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» Verticillium wilt has become an economically important disease of lettuce in California and other lettuce-growing regions.
» The fungal pathogens that cause Verticillium wilt of lettuce can survive in the soil for many years.
» Strategies for managing Verticillium wilt include minimizing the spread of the pathogen, lowering inoculum levels in the soil, and using disease-resistant lettuce varieties.
Until 1995, lettuce was thought to be immune to Verticillium wilt. Crops of lettuce had been grown in fields known to contain the pathogen for many years without any development of symptoms on lettuce. However, Verticillium wilt-like symptoms were observed on lettuce growing near Watsonville, California in 1994, and it was later determined that the symptoms were the result of infection by the fungus Verticillium dahliae. A few years later, the disease was observed in California’s Salinas Valley.1 Incidence levels of up to 80% have since been observed, and some fields have been abandoned before harvest due to the disease. A 2019 publication stated that Verticillium wilt of lettuce had not been detected in Arizona, but symptoms on lettuce in high tunnels and research field plots were observed in the state of Washington in 2010 and 2012.2,3 The disease on lettuce in Washington was found to be caused by a different species of Verticillium. Verticillium dahliae has a wide host range of about 400 identified plant species, including many crop, ornamental, tree, and weed species, and the pathogen commonly occurs in agricultural soils in temperate growing regions around the world.4,5 Verticillium wilt of lettuce has been found in northern Italy and on the Greek island of Crete.1
SYMPTOMS
The initial symptom on lettuce is a green, brown, or black discoloration of the vascular tissue in lateral root tips. On susceptible plants, this discoloration spreads into the taproot and crown (Figure 1). The vascular discoloration can look similar to discoloration caused by Fusarium wilt, corky root, and ammonia toxicity.4,5 The foliar symptoms of Verticillium wilt on lettuce appear about two weeks after the appearance of vascular discoloration, usually late in the season when the plants are near market maturity.1,4 Angular, necrotic lesions develop on the leaves, starting at the leaf margins (Figure 2A). Older, lower leaves become chlorotic, wilt, and then die (Figure 2B). The foliar symptoms progress inward, and eventually, the plant collapses. On crisp head varieties, outer leaves can collapse and die but remain tightly adhered to the head.4,5
Small, black structures called microsclerotia may develop andbecome visible along the veins of outer, affected leaves and in the roots of senescing plants.1,4 The symptoms progress to the center of the head, and the plant eventually dies. A study found that symptoms did not form on harvested heads, and that there was no additional post-harvest decline in quality. Therefore, the disease will likely have minimal impact on quality and marketability if heads are harvested before symptoms appear.4,6 All lettuce types are susceptible to Verticillium wilt, but the most damage usually occurs on crisphead varieties.1,4
PATHOGENS
There is little host specificity exhibited by V. dahliae, with isolates from lettuce infecting other host species and viceversa. However, two races (race 1 and 2) of V. dahliae have been characterized. Other Verticillium species, including V. isaacii, V. klehahnii, and V. tricorpus have also been found to cause wilt symptoms on lettuce. Verticillium tricorpus is the species identified on lettuce in Washington.3,4 These species appear to have narrower host ranges and cause less severe symptoms as compared to V. dahliae.
CYCLE AND CONDITIONS
Verticillium dahliae is a soilborne fungus that can persist in the soil indefinitely. The pathogen can survive as microsclerotia, which can remain viable in the soil for up to fourteen years.1,5,7 The pathogen can be spread within and between fields on anything that moves infested soil, including equipment, shoes, and wind. The pathogen has also been detected on lettuce seeds and the seeds of other crops. The rate of infection from inoculum on lettuce seed appears to be low, and it may require planting infested lettuce seed over several years to result in disease development in the field.1 However, there is good evidence that V. dahliae can be present on the seed of some common rotational crops, including other leafy green species, and that this seedborne inoculum might increase the risk of Verticillium wilt occurring on lettuce.1,7 Infection of lettuce occurs when the microsclerotia germinate and produce hyphae (fungal threads) that penetrate near the root tips. The fungus then grows into the xylem (waterconducting tissue) and spreads upward into the stem and leaves of the plant. Verticillium wilt is a monocyclic disease, meaning that there is no plant-to-plant spread during the growing season. Microsclerotia produced on the infected lettuce plants enter the soil as the crop debris decomposes and become available to infect future crops.4 The level of disease on a lettuce crop is proportional to the amount of Verticillium inoculum in the soil, more microsclerotia result in a greater number of infected plants. However, the threshold for infection of lettuce by V. dahliae is high compared to the thresholds for some other crops.
Infection of strawberry plants requires an inoculum density of one to two microsclerotia per gram of soil. For cauliflower, ten microsclerotia per gram of soil results in about a 50% incidence of infection and sixteen microsclerotia per gram results in near 100% incidence. The infection threshold for lettuce is around 100 to 120 microsclerotia per gram soil, and levels between 200 and 600 microsclerotia per gram soil are likely to result in total loss of the lettuce crop.1,7 Verticillium wilt can develop on lettuce in any of the seasonal plantings, but in California symptoms are often more severe and the highest yield losses are seen on fall crops because the warmer temperatures favor the growth and development of the pathogen.1,4
MANAGEMENT
It is very difficult to eradicate the Verticillium wilt pathogen once it is established in a field, and there are no rescue treatments for controlling the disease once lettuce plants are infected. Therefore, it is important to prevent the introduction of the pathogen into non-infested fields whenever possible.4,7
Preventing the spread of infested soil and crop debris is the primary strategy for preventing the introduction of V. dahliae into non-infested fields. Planting seeds (lettuce and rotational crops) that are not infested with the pathogen is also needed to help prevent the establishment of V. dahliae. Because of the longevity of microsclerotia in the soil, with the exception of planting broccoli, crop rotation to non-host crops is not usually effective for managing Verticillium wilt.4,5
Some lettuce varieties have some resistance to V. dahlia race 1, but no varieties have resistance to race 2. The single dominant race 1 resistance gene is also effective against V. isaacii and V.klehahnii. About 55% of V. dahliae isolates in California lettuce fields belong to race 1, and 45% belong to race 2. If the race 1 resistance gene is widely used in lettuce, it will likely result in an increase in the proportion of the population of race 2 isolates.1,4
Soil fumigants, such as chloropicrin + dazomet, can help reduce the population of V. dahliae microsclerotia in the soil. However, currently available fumigants will not completely eradicate the pathogen in a field.1 An alternative for reducing the number of microsclerotia in the soil is to rotate the field to broccoli. It has been shown that a planting of broccoli can help reduce the population of V. dahlia, potentially below the threshold level needed for the development of wilt on lettuce.
The lower return per acre for a crop of broccoli compared to lettuce has limited the use of this management strategy.1,7
SOURCES
1 Hayes, R. and Subbarao, K. 2011. Fifteen years of Verticillium wilt of lettuce in America’s salad bowl: a tale of immigration, subjugation, and abatement. Plant Disease 95:784-792.
3 Matheron, M. 2019.Comparison of two lettuce wilt diseases. Arizona Pest Management Center. University of Arizona. https://acis.cals.arizona.edu/agricultural-ipm/vegetables/vipm-archive/vipm-disease-view/comparison-of-two-lettuce-wilt-diseases.
3 Powell, M., Gundersen, B., Miles, C., Coats, K., and Inglis, D. 2013. First report of Verticillium wilt on lettuce (Lactuca sativa) in Washington caused by Verticillium tricorpus. Plant Disease, https://doi.org/10.1094/PDIS-12-12-1166-PDN.
4 Inderbitzin, P. and Subbarao, K. 2017. Verticillium wilt. In Subbarao, K., Davis, R., Gilbertson,
R., and Raid, R. (Eds.), Compendium of Lettuce Diseases and Pests (2nd ed. pp. 50-54) American Phytopathological Society, St. Paul. Ppg 50-54.
5 Koike, S. and Turini, T. 2017. Verticillium wilt. UC IPM Lettuce Pest Management Guidelines. https://ipm.ucanr.edu/agriculture/lettuce/verticillium-wilt/.
6 Puri, K. D., Vallad, G. E., Qin, Q.-M., Hayes, R. J., and Subbarao, K. V. 2019. Harvest of lettuce from Verticillium-infested fields has little impact on postharvest quality. Plant Disease,
103:668–676.
7 Carroll, C., Carter, C., Goodhue, R., Lawell, C., and Subbarao, K. 2018. A review of control options and externalities for Verticillium wilts. Phytopathology 108:160-171.
ADDITIONAL INFORMATION
For additional agronomic information, please contact your local seed representative. Performance may vary, from location to location and from year to year, as local growing, soil and environmental conditions may vary. Growers should evaluate data from multiple locations and years whenever possible and should consider the impacts of these conditions on their growing environment. The recommendations in this article are based upon information obtained from the cited sources and should be used as a quick reference for informationabout vegetable production. The content of this article should not be substituted for the professional opinion of a producer, grower, agronomist, pathologist and similar professional dealing with vegetable crops.
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5911_279851 Published 11/09/2023