HOW INSECTS TRANSMIT VIRUSES 23 Plants Are Grown and Infect Healthy Such Loss and for Many Other Features Plants Subsequently Grown Therein
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22 YEARBOOK OF AGRICULTURE 1953 carried through the winter for seed production. Ehmination of such sources of virus usually gives a high degree of control. The spread of X-disease on peach can be prevented by removing all infected chokecherries within 500 How Insects feet of peach orchards, and the virus diseases of raspberries can be con- trolled, in most instances, by destroy- Transmit ing all wild and escaped brambles in the immediate vicinity of plantings, provided the plandngs themselves are Viruses not already infected. Reducing the population of insect vectors by spraying or by other means L. M, Black has value in the control of some virus diseases. Usually it is not possible, Most viruses that cause plant disease however, to reduce the insect popu- are transmitted by insects, principally lations sufficiently or soon enough to those that have sucking mouth parts— obtain completely satisfactory results. aphids, leaflioppers, white flics, mealy- Some virus diseases can be partly con- bugs, and tingids. Leaf hoppers and trolled by destruction of the hosts of the aphids are the most important. insect vectors. Extensive reduction of Although many plant viruses are the weed hosts of the beet leaf hopper without known insect vectors, it is in the Western States would corre- generally expected that insect carriers spondingly reduce the amount of curly will eventually be discovered for most top virus carried from desert plants to of them. There are exceptions. To- cultivated fields. In much of this area, bacco mosaic virus and potato latent reduction in weed hosts comes about virus are two viruses that occur in naturally under systems of land man- high concentration in infected plants agement in which annual and peren- and are stable enough to be spread nial grasses and other nonhost plants readily from one plant to another by are allowed to replace the weed hosts almost any means that releases juice of the beet leafhopper. Fall spraying from a wound in an infected plant and to kill leaf hoppers on weeds in un- transfers the juice to a fresh wound in a cultivated areas has been resorted to healthy plant. Tobacco mosaic virus is also in the program to control curly thus transferred by the hands of men top. working with tobacco plants even Virus-free nursery stock is extrem.ely though the wounds may be only micro- important in the control of virus scopic in size. It can also be thus trans- diseases of straw^berry and raspberry. ferred by the mouth parts of grass- Natural virus spread often is not ex- hoppers. Potato latent virus can be tensive enough to cause serious dam- transferred from plant to plant when age during the life of plantings started the wind blows the leaves of diseased with virus-free nursery stock. That is plants against healthy ones so as to true also of some of the virus diseases injure both. The mystery about these of tree fruits. two viruses is not their transmission by such methods but why potato latent C. W. BENNETT is a pathologist of the virus is apparently not transmitted by division of sugar plant investigations^ sucking insects and why tobacco Bureau of Plant Industry, Soils, and Agri- mosaic virus is so poorly transmitted. cultural Engineering, who has had more A few viruses, such as wheat rosette than 50 years of experience in the study of virus and lettuce big-vein virus, con- plant viruses and virus diseases. taminate the soil in which diseased HOW INSECTS TRANSMIT VIRUSES 23 plants are grown and infect healthy such loss and for many other features plants subsequently grown therein. of aphid transmission. A virus trans- Just how inoculation takes place in mitted by aphids, particularly if it is such diseases is not known. Dodders of this nonpersistent type, is usually {Cuscuta species), parasitic flowering transmissible by several or, indeed, plants, can transmit plant viruses by many species. The virus of onion yel- means of the natural graft unions they low dwarf can be transmitted by more make with their hosts. Most plant vi- than 50 species of aphids, but not by ruses, however, dcpead on insects for thrips, mites, grasshoppers, beetles, their dispersal. caterpillars, or maggots. That kind of transmission is in con- APHIDS transmit more plant viruses trast to another type in which, follow- than any other group. Aphid-borne ing acquisition of virus, a latent period viruses induce in plants a great variety must elapse before the aphid is able of symptoms, the most important of to transmit. The aphid may then, do which are the mosaics. One of the so for many days without fresh access most efficient aphid vectors is the green to virus. A minority of viruses trans- peach aphid, Myzus persicae^ which mitted by aphids are spread in that transmits more than 50 different plant manner. One of them, the virus of po- viruses. tato leaf roll, is not transmitted by Much remains to be learned about the aphid until 24 to 48 hours have what actually occurs during trans- elapsed after acquisition. The virus mission by aphids. Many aphid vec- may then h^ retained by the insect for tors transmit virus after very brief 7 to 10 days, even through molts, with- feedings on diseased plants. Studies on out fresh access to virus from plants. this type of transmission have reached That an aphid may transmit one a point where the feeding intervals of virus in the persistent manner and individual aphids arc closely observed another in the non persistent from the and timed by a stop watch. For same host plant clearly indicates that example, the vector of beet mosaic persistence or nonpersistence is de- virus requires an acquisition feeding termined by the virus. of only 6 to 10 seconds. During subse- Why do we concern ourselves with quent consecutive inoculation feediiigs such minute details of transmission? of 10 seconds each on healthy plants, Simply because such knowledge may the virus is gradually lost by the aphids make the difl'ercnce between success and fewer than 2 percent of them can and failure in finding a vector of a transmit to more than four plants virus. For instance, no transmission without fresh access to virus. This virus of a certain mosaic virus could be o]> is said to be nonpersistent in the vector. tained after extensive trials with three Usually such a virus is lost more species of aphids when they were fed rapidly from feeding aphids than from one day on diseased plants and then fasting ones—the virus of cucumber transferred to healthy ones. When the mosaic disease, for instance, is lost by aphids were fasted for 30 minutes be- the aphid within 6 to 8 hours when fore an acquisition feeding of 5 to 10 fasting, but within 10 to 20 minutes minutes and were then allowed an when feeding on healthy plants. In inoculation feeding of 5 to 10 minutes, other instances this relation may be however, transmissions were obtained. reversed. The loss of virus from aphids during feeding may be due to a virus- LEAFHOPPERS, next to aphids, are inactivating enzyme secreted by the the most important vectors of plant aphids while feeding but not while viruses. Experiments by a Japanese fasting. Such a substance has not been grower in 1884 demonstrated a con- demonstrated, however, and in reality nection between rice stunt and leaf- we do not know the explanation for hoppers. That might be considered 24 YEARBOOK OF AGRICULTURE 1953 the first virus shown to be insect- viruses multiply to an infective con- transmitted, but actually it was not centration in their vectors. realized until more than 20 years later Although most leafhopper vectors do that the causal agent 01 the disease not transmit virus to their progeny was not the leafhopper but some au- through the egg, certain exceptions tonomous agent carried by the insect. exist. Rice stunt virus and clover club Viruses transmitted by leafhoppers leaf virus may be passed to 95 or 100 cause a variety of symptoms in plants, percent of young insects through the including chlorotic streaking of leaves eggs of the vectors. A single female (as in corn streak), necrosis or death of leafhopper carrying clover club leaf tissues (as in elm phloem necrosis), virus has originated at least 21 genera- tumors (as in wound-tumor disease), tions of infective progeny during a and yellows (as in aster yellows). All 5-year period without fresh access to known vectors for virus diseases with virus from plants. The virus in the a symptom picture like that of aster original female had been diluted yellows arc leafhoppers. at least 100,000,000,000,000,000,- Although many aphid-borne viruses 000,000,000 times. That would be can be transmitted by rubbing leaves impossible had not the virus multi- with juice from diseased plants, only plied in the leafhopper. two leafhopper-borne viruses have There may be, then, two main types been so transferred. In other cases of transmission of virus by leafhoppers. transmission has been accomplished One type, exemplified by curly top only by the use of insects, dodder, or virus, may be characterized by a very grafting. This has accordingly made short incubation period and no multi- the study of the viruses themselves plication in the vector; the other type very diihcult. For such researches it by a long period of incubation and has been necessary to permit the leaf- multiplication in the vector. hoppers to feed on virus solutions Once infective, leafhoppers tend to through membranes or to inject the remain so for many days without fresh virus solution into leafhoppers.