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Wine Science: Principles and Applications, Third Edition - Vinum Vine

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198 4. <strong>Vine</strong>yard Practice<br />

Because grapevine roots do not form natural grafts,<br />

vine-to-vine transfer does not occur. Although transferable<br />

to other plants, the disease is limited to grapevines<br />

under fi eld conditions. This may result from the limited<br />

host range of its nematode vectors.<br />

The impact of infection varies widely, depending on<br />

cultivar tolerance <strong>and</strong> environmental conditions. Tolerant<br />

cultivars are little affected by infection, whereas susceptible<br />

varieties show progressive decline. Yield losses<br />

may be up to 80%, with the fruit of poor quality.<br />

Infected vines have a shortened life span, increased sensitivity<br />

to environmental stress, <strong>and</strong> reduced grafting<br />

<strong>and</strong> rooting potential. Three distinctive syndromes have<br />

been recognized, based on particular strain–cultivar<br />

combinations. These are malformation, yellow mosaic,<br />

<strong>and</strong> vein-b<strong>and</strong>ing.<br />

Fan-shaped leaf malformation is the most distinctive,<br />

but not only, foliage expression. Chlorotic speckling <strong>and</strong><br />

a leathery texture commonly accompany leaf distortion.<br />

Shoots may be misshapen, showing fasciation (stem<br />

fl attening), a zigzag pattern at the nodes, atypically variable<br />

internode lengths, double nodes, <strong>and</strong> other aberrations.<br />

Fruit set is poor <strong>and</strong> bunches are reduced in<br />

size. The yellow mosaic (chromatic) syndrome develops<br />

as a strikingly bright-yellow mottling of the leaves,<br />

tendrils, shoots, <strong>and</strong> infl orescences in the early spring.<br />

Discoloration can vary from isolated chlorotic spots to<br />

uniform yellowing. The third expression, veinb<strong>and</strong>ing,<br />

develops as a speckled yellowing on mature leaves, bordering<br />

the main veins in mid- to late-summer. In both<br />

discoloration syndromes, leaf shape is normal, but fruit<br />

set is poor, with many shot berries (“hen <strong>and</strong> chickens”<br />

appearance). Fanleaf degeneration also shows a characteristic<br />

intracellular development of trabeculae. These<br />

appear as str<strong>and</strong>s of cell-wall material spanning the<br />

lumen of xylem vessels.<br />

Where both vector <strong>and</strong> virus are established, planting<br />

vines grafted on rootstock resistant to the virus is usually<br />

required (tolerance to the vector can still permit infection).<br />

Fumigation of the soil with nematicides can, to<br />

varying degrees, reduce but not eradicate nematode populations.<br />

Allowing the l<strong>and</strong> to lie fallow can be useful in<br />

reducing nematode populations, but the strategy often<br />

requires 6–10 years. The long requisite fallow period<br />

probably results from nematode survival on undislodged<br />

roots. These occasionally remain viable for up to 6 years<br />

following vine uprooting.<br />

Leafroll Leafroll is a widespread virus-like disease<br />

associated with one or more of up to eight or nine grapevine<br />

leafroll-associated viruses (GLRaVs) (Boscia et al.,<br />

1995). GLRaV-1 <strong>and</strong> GLRaV-3 are the most widespread<br />

<strong>and</strong> economically signifi cant. They are all members of<br />

the closterovirus group. Two vitiviruses (GVA <strong>and</strong> GVB)<br />

have also been associated with leafroll, although they are<br />

more commonly associated with Kober stem grooving<br />

<strong>and</strong> corky bark, respectively. Symptomatic expression<br />

varies considerably, but it does not generally lead to vine<br />

degeneration. Many scion <strong>and</strong> rootstock cultivars are<br />

symptomless carriers of the infectious agent(s). As with<br />

fanleaf degeneration, the pathogens probably originated<br />

in Europe or the Near East; feral North American grapevines<br />

are not infected.<br />

The spread of the causal agents depends primarily<br />

on graft transmission. Nevertheless, insect vectors may<br />

occasionally be involved. For example, GLRaV-1, -3, -5<br />

can be transmitted by mealybugs or soft scale insects<br />

(Sforza et al., 2003). Several species of mealybugs have<br />

been reported to transfer grapevine viruses A (GVA)<br />

<strong>and</strong> B (GVB). Nevertheless, healthy <strong>and</strong> infected vines<br />

often coexist side by side without transmission. That<br />

the mealybug, Planococcus fi cus, has been found on<br />

the roots of grapevines (Walton <strong>and</strong> Pringle, 2004) is of<br />

concern since most control measures have been aimed<br />

at aboveground insects. It is a key pest in South Africa,<br />

the Mediterranean, Argentina, <strong>and</strong> now occurs in some<br />

regions of California.<br />

The disease complex derives its name from a marked<br />

down-rolling of the basal leaves that occurs late in the<br />

season in some varieties. The interveinal areas of leaves<br />

also may turn pale yellow or deep red, depending on the<br />

cultivar, whereas the main veins remain distinctly green.<br />

Infected vines can occasionally be detected by their<br />

retention of leaves much longer than adjacent healthy<br />

vines. In addition, leaf blades may fall, leaving petioles<br />

still attached to the cane. Whole vines, as well as individual<br />

shoots <strong>and</strong> leaves are dwarfed in comparison to<br />

healthy plants. Fruit production may be depressed by<br />

up to 40%, <strong>and</strong> berries may show delayed ripening,<br />

reduced sugar levels, <strong>and</strong> altered pigmentation.<br />

Control is dependent on the destruction of infected<br />

vines <strong>and</strong> replacement by disease-free stock. Disease-free<br />

nursery stock may be generated by thermotherapy or<br />

by micropropagation. Because the identity of all causal<br />

agents still remains unestablished, confi rmation of elimination<br />

of the infectious agent(s) is performed by grafting<br />

to sensitive cultivars (indexing).<br />

Yellow Speckle Yellow speckle is a widespread but<br />

relatively minor viroid disease of grapevines. Other<br />

viroids occur in grapevines, but their economic signifi -<br />

cance <strong>and</strong> relationship to recognized grapevine diseases<br />

are unclear. Symptoms of infection by grapevine yellow<br />

speckle viroids (GYSVd1 <strong>and</strong> GYSVd2) are often shortlived<br />

<strong>and</strong> develop only under special climatic conditions.<br />

Foliar symptoms generally develop at the end of the<br />

summer <strong>and</strong> consist of leaf spotting. When suffi ciently<br />

marked, the scattered chlorotic spots may resemble the

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