RUST DISEASES of APPLE 659 Miles from Redcedars at Altitudes up Irregular in Shape Than the Apple Rust to One-Half Mile
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658 YEARBOOK OF AGRICULTURE 1953 destructive on quince. Its native host plants include hawthorn, Amelanchier, and wild apples. It also infects the fruit of some varieties of apple. The fungus is G. clavipes. The three fungi were studied early in the nineteenth century. At first the forms on the cedar and on the apple Rust Diseases were considered to be different. By 1886, however, more study showed that the spores produced by the fungus on cedar of Apple trees would infect apples and that the spores produced on apples were re- sponsible for new infections on the ce- D, H, Palmiter dar. The rust diseases cannot spread from cedar to cedar nor from apple to Many a fruit-growing district in the apple but must alternate between the East is "a land of red apples and red two hosts. cedars.'' In the Midwest where cedars The fungus that causes cedar-apple did not occur naturally the pioneers rust spends almost 2 years of its life often planted them for windbreaks cycle on the cedar trees. Gedar leaves near orchards. The combination would are infected between July and April be felicitous were it not that some fungi by aeciospores blowm from lesions that require both apples and cedars for develop on apple leaves. Small brown their development. galls, called cedar apples, appear dur- Before our cultivated apples, quince, ing the sumxner but do not mature until and other pome fruits w^ere introduced the following spring, when they may from Europe, three such fungi, natives, be 2 inches in diameter. After a few infected the leaves or fruit of native warm spring rains, about the time crab apples, hawthorn, serviceberry, apple blossoms are in the pink stage, and sorbus, all members of the rose the galls increase in size and extrude family. They could not overwinter on gelatinous tendrils from round de- those hosts, how^ever, and used the ce- pressions on all sides of the galls. The dar trees Juniperus virginianae, J. scopu- tendrils—long, thin, and bright or- ¿orum, J, horizontalis^ and J. communis as ange—form a mass several times that winter hosts. of the original brown galls. The ten- Since three different fungi are re- drils, or horns, consist of many two- sponsible for three different rust dis- celled teliospores, which germinate in eases of apples, we must know some- the tendrils by producing four sporidia thing of their life history and symptoms from each of the two cells. All telio- on both apple and cedar before w^e can spores do not germinate at the same control them. time. With each rain the horns push The best known of the three diseases out farther and expose more spores. is cedar-apple rust. It produces yellow When the supply of teliospores is ex- lesions on the leaves and fruit of wild hausted the galls dry up and may drop. and cultivated apples and is caused by If the twigs bearing the galls are killed, the fungus Gymnosporangium juniper i vir^ the dead galls often remain hanging ginianae, on the cedars through the next year. A similar disease is hawthorn rust. After the small sporidia are pro- Its common native host is hawthorn. duced, a decrease in humidity causes It also infests apple leaves but not the them to be discharged forcibly into the fruit. It is caused by the fungus G. air. Air currents carry them consider- globosum. able distances. Viable spores have been The third disease is quince rust. It is collected by airplane traps several RUST DISEASES OF APPLE 659 miles from redcedars at altitudes up irregular in shape than the apple rust to one-half mile. They may eventually galls. The teliospores are extruded as settle on apple leaves or fruit. If a film dark-orange, wedge-shaped protru- of moisture is present and the temper- sions during spring rains. Sporidia may ature is between 56° and 61° F., the be discharged for 2 or 3 days while sporidia germinate quickly and pene- the galls are drying. Unlike the apple trate the host tissue in i to 3 hours. rust galls, they persist and produce Little germination occurs at tempera- spores again another year. tures below 47° or above 85°. Either The hawthorn rust fungus does not leaf surface may be infected. Fruit le- cause lesions on apple fruit. The lesions sions are usually near the blossom end. on the upper surface of apple leaves The yellow rust spots appear on the are similar to those of the apple rust upper surface of the apple leaves in i fungus but smaller. On the lower leaf to 3 weeks, depending on the tempera- surface, the two fungi look quite dif- ture and the susceptibility of the ferent. With the hawthorn rust the variety. As the spots increase in size, a aecia are relatively few in number and sticky exúdate containing pycniospores are at the center of the lesion and sur- appears. The true function of this rounded by a region of orange-col- spore stage was not known until 1933, ored leaf tissue. The peridial tubes, in when investigations by J. G. Liu at which the spores are formed, are per- the University of Wisconsin showed sistent and long and spread apart to that insects are attracted by this ex- release the aeciospores. They do not údate. They carry spores from one curl back, as do those of the apple rust spot to another and thus fertilize rust fungus. the fungus. Thus it continues growth The aeciospores of the hawthorn rust and completes its life cycle on the are mature in the fall. The wind dis- apple tree by producing the final spore tributes them. They do not germinate stage. well until cold weather approaches. After fertilization, the fungus grows Some spores perhaps do not germinate through the leaf and produces fruiting until the following spring, when new bodies, called aecia, on the lower sur- galls on the cedars are started. If no face of the leaf. Fruit of susceptible apple orchards are nearby or if or- apple varieties also may be infected chards are well sprayed, aeciospores and spores may be produced. Aecio- from infected hawthorn trees may spores are produced in thin-walled serve to reinfect the cedars. tubes. In July and August the spores mature and are released by the split- THE QUINCE RUST fungus infects ting of the tube walls. They are carried twigs, branches, or trunks of the cedar, about by the wind. Those that land on but not the leaves. It produces elon- cedar leaves may germinate at once to gated, swollen, rough cankers, which start galls or may remain dormant turn red as the teliospores begin to until the following spring, when they form in the spring. Jellylike masses of germinate to infect the new growth. yellowish-orange teliospores later are Now that better control methods are exuded during the spring rains about available, few spores are produced on the time apple trees are in bloom. orchard trees. Infection of cedar trees The cankers remain active and in- is largely from spores produced on wild crease in size year after year unless the or seedling apple trees growing near twig or branch dies. Spore forms are the cedars. like (but larger than) those of the other apple rust fungi. Infections of THE LIFE HISTORY of the hawthorn apple leaves result in flecks or abortive rust fungus is about the same as that lesions. The fruit of many apple of the cedar-apple rust fungus. The varieties may be infected, but few if galls produced on cedar trees are more any aecia are produced on most 66o YEARBOOK OF AGRICULTURE 1953 varieties. The fungus sporulatcs abun- county. Complaints of severe damage dantly on quince and hawthorn fruits from rust diseases w^ere also reported and twigs, however. They are covered from Iowa, Wisconsin, and New York. by a mass of bright-orange tubes, in Another serious outbreak occurred in which the spores are formed. The 1927 and 1928. Considerable loss of spores formed on infected quince or fruit occurred in New York and Iowa. hawthorn fruit are then able to Rust diseases injure apples in several reinfect cedar trees. ways. The infected fruit is reduced in Many of the apples infected by grade, so that most of them are culls. quince rust drop in June. Those that Quince rust tends to cause a high mature are misshapen and may have proportion of Mclntosh fruit to drop dark-green sunken lesions, usually in June, and fruit of Cortland and some near the blossom end. Some varieties, other varieties tends to crack. Con- such as Cortland, tend to crack open. siderable loss from secondary rots may Infection becomes visible 2 or 3 weeks follow rust infection. Fruit size is after bloom, and the lesions often reduced by heavy foliage infection, so appear as dark-green raised areas near that even uninfected fruits have little the blossoxTi end. As the fruit grows, the value. Infected trees are greatly weak- diseased parts develop slowly and ened by defoliation and the reduction finally appear as sunken areas. The of active leaf surface. As few as 10 rust tissue under the sunken area is hard spots on a leaf may cause leaves of some and extends to the apple core. varieties to fall by midsummer. Such trees may suffer winter injury and MOST OF THE APPLE plantings in the often fail to set fruit buds for the United States before 1850 were seed- following season. Young trees in ling trees. No two of them were alike.