can serve as alternate hosts for Cronartium ribicola , the causal agent of white pine blister rust (WPBR). This disease was introduced into North America over 100 years ago and has caused major mortality to native five-needle pines. Once the Ribes
Todd A. Burnes, Robert A. Blanchette, Jason A. Smith, and James J. Luby
Bohun B. Kinloch Jr.
After a century since introduction to North America from Europe, white pine blister rust, caused by Cronartium ribicola J.C. Fisch., is recognized as one of the catastrophic plant disease epidemics in history. It has not yet stabilized and continues to spread and intensify. Its nine native white pine hosts comprise major timber producers, important watershed protectors, keystone ecological species, and the oldest trees on earth. All are highly susceptible and some have been damaged severely in parts of their native range, as well as where they have been planted as exotics. Resistance, the most promising approach to control, requires understanding of genetic interactions between hosts and pathogen, a quest that has been ongoing for half a century. Unlike other hosts of spectacular exotic diseases, such as chestnut blight [caused by Cryphonectria parasitica (Murrill) M.E. Barr] and dutch elm disease [caused by Ophiostoma ulmi (Buisman) Nannf.], white pines (Pinus L.) exhibit a surprising number of resistance mechanisms to blister rust, if at only low frequencies. There are three main kinds:
Kim E. Hummer and Stan Pluta
In the late 1800s a European disease called white pine blister rust, Cronartia ribicola Fisher, was introduced into the United States. By 1937 this disease had naturalized and was firmly established in native Ribes across the country. White pine blister rust causes economic damage to white pines and infects leaves of some Ribes late in the summer after harvest. Ribes serve as obligate alternate hosts for this disease. Our objective was to determine which Ribes species were susceptible to white pine blister rust under field conditions in Corvallis, Ore., where inoculum is naturally present. In 1995 and 1996, 57 Ribes taxa from North and South America, Europe, and Asia, were evaluated in mid-August and mid-September for presence of white pine blister rust. Susceptibility was determined by the rust infection of the abaxial leaf surfaces. Rust infection was rated on a scale from 1, no infection observed, to 9, severe infection covering almost the entire surface of at least three or more leaves. Data from 1995 indicated that 22 Ribes taxa were susceptible to white pine blister rust, while 35 others had no infection. The 1996 data will be reported. Species without infection may offer resistance genes to breeders who wish to develop rust-resistant commercial fruit cultivars.
Recent interest in expanding commercial currant and gooseberry (Ribes L.) plantings in the United States has put pressure on the states with Ribes restrictions to review their regulations. A meeting on 9 January 1998 initiated discussion between the state agriculture regulatory agencies, forest pathologists, and horticulturists. Since then a white pine blister rust (WPBR), Cronartium ribicola J.C. Fischer) World Wide Web (Web) site (McKay, 1998) and list serve have been activated to facilitate communication. Vermont is a state that has no regulations on the books at this time. Connecticut and New York also have mentioned that infection rates are low. Maine retains a Ribes reduction program, and Massachusetts is strictly enforcing their regulations. The following summarizes the general consensus among the majority of regulating states: 1) It is desirable to find a way for both white pines (Pinus L.) and commercial Ribes plantings to coexist. 2) More research is needed to survey existing Ribes and pines, the potential impact of commercial plantings versus the impact of existing Ribes, and the potential impact of escape /volunteer seedlings from immune Ribes cultivars. 3) There is interest in permitting immune Ribes cultivars to be planted. 4) There is interest in having consistency in regulations from state to state.
Richard A. Sniezko, Andrew Bower, and Jude Danielson
help with the rust assessment in summer of 1999. The cost of publishing this paper was defrayed in part by the payment of page charges. Under postal regulations, this paper therefore must be hereby marked advertisement solely to indicate this fact.
Stanislaw Pluta and Agata Broniarek-Niemiec
Field resistance to white pine blister rust (WPBR) (Cronartium ribicola J. C. Fischer) was investigated on 53 black currant (Ribes nigrum L.) genotypes (cultivars and breeding selections) in 1998 and 1999. Uredia did not form on the black currant `Titania' and 17 advanced selections during field evaluations made at the Experimental Orchard at Dabrowice, near Skierniewice, Poland.
R.S. Hunt and G.D. Jensen
For the white pine blister rust disease (WPBR), reports conflict concerning the time of year the pathogen, Cronartium ribicola J.C. Fisch., infects western white pine (Pinus monticola D. Don) and what needle age increments are most susceptible. To determine timing of infection, western white pine seedling were placed under infected currants (Ribes nigrum L.) for 1-week periods from May to November. Needles became spotted and stems cankered after exposure to diseased currants from early summer until leaf drop in November. To determine what foliage age increment was most susceptible, 5-year-old seedlings were placed in a disease garden, and older trees were inoculated in situ. All age increments of pine foliage were susceptible to infection. For young seedlings, all age increments were about equally susceptible, but on some older seedlings and trees, the current year's foliage appeared more resistant than older foliage.
John A. Muir and Richard S. Hunt
Introductions of white pine blister rust (WPBR, causal fungus: Cronartium ribicola J.C. Fischer) to eastern and western North America before 1915 caused such extensive damage that western white pine (Pinus monticola D. Don) was essentially abandoned as a manageable forest tree species for over 60 years. Recent results from WPBR resistance selection and breeding programs, and from field trials of tree spacing, pruning and bark excision treatments have supported efforts to increase establishment and to intensively manage western white pine. Western white pine is a desirable component in many forested areas because of its faster growth and much higher value compared to many other associated tree species. It also has a low susceptibility to armillaria root disease caused by Armillaria ostoyae (Romagnesi) Herink and laminated root rot, caused by Phellinus weirii (Murr.) Gilb. Some regulations, e.g., Forest Practices Code of British Columbia (BC) Act, require anyone who harvests timber on provincial forestland and uses western white pine for reforestation to either plant genetically resistant western white pine stock or prune susceptible young trees for protection. Risks of increased WPBR associated with increased commercial cultivation of gooseberries and currants (Ribes L.) have yet to be determined. However, major threats appear to include 1) increase in local amounts of spores for nearby infection of pines; and 2) possible introductions or development of new, virulent races of C. ribicola, particularly from eastern to Pacific northwestern North America. In view of these possible threats, we recommend that existing regulations and legislation should be amended, or possibly new measures enacted, to permit propagation and commercial cultivation only of varieties of Ribes that are immune or highly resistant to WPBR.
Kim E. Hummer
The center of diversity for white pine blister rust (WPBR) (Cronartium ribicola J.C. Fischer) most likely stretches from central Siberia east of the Ural Mountains to Asia, possibly bounded by the Himalayas to the south. The alternate hosts for WPBR, Asian five-needled pines (Pinus L.) and Ribes L. native to that region have developed WPBR resistance. Because the dispersal of C. ribicola to Europe and North America occurred within the last several hundred years, the North American five-needled white pines, Pinus subsections, Strobus and Parya, had no previous selection pressure to develop resistance. Establishment of WPBR in North American resulted when plants were transported both ways across the Atlantic Ocean. In 1705, Lord Weymouth had white pine (P. strobis L.), also called weymouth pine in Europe, seed and seedlings brought to England. These trees were planted throughout eastern Europe. In the mid-1800s, WPBR outbreaks were reported in Ribes and then in white pines in eastern Europe. The pathogen may have been brought to Europe on an infected pine from Russia. In the late 1800s American nurserymen, unaware of the European rust incidence, imported many infected white pine seedlings from France and Germany for reforestation efforts. By 1914, rust-infected white pine nursery stock was imported into Connecticut, Indiana, Massachusetts, Minnesota, New Hampshire, Ohio, Pennsylvania, Vermont, and Wisconsin, and in the Canadian provinces of Ontario, Quebec, and British Columbia. The range of WPBR is established in eastern North America and the Pacific Northwest. New infection sites in Nevada, South Dakota, New Mexico and Colorado have been observed during the 1990s.
Geral I. McDonald
Frequency of infection, main effects and interactions among four geographic sources of white pine blister rust (WPBR) (Cronartium ribicola J.C. Fisch.) and 10 Ribes-sites (Ribes L. specie × site combinations) was investigated using leaf disk assay. Two clones, one of R. hudsonianum (Dougl.) Jancz. and one of R. viscosissimum Prush., were not infected by any sources of WPBR. One clone of R. viscosissimum that was not infected by two sources of WPBR was susceptible to the other sources. Highly significant WPBR sourc × Ribes-site analysis of variance interaction for incubation period and infection efficiency precluded testing main effects. Profile plots of incubation period interaction means showed orderly interaction by all WPBR sources and plots of infection efficiency showed that aggressively virulent WPBR from Oregon (Champion Mine) ranked near or at the bottom for infection efficiency for all Ribes-sites. Meanwhile, aggressively virulent WPBR from Idaho (Merry Creek) ranked near the bottom for infection efficiency when inoculated onto Ribes obtained from the Cascade Mountains but switched to the highest ranking when inoculated onto Ribes obtained from Idaho. Geographic interaction of white pine blister rust and Ribes for incubation period and infection efficiency may help to explain geographic patterns recently observed in WPBR molecular markers.