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RAPD Assessment of California Phylloxera Diversity. Molecular Ecology 4:459464.)

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RAPD Assessment of California Phylloxera Diversity. Molecular Ecology 4:459464.) California grape phylloxera more variable than expected Jeffrey Granett 0 Andrew Walker P John De Benedictis GenineFong o Hong Lin P Ed Weber Many strains of grape phylloxera phylloxera’s biology, its life cycle and now have been identified in Cali- how grape species and rootstocks re- fornia vineyards. This variability sist its feeding. may be the result of multiple intro- This paper reviews our recent dis- ductions of this pest or of evolu- coveries that the DNA and feeding f tion of new strains on susceptible behavior of different collections of -G* or weakly resistant rootstocks. California phylloxera are relatively di- 8 1 verse. The discovery of diversity in 0 Thus own-rooted vines, weakly re- -J sistant rootstocks and those with biological populations is not novel, but the amount we found in California Grape phylloxera on rootstock. The minute V. vinifera parentage should not aphidlike insects feed on the root, causing a was unexpected. Because phylloxera be used in phylloxerated areas. In gall to form. The expanding and cracking gall were supposedly introduced into Cali- limits the root’s ability to take up water and addition, because of the observed fornia a relatively short time ago, and nutrients and allows rot organisms to invade. variability, quarantines are inef- they only reproduce asexually on the fective in preventing the occur- roots, we expected a relatively low are rarely damaged by its feeding. rence of biotype B phylloxera, as level of diversity. Other Vitis species, from parts of the it appears to evolve independently The variability in phylloxera’s abil- world where phylloxera are not na- in different areas. ity to feed on different grape species tive, did not evolve this resistance and and cultivars is of central importance are highly susceptible to damage. Vitis Grape phylloxera, Daktulosphaira to the use of rootstocks. Vitis species vinifera, the European grape species vitifoliae (Fitch), is one of the worlds and the rootstocks bred from them are from which most of our cultivated most important viticultural pests. This not uniformly resistant to grape phyl- grapes are derived, is one such suscep- minute aphidlike insect lives on grape loxera, nor are grape phylloxera types tible species. roots and foliage, and is most damag- uniformly aggressive on a given spe- Phylloxera were unintentionally ing to grape species that are suscep- cies or rootstock. As is common imported to Europe in the mid-1800s. tible to its root feeding. This feeding throughout biology, there is variability Vineyards before and during this pe- causes a swelling of the root that de- in both the plant and the insect. Many riod were planted on their own roots, velops into a gall and nourishes the questions have emerged: What gener- and phylloxera soon decimated them. insect. As the galling develops it ex- ated the diversity? Are other root- Numerous insecticides and other con- pands and cracks, wounding the root stocks in imminent danger from a trol measures were attempted, but and potentially allowing the entry of damaging biotype? How does this di- none proved effective at limiting rot organisms. The combination of versity affect the breeding and selec- phylloxera’s rapid spread. By the phylloxera’s limiting effect on the root tion of new rootstocks? 1870s, French viticulturists observed system’s ability to take up water and To begin addressing these ques- that the American grape species nutrients and the effect of rot organ- tions, we describe experiments to were not susceptible to phylloxera. isms eventually kills infested vines. evaluate the variability of grape phyl- Planchon and Laliman developed the Phylloxera can devastate a susceptible loxera in California and elsewhere. We idea that these root systems could be vineyard in just a few years. also review what is known about phyl- grafted with V. vinifera scion cultivars Grape phylloxera have been de- loxera aggressiveness from other to produce a perfect plant - one stroying vineyards around the world viticultural regions. whose root system was resistant to for the past 140 years. Fortunately, phylloxera, but whose top produced phylloxera-resistant rootstocks have Phylloxera native to the Americas commercially desirable grapes. Breed- been available and have effectively de- Grape phylloxera are native to the ers were soon crossing American Vitis fended vineyards against this pest for Americas, where Vitis species serve as species to produce rootstocks with about 110 years. Despite this long his- hosts. These American Vitis species phylloxera resistance, good horticul- tory, many questions remain about have coevolved with phylloxera and tural characters (such as rooting and CALIFORNIA AGRICULTURE, JULY-AUGUST 1996 9 grafting ability) and adaptation to Eu- ropean soils and climates. Hundreds of rootstocks were produced, tested and deployed throughout Europe. As phylloxera was spread to the rest of the viticultural world, these rootstocks followed in the insect’s devastating wake and are still effectively used today. AxR#1 was one such rootstock de- veloped in the late 1800s. It is a hybrid between the V. vinifera cultivar Aramon and the resistant grape spe- cies V. rupestris. This combination was made to produce a rootstock with ex- cellent drought and lime tolerance, the easy propagation of V. vinifera and the phylloxera resistance of V. rupestris. AxR#l was used in Europe and South Africa after its release, but its V. vin- ifera parentage caused it to succumb to phylloxera after 10 to 30 years of use. Phylloxera was first found in Cali- fornia in the 1860s, and viticulturists here used the European rootstocks to combat the pest. Rootstock trials were initiated in 1905 to determine which rootstocks were best suited to California’s grape-growing regions. By Vines on the left are grafted to the rootstock llOR, which is resistant to phylloxera 1958 the collected data were used to biotype B. Vines on the right are grafted to AxR#l. recommend AxR#l as well adapted to most of California’s vineyard soils, cli- mate, water conditions and scions. replanting this acreage using root- tional designations based on the be- Based on this recommendation, AxR#l stocks more resistant to phylloxera havior of phylloxera, not necessarily was used in 60 to 70% of the Napa and than AxR#l. on genetic differences. Sonoma County plantings that oc- In addition to biotypes A and B, curred in the 1960-1980 planting Phylloxera biotypes and diversity other types of phylloxera have been boom. Biotype B phylloxera were studied found on rootstocks other than AxR#1. However, AxR#1 began to collapse by comparing their survival and re- Some of these phylloxera developed under phylloxera pressure in 1983, production to those of the less aggres- relatively high populations, but did and by 1989 it was clear that AxR#l sive biotype A in laboratory-based as- not appear to affect the vines. These was not adequately resistant to Cali- says (bioassays) using root pieces of phylloxera behave differently than fornia phylloxera (see California Agri- AxR#l and Cabernet Sauvignon (see biotypes A and B in the root bioassays. culture March-April 1991). The decline California Agriculture Jan-Feb 1987). of AxR#1 was attributed to a new type The two biotypes were defined by of phylloxera, designated biotype B to their performance in these bioassays: distinguish it from phylloxera inca- biotype A phylloxera are able to grow pable of killing AxR#l, termed bio- and reproduce well on V. vinifera type A. roots, but do not perform well on Biotype B has been found through- AxR#l. In contrast, biotype B phyllox- out Napa and Sonoma counties and in era are able to grow and reproduce Lake, Mendocino, San Joaquin, Sacra- well on both root types. In all cases, mento, Santa Clara and Alameda phylloxera evaluated from declining counties. By the end of 1995, over AxR#l vineyards have been biotype B 12,000 acres of vineyards in Napa and Neither biotype A nor B is able to Sonoma counties had already been re- grow well on rootstocks such as Har- Fig. 1. Yearly total of AxR#l vineyard acre- moved as a direct result of biotype B mony, 5C, 3309, St. George, 110R and age removed due to biotype B phylloxera phylloxera (fig. 1).Grape growers are many others. Thus biotypes are func- infestation, Napa and Sonoma counties. 10 CALIFORNIA AGRICULTURE, VOLUME 50, NUMBER 4 feeding on mature roots is most dam- aging in the field. DNA was extracted from 13 of our lab-based phylloxera colonies repre- senting biotypes A and B and several strains. Extractions were made from 200 to 300 phylloxera eggs per colony, which were surface sterilized and sub- jected to PCR-RAPD reactions (poly- merase chain reaction-randomly am- plified polymorphic DNA). RAPDs depend on the ability of randomly generated 10-base-pair DNA se- quences (primers) to match with the corresponding sequences in the phyl- loxera DNA. The lengths of DNA be- tween attachment locations for the primers are spliced out and replicated many times. RAPD results are viewed on agarose gels as bands of different- sized DNA, the smaller bands moving further in the gel (fig. 2). Bands are quantified by their position and their presence or absence at each possible location. When analyzed, the results estimate how closely related (or unre- lated) the phylloxera DNAs are to each other (Fong et al. 1995). Most of the vines in this vineyard are grafted to AxR#l. The healthy vine in the center is RAPD analyses using 10 different grafted onto St. George, a rootstock that is resistant to biotype B. primers were run three times on the 13 DNA samples to ensure reaction re- producibility. Clear RAPD bands from Because they were not killing their Phylloxera development was exam- the three replicates were then scored host rootstocks, but instead appeared ined in petri dishes by placing single as either present or absent and com- to have adapted to them, the new eggs onto cleaned root pieces (about pared.
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