most important compounds contributing to mango fl avor (Pino and Mesa 2006 ). The acids, esters, and lactones found were considered to be produced by the lipid metab-olism in the fl avor development of mango fruit during ripening.

Papaya

Papaya possesses a characteristic aroma, which is due to several volatile components such as alcohols, esters, aldehydes, and sulfur compounds (Chan et al. 1973 ). Fifty - one volatile compounds from intact “ Hawaiian ” papaya at different ripening stages were detected. Linalool, followed by linalool oxide A, linalool oxide B, ethyl acetate, phenylacetonitrile, and benzyl isothiocynate, was the major compound in the fully ripe fruits (Flath et al. 1990 ). Other work indicated the esters as the predominant compounds among the volatiles of papayas from Sri Lanka and Colombia (Heidlas et al. 1984 ; Macleod and Pieris 1984 ). In addition, methyl butanoate, ethyl butanoate, 3 - methylbutanol, benzyl alcohol, α - terpineol, and butanol are found to be important volatiles in papaya fruit (Almora et al. 2004 ; Pino et al. 2003 ).

Pineapple

More than 280 volatile compounds have been found in pineapple fruit (Tokitomo et al. 2005 ). The major volatile compounds are identifi ed as 4 - methoxy - 2,5 - dimethyl - 2(H) - furan - 3 - one, 2 - propenyl hexanoate, sesquiterpene hydrocarbons, 1 - ( E , Z ) - 3,5 - undecatriene, 1 - ( E , Z , Z )3,5,8 - undecatetraene, 2 - propenyl n - hexanoate ethyl, para - allyl phenol, γ - butyrolactone, γ - octalactone, acetoxyacetone, methyl esters of β - hydroxybutyric, and β - hydroxyhexanoic acids. Monoterpene alcohols (linalool, α - terpineol, and terpinen - 4 - ol) and sesquiterpenes were also identifi ed (Berger et al. 1985 ; Flath and Forry 1970 ). In addition, the sulfur compounds such as methyl 3 - (methylthio) - ( E ) - 2 - propenoate, methyl 3 - (methylthio) - ( Z ) - 2 - propenoate, ethyl 3 - (methylthio) - ( Z ) - 2 - propenoate, ethyl 3 - (methylthio) - ( E ) - 2 - propenoate, methyl 5 - hexenoate, methyl ( E ) - 4 - hexenaote, methyl 4 - (methylthio) - butanoate, nonanol, and ethyl 4 - methylthiobutanoate, were reported as impact - fl avor compounds in fresh “ Hawaiian ” pineapple (Takeoka et al. 1991 ).

Plum

Approximately 75 volatile compounds have been identifi ed in plum juices (Maarse 1991 ). Lactones from C 6 to C 12 are major classes of volatile compounds in plums (Horvat 1992 ), but the key fl avor compounds in fresh plum fruit are not yet identifi ed.


CONCLUDING REMARKS

The diversity of varieties of fruit for today ’ s human consumption has resulted from a long history of natural development, selection, and scientifi c breeding. Fruits play important roles in human nutrition and diet. However, they are perishable due to natural ripening, senescence, and pathological decay. Fruit quality attributes, such as texture, appearance, fl avor, and nutrition, signifi cantly change during ripening, but they have not been understood fully. Thus, the further development of modern technologies of breeding, production, and postharvest handling will enable con-sumers to enjoy fruits and their products without limitations of seasons and geographic locations.

Fruit fl avor is an important aspect of quality. Many compounds are responsible for the fruit aromas that have strong penetration odors with low threshold values. Advances in identifying and quantifying volatile compounds by improved analysis techniques in various fruits have greatly increased our knowledge about fruit fl avor (Br ü ckner 2008 ; Song 2007 ; Song and Forney 2008 ; Tholl et al. 2006 ; Tzortzakis 2007 ). Advances in the biogenesis of volatile compounds in fresh fruits have also improved our current understanding; however, knowledge of the biochemical pathways and key regulating steps of the synthesis of these volatile compounds is still incomplete. A fuller understanding of the fl avor chemistry and biology of volatile compounds of fruits is important to improve the fl avor quality of fresh fruit that complies with the consumer needs for better quality. Furthermore, employing state - of - the - art genomic, proteomic, and microscopy tools to study fundamental metabolism, and combining these results with direct measurement of chemical and sensory properties (Baldwin 2002a ; Bood and Zabetakis 2002 ; Raab et al. 2006 ; Song and Forney 2008 ) will lead to a better understanding of how to optimize and retain fruit fl avor quality in the market places for the benefi t of both consumers and fruit industry.

ACKNOWLEDGMENTS

This work was supported by the National Natural Science Foundation of China (Grant Nos. 30425040 and U0631004) and Guangdong Provincial Natural Science Foundation (No. 06200670).

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