Table Of ContentTHE SCIENCE OF GRAPEVINES
This page intentionally left blank
THE SCIENCE
OF
GRAPEVINES:
ANATOMY AND
PHYSIOLOGY
M K
ARKUS ELLER
IrrigatedAgricultureResearch andExtensionCenter,
WashingtonStateUniversity,Prosser,WA
AMSTERDAM (cid:129) BOSTON (cid:129) HEIDELBERG (cid:129) LONDON
NEW YORK (cid:129) OXFORD (cid:129) PARIS (cid:129) SAN DIEGO
SAN FRANCISCO (cid:129) SINGAPORE (cid:129) SYDNEY (cid:129) TOKYO
Academic Press is an imprint of Elsevier
Academic PressisanimprintofElsevier
30CorporateDrive,Suite400,Burlington,MA01803,USA
525BStreet,Suite1900,SanDiego, California92101-4495,USA
84Theobald’sRoad,London WC1X8RR,UK
Copyright#2010MarkusKeller, PublishedbyElsevierInc.Allrightsreserved.
Nopartofthispublication maybe reproducedor transmitted inanyformor byanymeans,electronicor
mechanical,includingphotocopy,recording,or anyinformationstorageandretrievalsystem,without
permission inwriting fromthepublisher.
Permissionsmaybesoughtdirectly fromElsevier’sScience&TechnologyRightsDepartmentinOxford,
UK:phone:(þ44)1865843830,fax:(þ44)1865853333,E-mail:[email protected]
completeyour requestonlinevia theElsevierhomepage(http://elsevier.com),byselecting“Support&
Contact”then“CopyrightandPermission”andthen“ObtainingPermissions.”
Library ofCongress Cataloging-in-Publication Data
Keller, Markus.
The scienceofgrapevines:anatomyandphysiology/Markus Keller.
p.cm.
Includesbibliographical referencesandindex.
ISBN978-0-12-374881-2(hardcover:alk.paper) 1.Grapes–Anatomy.
2.Grapes–Physiology.I.Title.
QK495.V55K442010
634.8–dc22 2009033159
British Library Cataloguing-in-PublicationData
Acatalogue recordfor thisbookisavailablefrom theBritishLibrary.
ISBN:978-0-12-374881-2
Forinformation onallAcademic Presspublications
visitourWebsiteat www.elsevierdirect.com
Printed inChina
09 10 11 9 8 7 6 5 4 3 2 1
Contents
4.4 Respiration 118
About the Author vii
4.5 FromCellstoPlants 121
Preface ix
Acknowledgments xi 5. Partitioning of Assimilates 125
1. Botany and Anatomy 1 5.1 PhotosynthateTranslocationandDistribution 125
5.2 Canopy–Environment Interactions 140
1.1 BotanicalClassification andGeographical 5.3 Nitrogen Assimilationand InteractionwithCarbon
Distribution 1 Metabolism 158
1.2 Cultivars,Clones,andRootstocks 9
1.3 MorphologyandAnatomy 20 6. Developmental Physiology 169
2. Phenology and Growth Cycle 49 6.1 YieldFormation 169
6.2 GrapeCompositionand FruitQuality 178
2.1 Seasonsand DayLength 49 6.3 SourcesofVariation inFruitComposition 209
2.2 VegetativeCycle 53
2.3 ReproductiveCycle 68 7. Environmental Constraints and Stress
Physiology 227
3. Water Relations and Nutrient Uptake 85
7.1 ResponsestoStress 227
3.1 Osmosis,WaterPotential, andCellExpansion 85 7.2 Water:TooMuchorTooLittle 231
3.2 Transpirationand StomatalAction 89 7.3 Nutrients:Deficiencyand Excess 243
3.3 WaterandNutrient UptakeandTransport 92 7.4 Temperature:TooColdorTooWarm 274
7.5 LivingwithOtherOrganisms:Defense and
4. Photosynthesis and Respiration 107
Damage 289
4.1 LightAbsorptionand EnergyCapture 107
References 311
4.2 CarbonUptake andAssimilation 111
4.3 Photorespiration 117 Index 369
v
This page intentionally left blank
About the Author
Markus Keller is the Chateau Ste. Michelle University in Wagga Wagga, Australia, before
Distinguished Professor of Viticulture at coming to eastern Washington. Dr. Keller is
Washington State University’s Irrigated Agri- the author of numerous scientific and technical
culture Research and Extension Center in papers and industry articles in addition to
Prosser, Washington. He received his master’s being a frequent speaker at scientific confer-
degree in agronomy (plant science) in 1989 ences and industry meetings and workshops.
andadoctorateinnaturalsciencesin1995from He also has extensive practical experience in
the Swiss Federal Institute of Technology in both the vineyard and the winery as a result
Zu¨rich. He has taught and conducted research of work in the family enterprise, and he was
in viticulture and grapevine physiology on awarded the Swiss AgroPrize for innovative
threecontinents,beginningattheSwissFederal contributions to Switzerland’s agriculture
Research Station for Fruit-Growing, Viticulture industry. His current research focuses on
and Horticulture in Wa¨denswil (now Agro- environmental factors and management prac-
scope Changins-Wa¨denswil), Switzerland, and tices as they influence crop physiology and
then moving to Cornell University in Geneva, production of wine and juice grapes.
New York, and from there to Charles Sturt
vii
This page intentionally left blank
Preface
Grapes were among the first fruit species to stomatalporesmaybeanexception)arerooted
be domesticated and today are the world’s in biochemistry. They are driven or at least
most economically important fruit crop. facilitated by enzymes, which in turn are built
According to 2009 statistics by the Food and based on blueprints provided by genes. I have
Agriculture Organization of the United therefore taken it for granted that it is
Nations, grapevines were planted on almost understood that a developmental process
7.3 million hectares producing more than 67 or change in chemical composition implies a
million metric tons of fruit in 2007. This makes change in enzyme activity, which implies
grapes the number 25 food crop in terms of a change in the activity of one or more genes.
planted area and number 16 in terms of ton- Thisdoesnotimply,asusedtobethought,that
nage. More than 70% of this crop was used to “one gene makes one enzyme,” nor that “one
make wine, 27% consumed as fresh fruit (table enzyme makes one chemical,” but it merely
grapes), 2% as dried fruit (raisins), and less means that all enzymatic processes have a
than 1% was processed to grape juice or dis- genetic basis.
tilled to brandy. Manybiochemicalandbiophysicalprocesses
This book is an introduction to the physical apply to many or even all plants. Perhaps no
structure of the grapevine, its various organs process is truly unique to grapevines. Chances
and tissues, their functions, their interactions are that if grapes do it, some or many other
with one another, and their responses to the species employ the same solution to a survival
environment.Itfocusesessentiallyonthephys- issue because they share a common ancestor
ical and biological functions of whole plants that invented the trick a long time ago. For
rather than the metabolism and molecular example, microbes hit upon photosynthesis
biology of individual cells. It is nonetheless and respiration long before these discoveries
necessary to review some fundamental enabled some of them to combine forces and
processes at the cell, tissue, and organ levels evolve into plants. Consequently, although this
in order to build up an appreciation of whole- book is about grapevines, and primarily about
plant function. The book covers those elements thewinegrapespeciesVitisvinifera,Ihavebor-
of physiology that will enhance our under- rowed heavily from research done with other
standing of grapevine function and how they plant species, both wild and cultivated, peren-
relate to grape production. Although of neces- nial and annual, woody and herbaceous,
sity the text contains a plethora of technical including the “queen of weeds”—at least in
terms and details, I have tried to resist the thefast-pacedworldofmodernmolecularbiol-
temptation to dwell in biochemical and molec- ogy—Arabidopsis thaliana, the otherwise incon-
ular biological jargon. Most physiological spicuous thale cress. I have even taken the
processes (water movement through the vine’s liberty to borrow insights gained using micro-
hydraulic system and evaporation from the organisms, such as the yeast Saccharomyces
ix
