Anatomy and Physiology
E-Book, Englisch, 522 Seiten
ISBN: 978-0-12-420008-1
Verlag: Elsevier Reference Monographs
Format: PDF
Kopierschutz: Adobe DRM (»Systemvoraussetzungen)
Dr. Keller received his master's degree in agronomy (plant science) and doctorate in natural sciences from the Swiss Federal Institute of Technology in Zurich. He has taught and conducted research in viticulture and grapevine physiology in three continents and is the author of numerous scientific and technical papers and industry articles in addition to being a frequent speaker at scientific conferences and industry meetings and workshops. He also has extensive practical experience in both the vineyard and winery as a result of work in the family enterprise and was awarded the Swiss AgroPrize for innovative contributions to Switzerland's agricultural industry.
Autoren/Hrsg.
Weitere Infos & Material
1;Front Cover;1
2;The Science of Grapevines;4
3;Copyright Page;5
4;Contents;6
5;About the Author;10
6;Preface to the Second Edition;12
7;1 Botany and Anatomy;14
7.1;1.1 Botanical Classification and Geographical Distribution;15
7.1.1;Domain Eukaryota;16
7.1.2;Kingdom Plantae;16
7.1.3;Division (phylum) Angiospermae (synonym Magnoliophyta);17
7.1.4;Class Dicotyledoneae (synonym Magnoliopsida);17
7.1.5;Order Rhamnales (Vitales according to the Angiosperm Phylogeny web);17
7.1.6;Family Vitaceae;17
7.1.7;Genus Muscadinia;18
7.1.8;Genus Vitis;18
7.1.9;American group;20
7.1.10;Eurasian group;22
7.2;1.2 Cultivars, Clones, and Rootstocks;23
7.2.1;1.2.1 Variety Versus Cultivar;23
7.2.2;1.2.2 Cultivar Classification;30
7.2.3;1.2.3 Clones;32
7.2.4;1.2.4 Rootstocks;34
7.3;1.3 Morphology and Anatomy;37
7.3.1;1.3.1 Root;39
7.3.2;1.3.2 Trunk and Shoots;46
7.3.3;1.3.3 Nodes and Buds;54
7.3.4;1.3.4 Leaves;57
7.3.5;1.3.5 Tendrils and Clusters;63
7.3.6;1.3.6 Flowers and Grape Berries;65
8;2 Phenology and Growth Cycle;72
8.1;2.1 Seasons and Day Length;72
8.2;2.2 Vegetative Cycle;74
8.3;2.3 Reproductive Cycle;96
9;3 Water Relations and Nutrient Uptake;114
9.1;3.1 Osmosis, Water Potential, and Cell Expansion;114
9.2;3.2 Transpiration and Stomatal Action;119
9.3;3.3 Water and Nutrient Uptake and Transport;123
10;4 Photosynthesis and Respiration;138
10.1;4.1 Light Absorption and Energy Capture;138
10.2;4.2 Carbon Uptake and Assimilation;143
10.3;4.3 Photorespiration;149
10.4;4.4 Respiration;151
10.5;4.5 From Cells to Plants;154
11;5 Partitioning of Assimilates;158
11.1;5.1 Photosynthate Translocation and Distribution;159
11.1.1;5.1.1 Allocation and Partitioning;169
11.2;5.2 Canopy–Environment Interactions;176
11.2.1;5.2.1 Light;178
11.2.2;5.2.2 Temperature;187
11.2.3;5.2.3 Wind;190
11.2.4;5.2.4 Humidity;192
11.2.5;5.2.5 The “Ideal” Canopy;193
11.3;5.3 Nitrogen Assimilation and Interaction with Carbon Metabolism;195
11.3.1;5.3.1 Nitrate Uptake and Reduction;196
11.3.2;5.3.2 Ammonium Assimilation;198
11.3.3;5.3.3 From Cells to Plants;200
12;6 Developmental Physiology;206
12.1;6.1 Yield Formation;207
12.1.1;6.1.1 Yield Potential and Its Realization;210
12.2;6.2 Grape Composition and Fruit Quality;218
12.2.1;6.2.1 Water;222
12.2.2;6.2.2 Sugars;225
12.2.3;6.2.3 Acids;229
12.2.4;6.2.4 Nitrogenous Compounds and Mineral Nutrients;232
12.2.5;6.2.5 Phenolics;236
12.2.6;6.2.6 Lipids and Volatiles;249
12.3;6.3 Sources of Variation in Fruit Composition;256
12.3.1;6.3.1 Fruit Maturity;257
12.3.2;6.3.2 Light;258
12.3.3;6.3.3 Temperature;262
12.3.4;6.3.4 Water Status;267
12.3.5;6.3.5 Nutrient Status;270
12.3.6;6.3.6 Crop Load;274
12.3.7;6.3.7 Rootstock;277
13;7 Environmental Constraints and Stress Physiology;280
13.1;7.1 Responses to Abiotic Stress;281
13.2;7.2 Water: Too Much or Too Little;285
13.3;7.3 Nutrients: Deficiency and Excess;300
13.3.1;7.3.1 Macronutrients;306
13.3.1.1;Nitrogen;306
13.3.1.2;Phosphorus;311
13.3.1.3;Potassium;313
13.3.1.4;Sulfur;316
13.3.1.5;Calcium;317
13.3.1.6;Magnesium;319
13.3.2;7.3.2 Transition Metals and Micronutrients;321
13.3.2.1;Iron;321
13.3.2.2;Zinc;324
13.3.2.3;Copper;325
13.3.2.4;Manganese;326
13.3.2.5;Molybdenum;327
13.3.2.6;Boron;327
13.3.2.7;Nickel;329
13.3.2.8;Silicon;329
13.3.3;7.3.3 Salinity;330
13.4;7.4 Temperature: Too Cold or Too Warm;334
13.4.1;7.4.1 Heat Acclimation and Damage;335
13.4.2;7.4.2 Chilling Stress;339
13.4.3;7.4.3 Cold Acclimation and Freeze Damage;342
14;8 Living with Other Organisms;356
14.1;8.1 Biotic Stress and Evolutionary Arms Races;356
14.2;8.2 Pathogens: Defense and Damage;365
14.2.1;8.2.1 Bunch Rot;366
14.2.2;8.2.2 Powdery Mildew;370
14.2.3;8.2.3 Downy Mildew;372
14.2.4;8.2.4 Bacteria;374
14.2.5;8.2.5 Viruses;377
15;Abbreviations and Symbols;382
16;Glossary;384
17;References;394
18;Internet Resources;502
19;Index;504
Chapter 2 Phenology and Growth Cycle
The annual growth cycle of fruiting grapevines is divided into a vegetative and a reproductive cycle. Fruit production extends over 2 years: buds formed in the first year give rise to shoots bearing fruit in the second year. Seasonal growth is driven by day length and temperature, and alternates with winter dormancy. The transition from dormancy to active growth in spring is marked by bleeding of xylem sap from pruning wounds due to root pressure. Root pressure restores xylem functionality and rehydrates the dormant buds. Increasing temperature then leads to budbreak and shoot growth that is marked by apical dominance. Shoots and roots grow as long as the environment permits. The shoots form brown periderm when the days shorten in late summer, enter dormancy, and shed their leaves in autumn. Chilling temperatures release dormancy to resume growth in spring. Flower clusters are initiated in the buds in early summer, and flowers differentiate after budbreak the following spring. Double fertilization during bloom initiates the transition of flowers to berries. Berry growth follows a double-sigmoid pattern of cell division and expansion, seed growth, and final cell expansion concomitant with fruit ripening. Seedless berries have less discernible growth phases. Ripening makes berries attractive for seed dispersers to spread a vine’s genes. Keywords
Anthesis; apical dominance; berry fruit; differentiation; dormancy; phenology; reproductive growth; ripening; root pressure; vegetative growth Chapter Outline 2.1 Seasons and Day Length 59 2.2 Vegetative Cycle 61 2.3 Reproductive Cycle 83 2.1 Seasons and Day Length
Phenology (Greek phainesthai=to appear, logos=knowledge, teaching) is the study of natural phenomena that recur periodically in plants and animals and of the relationship of these phenomena to climate and changes in season. In other words, it is the study of the annual sequence of plant development. Its aim is to describe the causes of variation in timing of developmental events by seeking correlations between weather indices and the dates of particular growth events and the intervals between them. Phenology investigates a plant’s reaction to the environment and attempts to predict its behavior in new environments. In viticulture, phenology is mainly concerned with the timing of specific stages of growth and development in the annual cycle. Such knowledge can be used for site and cultivar selection, vineyard design, planning of labor and equipment requirements, and timing of cultural practices as part of vineyard management (Dry and Coombe, 2004). Grapevines, like other plants, monitor the seasons by means of an endogenous “clock” (Greek endon=inside, genes=causing) in order to prevent damage by unfavorable environments. Every plant cell contains a clock (perhaps even several clocks), and the clocks of different cells, tissues, and organs act autonomously (McClung, 2001, 2008). The clock is driven by a self-sustaining oscillator consisting of proteins that oscillate with a 24 h rhythm in response to light detected by phytochromes (see Section 5.2) and other light-sensitive proteins that translate the light signal into a clock input that uses this time-encoded information to regulate physiological functions (Fankhauser and Staiger, 2002; Spalding and Folta, 2005). Additional input is provided through feedback loops that arise from various compounds produced by the plant’s metabolism (McClung, 2008). Just how plants manage to integrate and transform this environmental and metabolic information into daily and seasonal functions and how they avoid being “fooled” by bright moonlight is still mysterious, but it could involve fluctuations in calcium concentration of the cytosol. Cytosolic Ca2+ also oscillates with a 24 h period with a peak before dusk, and it participates in the “translation” of many signals, both internal and external (Hotta et al., 2007; McClung, 2001; Salomé and McClung, 2005; see also Section 7.3). In contrast to the temperature dependence of most biochemical processes, the period of the rhythm is temperature compensated so that it remains constant at 24 h over a wide range of temperatures. The gradual shift in time of sunrise (actually the change between dawn and dusk) as the seasons progress serves to reset the phase of the clock every day and enables plants to “know” when the sun will rise even before dawn (Fankhauser and Staiger, 2002). This synchronization of the circadian (Latin circa=about, dies=day) clock to the external environment is termed entrainment. It translates day length (i.e., photoperiod) into a vine’s internal time as an estimate of both the time of day and the time of year, enabling it to anticipate and prepare for daily and seasonal fluctuations in light and temperature. For example, the genes responsible for “building” enzymes involved in the production of ultraviolet (UV)-protecting phenolics are most active before dawn, photosynthesis-related genes peak during the day (and opening and closing of the stomata can anticipate dawn and dusk), and genes associated with stress are induced in late afternoon (Fankhauser and Staiger, 2002; Hotta et al., 2007). On a seasonal scale, flowering, onset of fruit ripening, bud dormancy, leaf senescence (Latin senescere=to grow old), and cold acclimation are typical responses to day length (the so-called photoperiodism), although each of these developmental processes is also modulated by temperature, and some can be altered by stress factors such as drought (see Section 7.2), nutrient deficiency or excess (see Section 7.3), or infection by pathogens (see Section 8.2). In general, higher temperatures accelerate plant development and advance grapevine phenology (Parker et al., 2011). For example, the time of budbreak of a given V. vinifera cultivar grown in a specific location can vary by about 1 month from one year to another (Boursiquot et al., 1995). Even very high temperatures (e.g., average daily mean 35°C) still shorten the time from budbreak to bloom compared with lower temperatures (Buttrose and Hale, 1973). Therefore, one of the most conspicuous consequences of climate change associated with the current and future increase in temperature due to the man-made rise of atmospheric CO2 is a shift of phenological stages to earlier times during the growing season (Chuine et al., 2004; Wolfe et al., 2005). Because day length during the growing season is longer at higher latitudes, the effects of day length and latitude on the internal clock are very similar (Salomé and McClung, 2005). In tropical climates, with little or no change in day length (and temperature at lower elevations) during the year, grapevines behave as evergreens with continuous growth and strong apical dominance, and often continuous fruit production, throughout the year (Dry and Coombe, 2004; Lavee and May, 1997; Mullins et al., 1992). With appropriate pruning strategies (and sometimes deliberate defoliation by hand, often in combination with imposed water deficit and sprays of ethephon or other chemicals to induce near-dormancy, followed by applications of hydrogen cyanamide to induce uniform budbreak), two crops can often be harvested each year in the tropics (Bammi and Randhawa, 1968; Favero et al., 2011; Lin et al., 1985). In temperate climates (and at high elevations in the tropics), where winter conditions prevent the survival of leaves, grapevines have a discontinuous cycle with alternating periods of growth and dormancy. Under these conditions, the time of active growth generally occurs from March/April to October/November (Northern Hemisphere) or September/October to April/May (Southern Hemisphere). Several distinct developmental stages or key events have been identified (Figures 2.1 and 2.2) and have been given a variety of names. These stages include dormancy, budbreak (budburst), bloom (anthesis or flowering), fruit set (berry set or setting), veraison (French vérer=to change: color change or onset of ripening), harvest (ripeness or maturity), and leaf senescence and subsequent fall (abscission). Although by definition no visible growth occurs during dormancy, metabolism does not rest completely, but high concentrations of the dormancy hormone abscisic acid (ABA) keep it at a minimum necessary for survival of the buds and woody tissues. Although the division of meristematic cells in the buds and cambium is blocked, chromosome duplication and protein synthesis resume during the later stages of dormancy in preparation for the activation of growth when temperature and soil moisture become favorable in spring.
Figure 2.1 Grapevine growth stages according to Baillod and Baggiolini (1993).
Figure 2.2 Grapevine growth stages according to Eichhorn and Lorenz. Reproduced from Coombe, B.G. (1995). Adoption of a system for identifying grapevine growth stages. Aust. J. Grape Wine Res. 1, 104–110, with permission by Wiley-Blackwell. The annual growth cycle of mature, fruiting grapevines is often divided into a vegetative and a reproductive cycle. 2.2 Vegetative Cycle
In late winter or early spring, grapevines often exude xylem sap from pruning...
