Bulletin STATE OF ILLINOIS DEPARTMENT OF REGISTRATION AND EDUCATION DIVISION OF THE ATURAL HISTORY SURVEY STEPHEN A. FORBES. Chief Vo!' XIII. BULLETIN A rticle IX. OOT-ROT DISEASE OF WHEAT­ H1STORICAL AND BIBLIOGRAPHIC BY F. l.. STEV EN S PR INTED BY AUTHOIUTY OF THE STATE OF:.ILLlNOIS URBANA. ILLINOIS October, 1919 ERRATA Page 97, line 17, for first larval read puital. Page 112, in legend, for jonessi read jonesii. Page 114, in legend, for or read of. Page 125, line 4, for Bonosa read Bonasa. Page 131, in legend, for Jiirundinaceus read hirudinaceus. Page 138, last line, for coccoon read cocoon. Plate XII, explanation page, next to last line, for acrivora read aerivora. Plate XIII, explanation page, next to last line, for WMte-gruhs read White-gruh. Page 293, Figure 5a was reversed in printing, and the two items of the legend should change places. Page 515, second table, for Pelocoris femorata read Pelocoris femoratus. Article IX. — Foot-rot Disease of Wheat—Historical and Biblio- yraphic. By F. L. Stevens. In April, 1919, a serious wheat disease was noted in Madison and other counties in Illinois ; al.?o in several counties in Indiana. Its most constant character is a darkeninjj, and in severe cases a rotting. Fii.. 1. Wheal plants from Madison County, Illinois, showing typical foot-rot. Three of the plants show the char- acteristic development of extra shoots. of the l)asal ]i(irtion of the stem.* By various means—telegram, mimeographed letters and circulars, and public notice— the U. S. De- jiartment of Agriculture, and its agents, have announced the disease as almost certainly "take-all" (180) (181) (182) (183) (184) (LS7). Take-all and diseases of similar character—known also as white heads, * other characters will be discussed in a later paper. 260 black-foot, black-stem, stalk disease, foot disease, root disease, foot-rot, wheat-stem killer, straw blight, pietin, pied noir des cereales, mal del piede, mal do pe do trigo, briseur de chaiimes, brusone, fogheta, pietin du ble, maladie du pied. Fusskrankheit—occur in Italy, France, Germany, Belgium, Australia, Sweden, Finland, United States, Mexico, England, New South Wales, Holland, Russia, Brazil (?), and Japan, and have been attributed to several different fungi, but most prominently to Ophi- obolus. In connection with a study of the Illinois disease it was necessary to glean as many facts as possible from the literature concerning foot- rot, or take-all. and related diseases, and the large amount of material resulting suggested the utility of putting some of it into permanent form as a convenience to students of the subject. The following chrono- logical account presents facts and opinions selected from important articles cited in the appended bibliography. Disease of the nature of foot-rot is said by Mangin (158) to have been known in France as early as 1840. In 1833 (1), 1845 (2), 1856 (3), 1863 (4), 1875 (7), 1884 (11), 1887 (14), and 1897 (44) Ophi- obolus was noted or collected on cereals, not, however, with reference to its pathogenic relation to the plant. The disease is said to have been noted in South Australia in 1852 (84). In 1864 Parliament appropriated £200 for the study of take-all and other cereal diseases (89), and in 1868 a report of a Commission on Diseases of Cereals was published in Australia (5). In this they say of take-all, "Its ravages are irrespective of climatic influence." In 1870 (6) a fungus was observed in the lesions. In 1878 (8), in France, the first definite description of pietin was published, the disease being attributed to an excess of moisture and a mild winter. In 1880 (9) the disease was noted in Italy. In 1883 Saccardo (10) mentions Hendersonia as the conidial stage of Ophi- obolus. In 1884 the first reference in England occurs tmder the name "straw blight" (12). Prillieux and Delacroix (16) in 1890 emphasize the importance of a brown mycelium and attribute the disease to Ophi- obolus graminis Sacc. In 1893 Schribaux (19) noted that early varie- ties are most susceptible, and tried various control measures. Loverdo (20) reports the disease on Cynodon and Agrostis. Cobb, in 1892 (22), in a rather extensive article, discusses the dis- ease knovi'n as take-all as found in Australia and attributes it to Clado- sporium herbarum. He advises rotation, drainage, good tillage, burning stubble, seed treatment, and resistant varieties. The disease was also noted as occurring in oats. Frank (23) (24) (35) (36) (27) in 1894 discusses foot-rot in Germany, as does also Hiltner (28) (29). In 1895 Saccardo and Berlese (34) note foot-rot in Italy and attribute it to Sphae- roderma damnosum. Frank in 1895 (30) notes Leptosphaeria hcrpo- trichoides on rye and Ophioholns herpotriclius on wheat. The mycelium grows into the roots, which become black and die. No conidial form was observed. In 1897 (43) the disease was noted on wheat, rye, and barley in France. Prillieux {4:2) mentions the mycelium in the vessels, also in a brown surface development in flecks. AIcAlpine in 1898 (46) lists nurnerous fungi on diseased wheat, among them being Cladosporium hcrbaruin, Macrosporium graminum and M. ccreaUum, and Epicoccum neglectum. He refers to Dr. Cobb's conclusion and says "It is premature to speak of it {Cladosporium hcrbariim] as the take-all fungus." In 1898, 1899, and 190:2 Mangin (47) (56) (70) contends that Leptosphaeria is the main cause of the disease and that Ophiobolus is secondary, as are also several other fungi which were present. Inocu- lations with spores in w-ater suspension gave positive results. Frank in 1900 (64) describes Ophiobolus herpotrichus as the cause of the disease, while Delacroix (67) in 1901 asserts, on the basis of inoculation ex- periments, that both Leptosphaeria and Ophiobolus are parasitic, the disease a])j)earing three or four months after inoculation. In 1901 Ophi- obolus was first recorded in Australia (68). In 1902 Mangin (70) con- tends that Leptosphaeria is the primary cause and Ophiobolus of sec- ondary importance. In 1903 Cordley (75), in Oregon, notes a disease which from the description is indistinguishable from that now in Illinois. In 1904 Mc- Alpine (84) claims to prove that Ophiobolus is the cause of take-all, since wheat planted in plots with diseased stubble became diseased and bore spores of Ophiobolus within fifty-two days. It is to be noted that McAlpine did not employ pure cultures, and it is not clear how he recog- nized with certainty "the young mycelium of Ophiobolus." In 1906 take-all was noted on barley and other grasses in Australia (89). Kirch- ner (90) holds that the disease follows weakening of the plant from other causes. Robinson, 1907 (03), notes perithecia in the sheaths, and sa3'S that wheat sown in diseased stubble soon becomes sick and in two months may all be dead and covered with perithecia. In 1908 Krilger ( 95 ) holds that fungi are not the immediate cause but that the disease is due to weakening of the plant. Richardson, in 1910 (111), in Austra- lia, found upon close examination that the Ophiobolus vegetative myce- lium was present in every case. "Swollen roots", he says, "were im- pregnated with a black sooty incrustation." He also notes the disease on Hordeum, and says that a good stubble-burn gives good crops follow- ing. Pridham, 1911 (112) describes the disease as negligible in favor- able years, and adds that in all cases of disease there is a blackening of the stalk at the foot. Fron, in 1913 (120), notes the presence of felt- like brown masses and finds Leptosphaeria on all specimens ; Ophiobolus, rarely. From infection experiments he concludes that Leptosphaeria is the cause. Eriksson, in 1912 (121), refers to Coniosporium as the conidial stage of the causal fungus. Averna-Sacca (124) writes in Brazil of the disease. Stormer and Kleine in 1912 (129) sav the disease can also be caused by Typhula. Voges in 1912 (130) says that fungi are not the primary cause, but that the disease is due to lowering of the vitality of the wheat plant bv environmental conditions. SchafFnit in 1912 (131) attributes a foot-rot of rye to Fusarium nivalc. 362 Massee, 1912 (137), say; that the mycelium forms a thin felt on the stem and sheaths and that a black superficial mycelium is also char- acteristic. Oats and wheat inoculated when about an inch high with Ophiobolns graminis were yellowed and drooping in six weeks, with characteristic mycelial infection. Bromus and Agropyron are also in- fected. Prunet, 1913 (138), concludes that either Ophiobolus graminis, O. hcrpotrichus, or Leptosphaeria herpotrichoides can cause the disease, and says that the name pietin is thus applied to three diseases with the common character of stem-blackening, and that each should be studied separately. In 1913 Guerrapain and Demolon (139) (110) call particular at- tention to this disease following a mild winter. Voges, 1913 (143), in a very extensive article, concludes that the Ophiobolus ascospores can give rise to two kinds of mycelium, one of which is Fiisarium rubigino- sum A. & W. ; that the Ophiobolus mycelium is not characteristic of foot disease and that the specific cause is not Ophiobolus ; and that the dis- ease has various causes, for example frost, the Ophiobolus appearing only after weakening due to other causes. Mangin, 1911 (158), says that both Leptosphaeria and Ophiobolus give a brown or black tint to the stem and roots. The presence of several Fusaria is also noted in connec- tion with this disease. Berthault, 1914 (161), says that Fusarium riibi- giiiusiiui is often present and perhaps causal. Voges, 1914 (162), says that the conidial form is not Fusarium but is Acremonium. Spafiford, 1917 (175), suggests the name "wheat stem killer", and says that the disease may be recognized at the base of the stem, which is "covered with a brownish powder or stained brown." He regards this character as im- portant in distinguishing this disease from the results of climatic action due to drying out, in which cases the plant is yellow throughout. He states that if wheat gets a good start, even in infected soil, but little injury follows. In 1917 (176) Ophiobolus graminis was reported on rice in Japan. In ]919 Brittlebank (179) lists as hosts, wheat, oats, barley, Hordeum niurinum, Bromus niolUs, B. sterilis, and Agropyron scabrnm. In 1919 the latest paper available from Australia (17S) is less definite as to etiology than are earlier Australian papers. In the voluminous mass of evidence presented in these articles cer- tain points stand out clear, others are far from clear. Points of Agreement 1. A serious loss to the wheat crop in many countries arises from a diseased condition of the wheat plant which is aptly called foot-rot, black-foot, or black-stem, or, by the French, pietin, though known by many other names. 2. This disease is characterized by a darkening, blackening, or rot- ting of the lower stem, usually the first internode above the roots. 3. The disease always occurs in spots in the field. Beyond these three facts there is practically Httle or nothing in which there is entire accord. Points oi- Disagreement These may be summarized as follows : Symptoms.—Many observers describe as an essential symptom of foot-rot the presence of a brozvn felt-like mycelial zvcjt on the diseased stem and adjacent sheath. Sonie describe it as a black crust. Others fail to find this, or at least do not mention it, while still others definitely assert that it is not an essential symptom. Some describe the roots as swollen, blackened, or stunted, though most writers mention no root symptoms. Most writers assert that if land on which the disease occurs is cropped with wheat the following year the disease will again appear, and often on a largely increased area. Some show by experiment that wheat planted in diseased stubble will become diseased ; others say that succeeding crops may be healthy, and that the recurrence of the disease is dependent upoti various factors, chief among which is the weather. Etiology.—Most writers on foot-rot say that the causal fungus is Ophiobolii.s graminis; a smaller number that it is O. Iicrpotrichus. Some merely report the presence of one or both of these fungi, while other in- vestigators claim to have demonstrated their actual parasitism by inocu- lations with pure or nearly pure cultures of one or another of these fungi. Other writers claim that Leptosphaeria hcrpotrichoides, Typhula graminis, and Cladosporiiim hcrbanmi are causal agents of foot-rot, while some assert that any one of several of the five fungi mentioned above may be the cause of the disease. Some hold that these fungi are saprophytes or but weak parasites that can attack the wheat plant only when it is weakened by other causes, chiefly by cold. Conidial stages of Ophiobohis.—Various claims have been made as to the conidial form of Ophiobolus. Some state that the fungus is usually sterile in its young stages. Some find as the conidial form a Coniosporium, an Acremonium, a Fusarium, or a Hendersonia. Various prophylactic measures, as burning stubble, rotation, good culture, etc., have been suggested, but they are based on general grounds rather than specific knowledge of this disease, and will not be discussed. In view of these conflicting observations and conclusions the most reasonable attitude is that of Prunet, Voges, and others, namely, that we have here several distinct diseases due to as many separate causes. Clearly science will be best served by such a conception. So far as I can judge, the disease now present in Madison county, Illinois, is indistinguishaJDle from that descriljed by Cordley in Oregon, U. S. A., in 1902 (75), the cause of which he did not determine. I am not yet sure what the cause of the Madison county disease really is, but 264 as to the three points concerning which we have noted universal agree- ment it is no exception to the rule. I do not find in it the brown felt de- scribed by so many as characteristic of Ophiobolus, nor do I find con- stantly present any of the fungi mentioned above as the cause of foot- rot. Further long and patient investigation must determine which foot- rot, or which of its several types, is now interesting us in Illinois—if indeed it is any one of those yet studied. The name take-all is linked in the literature with Ophiobolus, and if the disease in Illinois is caused by Ophiobolus this name may be used properly in connection with it, but unless this is the case I prefer to employ the more comprehensive general name, foot-rot. Until definite knowledge is secured as to the cause of the disease it is impossible to predict its development, or to state whether it will recur in the same fields and spread to others. If it is identical with the Oregon disease which appeared in 1902 and apparently has attracted no attention since then ('75), it may here lapse into a similar insignificant position. It should be borne in mind that the winter of 1918-19 was a very excep- tional one and of the type that is most favorable to foot-rot in Europe, and that in the absence of such winters the foot-rot may do little harm.* In our present state of ignorance as to the precise cause of this disease in Illinois it is impossible to give specific advice as to its preven- tion. A wise course, however, will be to avoid planting wheat for one year or, better, for two years, on lands where the disease occurred this year. Oats may probably be safely substituted. Burning over the stubble after harvest will also be a safe precaution. Annotated Bibliography explanatory The asterisks used in connection with the entries indicate that the articles or books or pertinent passages cited have been carefully read in the original. My acquaintance with those not so marked has been gained through reviews or abstracts. The notes do not indicate the entire con- tent of the articles, but call attention to points most important in con- nection with the Illinois disease or in the general history of foot-rot. Information given in the preceding pages is not, as a rule, repeated in the notes. Quite a number of the entries lack the author's name, since they either appeared anonymously or in secretarial or journal reports. Other entries are without title because they so appeared in foot-notes of articles read. It seemed ill-advised to omit them, though I was unable to verify the citations. For the sake of brevity the abbreviation E. S. R. is used to indicate the "Experiment Station Record" published by the U. S. * Since the above was written. Dr. A. D. Cotton, of the Pathological Laboratory of the Kew Botanic Gardens, has examined the foot-rot in Illinois, and he tells me that the specimens he examined lacked the distinctive characters of take-all as that disease is understood in England. 265 Department of Agriculture; and Zeit. f. Pfk., for Sorauer's "Zeitschrift fiir Pllanzenkrankheiten." ACKNOWLEDGMENTS I am indebted to Miss C. R. Bemiett, of the Library of the U. S. Department of Agricuhure, for four titles which I should not have had but for her courtesy ; to Mr. P. L. Windsor, Librarian of the University of Illinois, who secured for me very many original articles ; and to Mrs. E. Young True for assistance in the preparation of the manuscript. Fries, E. *(1) 1823. Systema mycologicum, 2:504. Hyhavria herpotricha listed. Mougeot, A. (2) 18-15. Champignons des Vosges, Epinal, p. 102. Marchand. (3) 1856. Sylloge—p. 251. Tulasne, L. and C. *(1) 1863. Selecta fungorum carpologia, 2:355. Paris. Mentions Opliiobolus under . name Rhaphidophora herpo- tricha. Commission on Diseases of Cereals (5) 1868. Report on diseases of cereals. Adelaide, S. Austral. Muecke, Carl (6) ISTO. The take-all {Xenodochus cerealimn). Prize essay, published under the authority of the Board of Agriculture of Victoria. Observed a fungus in the disease lesions. Estimated loss in South Australia at £6,000,000. Saccardo, P. A. (7) 1875. Fungi Veniti novi vel critici. Nuovo Giorn. Bot. Ital. Vol. 7. (S) 1878. Sur I'etat de la recolte du ble et sur la maladie du pietin. Bui. des seances de la Soc. Centrale d'Agr. de France, 38:368. First adequate description of the disease; "la cause du mal soit generalement attribuee a I'humidite." Cugini (9) 1880. Supra una malattia del frumento recentmente composa nella Provincia di Bologna. Giorn. Agr. Ital. 14 : No. 13. Noted the disease as serious near Bologna. 266 Saccardo, P. A. *(10) 1883. Sylloge fungorum omnium hucusque cognitorum, 2:352. Says conidial form of Ophiobolus herpotrichus is a Hender- soaia. Brunaud (11) 1884. Description des Sphaeriacees des environs de Saintes, La Rochelle, p. 213. Smith, W. G. *(12) 1884. Diseases of field and garden crops, p. 69. DescrHjes lesion at base of the stem and fungus present there. No spores known. Morini (13) 1886. . Nuova Giorn. Bot. Ital. 18:32. Noted Oiihiobolus Tierpotrichus, but regards it as a sapro- phyte. Winter, G. *(14) 1887. Die Pilze Deutschlands, Oesterreichs und der Schweiz. Rabenhorst's Kryptogamen-Flora von Deutschland, Oes- terreich und der Schweiz, 1 (Abth. 2) : 523, 524. Pearson, A. N. *(15) 1888. Blight in wheat. Dept. Agr. Victoria, Bui. 1:38. Says there is a general consensus of opinion that this is a poverty disease, and that the fungus which causes talte-all attacks mainly such crops as are Insufficiently nourished. Prillieux and Delacroix *(16) 1890. La maladie du pied du ble causee par VOphiobolus graminis Sacc. Bui. Soc. Myc. de France, G:110. Authors note brown mycelium in flecks; perithecia in winter; and mycelium in all browned tissues. Cugini (17) 1890. . . Bol. Staz. Agr. Modena, 9:46. McAlpine, D. (18) 1891. . Dept. Agr. Victoria, Bui. 14. Cited by Mc- Alpine in (46). Schribaux, E. *(19) 1892. Le pietin ou maladie du pied des cereales. Jour. d'Agr. Prat. 2 : 317-320. Accepts Ophiobolus as the cause of the disease. Loverdo, J. *(20) 1892. Les maladies cryptogamiques des cereales, p. 231, 236. Reports Ophiobolus herpotrichus on Cynodon and' Agrostis. Tepper, J. G. O. *(21) 1892. "Take-all" and its remedies. Agr. Gaz. N. S. Wales, 3 : 69-72. Says that "Take-all is nothing else than starvation of the crop." 267 Cobb, N. A. *(22) 1892. Plant diseases and how to prevent them. Agr. Gaz. N. S. Wales, 3 : 991-1001. Notes false theories as to take-all: due to insects, weeds, nematodes, soil deficiency, poor drainage, poor tillage. Char- acters: occurs in patches; plants in a majority of cases dry up when young; much Chidospoi-ium herlxinnii and i