Color profile: Disabled Composite 150 lpi at 45 degrees Postharvest Physiology and Hypobaric Storage of Fresh Produce A4729 - Burg - Prelims.prn 656 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees A4729 - Burg - Prelims.prn 657 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees Postharvest Physiology and Hypobaric Storage of Fresh Produce Stanley P. Burg Consultant Miami, Florida USA CABI Publishing A4729 - Burg - Prelims.prn 658 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees CABIPublishing is a division of CAB International CABIPublishing CABIPublishing CAB International 875 Massachusetts Avenue Wallingford 7th Floor Oxfordshire OX10 8DE Cambridge, MA 02139 UK USA Tel: +44 (0)1491 832111 Tel: +1 617 395 4056 Fax: +44 (0)1491 833508 Fax: +1 617 354 6875 E-mail:[email protected] E-mail:[email protected] Web site:www.cabi-publishing.org ©S.P. Burg 2004. All rights reserved. No part of this publication may be reproduced in any form or by any means, electronically, mechanically, by photocopying, recording or otherwise, without the prior permission of the copyright owners. A catalogue record for this book is available from the British Library, London, UK. Library of Congress Cataloging-in-Publication Data Postharvest physiology and hypobaric storage of fresh produce / edited by Stanley P. Burg. p. cm. Includes bibliographical references. ISBN 0-85199-801-1 (alk. paper) 1. Fruit--Postharvest physiology. 2. Vegetables--Postharvest physiology. 3. Fruit--Storage. 4. Vegetables--Storage. I. Burg, Stanley P., 1933- II. Title. SB360.P668 2004 ′ 631.568--dc22 2003018164 ISBN 0 85199 801 1 Typeset by AMADataSetLtd, UK. Printed and bound in the UK by Biddles Ltd, King’s Lynn. A4729 - Burg - Prelims.prn 659 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees Contents Preface xiv Acknowledgements xvi 1 Introduction 1 1.1 Causes of Postharvest Losses 1 1.2 Current Status of LP 1 1.3 Comparison of LP and CA 3 1.4 History of CA and LP Storage 3 1.5 LP Research and Development 4 1.6 Is LP More Complicated than CA? 4 1.7 LP Patent Protection and Trade Secrets 6 2 Origins of the LP Concept 9 2.1 The LP Apparatus 9 2.2 Evidence that Gas Exchange in Fruits is Limited by an Air Phase 10 2.3 Testing for Ethylene-controlled Developmental Processes 13 2.4 Involvement of Other Gases and Volatile Substances 15 2.5 Converting the LP Test Apparatus into a Storage System 15 2.6 Who Invented Hypobaric Storage? 15 3 GasandVapourMassTransport 18 3.1 Resistance Network 18 3.2 Effect of pH and Temperature on Gas Exchange through Bio-membranes 20 3.3 Cytosol Resistance 23 3.4 Permeability of the Cell Wall 23 3.5 Resistance of the Mitochondria 23 3.6 Total Gas Transport Resistance 24 3.7 Kinetics of Gas Equilibration between Cells, the Intercellular System and Atmosphere 24 3.8 Measuring the Air Phase Resistance Coefficient 27 3.9 O , CO and Ethylene Exchange between Cell Fluid and Intercellular Air 28 2 2 3.10 Dependence of Ethylene Action on the IEC 29 3.11 NH Exchange between the Apoplast and Intercellular Air 35 3 3.12 Jamin’s Chains: the Effect of Liquid Water on Intercellular Gas Transport 37 v A4729 - Burg - Prelims.prn 660 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees vi Contents 3.13 Cell Leakage during Senescence and Ripening 39 3.14 Measuring the Intercellular Gas Concentrations 44 3.15 Vacuum-extraction Errors 46 3.16 Intercellular Gas Transport 48 3.17 Axial Diffusion of Intercellular Gases 51 3.18 Stomates and Lenticles 52 3.19 Pedicel-end Stem Scar 56 3.20 Cuticular Gas Exchange 60 3.21 Effect of LP on NH Efflux 62 3 3.22 Changes in Gas Exchange during Storage 63 3.23 Resistance of Boxes and Wraps 63 3.24 Effect of LP on Mass Transport through Air Phases 64 3.25 Effect of LP on Vapour Mass Transport 65 3.26 LP Does Not Remove Flavour and Aroma Volatiles from Fruits 66 3.27 Examples 67 4 Oxygen, Carbon Dioxide, Ammonia and Cyanide 81 4.1 Measuring Respiration and Ethylene Production under Hypobaric Conditions 81 4.2 Effect of O on Respiration 83 2 4.3 Low [O ] and High [CO ] Injury 91 2 2 4.4 Effect of O on Ethylene Production 93 2 4.5 Effect of O on Ethylene Action 94 2 4.6 Effect of O on Ascorbic Acid Oxidation 95 2 4.7 Effect of Hypoxia on Ethylene Production, Fruit Softening, ACC Accumulation and Ethylene-induced Enzyme Activity 97 4.8 Advantages of a CO -free Atmosphere 97 2 4.9 Effect of CO on Respiration 98 2 4.10 Effect of CO on Ethylene Production 98 2 4.11 CO as an Activator, Regulator and Inhibitor of Enzymatic Reactions 100 2 4.12 Effect of CO on Ethylene Action and Metabolism 101 2 4.13 Effect of CO on Succinic Acid Accumulation 106 2 4.14 Effect of CO on Ascorbic Acid Content 108 2 4.15 Effect of CO on Stomatal Action 109 2 4.16 Effect of CO on Cellular pH 113 2 4.17 Ammonia Toxicity 120 4.18 Ammonia Incorporation into Organic Compounds 121 4.19 Ammonia Formation 122 4.20 Ammonia Production during Flower Fading and Leaf Senescence 123 4.21 Cyanide 125 4.22 Examples 127 5 Ethylene 132 5.1 Biosynthetic Pathway 132 5.2 Effect of Temperature on Ethylene Production and Perception 135 5.3 Cellular Localization of EFE 137 5.4 Effect of Ethylene on Respiration 140 5.5 Climacteric and Non-climacteric Fruits 141 5.6 Systems 1 and 2 Ethylene Production 145 5.7 Auto-inhibition of System 1 Ethylene Production 145 5.8 Auto-stimulation of System 2 Ethylene Production 147 5.9 Significance of the Pre-climacteric IEC 148 A4729 - Burg - Prelims.prn 661 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:00 PM Color profile: Disabled Composite 150 lpi at 45 degrees Contents vii 5.10 Effect of Harvest 154 5.11 Ethylene Receptor 164 5.12 Ethylene Transduction Pathway 172 5.13 Regulation of Ethylene Sensitivity 173 5.14 Initiation of Ripening in Climacteric Fruits 177 5.15 Ethylene-independent Developmental Regulation of Ripening in Climacteric Fruits 181 5.16 Ripening in Non-climacteric Fruits 186 5.17 Circadian Rhythms of Ethylene Production 188 5.18 Stress-induced Ethylene Production 193 5.19 Auxin-induced Ethylene Production 194 5.20 Effect of Ethylene on Polar Auxin Transport 199 5.21 Effect of Ethylene on Auxin Synthesis and Metabolism 199 5.22 Gravity-dependent Ethylene Action 200 5.23 Effect of Ethylene on Auxin Gravitransport 205 5.24 The ‘Triple Response’ of Seedlings 210 5.25 Ethylene-induced Epinasty 213 5.26 Effect of Ethylene on Correlative Inhibition 215 5.27 Effect of Ethylene on Microfibrillar and Microtubular Orientation 219 5.28 Ethylene-induced Leaf Senescence 221 5.29 Role of Ethylene in Abscission 224 5.30 Role of Ethylene in Flower Fading 225 5.31 Induction of Physiological Disorders by Ethylene 231 5.32 Effect of Ethylene on Stomatal Opening 232 5.33 Effect of Ethylene on Ascorbic Acid Content 232 5.34 Examples 232 6 Heat Transfer and Water Loss 240 6.1 Dependence of Water Loss on Respiratory Heat 241 6.2 The Cuticle 248 6.3 Dependence of Transpirational Resistance on Relative Humidity 250 6.4 Water-retentive Packaging 252 6.5 Dependence of Transpirational Resistance on Pressure 253 6.6 Radial Transport of Respiratory Heat 256 6.7 Cellular Water Conductance 258 6.8 Apoplastic Water Transport 259 6.9 Symplastic Cell-to-cell Water Transport 259 6.10 Water Vapour Transport through the Intercellular System 260 6.11 Skin Shrivel and ‘Peel Puffing’ 263 6.12 Dimensionless Groups used for Convective Heat Transfer Calculations 265 6.13 Heat Transfer Coefficient for Free Convection at Atmospheric Pressure 267 6.14 Radiative Couplings 268 6.15 Evaporative Couplings 270 6.16 Heat Transfer in CA Storage at Atmospheric Pressure 270 6.17 Controlling Water Loss at Atmospheric Pressure 272 6.18 LP Heat Sources 273 6.19 Humidity Control in LP 273 6.20 LP Stacking Arrangements 275 6.21 Forced Convection in LP 279 6.22 Effect of LP on Free Convective Heat Transfer 280 A4729 - Burg - Prelims.prn 662 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:01 PM Color profile: Disabled Composite 150 lpi at 45 degrees viii Contents 6.23 Thermal Balance under Hypobaric Conditions 282 6.24 Cool down under Hypobaric Conditions 286 6.25 Examples 289 7 Postharvest Diseases and Physiological Disorders 302 7.1 Direct Effects of O on Microbial Growth 302 2 7.2 Direct Effects of CO on Microbial Growth 305 2 7.3 Combined Direct Effects of CO and O on Microbial Growth 307 2 2 7.4 Microbial Development in LP 308 7.5 Ripening, Ageing and Senescence Influence the Host’s Susceptibility to Disease 313 7.6 Ethylene 313 7.7 Phytoalexins and Anti-fungal Compounds 314 7.8 Water Condensation 317 7.9 Hypochlorous Acid Vapour 318 7.10 Chilling Injury 321 7.11 Physiological Disorders other than Chilling Injury 327 7.12 Examples 329 8 Insect Quarantine 333 8.1 Effect of High Temperature 333 8.2 Effect of Low Temperature 336 8.3 Dependence of Insect O Consumption on Temperature, [O ] and [CO ] 336 2 2 2 8.4 Lethal Effects of High [CO ], Low [O ] and Desiccation 337 2 2 8.5 The Gas Exchange System of Insects 339 8.6 Comparison of the Gas Exchange Systems of Insects and Plants 344 8.7 Effect of a Low Pressure on Insect Mortality 345 9 Technical Difficulties Associated with Laboratory Hypobaric Research 350 9.1 Does the Inability to Provide CO Detract from LP Storage? 351 2 9.2 Factors Influencing Weight Loss in a Laboratory Hypobaric Experiment 351 9.3 Factors Determining the Steady-state Relative Humidity 353 9.4 Measuring the Relative Humidity 357 9.5 Other Sources of Moisture 357 9.6 Chamber Leak Rate 358 9.7 Chamber Temperature Uniformity 359 9.8 Elevating the Humidity 362 9.9 Mylar Radiation Shielding 363 9.10 Air Change Rate 365 9.11 Design of a Laboratory Apparatus 365 9.12 Measuring the Pressure 365 9.13 Examples 366 10 Horticultural Commodity Requirements 371 FRUITS 372 × 10.1 Apple (Malus domesticaBorkh.), American Summer Pearmain, Jonathan, Cox Orange, Red King, Golden Delicious, Boskoop, Schweizer Glockenapfel, McIntosh, Red Delicious, Ida Red, Spartan, Staymen, Northern Spy, Red Rome, Lodi, Empire, Ontario 372 10.2 Apricot (Prunus armeniacaL.), var. Large Early Montgament 375 10.3 Avocado (Persea americanaMill.), Choquette, Waldin, Booth 8, Lula, Hass 376 A4729 - Burg - Prelims.prn 663 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:01 PM Color profile: Disabled Composite 150 lpi at 45 degrees Contents ix 10.4 Banana (Musa paradisicavar. Sapientum), Lacatan, Valery, Gros Michel and Dwarf Cavendish 378 10.5 Blueberry (Vaccinium corymbosumL.), var. Jersey 380 10.6 Cantaloupe (Cucumis meloL.), cv. Earl’s Favourite 381 10.7 Carambola (Averrhoa carambolaLinn.) 381 10.8 Cherimoya (Annona cherimolaMiller) 381 10.9 Cherry, Sweet (Prunus aviumL.) cvs Bing, Lambert, Grosse Schwarze Knorpelkirsche, Emperor Francis, Picota 382 10.10 Cherry, Barbados (Malpighia glabraL.) and Surinam (Eugenia unifloraL.) 383 10.11 Cranberry (Vaccinium macrocarponAit.) 383 10.12 Currant (Ribes vulgareLam.), Red, White and Black 383 10.13 Grape (Vitis viniferaL.), American, European and var. Red Emperor 384 × 10.14 Grapefruit (Citrus paradisiMacfad.), Florida Ruby Red, Florida Marsh White, Texas Ruby Red 384 10.15 Guava (Psidium guajavaL.) 385 10.16 Honeydew Melon (Cucumis meloL.) 386 10.17 Lime, Persian [Citrus aurantifolia(Christm.) Swingle], var. Tahiti 386 10.18 Mango (Mangifera indicaL.), cvs Haden, Irwin, Tommy Atkins, Kent, Keitt, Maya, Okrang 387 10.19 Nectarine (Prunus persicaL.), var. Nectarina 389 10.20 Orange (Citrus sinensisOsbeck), Valencia 389 10.21 Papaya (Carica papayaL.), var. Solo 390 10.22 Peach (Prunus persicaL. Batsch.), cvs White Okubo, Cardinal, Red Haven, Gleason Early Elberta 392 10.23 Pear (Pyrus communisL.), cvs Bartlett, Clapp, Commice 393 10.24 Pineapple (Ananas comosusL. Merr.), var. Smooth Cayenne 393 10.25 Prune (Prunus domestica), cv. Richards Early Italian 395 × 10.26 Strawberry (Fragaria ananassaDuch.), cvs Tioga, Shasta, Florida 90, Donna, Z-5A, Long Island Jerseybelle, Cambridge Favourite 395 10.27 Tomato (Lycopersicon esculentumMill.), vars Homestead 2, Jupiter, Michigan-Ohio Hybrid, Heines Hochzucht, NM Super II, NM Milo I, Waldin, Ace 397 VEGETABLES 399 10.28 Asparagus (Asparagus officinalisL.) 399 10.29 Beans (Phaseolus vulgarisL.), Pole, Snap and Bush; cvs Improved Tendergreen, ‘McCaslan 42’, Sprite 400 10.30 Beet Leaf (Beta vulgarisL.), var. Conditiva Alef 402 10.31 Broccoli (Brassica oleraceaL. – Italica Group) 402 10.32 Brussels Sprouts (Brassica oleraceaL. – Gemmifera Group) 403 10.33 Cabbage (Brassica oleraceaL. – Capitata Group), cv. Quick Green Storage 403 10.34 Carrot (Daucas carotaL.) 404 10.35 Cauliflower (Brassica oleraceaeL. – Botrytis Group) 404 10.36 Celery, Transplants (Apium graveolensL.), cv. Utah 52–70 405 10.37 Corn, Sweet (Zea maysL.), cvs Iobelle, Florida Sweet, Wintergreen 405 10.38 Cress (Lepidium sativumL.) 406 10.39 Cucumber (Cucumis sativusL.), var. Poinsett 407 10.40 Kohlrabi (Brassica caulorapa) 408 10.41 Leek (Allium ampeloprasumL.) 408 10.42 Lettuce (Latuca sativaL.), var. Iceberg 408 A4729 - Burg - Prelims.prn 664 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:01 PM Color profile: Disabled Composite 150 lpi at 45 degrees x Contents 10.43 Mushroom [Agaricus bisporrus(Lge.)], cvs Golden Light, White Button, Shitake, Enokitake, Matsutake 410 10.44 Onion, Green (Allium cepaL.) 412 10.45 Onion, Dry (Allium cepaL.), cvs Autumn Spice, Rocket, Trapps #6 412 10.46 Parsley [Petroselinum sativum(Mill.) Nym.] 413 10.47 Pepper, Sweet (Capsicum annumL.), Neusiedler Ideal, Bellboy 414 10.48 Potato (Solanum tuberosumL.) 415 10.49 Radish (Raphanus sativaL.), var. Sativus 415 10.50 Spinach (Spinacia oleraceaL.) 415 10.51 Squash (Curcurbita maximaDuch.), var. Acorn; (Curcurbita pepo L. var. Melopepo Alef.); and Yellow Crookneck ‘Summer’ Squash 416 10.52 Turnip (Brassica campestrisL. – Rapifera Group) 416 CUT FLOWERS, CUTTINGS AND POTTED PLANTS 417 10.53 Diseases of Cut Flowers 417 10.54Alstroemeria, Cut Flower (Alstroemeriasp.) 417 10.55Anthurium, Cut Flower (Anthuriumsp.), var. Ozaki 417 10.56 Aster, China, Cut Flower (Callistephus chinensis) 418 10.57 Azalea, Potted Plant (Rhododendronsp.) 418 10.58 Carnation, Cut Flower (Dianthus caryophyllusL.), cvs White Sims (White), Linda (Pink), Scania (Red), Laddie and Various Other Colours 418 10.59 Chrysanthemum, Rooted and Non-rooted Cuttings (Chrysanthemum morifolium=Dendranthema morifolium), cvs Delaware, Neptune, Yellow Delaware, Blue Marble, Golden Anne, Bright Golden Anne, Regal Anne, Yellow Snowdown, Hurricane, Yellow Mandalay, Polaris, Fandango, Super White, Heyday, Rose Chip 422 10.60 Chrysanthemum, Cut Flower (Chrysanthemum morifolium= Dendranthema morifolium) 423 10.61 Chrysanthemum, Potted Plant (Chrysanthemum morifolium= Dendranthema morifolium) 424 10.62 Cuttings (Foliage and Woody) 425 10.63 Foliage Plants, Potted 426 10.64Freesia, Cut Flower (Freesiasp.) 427 10.65 Fynbos, Cut Flower (Erica sparsa,Stoebe plumosa,Phylica imberbus) 427 × 10.66 Geranium, Cutting (Pelargonium hortorumBailey) 427 × 10.67 Geranium, Potted Plant (Pelargonium hortorumBailey), Red 427 10.68 Ginger, Red, Cut Flower (Alpinia purpuata) 428 10.69 Ginger, Wax, Cut Flower (Alpinia speciosa) 429 10.70Gladiolus, Cut Flower (Gladiolussp.) 429 10.71Gypsophila, Cut Flower (Gypsophilasp.) 429 10.72Heliconia, Cut Flower (Heliconia humilis,Heliconia latispatha) 430 10.73Leucadendron, Cut Flower (Leucadendron argenteum), vars Pisa, Jubilee Crown 430 10.74Liatris, Cut Flower (Liatrissp.) 430 10.75 Lily, Easter, Cut Flower (Lilium longiflorum speciosum rubrum) 430 10.76 Lily, Easter, Potted Plant (Lilium longiflorum speciosum rubrum) 430 10.77Narcissus, Cut Flower (Narcissussp.) 431 10.78 Orchid, Cut Flower (Vanda Agnes Joaquim) 431 10.79 Poppy, Cut Flower (Papaversp.) 431 10.80Protea, Cut Flower (Proteasp.), vars Eximia, Cardinal, Sylvia, Pink Ice, Renata, Repens 432 10.81 Rapeseed, Cut Flower (Brassica napus) 433 A4729 - Burg - Prelims.prn 665 Z:\Customer\CABI\A4729 - Burg\A4729 - Burg - First Revise - VP10 #L.vp Thursday, May 20, 2004 12:29:01 PM