栎树猝死病菌检疫鉴定方法

根据栎树猝死病菌的形态学、生物学培养性状和分子生物学等方面的特征，确立了检疫鉴定栎树猝死病菌的各项技术要求。     

Managing Phytophthora ramorum in the Netherlands1

Phytophthora ramorum came to the Netherlands in 1993. Despite initially not seeming to pose a high risk, findings in California showed its potential destructive impact on ecosystems. A programme began in the Netherlands to eliminate P. ramorum from nurseries and surveys in the natural environment were held to obtain information to determine a strategy for dealing with the disease. About 1100 nurseries are inspected annually by NAKtuinbouw under the auspices of the Plant Protection Service and measures are taken according to EC directives. The percentage of infected nurseries decreased steadily during recent years, from 4% in 2002/2003 to 0.5% in 2004/2005. Surveys in the natural environment show that P. ramorum occurs on 2% of the sites with Rhododendron and therefore it was concluded that an elimination scenario is not realistic. A programme based on containment measures supported by an extension programme was put into place with its effects being monitored by the Plant Protection Service. 12 years of observing P. ramorum show that the risk for indigenous trees and shrubs in the Netherlands is very limited. Spread from infected rhododendrons to other potential hosts, even at heavily infected Rhododendron sites, hardly takes place although some infected Quercus rubra trees have been found. Recently several new Phytophthora species were found in natural environments in Europe and California, mainly as a result of intensive P. ramorum surveys. As well as P. ramorum, the Phytophthora spp. P. kernoviae, P. numerosa and P. pseudosyringae pose risks, indicating the need for a more general approach against Phytophthora diseases. As a result, a new protocol for detection and identification of Phytophthora spp. both as a group and individually is being developed and workers are asking whether these Phytophthora species could be managed together.     

Phytophthora ramorum: the situation in Scotland1

In Scotland inspections for Phytophthora ramorum on plants in the horticulture nursery trade started in July 2001 and are currently carried out four times per year. In addition, approximately 130 established gardens have been inspected for the disease during the years 2003 and 2004. Phytophthora ramorum has been found on Rhododendron, Viburnum, and lilac (Syringa vulgaris); the most important host plant is Viburnum tinus. The pathogen is confined to nurseries and garden centres with the exception of one private garden. The first finding was in April 2002 with 17 more outbreaks the same year. Since then the number of outbreaks per year has declined dramatically to 6 in 2003, 5 in 2004 and 3 until November 2005. Altogether, there have been 21 different outbreaks sites since the first finding, some with repeated occurrences of the disease.     

Fitness characteristics of the European lineages of Phytophthora ramorum

As an introduced pathogen, Phytophthora ramorum exists as four near-clonal evolutionary lineages, of which only EU1 and EU2 are established in the UK. EU1 has become widespread since the first findings in 2002 whereas EU2, detected in 2011, has a more limited distribution. Both lineages are epidemic in plantation-grown larch, sporulating asexually on needles, but also causing heavy dieback and mortality. To understand whether EU1 and EU2 pose different threats to forest health, we compared their growth characteristics on agar, pathogenicity on several hosts, and sporulation on Japanese larch needles. When pathogenicity was evaluated by measuring colonization at 20 °C in mature bark (phloem) of Japanese and European larch (Larix kaempferi and L. decidua), English oak (Quercus robur), and beech (Fagus sylvatica), Japanese larch was the most susceptible and oak the least susceptible. On average, EU2 isolates produced significantly larger lesions than EU1 isolates in Japanese larch and oak although not in the other hosts. With tests using young saplings of Japanese and European larch, damaging bark lesions formed at both 10 °C and 20 °C, but EU2 was significantly more pathogenic at 20 °C on both hosts compared with EU1. In contrast, both lineages caused similar amounts of necrosis on inoculated leaves of rhododendron (Rhododendron ponticum). Moreover, EU2 isolates usually sporulated less abundantly on larch needles compared with EU1 isolates, suggesting a trade-off in pathogenicity and sporulation between lineages. As EU2 tends to have smaller sporangia than EU1, this could also reduce the inoculum potential of EU2.     

Surveys for Phytophthora ramorum in the Czech Republic1

The State Phytosanitary Administration (SPA) of the Czech Republic has been conducting an official survey for Phytophthora ramorum since 2003. A single outbreak of P. ramorum was detected by the SPA on guelder rose (Viburnum bodnantense) in autumn 2003 during a phytosanitary inspection in an ornamental nursery made on the request of owner of the nursery as he had noticed plants of Viburnum bodnantense withering. The infested lot consisted of 15 plants, which were imported from Italy into the Czech Republic at the end of April in 2003. A post‐entry inspection was carried out at the place of destination just after arrival and no visual symptoms were seen at that time. The lot was destroyed by the owner of the nursery immediately after the sampling. A survey was carried out by the SPA in all nurseries in which host plants coming from the same origin as the infested plants had been planted. No other outbreak was found. Phytosanitary inspections were carried out in ornamental nurseries and garden centres, public parks and forestry sites. No outbreak was detected in 2004 (451 visual inspections) and 2005 (522 visual inspections). The status of Phytophthora ramorum in the Czech Republic is: absent, one isolated outbreak linked to an imported consignment was eradicated.     

Phytophthora ramorum and Phytophthora kernoviae: the woodland perspective

Phytophthora ramorum and P. kernoviae were first identified in Great Britain (GB) in 2002 and 2003 respectively, although both are believed to have been present for perhaps 10–20 years. Public gardens have been badly affected by these pathogens, but British woodlands are also considered at risk as a number of tree species have been found to be susceptible to the pathogens. The principal host of both Phytophthoras in GB is Rhododendron ponticum , which is arguably our most invasive weed species. It is found extensively in GB, especially along the west coast which has ideal climatic conditions for its growth and spread. In 2004 the Forestry Commission surveyed 1479 woodlands where R. ponticum was present, and in 2008 both infected and non-infected R. ponticum were cleared from approximately 1200 ha of woodlands in England and Wales where P. ramorum had been found. No clearance to date has taken place in Scotland. In the case of a positive finding in a garden or woodland, all woodlands with rhododendron within a 3 km radius of such a finding are to be surveyed. As both pathogens are notifiable any positives must be eradicated or contained through the serving of an official phytosanitary notice. Ministerial approval has now been given to a joint Defra and Forestry Commission initiative to undertake inter alia the removal of host plants (primarily rhododendrons) infected with these sporulating pathogens in woodlands and the wider environment in England and Wales. A similar programme will be effected in Scotland where the Forestry Commission is working with the Scottish Government's Rural Payments and Inspections Directorate.     

Transmission of Phytophthora ramorum in Mixed-Evergreen Forest in California

ABSTRACT During 2001 to 2003, the transmission biology of Phytophthora ramorum, the causal agent of sudden oak death, was studied in mixedevergreen forest, a common forest type in northern, coastal California. Investigation of the sources of spore production focused on coast live oak (Quercus agrifolia) and bay laurel (Umbellularia californica), dominant hosts that comprised 39.7 and 46.2% of the individuals at the study site, respectively. All tests for inoculum production from the surface of infected coast live oak bark or exudates from cankers were negative. In contrast, sporangia and chlamydospores were produced on the surface of infected bay laurel leaves. Mean number of zoospores produced from infected bay laurel leaves under natural field conditions during rainstorms was 1,173.0 +/- SE 301.48, and ranged as high as 5,200 spores/leaf. P. ramorum was recovered from rainwater, soil, litter, and streamwater during the mid- to late rainy season in all 3 years of the study. P. ramorum was not recovered from sporadic summer rains or soil and litter during the hot, dry summer months. Concentrations of inoculum in rainwater varied significantly from year to year and increased as the rainy season progressed for the two complete seasons that were studied. Potential dispersal distances were investigated for rainwater, soil, and streamwater. In rainwater, inoculum moved 5 and 10 m from the inoculum source. For soil, transmission of inoculum was demonstrated from infested soil to bay laurel green leaf litter, and from bay laurel green leaf litter to aerial leaves of bay laurel seedlings. One-third to one-half of the hikers tested at the study site during the rainy season also were carrying infested soil on their shoes. In streamwater, P. ramorum was recovered from an unforested site in pasture 1 km downstream of forest with inoculum sources. In total, these studies provide details on the production and spread of P. ramorum inoculum in mixed-evergreen forest to aid forecasting and managing disease transmission of this environmentally destructive pathogen.     

Sources of inoculum for Phytophthora ramorum in a redwood forest

ABSTRACT Sources of inoculum were investigated for dominant hosts of Phytophthora ramorum in a redwood forest. Infected trunks, twigs, and/or leaves of bay laurel (Umbellularia californica), tanoak (Lithocarpus densiflorus), and redwood (Sequoia sempervirens) were tested in the laboratory for sporangia production. Sporangia occurred on all plant tissues with the highest percentage on bay laurel leaves and tanoak twigs. To further compare these two species, field measurements of inoculum production and infection were conducted during the rainy seasons of 2003-04 and 2004-05. Inoculum levels in throughfall rainwater and from individual infections were significantly higher for bay laurel as opposed to tanoak for both seasons. Both measurements of inoculum production from bay laurel were significantly greater during 2004-05 when rainfall extended longer into the spring, while inoculum quantities for tanoak were not significantly different between the 2 years. Tanoak twigs were more likely to be infected than bay laurel leaves in 2003-04, and equally likely to be infected in 2004-05. These results indicate that the majority of P. ramorum inoculum in redwood forest is produced from infections on bay laurel leaves. Years with extended rains pose an elevated risk for tanoak because inoculum levels are higher and infectious periods continue into late spring.     

Phytophthora ramorum and Phytophthora kernoviae on naturally infected asymptomatic foliage

Phytophthora ramorum and Phytophthora kernoviae are recently discovered invasive Phytophthoras causing leaf necrosis and shoot tip dieback mostly on ornamental and forest understorey species, but also cause bleeding cankers on stems of a wide range of tree species. Sporulation occurs only on infected shoots or fruits and foliage so foliar hosts are central to the disease epidemiology. In field trials to assess infection in trap plants exposed to natural inoculum of P. ramorum and P. kernoviae on rhododendron in south west England, it was discovered that leaves of the trap plants ( Rhododendron 'Cunninghams White') and holm oak ( Quercus ilex ) were asymptomatically infected and supported sporulation of both pathogens. More than half the rhododendron trap plants exposed to inoculum of P. kernoviae became infected compared with approximately a third of those exposed to P. ramorum in a natural situation. Approximately one third of the infections were detected from asymptomatic foliage for both pathogens. The significance of these findings for plant health regulation based on visual inspection as a measure to prevent introduction and dissemination of both these pathogens is explored and research gaps identified.     

Forest type influences transmission of Phytophthora ramorum in California oak woodlands

The transmission ecology of Phytophthora ramorum from bay laurel (Umbellularia californica) leaves was compared between mixed-evergreen and redwood forest types throughout winter and summer disease cycles in central, coastal California. In a preliminary multisite study, we found that abscission rates of infected leaves were higher at mixed-evergreen sites. In addition, final infection counts were slightly higher at mixed-evergreen sites or not significantly different than at redwood sites, in part due to competition from other foliar pathogens at redwood sites. In a subsequent, detailed study of paired sites where P. ramorum was the main foliar pathogen, summer survival of P. ramorum in bay laurel leaves was lower in mixed-evergreen forest due to lower recovery from infected attached leaves and higher abscission rates of infected leaves. Onset of inoculum production and new infections of bay laurel leaves occurred later in mixed-evergreen forest. Mean inoculum levels in rainwater and final infection counts on leaves were higher in redwood forest. Based on these two studies, lower summer survival of reservoir inoculum in bay laurel leaves in mixed-evergreen forest may result in delayed onset of both inoculum production and new infections, leading to slower disease progress in the early rainy season compared with redwood forest. Although final infection counts also will depend on other foliar pathogens and disease history, in sites where P. ramorum is the main foliar pathogen, these transmission patterns suggest higher rates of disease spread in redwood forests during rainy seasons of short or average length.     

Insights into Phytophthora ramorum sporulation: epidemiological and evolutionary implications1

Two aspects on the epidemiology of Phytophthora ramorum are discussed. Firstly, in order to elucidate its spatial scale of dispersal, its modes of asexual sporulation in culture and in planta are examined and compared to those of the also aerially dispersed P. infestans. On agar media, P. ramorum sporangia are slightly sticky, tend to aggregate in clusters, and are not shed into the air even when violently shaken or subjected to strong air humidity changes. After inoculating the underside of leaves from an assortment of Mediterranean woody species, no evidence of any adaptations for wind dispersal was observed. On some hosts, it formed sporangiomata (clusters of sporangia) on the upperside. Our observations agree with previous field research in that P. ramorum is rain‐splash rather than wind dispersed, as opposed to P. infestans. Secondly, in Petri dish cultures both species sporulate heavily. However, a steep reduction in sporangial density (sporangia cm^?2) was found on leaves of susceptible Mediterranean species for P. ramorum and a progressive one for P. infestans when this was inoculated on leaves of susceptible potato and other members of the Solanaceae. Furthermore, there was a negative correlation between sporangial density and virulence. We propose a similar pattern of a trade‐off between spore production and virulence to that observed by Thrall & Burdon (2003 ) for the Linum‐Melampsora plant‐pathogen system. Epidemiological and evolutionary implications are discussed.     

Susceptibility of Iberian trees to Phytophthora ramorum and P. cinnamomi

The capacity of Phytophthora ramorum to colonize the inner bark of 18 native and two exotic tree species from the Iberian Peninsula was tested. Living logs were wound-inoculated in a growth chamber with three isolates belonging to the EU1 and two to the NA1 clonal lineages of P. ramorum . Most of the Quercus species ranked as highly susceptible in experiments carried out in summer, with mean lesion areas over 100 cm² in Q. pubescens , Q. pyrenaica , Q. faginea and Q. suber and as large as 273 cm² in Q. canariensis , ca . 40 days after inoculation. Quercus ilex ranked as moderately susceptible to P. ramorum , forming lesions up to 133 cm² (average 17·2 cm²). Pinus halepensis and P. pinea were highly susceptible, exhibiting long, narrow lesions; but three other pine species, P. pinaster , P. nigra and P. sylvestris , were resistant to slightly susceptible. No significant difference in aggressiveness was found between the isolates of P. ramorum . In addition, there was evidence of genetic variation in susceptibility within host populations, and of significant seasonal variation in host susceptibility in some Quercus species. The results suggest a high risk of some Iberian oaks to P. ramorum , especially in forest ecosystems in southwestern Spain, where relict populations of Q. canariensis grow amongst susceptible understory species such as Rhododendron ponticum and Viburnum tinus . One isolate of P. cinnamomi used as positive control in all the inoculations was also highly aggressive to Iberian oaks and Eucalyptus dalrympleana .