Smuts That Parasitize Grasses
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28o YEARBOOK OF AGRICULTURE 1953 rusts of importance on grasses that can thus be controlled to any practical degree. Elimination of the alternate host has another advantage: It is on that host that hybridization between strains of rust takes place, often with Smuts That the result of new^ and more virulent strains capable of attacking varieties of grasses and cereals that previously Parasitize w^ere resistant. The development or selection of grasses resistant to rust has lagged Grasses behind such projects in the cereals. Strains have been noted in timothy, orchard grass, crested wheatgrass, blue George W. Fischer bunchgrass, big blucgrass, and others that are resistant to their particular Nearly 140 different species of smuts rusts. It is usually true, however, that attack approximately 300 species of those resistant strains are valid only in grasses in the United States. restricted regions and arc not generally The common name smut derives from resistant to the various and numerous words meaning to besmirch or smudge races or strains of rusts in all localities. and refers to the presence of sooty, The individual State colleges can best dirty, black, or brownish masses on the give advice on such local problems. affected plants. The fact remains that genetic resist- All the smuts are plant parasites. ance does exist in strains of most of Most of them occur on grasses and our forage grasses and can be used cereals. As wdth the rusts, the average in breeding work w^hen necessary or person is much more familiar with the desirable. destruction wrought by smuts on ce- reals than on grasses. Familiar ex- GEORGE W. FISCHER, a native of amples of destructive cereal smuts are Indiana, has degrees from Butler Univer- bunt or stinking smut of wheat, oat sity^ Northwestern University^ and the smuts, barley smuts, sorghum smut, University of Michigan. From ipj6 to and boil smut on corn. In everyday ^945 h^ ^^^ plant pathologist with the practice, the name smut is used to division of forage crops and diseases, designate the disease as well as the Bureau of Plant Industry, Soils, and fungus responsible for the disease, Agricultural Engineering. In ig4^ he although the latter is more correctly became chairman of the department of spoken of as the smut fungus. plant pathology at the State College of The smut fungi have much less com- Washington, plicated life histories than do the rust fungi and generally are easier to con- trol. The familiar dusty black smut masses are made up mostly of millions of tiny cells or groups of cells—spores, w^hich serve the same purpose to the smut fungi as seeds do to the seed- bearing plants, namely, for reproduc- tion and dissemination. Some smuts destroy the flow^ering structure. Others are restricted only to certain parts of it. Some are con- fined almost exclusively to the stems Germinating ergot sclerotia. of grasses. Others produce galls or SMUTS THAT PARASÎTIZE GRASSES 281 tumorlike structures in various parts Some of the smut fungi are not of their host plants. limited to seedling infection. Apparent- As with the rust fungi, many smut ly they can infect any succulent or fungi exhibit a remarkable degree of rapidly growing part of their host specialization not only to certain plants and thereby make them smutty species of plants but also to certain in time. The time between infection varieties or strains within those host and the appearance of smut varies species. Likewise, there are often rather widely. In the stripe smut of strains or races of the smut fungi to grasses, for example, seedlings often contend with. The common stinking show smut within 6 weeks of the time smut, for example, has nearly 30 the smut-contaminated seed is plant- known strains or races, each capable ed. Other smuts are not evident until of attacking different varieties of wheat their hosts head out. One of the stem and different strains or varieties of smuts, to be described later, requires wheatgrasses and related grasses. 2 to 4 years after infection before smut The smut fungi are not entirely actually appears. obligate parasites as are the rust fungi. Spores of the smut fungi generally In fact, some smuts can easily complete are much more durable than are their entire life cycle on an artificial those of the rust fungi. It is hard to medium if it contains the nutrients maintain viability of rust spores for essential for growth. Some smuts more than a year; usually it is much apparently persist indefinitely in the less than that. In the case of the smut soil or in old manure piles. They are fungi, it is unusual for viability to be fully capable, however, of attacking maintained for less than a year. For their host plants when the plants are most smuts viability is maintained for available under the requisite condi- two to several years, especially if tions for infection. humidity is low. Some smuts have The smut fungi have a more adverse been known to retain at least some effect directly (and perhaps indirectly) viability for 25 years. on their hosts than do the rust fungi. The cultivation of grasses seems to The smuts that attack all or parts of have much to do with the develop- the flowering structures generally de- ment of smut in them: Often cultiva- stroy the seeds entirely. The leaf tion practices, including harvesting smuts and the stem smuts, while only and threshing, disperse the smut and occasionally involving the flowering contribute toward an increase in the structures, do nevertheless generally amount of smut in succeeding crops. suppress these structures and likewise Any of the grass smuts therefore is a result in a more or less complete loss potential threat to the welfare of its of seed on affected plants. Thesmuts grass host if the grass comes under that attack the vegetative structures cultivation. (that is, the leaf smuts and stem smuts) Some of the grass smuts are the same have a decidedly weakening effect on as those that are destructive parasites their host plants and make them more of our cereal crops. susceptible to other sinister factors in their environment. SEED SMUTS or bunt {Tilletia species) Many of the grass smuts are seed- are terms used here to designate the borne. The wind carries millions of smuts of grasses in which the normal smut spores, which [)ecome lodged in seeds are replaced by smut ''balls." or on the developing seeds of healthy The balls retain somewhat the shape plants. When the seeds germinate, of the normal seed and are even the smut spores also germinate and encased by the seed wall; actually, infect the young seedlings. Then the though, the inside of the seed is a solid plants that arise from the seedlings are mass of spores. smutted. About 25 of these grass seed smuts 202 YEARBOOK OF AGRICULTURE T953 occur in the United States. Several of the grasses, I)ut only a few pose an have actual or potential importance in economic problem, probably because cultivated grasses. One of the poten- grasses have been cultivated only to a tially most important is the one that on limited extent. wheat is commonly called stinking One of the most common of the smut or bunt. Three species of fungi head smuts of grasses is caused by are involved in it, including what is Ustilago búllala. It complicates the commonly known as dwarf smut. cultivation and production of several These smuts are well known on w^heat of our best forage grasses. More than because of the very extensive losses 60 species of grasses have been re- they have caused for a long time the ported as hosts to it, among them world over. Besides wheat, tall oat- wheatgrasses, bromes, fescues, wild- grass and several of the wheatgrasses ryes, and barley grasses. In 1946 it was are known to be susceptible, including thought that head snuit in mountain crested wheatgrass, slender wheatgrass, brome, one of our most valuable for- and intermediate wheatgrass. Care has age grasses, w^as under control through to l^e taken to keep those grasses from the release of a new variety. Bromar, falling prey to the cereal smuts. which was developed jointly by the The dwarf bunt is soil-borne. It has Department of Agriculture and the thus far presented a knotty problem Washington Agricultural Experiment of control except when resistant vari- Station. In repeated tests the new eties are used. The other two species, strain had remained resistant to the being mostly seed-borne, may be various races of the head smut fungus. controlled by seed treatment and the In 1950, however, an entirely new and use of resistant varieties. very virulent strain of head smut made Some of the i^entgrasses are suscep- its appearance and caused some very tible to another of the seed smuts, badly smutted fields of Bromar moun- especially on the east and west coasts tain brome. There are 12 known races where considerable smut infestation is or strains of the head smut fungus, each vSon"ictimes encountered in harvested of which is specialized to different seed crops. The life history of this smut species of the wheatgrasses, bromes, is not known. wild-ryes, a.nd barley grasses. None of - Various other groups of grasses, for the cereals is susceptible to it. example the bromes, fescues, wild- Several dozen other head smuts of ryes, hairgrasses, velvctgrass, and grasses constitute a potential problem others, are prey to similar seed smuts. among economically important forage or range grasses. Practically nothing is HEAD SMUT (species of Sorosporium^ known concerning the life history and Sphacelotheca, and Ustilago) applies methods of controlling most of them.