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| ISBN-13: | 9780080890487 |
|---|---|
| Publisher: | Elsevier Science & Technology Books |
| Publication date: | 02/05/2010 |
| Sold by: | Barnes & Noble |
| Format: | eBook |
| Pages: | 400 |
| File size: | 5 MB |
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THE SCIENCE OF GRAPEVINES
ANATOMY AND PHYSIOLOGYBy Markus Keller
Academic Press
Copyright © 2010 Markus Keller, Published by Elsevier Inc.All right reserved.
ISBN: 978-0-08-089048-7
Chapter One
Botany and Anatomy
OUTLINE
1.1. Botanical Classification and Geographical Distribution 1
1.2. Cultivars, Clones, and Rootstocks 9 1.2.1. Variety versus Cultivar 9 1.2.2. Cultivar Classification 13 1.2.3. Clones 15 1.2.4. Rootstocks 16
1.3. Morphology and Anatomy 20 1.3.1. Root 21 1.3.2. Trunk and Shoots 27 1.3.3. Nodes and Buds 33 1.3.4. Leaves 36 1.3.5. Tendrils and Clusters 41 1.3.6. Flowers and Grape Berries 43
1.1. BOTANICAL CLASSIFICATION AND GEOGRAPHICAL DISTRIBUTION
The basic unit of biological classification is the species. According to the "biological species concept," a species is defined as a community of individuals—that is, a population or group of populations, whose members can interbreed freely with one another under natural conditions but not with members of other populations (Mayr, 2001; Soltis and Soltis, 2009). In other words, such communities are reproductively isolated. Although each individual of a sexual population is genetically unique, each species is a closed gene pool, an assemblage of organisms that does not normally exchange genes with other species. Their genes compel the individuals belonging to a species to perpetuate themselves over many generations. Yet all life forms on Earth are interrelated; they all ultimately descended from a common ancestor and "dance" to the same genetic code, whereby different combinations of three consecutive nucleotides of each organism's deoxyribonucleic acid (DNA) specify different amino acids that can be assembled into proteins. Because they are thus interrelated, organisms can be grouped according to the degree of their genetic similarity, external appearance, and behavior. In the classification hierarchy, closely related species are grouped into a genus, related genera into a family, allied families into an order, associated orders into a class, similar classes into a division (plants) or a phylum (animals), related divisions or phyla into a kingdom, and, finally, allied kingdoms not into an empire but a domain. The "evolutionary species concept" recognizes this ancestor–offspring connection among populations that may follow distinct evolutionary paths to occupy separate ecological niches but may continue to interbreed for some time (Soltis and Soltis, 2009).
As is the case with many plants, the species of the genus Vitis are not very well-defined because of the extreme morphological variation among and within populations of wild vines (Currle et al., 1983; Hardie, 2000; Mullins et al., 1992). This implies the following: (1) All Vitis species are close relatives that share a relatively recent common ancestor, and (2) evolution is still at work, throwing up new variants all the time (see Chapter 2.3). Many vine species are actually semispecies—that is, populations that partially interbreed and form hybrids under natural conditions, which is in fact common among plants and may be an important avenue for the evolution of new species (Soltis and Soltis, 2009). Despite some hybridization where their natural habitats overlap, however, the various Vitis gene pools usually stay apart so that the populations remain recognizably different. Grapevines are a good example of the limits of taxonomic systems, demonstrating that there is a continuum of differentiation rather than a set of discrete, sexually incompatible units. As early as 1822, the Rev. William Herbert asserted that "botanical species are only a higher and more permanent class of varieties," and in 1825 the geologist Leopold von Buch postulated that "varieties slowly become changed into permanent species, which are no longer capable of intercrossing" (both cited in Darwin, 2004). Charles Darwin later expressed it clearly: "Wherever many closely allied yet distinct species occur, many doubtful forms and varieties of the same species likewise occur" and, furthermore, "there is no fundamental distinction between species and varieties," and, finally, "varieties are species in the process of formation" (Darwin, 2004). Indeed, modern genetic evidence indicates that the various Vitis species evolved relatively recently from a common ancestor so that they have not yet had time to develop the complete reproductive isolation that normally characterizes biological species. Thus, Vitis species are defined as populations of vines that can be easily distinguished by morphological traits, such as the anatomy of their leaves, flowers, and berries, and that are isolated from one another by geographical, ecological, or phenological barriers; such species are termed ecospecies (Hardie, 2000; Levadoux, 1956; Mullins et al., 1992). The following is a brief overview of the botanical classification of grapevines, starting with the domain at the top of the hierarchy and finishing with a selection of species at the base.
Domain Eukarya All living beings, making up the earth's biological diversity or biodiversity, are currently divided into the three great domains of life: the Bacteria, the Archaea, and the Eukarya. The Eukarya (eukaryotes; Greek eu = true, karyon = nucleus) include all terrestrial, sexually reproducing "higher" organisms with relatively large cells (10–100 μ) containing a true cell nucleus, in which the DNA-carrying chromosomes are enclosed in a nuclear membrane, and cell organelles such as mitochondria and plastids (Mayr, 2001). They evolved following injections of oxygen into the atmosphere caused by abiotic (i.e., nonbiological) factors such as plate tectonics and glaciation (Lane, 2002). The vast majority of life and the bulk of the world's biomass—the small (1–10 μm), single-celled prokaryotes (Greek pro = before) with cell walls composed of peptidoglycans (protein–polysaccharides)—is grouped into the other two domains. However, both the photosynthetic organelles (chloroplasts, from cyanobacteria) and the "power plants" (mitochondria, from proteobacteria) of eukaryotic cells have descended from (symbiotic) bacteria that "infected" other single-celled organisms (or were "swallowed" by them) over 1 billion years ago. These organelles still retain some of their own DNA (i.e., genes), although more than 95% of their original genes have since been and are still being lost or transferred (donated) to their host's nucleus (Timmis et al., 2004). Some bacteria cause diseases of grapevines; for example, crown gall is caused by Agrobacterium vitis and Pierce's disease by Xylella fastidiosa (see Chapter 7.5).
Kingdom Plantae The Eukarya comprise at least four kingdoms; the number changes as the relationships among organisms become better known. The Animalia comprise the multicellular animals with two sets of chromosomes, cells without cell walls, except in arthropods (insects, spiders, and the like), which have chitin cell walls, and are the domain of zoology. The Plantae or plants have a haplo-diploid life cycle and cell walls composed of cellulose, and they are studied in the field of botany. The Fungi include the haploid mushrooms, molds, and other fungi, with cell walls composed of glucans and chitin; they are studied in mycology. The Protista or Protoctista are a catch-all group of all other "higher order" organisms from single-celled microbes, including unicellular fungi, plants (green algae), and animals (protozoans), to large, multicellular seaweeds (algae and kelp). There are approximately 500,000 plant species, which are classified into 12 phyla or divisions based largely on reproductive characteristics. The vascular or higher plants, to which grapevines belong due to their water conduits, form the Subkingdom Tracheobionta. Whereas one group of fungi (singular fungus), the yeasts (especially Saccharomyces cerevisiae), turns grapes into wine through fermentation, other fungi cause diseases of grapevines (see Chapter 7.5); for example, gray rot is caused by Botrytis cinerea Pers.:Fr. and powdery mildew by Erysiphe necator [aka Uncinula necator (Schwein.) Burr.]. Animals can also be important pests of grapevines, especially certain insects (e.g., phylloxera, Daktulosphaira vitifoliae Fitch), mites, and nematodes.
Division (Phylum) Angiospermae (Synonym Magnoliophyta) The angiosperms or, in new terminology, the magnoliophytes are the flowering plants, which include approximately 270,000 species. They are believed to have evolved from a common ancestor that lived approximately 160 million years ago during the late Jurassic period, and they make up the most evolutionarily successful group of plants. Angiosperms are the plants with the most complex reproductive system: They grow their seeds inside an ovary (Greek angeion = pot, vessel) that is itself embedded in a flower. After the flower is fertilized, the other flower parts fall away and the ovary swells to become a fruit, such as a grape berry. Indeed, the production of fruits is what defines the angiosperms and sets them apart from the gymnosperms, with whom they are classed in the Superdivision Spermatophyta or seed plants.
Class Dicotyledoneae (Synonym Magnoliopsida) This class is large and very diverse, and its members are often called dicot plants. The vast majority of plants (~200,000 species), including most trees, shrubs, vines, and flowers, and most fruits, vegetables, and legumes belong to this group. Like all members of the Dicotyledoneae, grapevines start their life cycle with two cotyledons (seed leaves) preformed in the seed.
Order Rhamnales (Vitales According to the Angiosperm Phylogeny Web) Grapevines belong to the order Rhamnales, which gets its name from the genus Rhamnus, the buckthorns. The order has three families: Rhamnaceae (e.g., Ziziphus jujuba Mill., jujube tree), Leeaceae (the oleasters), and Vitaceae. Plants of the family Leeaceae are more recognizable as being related to grapevines than those belonging to the Rhamnaceae, being shrubs or trees with flowers aggregated in inflorescences, black berries, and seeds that resemble grape seeds, also named pips. Some taxonomists have now separated the Vitaceae (Jansen et al., 2006) and Leeaceae from the Rhamnales and placed them in the order Vitales.
Family Vitaceae The members of this family are collectively termed grapevines. The family contains approximately 1000 species assigned to 17 genera that are typically shrubs or woody lianas that climb by means of their leaf-opposed tendrils—hence the name Vitaceae (Latin viere = to attach). Although most species of this family reside in the tropics or subtropics, a single species from the temperate zones has become the world's leading fruit crop grown in almost 90 countries for wine and juice production or as fresh table grapes or dried grapes (raisins). Vitaceae roots are generally fibrous and well branched, and they can grow to several meters in length. The leaves are alternate, except during the juvenile stage in plants grown from seeds, and can be simple or composite. The fruits are usually fleshy berries with one to four seeds. All cultivated grapes belong to either the genus Muscadinia (2n = 40 chromosomes) or the genus Vitis (2n = 38 chromosomes). The former classification of Muscadinia and Euvitis as either subgenera or sections of the genus Vitis has fallen out of favor among taxonomists (Mullins et al., 1992). Because of the different numbers of chromosomes, crosses between these two genera rarely produce fertile hybrids. Key morphological characteristics of the two genera include the following:
Simple leaves
Simple or forked tendrils
Generally unisexual flowers—that is, either male (staminate) or female (pistillate)
Fused flower petals that separate at the base, forming a "calyptra" or cap
Soft and pulpy berry fruits
Genus Muscadinia Members of the genus Muscadinia usually have glabrous (hairless) leaves, simple tendrils, nonshredding bark, nodes without diaphragms, and hard wood (Currle et al., 1983; Mullins et al., 1992). Because they do not root from dormant cuttings, they are usually propagated by layering, although they do root from green cuttings. The "homeland" of this genus extends from the southeastern United States to Mexico. The genus has only three species, which are all very similar and may not even deserve to be classed as separate species (Currle et al., 1983; Mullins et al., 1992; Olien, 1990).
Muscadinia rotundifolia Small (formerly Vitis rotundifolia Michaux): A dioecious plant, although breeding has yielded perfect-flowered and female cultivars, such as Noble, Carlos, or Magnolia, known as "muscadines" that are grown as table, jelly, or wine grapes. The species is native of the southeastern United States. The musky flavor and thick skins of the fruit can be unattractive. The species has co-evolved with and therefore resists or tolerates the grapevine diseases and pests native to North America, including the fungi powdery mildew and black rot, the slime mold downy mildew, the bacterium causing Pierce's disease, the aphid phylloxera (Daktulosphaira vitifoliae), and the nematode Xiphinema index (which transmits the grapevine fanleaf virus), but is sensitive to winter frost and lime-induced chlorosis (Alleweldt and Possingham, 1988). Although usually incompatible in both flowering and grafting with Vitis species, it does produce fertile hybrids with V. rupestris, which allows it to be used in modern (rootstock) breeding programs.
Muscadinia munsoniana Small (Simpson): Native to Florida and the Bahamas, with better flavor and skin characteristics than M. rotundifolia, but not cultivated.
Muscadinia popenoei Fennell: Native to southern Mexico ("Totoloche grape"), relatively unknown.
Genus Vitis The genus Vitis occurs predominantly in the temperate and subtropical climate zones of the Northern Hemisphere (Mullins et al., 1992; Wan et al., 2008a). All members of this genus are perennial vines or shrubs with tendril-bearing shoots. This genus probably comprises 60–70 species (plus up to 30 fossil species and 15 doubtful species) spread mostly throughout Asia (~40 species) and North America (~20 species) (Alleweldt and Possingham, 1988; Wan et al., 2008b,c). The Eurasian species Vitis vinifera L. gave rise to the overwhelming majority of grape varieties cultivated today. Plants that belong to this genus have hairy leaves with five main veins, forked tendrils, bark that shreds when mature, nodes with diaphragms, and soft secondary wood. They all can form adventitious roots, which permits propagation by cuttings, yet only V. vinifera, V. riparia, and V. rupestris root easily from dormant cuttings. Although the ancestor of all Vitis species may have had perfect (i.e., bisexual or hermaphroditic) flowers (McGovern, 2003), the extant wild species are dioecious (Greek dis = double, oikos = house), containing imperfect male (i.e., female sterile or staminate) or female (i.e., male sterile or pistillate) flowers on different individual plants, whereas the cultivated varieties of V. vinifera have perfect or, in a few cases, physiologically female flowers (Negrul, 1936; Pratt, 1971; see also Figure 1.1). Members of this genus are very diverse in both habitat and form. Nevertheless, all species within the genus can readily interbreed to form fertile interspecific crosses called hybrids, which implies that they had a relatively recent common ancestor. Moreover, all species can be grafted onto each other. The genus is often divided into two major groups: the American group and the Eurasian group. The dominant species of the two groups differ greatly in their useful agronomic traits (Table 1.1), which makes them attractive breeding partners (Alleweldt and Possingham, 1988; This et al., 2006). Unfortunately, none of the many attempts and thousands of crosses that have been tested to date has truly fulfilled the breeders' hopes to combine the positive attributes while eliminating the negative ones contained in the natural genetic variation of the two groups. Perhaps the genes conferring disease resistance are coupled to those responsible for undesirable fruit composition. Indeed, hybrids have often been banned in European wine-producing countries because of their perceived poor fruit (and resulting wine) quality. The only unequivocal success story thus far has been the grafting of phylloxera-susceptible European wine grape cultivars to rootstocks that are usually hybrids of tolerant American Vitis species (see Chapter 1.2).
(Continues...)
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Table of Contents
About the Author vii
Preface ix
Acknowledgments xi
1 Botany and Anatomy 1
1.1 Botanical Classification and Geographical Distribution 1
1.2 Cultivars, Clones, and Rootstocks 9
1.3 Morphology and Anatomy 20
2 Phenology and Growth Cycle 49
2.1 Seasons and Day Length 49
2.2 Vegetative Cycle 53
2.3 Reproductive Cycle 68
3 Water Relations and Nutrient Uptake 85
3.1 Osmosis, Water Potential, and Cell Expansion 85
3.2 Transpiration and Stomatal Action 89
3.3 Water and Nutrient Uptake and Transport 92
4 Photosynthesis and Respiration 107
4.1 Light Absorption and Energy Capture 107
4.2 Carbon Uptake and Assimilation 111
4.3 Photorespiration 117
4.4 Respiration 118
4.5 From Cells to Plants 121
5 Partitioning of Assimilates 125
5.1 Photosynthate Translocation and Distribution 125
5.2 Canopy-Environment Interactions 140
5.3 Nitrogen Assimilation and Interaction with Carbon Metabolism 158
6 Developmental Physiology 169
6.1 Yield Formation 169
6.2 Grape Composition and Fruit Quality 178
6.3 Sources of Variation in Fruit Composition 209
7 Environmental Constraints and Stress Physiology 227
7.1 Responses to Stress 227
7.2 Water: Too Much or Too Little 231
7.3 Nutrients: Deficiency and Excess 243
7.4 Temperature: Too Cold or Too Warm 274
7.5 Living with Other Organisms: Defense and Damage 289
References 311
Index 369
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