как производится вино
Как получается вино? - «Биомолекула»
Как получается вино? ... Получить вино можно, добавив винные дрожжи к виноградному суслу. В ходе спиртового брожения сахар превращается в этиловый ...
«лисьего» привкуса
Решение проблемы пришло от ученых: привить европейскую лозу на американский подвой (растение или его часть, на которое прививают черенок или почку другого растения) и вывести новые гибридные сорта путем скрещивания V. vinifera и американских видов. Вышеупомянутый вид V. labrusca демонстрировал низкую устойчивость к филлоксере, наиболее эффективными в борьбе оказались V. aestivalis, V. rupestris и V. riparia. К началу XX в. большая часть французских виноградников была пересажена, а война с филлоксерой была выиграна. Хотя первые гибриды спасли виноделие, они не получили широкого распространения в Европе из-за неприятного <anchor>«лисьего» привкуса</anchor>. Гибриды V. vinifera и V. labrusca были выведены для получения сортов винограда, способных противостоять более холодному климату и давать вина, более похожие на европейские сорта. Так были получены сорта Isabella, Concord и Wilder [3].
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**phylloxera**

This small yellow root-feeding aphid has probably had a more damaging impact on wine production than any other vine pest (see vine pests), or any vine disease (see vine diseases). It attacks only grapevines, and kills vines by attacking their roots. For many years after it first invaded Europe there was no known cure.

The effects of phylloxera were first noted in France in 1863 (see below), just as the country was recovering from another great scourge of 19th century European viticulture: oidium, or powdery mildew, which was first noted in 1847. Like powdery mildew, and the other fungal diseases yet to arrive (downy mildew in 1878 and black rot in 1885), the phylloxera louse was an unwelcome import from America which devastated European vineyards until appropriate control measures were found. In the history of agriculture it rivals the potato blight of Ireland as a plant disease with widespread social effects. In France, for example, almost 2.5 million ha/6.2 million acres of vineyards were destroyed, the aphid making no distinction between the vineyards of the most famous chateaux and those of humble peasants. For individual French vine-growers from the late 1860s, the sight of their vineyards dying literally before their eyes was particularly traumatic, although the epidemic soon spread elsewhere. Phylloxera invasion had a major social and economic impact, involving national governments and local committees, and requiring international scientific collaboration. For a while the very existence of the French wine industry was threatened. (See Burgundy, modern history, for example.)

Phylloxera has had several scientific names. Initially called Phylloxera vastatrix (the devastator) by the French scientist J.-E. Planchon, and also Phylloxera vitifoliae (A. Fitch), it is now more correctly known as Dactylasphaera vitifoliae (H. Shimer). See below for more detail.

**Biology**

The female phylloxera is yellow coloured and about 1 mm/0.039 in long. Typically surrounded by masses of eggs, it is barely visible to the naked eye as it feeds on the roots. There are four to seven generations in the summer, each producing females capable of laying more eggs. As the eggs hatch, so-called \`crawlers' move to other roots of the vine, and some climb the trunk and can spread to other vines, or even vineyards, by the action of wind or machinery dislodging them from the foliage. Because of the movement of the crawlers through soil cracks, phylloxera tends to spread in a circle from the original infected vine. Wind-blown crawlers create secondary infections downwind.

It has a complex life cycle, existing in both root-living and leaf-living forms, and causes damage to vines by injecting saliva to produce galls, by feeding on the sap, and by causing root deformities.

In humid regions there is also an associated life cycle whereby the root-hatched nymph produces a winged form which can travel longer distances, and which lays male and female eggs. The hatching female in turn lays a winter egg which develops into the stem mother or fundatrix, which lays eggs in a leaf gall (these galls typically being produced only on the leaves of American vines). Nymphs hatching from leaf galls can begin new infections as egg-laying females on roots.

The principal agent for the spread of phylloxera is man. It is most commonly transported from one vineyard to another on the roots of rootlings (one-year-old dormant plants). However, phylloxera is easily moved in the soil that sticks to implements, and by irrigation water. Recent research by King and colleagues in New Zealand has shown that crawlers present in vine foliage lead to secondary spread over considerable distances; they are moved by machines such as foliage trimmers and harvesters which brush against foliage, and also by wind.

Phylloxera is native to the east coast of the United States, and so native American vine species have generally evolved with resistance. Studies have shown that the basis of resistance is the development of cork layers beneath the wound made by phylloxera feeding on the root. This stops the invasion of other microbes (bacteria and fungi) which eventually rot the root and kill non-resistant vines.

Phylloxera kills vines which have not developed this resistance, such as the European vinifera vine species from which most wine is made, by destroying the root system. When first present, the phylloxera numbers build up quickly on roots and there is little apparent root damage. After a few years the root rot affects the top growth, shoot growth is stunted, and leaves lose their healthy green colour. Normally a vine dies within several years of the first infection. Vines which are struggling for other reasons are more susceptible to phylloxera and succumb quickly. Those growing on deep, fertile soil can continue to produce economic crops for many years after phylloxera attack. Phylloxera does not survive well in sandy soil, so vines planted in sand (such as on the Great Plain of Hungary, those planted by Listel on the French Mediterranean coast, or Colares in Portugual) are immune from attack.

**History**

Attempts by early European colonists in North America to establish imported vinifera vines met with disaster, presumably substantially because of phylloxera, to which they had no resistance-although at this time phylloxera had yet to be identified, and other American vine diseases such as Pierce's disease in Florida, downy and powdery mildew in all regions, and the very cold winters doubtless played a part. See United States, history, for more details.

How did phylloxera come to Europe? In the mid 19th century there was a considerable importation of living plants into Europe. This trade was supported by wealthy people who could afford elaborate gardens, greenhouses, and conservatories, and encouraged by the Victorians' keen interest in botany. Plants could be imported dormant, or kept alive and protected from salt spray by the ingenious Ward case, a closed glass container mounted on the deck of the ship and not unlike today's terrariums. In 1865 alone, 460 tons of plants worth 230,000 francs were imported into France, and this trade had grown to 2,000 tons by the 1890s. In 1875 50 tons were imported from the USA and much of this was vines. Jules Planchon, Professor of Pharmacy at Montpellier University, noted that rooted American vines were imported in particularly significant quantities between 1858 and 1862, and sent to parts of Europe as far apart as Bordeaux, England, Ireland, Alsace, Germany, and Portugal. No doubt phylloxera was an unsuspected passenger on vine roots at the same time.

Just like the fungal disease powdery mildew, the first reporting of phylloxera was in England, in 1863, when Professor J. O. Westwood, a famous entomologist at Oxford University, received insect samples from a greenhouse in Hammersmith, a London suburb. Since the insects were in a leaf gall, the vine species was probably American, and these plants may even have been a source of introduction of the pest to England. In 1863 an unknown vine disease in France was being talked about, with two vineyards in the southern Rhone affected. The first printed report in France was a letter written by a veterinarian in 1867 about a vineyard planted in 1863 at St-Martin-de-Crau (only about 30 km/20 miles north east of the University of montpellier's modern phylloxera-free domaines on the Mediterranean coast, ironically), which developed unhealthy vines in summer 1866, failing to grow the following spring. It is likely that infection occurred several years beforehand. There were other reports from Narbonne in the Languedoc, and the Gard and Vaucluse departements in the southern Rhone in 1867, and from Bordeaux in 1869. (It was usually several years after the initial sighting that the louse's predations had a serious effect; the Medoc, for example, was not commercially compromised until the late 1870s.)

The first of many committees formed to resolve the phylloxera question was a Commission instigated by the Vaucluse Agricultural Society. On 15 July 1868 it began its investigation of affected vineyards in the southern Rhone. One member of the Commission was Jules Planchon, who had good training to work on phylloxera. He had spent a period at Kew Botanical Gardens in England, and subsequently became the brother-in-law of J. Lichtenstein, an amateur entomologist. Planchon noticed that dying vines had small yellow insects on their roots and noted the resemblance to an aphid Phylloxera quercus living on oak trees. He named the insect believed to be ravaging vineyards Phylloxera vastatrix. The form previously described by A. Fitch as living in the leaf galls of American vines was established by Planchon to be the same insect and therefore, according to the rules of priority, it became known as Phylloxera vitifolii (Fitch).

The Herault Commission made its findings public in August 1868 but these caused little interest in any but the local newspapers. There was a marked reluctance to accept that this little yellow insect could be causing such devastation. Other explanations current at the time were over-production, winter cold and other bad weather, weakening of the vineyards as a result of continued vegetative reproduction, soil exhaustion, and also God's wrath at contemporary vices (which may sound familiar to modern readers). Even eminent scientists of the time misdiagnosed the problem. The distinguished entomologist V. Signoret thought phylloxera was an effect not the cause, and Dr Guyot thought its presence was due to over-severe pruning (the opposite of overcropping)! The debate was ended by the 1869 Commission of the Societe des Agriculteurs de France headed by L. Vialla, who gently but firmly debunked all false theories.

Total French wine production fell from a peak of 84.5 million hl/2,200 million gal in 1875 to a mere 23.4 million hl in 1889. But even by June 1873 the French government was sufficiently alarmed by the spread of phylloxera to offer a large prize (300,000 francs) for a remedy, which was to be verified by experimentation carried out by the School of Agriculture at Montpellier. Up to October 1876, 696 suggestions were forwarded to Professors Durand and Jeannenot at Montpellier and between 1872 and 1876 1,044 treatments were evaluated. Among the deluge of suggestions submitted were those verging on the ridiculous, which included burying a living toad under the vine to draw the poison, and irrigating the vines with white wine. Entries were received from other countries as different as Denmark and Singapore. All of this work, however, produced little benefit, as only two treatments based on the chemical application of various forms of sulphides appeared to show much advantage. Surprisingly, carbon bisulphide failed in the evaluation even though it was later used extensively.

Early attempts at commercial control of phylloxera included flooding, which was studied by the Commission in 1873. It was found that flooding in winter for weeks on end controlled the pest, but of course few vineyards were near enough to water supplies or sufficiently flat for this to be a widespread solution (although it is still used in parts of Argentina). Vineyards on sandy soils were noted to be immune, although this offered no control. The injection of the liquid carbon bisulphide was to become widespread, so that by 1888 some 68,000 ha/168,000 acres had been treated. The insecticidal properties of carbon bisulphide were found in 1854 on grain weevils, and Baron Paul Thenard evaluated it in Bordeaux in 1869. The first experiments used too high a dose, severely affecting the vines, but subsequently the practice of injecting it, mixed with water, into the soil, using about 30,000 holes per ha, became widespread.

By the time of the International Phylloxera Congress at Bordeaux in 1881 two distinct schools of thought on how the industry might be saved had emerged. The chemists considered salvation lay in carbon bisulphide or related chemicals, or in flooding. In opposition were the \`Americanists', who advocated grafting desired varieties on to American vine species used as rootstocks. Gaston Bazille had suggested grafting in 1869 and Leo Laliman of Bordeaux drew early attention to the resistance of these species to the phylloxera. Laliman had studied the resistance of American species to powdery mildew in a collection in his vineyards since 1840. (He subsequently tried to claim the prize for controlling phylloxera.)

Following successful demonstrations of the ability of American vines to withstand phylloxera, and the visit of Planchon to America in 1873 where his study was guided by the noted scientist C. V. Riley, state entomologist of Missouri, the use of grafted vines began, rising from 2,500 ha in 1880 to 45,000 ha in 1885. It was Riley who positively identified the unknown French insect as identical to the American one, a critical step in its eventual control; he was one of the first to suggest grafting European varieties to American rootstocks, and his authority and expertise on phylloxera gave weight to the suggestion. Riley referred French experts to the nurserymen of Missouri, who provided most of the rootstocks that saved the vineyards of France and ultimately Europe. Millions of rootstock cuttings were shipped to France, especially between 1873 and 1876. Fortunately in Missouri there were many native American Vitis species: labrusca was found to lack resistance, but aestivalis, rupestris, and riparia were found to be most effective. Another American who helped the French by providing expertise and rootstock was one of the fathers of American viticulture, T. V. Munson of Texas.

The use of grafted vines began, rising from 2,500 ha in 1880 to 45,000 ha in 1885. All was not plain sailing, however. Many American species could not tolerate the calcareous soils of France (see chlorosis), the amount of grafting to be done was almost overwhelming, and also there was a concern that the use of these foreign rootstocks (whose own wine is so often reviled for its foxy flavour) might affect wine quality. The latter objection is voiced by the uneducated even today, and in Burgundy the importation of American vines was prohibited until 1887, although the clandestine activities of growers anxious to save their vineyards forced its repeal.

Eventually rootstock use became the established method for the control of phylloxera and this has had a dramatic effect on vine nursery operations world-wide. There was a period in the late 19th century, however, when it was thought that breeding hybrids of European with American vine varieties might produce vines with sufficient phylloxera resistance not to need grafting, whose wines were not marked by the undesirable foxy flavour of some American grapes. The class of varieties so created is loosely termed French hybrids or direct producers. These efforts were generally unsuccessful, however, in that the vines had inadequate phylloxera tolerance and produced lower-quality wine, although they became popular with growers because of their high yields and their tolerance of fungal diseases. Nevertheless, many of the rootstocks used today (see rootstocks for a detailed list) were bred in this period-although hybrids with vinifera as one parent have been generally found to have insufficient tolerance to phylloxera (see below).

**Geography**

Phylloxera is now widespread around the world, having been found in California (1873), Portugal (1871), Turkey (1871), Austria (1872), Switzerland (1874), Italy (1875), Australia (1877), Spain (1878), Algeria (1885), South Africa (1885), New Zealand (1885), and Greece (1898). There are few parts of the world free from the pest, although these include parts of Australia (on which a strict quarantine is imposed), parts of China, Chile, Argentina, India, Pakistan, and Afghanistan, and some Mediterranean islands such as Crete, Cyprus, and Rhodes.

There are also small sandy vineyards in otherwise affected wine regions (see Bollinger and Colares) which have never been affected by phylloxera either because of isolation or because of the soil composition.

For much of the late 20th century phylloxera has not been regarded as a serious problem, either because it was not known in a region, or because a wide range of rootstocks are available which are suited to different varieties, soil, and climate conditions along with tolerance to other pests such as nematodes. About 85 per cent of all the world's vineyards were estimated in 1990 to be grafted onto rootstocks presumed to be resistant to phylloxera.

Phylloxera has been noted in the 1980s, however, on ungrafted vines in parts of Greece, England, New Zealand, Australia, Oregon, and, most dramatically, on grafted vines in California where the so-called \`biotype B' strain of phylloxera was identified. The widely used rootstock AXR1 (see rootstocks for details) was found to offer insufficient phylloxera tolerance, and a significant proportion of grafted vineyards in Napa and Sonoma succumbed to phylloxera and were replanted in the 1990s.

Phylloxera is known to occur naturally as a number of related strains or \`biotypes'. King and Rilling exchanged phylloxera and vine cuttings between New Zealand and Germany, and confirmed evidence of the biotype differences between the two countries. Interestingly, the AXR1 rootstock was found not to be affected by phylloxera found in New Zealand, but it was affected by German phylloxera. New Zealand probably obtained phylloxera from California in 1885, which suggests that the biotype of phylloxera originally in California did not affect AXR1. With the introduction of other biotypes, probably on rooted vines imported from eastern American states at a time of nursery shortages in California, the demise of AXR1 began.

No doubt one of the most important effects of the phylloxera invasion of the world's vineyards was the inadvertent spread of virus diseases. As grafting became widespread towards the end of the 19th century, virus diseases were also spread as a common result of grafting, because virus-infected cuttings were used in grafting. A virus originally present only in the rootstock would spread to the fruiting vinifera variety after grafting. Rootstocks do not show virus symptoms, and virology in viticulture was not well understood until the 1950s. With known effects on fruit ripening (see leafroll virus), perhaps it was virus diseases more than grafting which led to the debate as to the relative merits of \`pre-phylloxera' and \`post-phylloxera' wines.

**Bibliography**

- Boubals, D., \`La Realite internationale relative au phylloxera', Progres agricole et viticole, 108 (1991), 494-6.

- Buchanan, G. A., and Amos, T. G., \`Grape pests', in B. G. Coombe and P. R. Dry (eds.), Viticulture, ii: Practices (Adelaide, 1992).

- King, P. D., and Rilling, G., \`Further evidence of phylloxera biotypes: variations in the tolerance of mature grapevine roots related to the geographical origin of the insect', Vitis, 30 (1991), 233-44.

- Ordish, G., The Great Wine Blight (2nd edn., London, 1987).

- Pinney, T., A History of Wine in America (Berkeley, Calif., 1989).

- Winkler, A. J., et al., General Viticulture (2nd edn., Berkeley, Calif., 1974).


**References**

 [American vine species](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=102) [black rot](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=334) [Bollinger](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=371) [Burgundy](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=491) [chlorosis](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=690) [Colares](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=761) [direct producer](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=953) [downy mildew](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=999) [flooding](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1204) [foxy](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1237) [French hybrids](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1260) [fungal diseases](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1288) [grafting](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1386) [Hungary](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1548) [hybrids](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1551) [leafroll virus](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=1755) [nematodes](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2192) [nursery](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2237) [Pierce's disease](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2428) [powdery mildew](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2519) [pre-phylloxera](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2527) [quarantine](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2579) [rootling](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2730) [rootstock](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=2731) [sulphides](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3093) [United States](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3341) [vine diseases](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3450) [vine pests](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3462) [vinifera](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3482) [virus diseases](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3507) [Vitis](/web/20080727104408/http://www.winepros.com.au/jsp/cda/reference/oxford_entry.jsp?entry_id=3516)

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