Cornell University Library OF THE Hew Work State College of Agriculture Ie 32342 =e 1820 SB 624.0384 ‘nui Cornell University Library The original of this book is in the Cornell University Library. There are no known copyright restrictions in the United States on the use of the text. http://www. archive.org/details/cu31924003214529 BULLETIN OF THE Ohio Agricultural Experiment Station NoumBer 214 ‘Marcu, 1910. A BRIEF HANDBOOK OF THE DISEASES OF ' CULTIVATED PLANTS IN OHIO By A. D. SELBY INTRODUCTION The idea of disease is not a simple one, though it may seem so before trying to define it. In reality the term “‘disease’’ as applied to plants, means any change in that plant toward reduced vigor, etc., from the ordinary or average behavior. To put it another way, a plant is said to be ‘‘diseased’”’ when it shows any deviation from the ordinary or average behavior of that plant in respect to appearance, growth, color of bark, foliage, fruitfulness, time of dropping leaves or length of life; in short, when the plant fails to conform to those averages which we have established by extended observation for the species and variety in question, we say it is diseased. Under such a general definition, variegated or purple hued sports would be included, although potentially rather than actually in diminished -vigor. Variegated sports succumb easily to parasitic attack and, as later investigations show, are really suffering from enzymatic troubles. The more usual symptoms of disease are marked by evident differences in the plant. The leaves become spotted, curled or discolored, or may even drop prematurely; the fruit may develop unevenly or be marked by decayed spots, or the twigs may blight, wilt or die. In all such cases we have a manifest loss of vigor and reduced profit. Yet we may not attribute all these to parasitic fungi or to parasitic insects; purely physical or chemical agencies may be at the bottom of certain troubles. Plants may be as- phyxiated by too much water which excludes the air supply; they may likewise, be strangled by escaping gases, especially in the case of city shade trees, or their protoplasm may be attacked by chemi- (307) vy 308 OHIO EXPERIMENT STATION: BULLETIN 214 cal agents such as strong acids and alkalis. Quick growing plants appear to fall in drought, as with cucumbers when started during a period of excessive rains. Plants, and especially trees, may be locally injured by winter freezing, by hail, by overbearing with ex- haustion of water supply, and by a variety of causes. While we must keep our minds open to these varying causes of impaired vigor, by far the larger number of the diseases described in this bulletin are directly attributable to parasitic fungi which attack the plant or host in some vital part and rob it of its substance. The conditions of injury arising from the attacks of insects alone are not included. These fungus parasites of particular plants are of differing sorts, which produce, each, its more or less particular effects. It must follow, therefore, that the diseases produced differ in nature and that the names applied will vary accordingly. The names are not simply blight, rust, etc., indiscriminately applied—they are given with reference both to the parasite and its effect on the host plant.* Parasitic fungi and bacteria which cause disease, being plants, though of lower class, have differences among themselves which may be clearly designated and defined. ‘The names applied to them are accompanied by specific and generic descriptions which mark off the sort as definitely as do the descriptions on higher plants such as ferns, flowering plants and trees. The extreme minuteness of the parts of parasitic fungi and bacteria make necessary the use of the microscope in their description and detection. ‘The parts called spores which reproduce these minute plants have special form, size, etc., by which these are recognized when found. The agencies for the spread of parasitic diseases are those operations in which we engage or those which surround and envelop the plantsas well as ourselves. Light spores will be carried by currents of air like particles of dust. Allspores or germs of these lower plants may be carried by numerous agencies such as insects, higher animals, and man. They willalso find entrance into plants by whatever openings exist at the time. The epidermis of a green leaf or stem has breathing pores or stomates init; the leaves of mustard plants have water pores in them and wounded plants have these freshopenings to invite the entrance of the disease conveying spores or germs. The remedies for plant diseases are based upon the character and life history of the particular parasitic growth with which we have to deal and upon the nature of the host plant itself—some hosts being very different from others in respect to permitting of sprays of fungicides or insecticides. Common sense inferences are * See naming of diseases. DISEASES OF CULTIVATED PLANTS 309 always of use in dealing with plant diseases. Ifthe soil is too wet, drain it; if late growth predisposes to winter injury, avoid such growth; if overbearing weakens plants, prevent, it by thinning the fruit. The philosophy of seed treatments is stated under diseases which infest the seed; that of soil treatments or disinfection, under soil infesting disease, and the general doctrines of sprays, fungi- cides, etc., under that heading further on. The progress made in plant disease prevention throughout the world during the period of about 26 years which has elapsed since the discovery of Bordeaux mixture in France shows how well adapted that discovery was to the needs of the times. The progress made in recent years in the study and control of plant diseases has been made possible by the agencies recently developed in the United States in the Agricultural Colleges, the Agricultural Experiment Stations and the United States Depart- ment of Agriculture. It is not expected that this advance in our knowledge of the diseases of plants or of the methods of disease control will soon wane. Efforts like the present one to present briefly the doctrines of disease and the philosophy of disease con- trol together with brief descriptions of prevailing diseases in our state, have for their purpose the wider dissemination of the body of present day knowledge int’ ¢ lines. Such a statement will not close the march of progr nor make less the need for more knowledge. Itis hopedt ‘cultivators of plants, whether farmers, gardeners, horticulturists or florists will find suggestive statements of information in the bulletin by which they can direct their own efforts to better advantage and correct or broaden their own in- ferences from observed conditions about them. All such results will not only increase the need for more knowledge, but will furnish impetus to the movements by which we will gain the desired information. In the preparation of the revised edition of the original Bulletin, No. 121, the general part immediately following this introduction has been considerably enlarged and brief discussions are now given concerning groups of plant diseases as well as those concerning parasitic fungi. It is fully apprehended that the host plant is the center of practical as wellas economic interest and these statements concerning enzymatic diseases as in the case of peach yellows and mosaic disease of tobacco, diseases transmitted in the seed, soil infesting diseases, and the relation of the spread of certain diseases to leaf biting insects are given as aids in mastering the principles involved. ‘The same aim has governed the discussions upon wounds 310 OHIO EXPERIMENT STATION: BULLETIN 214 and wound fungi so especially dangerous with orchard, shade and forest trees. Somewhat fuller discussion of atmospheric agencies as affecting the occurrence and spread of plant diseases, of remedies for diseased conditions and of the application of the latter in combatting diseases and a presentation of storage troubles has also seemed desirable. Special attention is called tothe host plant in the matter of breeding or selection for disease resistance and in the contrasts offered by American and European points of view in plant disease study. ACKNOWLEDGMENTS The illustrations in this bulletin have been drawn from wider sources than in the previous hand book. A large number, including perhaps, a larger portion of the cuts, are taken from previous pub- lications of this Station by Weed, Miss Detmers, and the writer; small cuts have been at times made from certain larger illustrations while with others only portions of the original cut have been used. A great many of the illustrations are new, and I am deeply in- debted to Messrs. J. M. Van Hook and Thos. F. Manns for many of the photographs from which these have been made. I am also -in- debted to Professors Halsted of New Brunswick, N. J., and Atkin- son of Ithaca, N. Y., and to the Bureau of Plant Industry of the U.S. Department of Agriculture for many favors in the matter of cuts which are used in the Bulletin. Figures 24, 25, and 26 are from Dr. Freeman’s ‘‘Minnesota Plant Diseases.” For permission to reproduce these, Iam indebted to Prof. Frederick E. Clements, of ‘the University of Minnesota. In all cases where it is not otherwise obvious, it has been the aim to state the source of the illustrations in the ‘descriptions. The same applies to illustrations reproduced from standard works. In many matters connected with recent investigations of the Department bearing upon diseases included in this present bulletin, and upon current examinations, I am under many obligations to Thos. F. Manns, Assistant Botanist, who has rendered very great assistance. GENERAL PART I CONCERNING PLANT DISEASES IN GENERAL As defined in the introduction, a plant is called diseased when it fails to show normal vigor and normal condition of its parts. ‘The manner of disease attack is extremely varied and the conditions set up as a result of disease are accordingly of many different kinds. We learn to recognize disease by the symptoms shown in the plant; these symptoms will at times be readily interpreted and on other occasions they will prove misleading. Nothing is plainer than the necessity for continuous observation of growing plants if one is to be in a position to interpret the symptoms of disease. Fig. 1. Roots of white burley tobacco plant attacked by broom-rape. Each of these masses attached to the root shows beginning of the plant which will grow upin larger dense form, and pro- duce an abundance of blossoms and seeds but no leaves. Each one of these must have started from a buried seed of the broom-rape, Orobanche Ludoviciana Nutt. Diseased conditions may be due to the very obvious attacks of certain parasitic seed plants which lack leaf-green or chlorophyll'in their tissues and must subsist on other plants somewhat after the manner of parasitic fungi. The dodders which attack the clovers, alfalfa, onions, etc., belong in the class of parasitic seed plants of the genus Cuscuta. Their seeds are liable to be harvested with the , (311) 312 OHIO EXPERIMENT STATION: BULLETIN 214 seeds of clover or alfalfa and to be present in the commercial seeds. While these have been treated in the weed manual they require mention here. The seedling plant of dodder first forms a root and sends upward a whitish stem which twines about the clover or other stem, and sends sucking branches into the stem interior. ‘These “‘haustoria’ extract food material from the clover stem—that is they rob it of its own substance. Upon the formation of such organs the root of the dodder dies off and the future ex- istence of these twining, strawlike stems is at the expense of the host plant. A similar state of parasitic existence is found in the broomrape tribe whose very small seeds are scattered through the soil. Such a broomrape is well known on hemp, and the same hemp broomrape also attacks tobacco in Kentucky and possibly in our state. We have found another broomrape attacking tobacco in one district of Brown county, Ohio, and the illustration shows its appearance on the tobacco roots. When the leaves of a plant are attacked these show the direct effects; the symptoms of parasitic leaf diseases are usually localized injury resulting in spotting and often in browning of the leaf parts. Leaves may dry up somewhat slowly and drop to the earth, and yet the leaf tissues are simply dried up. Such conditions may result from late frost as upon shade, fruit, or ornamental trees. A most interesting case was once studied upon catalpa as a result of a frost in May. In that case the drying up was none the less to be expected at that time. An even more interesting case of leaf drying and dropping was upon young catalpa trees ina nursery caused by the attacks of a root-rot fungus, Zhzelavia. Owing to the death of many of the rootlets and finer roots as a result of the root-rot trouble, the leaves of these young trees dried up prematurely in August and Septem- ber and the leaves all dropped off. Thus we may have leaf dropping as a result of frost, injury by hail, noOE impairment or localized parasitic attack. LEAF SPOT AND SHOT-HOLE EFFECTS Leaf-spot symptoms are everywhere abundant and are really of very diverse origin. In any example in which the leaf tissues are locally invaded by a parasitic fungus we may expect evident effects. In the downy mildew troubles there may be wet-rot symptoms when the weather is moist, as in the case of Phytophthora or late blight attacking potato or tomato leaves; after the leaves have be- come badly diseased they may appear to die very suddenly because DISEASES OF CULTIVATED PLANTS the gradual invasion of the areas has been overlooked. In many other leaf diseases no such rapid multiplication or reproduction of the parasite is possible and limited dead patches or spots are the result. The leaf-spot disease of alfalfa, the various leaf-spots of apple and the conspicuous leaf-spot of the strawberry, the beet, the pea, etc., will be recalled. In these while the leaves are im- paired as to usefulness they do not perish immediately and one may readily fail to estimate the injury at its real seriousness. In a few leaf troubles we have the spotting of the leaf followed by the for- mation of a separation layer in the leaf tissues between the para- sitized and the healthy tissues. This results in ‘“‘shot holes” in the leaves as is so very conspicuous in the shot-hole leaf disease of the plum and less conspicuously so on certain sour cherry. trees. These leaf troubles are commonly very evident during rainy seasons and are preventable by spraying the foliage of the diseased plants at repeated intervals, thus keeping a supply of the fungicide on the leaves to arrest renewed spore development. An interesting leaf-spot disease of the tomato is sometimes very damaging. This disease seems to have appeared in Ohio during the memory of many close observers. Like most leaf-spot troubles which are strictly due to parasitic fungi, this tomato disease has been worst in seasons of abundant rainfall. The same applies to the shot-hole disease of the plum and the allied leaf-spot of cherry. The explanation appears to lie in more favorable conditions for spore germination and for the growth or spread of the parasitic organisms which produce the diseased conditions. Biting or sucking insects also open the way for the entrance of parasitic diseases. (See later pages.) LEAF IMPAIRMENT THROUGH FUNGUS COVERINGS In addition to the leaf-spots or dead areas in leaves to which reference has just been made, we have most noticeable examples of the spread of the mycelium of certain powdery mildews over the leaf surfaces. Casual observers note that these spread over the leaves and stems of roses, over the leaves of lilac, of oak, of peach, of grape, of forcinghouse cucumbers, of bean and pea and upon other plants. While the development of these fungi or powdery mildews occurs often rather late in the season, they are nevertheless damaging to the host plant over which they spread. Above and beyond the interference with the leaf action the impairment of the photosynthetic or sunlight processes of the leaves of the plants by which all real increase in substance is made to the plants, these mildews develop sucking or penetrating organs of the threads of 314 OHIO EXPERIMENT STATION: BULLETIN 214 the mycelium. These organs called haustoria penetrate the leaf epidermis and must do this for the purpose of food extraction—it is needless to add that all food extraction from the plant acts as robbery. f Furthermore, the mildew-covered leaves drop to the ground in fall and there afford the fungus the ‘needed conditions for the development of the resting or winter stages of its course by which ‘it is again ready to attack the plants the following season. Because so largely external in development these powdery mildews are usually comparatively easy of control. WILT DISEASES—SEEDLING COLLAPSE The stems or branches of plants may suffer from localized attack by parasitic fungias wellas from hail, insect attack and _mechanical agencies. The symptoms which follow will be found characteristic. In certain ones as in the clover anthracnose and in the fusarium of clover stems, we have the lesions accompanied by discolorations in which the fungus occupies a subordinate place outwardly. ‘On the other hand the spots or sorz of the rusts upon grains and grasses and the spots caused by the anthracnose of wheat, oats, rye, etc. show commonly a crowded occupation of the area by the parasitic fungus. _ ‘There are many examples of the effects of such lesions. Fuller discussions will be found under the description of the par- ticular diseases. ‘The anthracnose of the bean as well as that of the pea are good illustrations where these attack seedlings. Even -clearer symptoms come out in potato rosette where the fungus parasite at early stages of growth may kill off the stem attacked, while in later attack will cause such impaired development of the plant that stem or axial lengthening is arrested and a “rosette” appearance results. A still more striking arrest of stem elongation takes place in lettuce rosette wherein the roots are destroyed so largely by the fungus in the soil. (See soil infesting diseases.) In cankers of branches upon orchard trees the final death of the immediate branch is preceded by a depressed area invaded by by the parasite. PLANT DISEASES NOT BEYOND EXPLANATION The old mystery attached to disease prevalence can scarcely be maintained in our day. We have worked out in recent years or had determined for us the causal relations between the ferment or parasite and the effects upon the host plant or crop. So far as we can now discover the reason for the spread of diseases, or of a DISEASES OF CULTIVATED PLANTS 315 particular disease, is found in the specific disarrangemen\. in the host plants. This discovery and announcement of these causal relations are undertaken that proper measures for the control of diseases may be finally devised and applied. We must always bear in mind that under favorable conditions plant diseases become epi- demic and their rapid spread is to be expected. The host plant, with its climatic adaptations and the parasites of our crops with their mutual adaptations to their hosts are biolog- ical factors which are capable of being influenced by prevailing atmospheric conditions. With cool,'rainy.weather we have brought about conditions favorable to certain parasitic diseases which will be inclined to spread while these continue. Other diseases spread under the conditions which favor them. The more rapid development of diseases of plants under these favoring circumstances is not beyond reasonable understanding; there is no mystery about it any more than in outbreaks of typhoid fever or diphtheria. By apprehending the differing conditions we may learn to separate the causal from the merely adventitious factors and thus be the better able to master the diseases which result. While we may properly look upon infection by microscopic or other parasites as the general and usual cause of plant diseases, there are diseases of wide importance which arise from internal or physiological disarrangements in the plant. (See Enzymatic. Dis- eases). In all cases whether of parasitic attack or of physiological disarrangement due to other causes, the host plant is weakened and predisposed to death. GROUPS OF PARASITIC DISEASES Parasitic diseases may be grouped ina way, according to the groups of fungi which cause them. ‘This is helpful to the plant pathologist, though of limited practical guidance, since it requires microscopic study to determine the causal organisms. A more useful, limited grouping as is hoped, is proposed below and consists in making such groups or classes of diseases, as are descriptive of the general behavior. Such areseedinfesting diseases, soil infest- ing diseases, root diseases, diseases of foliage, wound troubles, timber rots, etc. The great mass of diseases are treated under each host in the descriptive portion arranged alphabetically. The objects to be attained by this method of arrangement are obvious and call for no discussion. NAMING PLANT DISEASES Plant diseases are named with due regard to the symptoms and cause of the disease. In the case of enzymatic diseases wherein we have peculiar variations or yellowing of the leaves, the names given are mare ar lece deerrintive The same applies to the diseases that al 316 OHIO EXPERIMENT STATION: BULLETIN 214 Parasitic diseases are named with regard to the organisms which cause the disease, or to the effects they produce in the host parts, that is, those diseases which result from attacks of the rust fungi, (Uredineaae), are properly called rusts; also the smutty, dirty conditions resulting from the attacks of the smut fungi, (Usti/agineae), are known everywhere as smuts; these are well known and destructive upon grassesand cereals. Thus we have smuts of oats, corn, wheat, broom corn, sorghum, millet, blue-grass, etc. The anthracnoses are produced by a definite class of fungi, (Melanconiae). The name anthracnose is applied to a disease of a given host caused by an organism of this group and the host name is usually retained, as the anthracnose of wheat, the anthracnose of rye, the anthracnose of raspberry, wherein the dis- eases are caused by species of this group of parasitic fungi. However, in the case of attack. _Fig.2. Headorpan- Upon the fruit as in the anthracnose of apple, ¢ we ome ommret because of the cat kernels and many bitter taste given of their surrounding tg the fruit, we parts have been con- Q >, verted into black, sooty h ave the popular \ yy 1 (smutty) masses by the name bitter eo rot; Q Mi). i N loose smut fungus, oh . = Al\s c} A V ih Nee Ustilago. in a similar in- : d WARS >™ stance, viz., that CLI of the anthrac- Fig. 3. Section through an anthracnose nose of the grapeberry, the dis- spot (acervulus) of the cucumber anthracnose Re is fungus (Colletotrichnm lagenarium) showing colorations of fruit are SO _ thetong, dark hairs (setae) of whose office we characteristic that it is popularly Ca hp sesre essing Besneneniee “ = yphae) and the spores of this fungus. The called the birds-eye rot. With members of that division of the commoner wheat, oats, rye, etc, thename is @nthracnoses having setae in the acervuliare lied b ¢ je "referred to the genus Codletotrichum, while applie ecause of the organisms _ similar ones without setae bear the genus found. As statedin the preceding x 9mes G/ocosportum, Sphaceloma, etc. (See i anthracnoses of apple, grape, lettuce, wheat, pages, we describe most leaf ats, etc). Berane! infesting diseases with regard to the effects the parasites have upon the host; thus we have the leaf-spot dis- ease resulting from attacks of any one of a number of fungi, chiefly, however, belonging to the imperfect forms. The shot-hole fungus of the plum is a good illustration of the naming of atrouble from the symptoms produced. A considerable group of diseases are known as downy mildews. Among these we have the destructive potato late blight and rot, Phytophthora; also the cucumber disease, Plasmopara, as well as the DISEASES OF CULTIVATED PLANTS 317 grape downy mildew and the common white molds of the mustard family. ‘The powdery mildews by reason of the appearance upon the surface of parts attacked, are descriptively named ‘‘mildews.” A definite system has been followed in most cases of naming plant diseases and I trust the results will not be altogether disappointing. The differences between the species of parasitic or other fungi are as strongly marked as those of higher plants, even though microscopic examination is necessary to distinguish these char- acters; it shows, therefore, that a discriminative system of naming diseases has a secure foundation. THE PLANT OR HOST IN RELATION TO DISEASE As stated elsewhere only closely related plants are usually subject to attack by a para- sitic organism, thus it hap- pens that the tomato as well as the potato plants are attacked by the downy mildew or late blight fungus of the potato. In general the true parasites among our fungi are limited toa rather narrow range of host plants; thus we may expect the potato Phytophthora to attack several plants of the potato family (Sodanaceae). The writer proved thissame was true of the attacks of downy mildew (Plasmopara) upon a number of species belonging to the cucumber family (Curcurbitaceae). Since our cereal grains belong to the same great Fig. 4. Avportion of the epidermis from the upper 1 he grasses (Gram- surface of a cucumber leaf, showing the breathing pores family ast g ( “i (stomates) surrounded by guard cells containing chloro- -gneae), we expect, and find phyll grains, much magnified. These guard cells, which that there is a development control the opening and closing of the stomates, are the Z only epidermal cells that contain this green substance, of the same diseaSeS Upon _ the others being colorless. many of them and upon the ; grasses growing nearby. In this connection it must be remembered that clover and alfalfa are not grasses, but legumes. ‘The leaves of the host plant provided as they are with stomates or breathing pores, minute openings through the epidermal cover- ing of the leaf, will be attacked through these openings. The spores 318 OHIO EXPERIMENT STATION; BULLETIN 214 of parasitic fungi after germinating upon the leaf will likely gain entrance into the interior leaf tissues through these openings much more readily than by actual boring through the leaf epidermis. The illustration, Fig. 4, shows how these openings are distributed in the epidermis of a cucumber leaf. These stomates are present in _ the leaf covering upon the outside of all green leaves and in the epidermis of young growing shoots. In addition to these stomates certain classes of plants such as the plants of the mustard family (Cruciferae), as cabbage, cauliflower, turnip, also the grape, fuchsia, impat- iens, etc., are provided with water pores—mar- ginal openings through which the excess water of the plants isexcreted. ‘These water solutions of various materials offer a means of growth for organisms, especially of the minuter forms. From the culture drops thus formed the para- site enters the leaf by the water pores. One of the most destructive known diseases of plants is the black-rot of cabbage, cauliflower, turnip, ruta-baga, etc. This is due to a bacterium which gains entrance very largely through the water pores just described. So we must bear in mind that the very avenues of transpiration or excre- tion, so essential toplant growth, are madeameans of exposing the plant to the danger of parasitic in- vasion. ‘This is analogous to the exposure of human subject to diseases of the respiratory organs. Atevery turn we find convincing evi- Fig. 5. Margin of cab- bage leaf showing excreted water from water pores after cool night. These drops contain enough food for the growth of the black rot bacteria. The motile forms may swim through the water pores into leaf from such drops. Dead marginal areas on lower fragment show results of this bacterial infection. (After Smith). be maintained, dences of the mutual adaptation of parasitic fungi to their host plants, in nothing more strongly marked than in the limitation of the species of plants attacked by a given parasite as discussed in the beginning of this paragraph. In view of the fact that so long as the leaves of a plant continue to function as leaves, these natural openings will it will be seen that the risk of exterior infection from parasitic fungiis continuous for any given plant; it lasts for its whole growing period. THE PLANT'S PROTECTION AGAINST PARASITES In the case of woody growths we have the development of corky epidermis or bark which seems primarily designed to protect the interior, living layer from invasions of this sort. In a similar DISEASES OF CULTIVATED PLANTS 319 manner the external layer or bark of all growing plants, inciading herbs, is provided with a protective covering or epidermis. The skin of the apple or of the grapeand the covering of the potato stem are allfamiliar and serve this function of protection to the inner tissues. In young plants there is retained the power of protective growth in response or resistance to parasitic attack; thus it happens that the potato scab organism induces the growth of cork cells on the outside of the potato and makes a roughness. The roughness is is not the scab fungus but the corky growth of the tubers in response tothe scab attack. Inasimilar manner the attack of the scab fungus upon the apple results in the roughening of the apple skin through the development of more protective or wound cork. The most remarkable example of this multiplication of protecting or outer cells in response to the attacks of parasitic fungi is found in “eaf-curl” of the peach and in the pockets or “bladders” of the plum, where we have such a rapid multiplication of cells in response to the stimulus of the fungus as to bring about an entire transformation in the form and structure of the parts. While we may think of this abnormal development as the result ‘of fungous growth, it is only indirectly so. It isin fact a response of the host to the stimulus of the invading fungus. The nature of the stimulus or excitation exerted by particular parasitic fungi isa highly interesting subject for investigation. DISEASE RESISTANCE IN PLANTS Disease resistance and disease susceptibility are as yet imper- fectly understood. ‘The cause of the inherent differences in the tendency of this or that variety to suffer, as with the leaf-curl in the Elberta variety of peach, the apple scab predisposition in White Pippin, Winesap and others, may become in practice, varietal weaknesses. Yetsuch is the commercial super- iority of some such varieties that they increase in public favor despite these weaknesses. The great differences among varieties of fruit in susceptibility tothe diseases which prevail under certain conditions, is a matter of observation and experience, From the difficulties involved in breeding a less susceptible or more resistant type of tree fruit belonging to any commercial variety, increased resistance is not yet within reach. This applies to established varie- ties and yet leaves the field open for new sports to be discovered or for its occupation by less desirable sorts which do not suffer so severely from disease. This actually happens in the growing of pears outside of certain favored districts; owing to the ravages of fire blight, a bacterial disease, the ordinary grower selects less popular but more resistant varieties for culture. 320 OHIO EXPERIMENT STATION: BULLETIN 214 In the study of disease susceptibility it has been shown that other features being the same, the percentage of water is an index: thus, parts having the higher water content are attacked more read- ily than those with lower water content. With annual plants or those reproduced each year by tubers or seed, the opportunities to breed resistant strains are extremely good and the results obtained are highly promising. Physiological weak- ness in plants may often be translated in terms of disease suscept- ibility; this holds with emphasis in vegetables and grains. Apparent physiological vigor may arise from various causes, and when expressed in terms of more rapid growth or higher water content or succulence of the parts may be indeed a source of weakness in the midst of disease. Selections made for the purpose of securing resistance to disease are made under conditions of dzsease prevalence with highest promise. This field of breeding for disease resistance is one of fruitful promise. Studies in this line have been made by the Horticultural Depart- ment of the Station in respect to resistance of potato plants to the early blight disease. By selection of hills that withstood early blight attack and planting tubers therefrom and subsequent repeti- tion of this work (See Bulletin 174) early blight resistant strains weresecured. The differences between these strains and non-se- lected tubers in 1908 during the marked prevalence of early blight was very striking and clearly showed thata tangible resistance capable of reproduction has been secured. Owing to the wide extent of this field with vegetables and grains, much may properly be expected from breeding for disease resistance in the future. Much progress has been made with cotton resistant to wilt and with musk melons resistant to leaf blight. For the present other remedial measures will also need to be pushed. CONCERNING PARASITIC FUNGI A fungus (plural, fungi) is a plant, a member of the class called fungi. The fungi are low in the scale of plant life, being classed with the alge and other similar plant forms. They are lower still in the life scale than the mosses and liverworts; above the mosses come the fern-plants, and above these the seed plants, suchas grasses grains, clovers, trees, shrubs, herbs and the like, with which we come in contact every day. The fungi are distinguished from higher plants as wellas from their nearer relatives, the alge, by the absence: of green color, and for that reason, we may assume, by the lack of power to prepare their own food from the mineral substances dis- solved in water, and from the gases contained in the atmosphere. DISEASES OF CULTIVATED PLANTS 321 Herein they are marked off from most groups of plants: the fungi must live upon the substance of living or dead plants or animals. If they ever possessed the power of utilizing the same foods as most other plants, this ability has been lost. Parasitism is usually taken to indicate degeneracy in character. One way of regarding the fungi is as alge without chlorophyll, to which the latter owe their green color. As above stated, the fungi are, in the absence of chlorophyll, forced to live upon the dead remains of plants or animals, or to prey upon the living organisms. CLASSES OF FUNGI Such fungias subsist upon living plants or animals are called parasitic fungi. A parasite is one who eats at another’s table and the adjective ‘‘parasitic” comes from this word, parasite. It is the parasitic fungi especially of which we must learn, since this class produce diseases when they attack other plants. The plant attacked is the “host” plant, however unwilling the entertainment of the sycophant. Most fungi are very minute in size and require the use of a microscope to study their parts; certain ones, however, such as the mold upon bread or other foods, may be seen very easily to consist of fine, thread- . a . Fig. 6. Mycelium of the common mold (Mxcor Mucedo). like g rowths interwov- From the spore lying near the middle of the figure, and strongly en toge ther, and __ swollen, one sees the thick threads of the mycelium arise; these A < inturn become richly branched. There are nodivisionsin the bearing certain round- mycelium. From the level of the mycelium arise three vertical ed parts upon erect fertile hypae, 2, 4, ¢, of which @ is still very young and that at 4 . is already producing a sporangium containing manyspores. All branches. Some idea highly magnified. (After Zopf, from Reinke). of fungus-structure may be obtained by studying these common molds; that ona dis- carded melon rind will shows the parts above described, and by the use of a microscope we may learn that the rounded, ball-like enlarge- ments just mentioned consist chiefly of small bodies that are capable of growing into other fungus-threads. (Fig. 6). Such min- ute parts capable of germinating and again producing the fungus are called spores. Most sporesare very minute and are not heavier 322 OHIO EXPERIMENT STATION: BULLETIN 214 than the other dust particles carried by the wind. The spores of fungi are the means by which these are most commonly reproduced: somewhat after the 7% manner that the | higher plants about / us are reproduced by their seeds. While we have cited the bread mold eng as a good illustration heer to show the struc- “=a ~—s« ture of a fungus, it 6 is not. a parasitic -/ fungus; a mold or 48 like growth which’ Fig. 7. 72. A portion of leaf of pea showing breathing pores and parasitized by powdery mildew; the horizontal threads (steréle hyphae) and summer spore bearing parts of the mildew fungus (fertile hyphae) are distinctly shown. In these latter the septa are evident. 74, A spore sac (asczs) of the same fungus, 4, 5, 6, show the sucking organs (azszoria) of the sterile hypae of this fungus; these penetrate the epidermis of the leaf. 10 shows the spores of the rose mildew germinating. All highly magnified. (After Tulasne). » Note—The stomate in foreground is distorted. See Fig. 2. lives upon decaying material is called a sapro- phytic fungus. To this same belong the mush- rooms or toadstools that may be found in manure piles, in the woods and in orchards; the fact that we find them in such places shows that © there is decaying organic substance at ihat point, upon which these plants may subsist. A like condition is found in the shelf-fungi on old logs and stumps, onthe under surface of which we may write our names. Yet if we will usea hand lens we may often discover this under surface to be but a network filled with small openings or pores from which the spores of the fungus will in time escape. In like measure the spores of mushrooms are found in similar canals or upon the sides of the gills beneath the cap of this sort of fungus. The bacteria, or fission fungi, are one-celled plants multiplying by divi- e Fig. 8. Fertile hyphae ‘conidiophores) of the downy mildew fungus on Cardamine, a mustard pro- truding from astomate; the one shown in full, bearing spores at the end of its branches. Highly magni- fied. Very similar to this are the downy mildews of rape, cucumber, lettuce and some others. (After Zopf). sion and by spore production; with bacteria evident mycelium is lacking and they are structurally lower in the scale of plant life than fungi provided witha mycelium. Bacteria are both parasitic and saprophytic. But to return to parasitic fungi: . DISEASES OF CULTIVATED PLANTS 323 PARTICULAR FACTS ABOUT PARASITIC FUNGI Like the bread mold, or the other fungi just mentioned, parasitic fungi consist of a growth of threads or hyphae (singular, hypha) which-do the necessary work of getting food for the parasite; these also in due time give out certain branches destined to bear spores, somewhat after the manner that the pear tree has flower clusters, or the wheat plant forms its dense spike of bloom, both of which are especially designed to produce seeds from which wheat plant and pear tree may inturn be grown. The essential parts of a parasitic fungus are these threads, or hyphae, and the spores produced by them. ‘The hyphae of the fungus taken collectively are called the mycelium, which consists of threads that produce no spores (sterile hyphae) and of those destined for spore production (fertile hyphae. (Figure 7). Itis to the food getting qualities of the hyphae that the fungus owes its continual existence, and they in turn arise from a spore or directly by the growth of some fragment of fungus-thread, as the Carolina poplar may be grown fromacutting. Yet, while all parasitic fungi are made up of these few parts, the differences in form and apparent structure among the several groups are very marked; differences exist as to the thickness of the hyphae whether or not the threads are divided into separate cells by divisions like those at the joints of a bamboo rod, as wellas in the manner of spore formation and in the size, color, form markings and structure of the spores themselves. It is almost hopeless to undertake to illustrate types of spore production and spore forms, since these are so varied and may differ so much at different stages of the development ofa single given species of fungus, yet we may cite a few examples: Fig.9. Showing the common rust of oats and rye. At Aasmall fragment of rye leaf with several orange-red, rust sori breaking through the epidermis; these are of the earlier summer spores (Uredo) or red rust of popular speech. At B asmall fragment of a rye leaf with several black, rust sori, elongated in form, breaking through the external covering; these are of the later summer or winter spores (Z¢lewtospores), AandCslightly magnified. At C section through thg uredosorus of 4; on the slender stalks (basidia) the rough one-celled uredo spores, and between them a young, two-celled teleutospore, which later alone form the sorus. ¢, ¢; epi- dermal cells; /, 2, cells of the leaf interior through which runs the mycelium of the fungus. At D a teleutospore from the black sorus of B; this is divided by a septum into two cells. Similar uredospores are found in most rusts; similar teleutospores occur in corn rust, wheat rust, etc., and in the spores of the cedar apple fungus. C and D considerably magnified. (After Zopf, 324 OHIO EXPERIMENT STATION: BULLETIN 214 Fungus spores may be produced as single spores or in naked clusters attached to certain branches. We find this sort in the downy mildew of the cucumber and its relatiye the peronospora of mustards (Fig. 8); in potato early blight; in fruit rot of plum, cherry, peach, etc., and later in the spores of apple scab. ‘They may also be found in dense clusters breaking through the skin of the plant like the many tubers of a potato breaking through the earth-crust; such without further conspicuous covering are found in the rust spots, in the anthracnoses and the like. (Figs. 3and 9). These dense clusters may arise beneath a special covering resembling nothing so much as the traditional beehive, but are usually ejected forcibly from a specially provided opening at the top of the cone or half-ball. (Fig. 10). A yet more interesting class is that in which the spores are packed so many to a sac (usually eight) and a large number of these crowded into a ball-like, hollow spore- case, such as we find in black-Ixnot, strawberry leaf- spot, the powdery mildews nae A an cam reid ate tet al-pet and in some other instances. flexuous mass of spores, ejected from the pycnidium. 4, (Fig. 11). ‘There is yet an- section of a pycnidium, seated in the leaf tissues and filled other sort in which the with spores. ¢,agroupofthe spores. All highly mag- Nified. (After Allescher from Delacroix). Spore sacs are abundant near the surface of the dis- eased part, as in leaf-curl of the peach, where the maturity of the fungus is shown by the change in color of the affected leaf surfaces. Other gradations will be found as one proceeds in this study. 9 008° THE SURVIVAL OF PARASITIC FUNGI Further, respecting parasitic fungi we must realize that they are all derived by specific processes of reproduction peculiar to the fungus in question; in other words spontaneous generation does not find support among the students of plant diseases. The presence of any given fungus leads us at once toinfer the previous existence, somewhere within reach, of a fungus of like species from which this one was derived by definite methods of reproduction. Likewise, the destructive prevalence of a parasitic fungus in any given time and at any given place, assures us of the necessary supply of spores to start the trouble again under favorable conditions. In fact, all our study leads us to look through mere phenomena, mere evidences of disease, to find the specific parasitic growth which causes them and the favoring conditions under which DISEASES OF CULTIVATED PLANTS 325 these develop. The spores of fungi serve for them the same purpose as do the seeds in higher plants; by reason of the extreme smallness of the spores they are easily transported by the wind and become deposited like dust particles upon exposed surfaces. Certain resting spores survive on the fallen leaves or other parts and will be destroyed if these parts are burned. (See black-knot). The survival of organisms capable of infecting the new crop is certainly to be expected in plant diseases as in epidemic disorders among people. Some fungi which produce disease survive by their thread-like parts (mycedium) ina manner similar to the survival of Canada thistle quack-grass and the mints among troublesome weeds by their visible underground stems. A good illustration of this form of survival is found in the case of potato rosette; in this disease the masses of mycelium (sc/erotia) remain upon the surface of the potato tubers and unless destroyed by treatment of the seed will be ready for immediate attack upon the growing plants (sprouts), even before these have reached the outer air and taken on a green color. Similar survival may occur in cultivated soils, especially where the same or closely allied crops are grown in succession. ‘Thus the same fungus as that,of the potato disease first named, survives in greenhouse soils or in celery soils outdoors. RESTING FORMS AMONG FUNGI The active parasitic phases of fungi necessarily coincide with the activity of the host plants; it, therefore, follows in our temp- erate climates with alternating periods of activity and rest of growth and practical somnolence, that the parasites require to be mutually adapted to intermittent activity. Some spores will survive the brief rest period between harvest and seed time, as in a number of the various grain smuts and in grain anthracnoses. Here they are found simply adherent to the seed grain. Seed infesting parasites like the loose smut of wheat, the anthracnose of pea and bean, and a variety of other vigoraus species survive as resting mycelium, which remains virtually inactive so long as the parasitized seed is not exposed to conditions of moisture and temperature such as bring about germination. ' /There are endless gradations between these instances of “rest- ing’? mycelium and the protected fruit cases of the higher type of fungi. Thus the perithecia or closed fruit bodies of the wheat scab fungus, develop shortly after harvest upon the infected glumes or culms of wheat, and may be observed by the unaided eye, as black bodies seated upon the pink mass of one summer form. ‘These fruit bodies in this case are the kind called ‘ ‘perithecia,’ which contain 326 OHIO EXPERIMENT STATION: BULLETIN 214 within them spore-sacs of a nearly fixed number and each sac con- tains a fixed number of spores of definite form for each species. A great many fungi develop these ‘housed’? or protected forms during the dormant period, and indeed, spore development may proceed in the periods of lower temperature. With the perithecial or sporehouse form of wheat scab, (Gzbber- ella), the spore sacs are formed during the later summer, in our latitude, and these spore sacs disappear before midwinter. For each genus or species under study, peculiar time relations of development may be discovered. The perithecial or spore sac (acsigerous) form just described, or some comparable development of the spores under a definite cover-form, is viewed asa more or less ultimate stage in the development of the higher fungi— the summit in the cycle of their development. The rot of stone fruits, such as peach, plum, cherry and the like, is commonly known only in its conidial development Fig. 11. Section through a spore case (ferithecium), late winter stage of black-knot fungus, showing spore sacs (asc?) called Botrytis. Recent- within. Beside it, three asci containing winter spores or asco- : spores, eight in each sac, arranged in adefinite manner. Along ly Norton has discov- with these are thread-like hyphae known as paraphyses ered the Sclerotinia or ascigerous stage devel- oped from the mummy fruits in which the fungus lay dormant fora time awaiting spring or summer conditions. The bitter-rot of apple and its cycle of development not long since brought to light in Illinois, also shows the relation of the apple mummies, decayed by attacks of this anthracnose, to its survival. The fungus lives over in the old rotted fruits, acted upon by bitter- rot alone, which hang upon the trees. The fungus may also survive in branch cankers upon the tree adjacent to mummies of the bitter- rot. Inthese branch cankers the spore sac or perithecial stage of the fungus is developed. Upon the coming of warm showery weather about early June, new spores are produced from either mummies or cankers and new infection may occur upon the new fruits. The problem of the control of this disease, therefore involves a knowiedge of its manner of survival. DISEASES OF CULTIVATED PLANTS 327 ALTERNATION OF HOSTS IN FUNGUS SURVIVAL This relation of alternating forms in the life cycle of a given parasitic species, to its survival, has been mentioned in wheat scab wherein we have the Fusarium or pink mold and the Gibberedlla forms; in rot of stone fruit where we find Botrytis and Sclerotinia forms, and in apple anthracnose or bitter-rot where we discover the Gloeosporium followed by the Glomorella ascospores. In these instances there seems no real need for the advent of another host plant. In other groups of fungi, notably among the Uvredinee or rusts, we discover in certain species, that survival is accompanied by a necessary change of host plant. The apple rust is known in summer to attack the leaves and fruit of apple, thorn apple (Crataegus), june-berry and mountain ash. This is the aecidial or cluster-cup stage of the apple rust and has its counterpart in the aecidiospores or cluster-cups of the wheat rust upon barberry as well. With apple rust we climb far on the plant ladder and find the teleutospores of rust survive upon the cedar treesas branch enlarge- ments called cedar apples (Gymnosporangium). 'The dry looking apples upon the cedar trees take onanew form during spring showers when they become great, jelly-like masses which emit the teleutospores of the rust, to be carried to apple, juneberry and crataegus leaves by whatever agency is available. The relation of cedar trees to the prevalence of apple rust isa practical matter for each orchardist. It may be better to make firewood of the cedar trees than to combat the apple rust in his orchards. A similar problem as between the barberry hedges which adorn rural England, and the virulence of wheat rust in their grain fields, may also be raised. With us we have plenty of grain rust in the absence of barberry hedges. An adaptive form of survival apparently takes the place of the alternating hosts, and we still have the wheat rust. The instances given are simply illustrative and the student of plant pathology will discover many more in the course of his study. Likewise a careful perusal of the special part of this bulletin will show other instances of survival under many various and instructive conditions. HOW THESE PARASITES ROB THE HOSTS There is an old saying about the stable door and the stolen horse; similar application may be made for plants-and parasitic fungi in a manner which we shall presently perceive. To obtain food we must reach the source of supply; the manner of reaching it is less important than the result. Now it occurs that cultivated and wild 328 OHIO EXPERIMENT STATION: BULLETIN 214 plants of the higher classes are wrapped about by a covering of skin or bark, and the food-filled juices are within; to feed upon any living host the parasite must gain access to the internal tissues of that host. Itso happens that there are minute openings or stomates (breathing pores) through the skin of leaves and of young green stems; these openings are as necessary as the stable door, and through them the thief may enter. (See Fig 4). Were these open- ings to become entirely closed the plant would languish, and remain- ing open, they constantly offer a way for the tender tip of the growing germ thread of a fungus to push its way through the plant covering and to luxuriate within the host upon the substance of the plant. Once within, the fungus thrives, rapidly multiplies its branches, and if in summer, commonly thrusts its fertile threads through some of these breathing pores to bear its spores outside where they may become more widely distributed than if remaining within the tissues of the host plant. Should, however, the Fig. 12. Haustoriaof the fungus of the grape downy mildew penetrating cells of grape stem. ‘The shaded portion shows the mycelium of the fungus growing be- tween the cells, sending haustoria, ¢ a, into the interior ofthe cells. (After Scrib- ner from Farlow). Note: In this figure the lower row of cells have the form of empty epidermal cells in which the fungus. would find little winter season be near, resting spores may be formed, or their formation be provided for within the leaves, or dis- eased parts, as in grape downy mil- dew, elm-leaf disease and in black-knot of plumandcherry. ‘Thus the cycle. of development continues indefinitely to subsist upon. Farlow’s original figure does not give these cells such form. unless some agency intervene to destroy the spores, to prevent their germination, or the parasite itself so exhaust the host plantas to destroy it entirely and the fungus perish for lack of suitable nidus. However, this rarely occurs, not perhaps, so often as men are guilty of killing the goose which lays the golden egg. Herein, we meet another fact, namely, that parasitic fungi of a given kind are limited toa particular host plant of a certain species, or toa small number of related plants, so that if a congenial host is lacking the fungus will not thrive. The fungus threads growing within any plant will not flourish if simply passing between the cells of the host; penetrating organs pierce the cell-walls and are able to absorb nutriment from the cell interior. (Fig.12). The diverse forms of sucking organs, and the peculiar structures of fungus threads in these situations would in themselves require much study and investigation to present them properly. We must further conceive that a fungus may often DISEASES OF CULTIVATED PLANTS 329 penetrate the bark of a tree for example, if aided by rifts caused by freezing or similar disturbances, to say nothing of the openings offered by wounds, the breaking of branches, etc. A recent illustra- tion of the danger of rifts in the bark of trees is offered by the chestnut disease which is proving so destructive near New York City. Few parasitic fungi have that penetrating power of thrusting the haustoria through the plant covering such as we find in the case of the dodder that twines about and robs the wild herbs and shrubs of the woods and fields as well as the cultivated flax and clovers. HOW PARASITIC FUNGI AFFECT THE HOST We know the cumulative effects of insufficient food supply; these effects must hold for plants attacked by parasitic fungi. Aside from the nutriment diverted to the parasite, there is reduced functional vigor of leaf, stem or root, and the loss becomes increased in this way. Letallthe leaves be parasitized, or let even three-fourths of them be entirely soattacked, and we may look for great loss of foliage, possibly entire loss of fruit and the detailed effects of diminished vigor, unripened wood, or by repetition, entire destruction of the host. Usually the effects are of many gradations, but in all cases of leaf-parasites the entire plant must suffer. We have learned that bacteria may, in a suitable medium, destroy themselves by the formation or emission of poisonous products. which are fatal alike to the bacteria and to animals, or even man; that such takes place within plants parasitized by fungi remains in doubt, and may be disre- garded for the present. The results of impaired function in the parts are serious enough to demand our attention. It is altogether ‘probable that future investigations will modify our views upon some points. There are many curious transformations and malformations resulting from the attacks of parasitic fungi, simply by the multi- plication of cells of wound cork or other tissues in the effort of the host to shut off the fungus, not because the fungus consists of such amass of tissues. (See leaf-curl of peach). While exceedingly interesting to trace the effects of the white mold on shepherd’s purse and on the garden purslane, as well as the effects of bramble rust, cabbage club-root and a number of others, the principle above pointed out will be found generally applicable, and it is to the reactions of the host plant that the excrescences or malformations are chiefly attributable. It may further be stated that artificial cultures of parasitic fungi, either upon culture media or living plants are constantly adding to our knowledge in these lines. BENEFICIAL ORGANISMS: ROOT NODULES, ETC. While realizing the losses caused by parasitic fungi and bacteria we may not hastily condemn all fungi and bacteria. One of the most profound influences of aging culture of the soil is the beneficial 330 OHIO EXPERIMENT STATION: BULLETIN 214 effects in nitrogen fixing, exerted by the root nodule bacteria of leguminous plants. The well known beneficial effects of the root nodule bacterium upon clover has made rotation in clover an agricul- tural necessity. The species or forms of root nodule bacteria required on alfalfa, cowpeas, vetches, etc., have become recognized as factors of consequence in our efforts at seeding and new species of legumes on the farm. A less understood relation between certain fungi which develop as mycorrhiza upon the roots of some deciduous trees and notably on conifers may not be passed. Herein we may find an explanation of rotation in forest species when reforestation crops are to be grown. THE PROOF OF PARASITIC CAUSE IN PLANT DISEASE The mere presence of a fungus, determined by the microscope in diseased tissues of the plant, does not prove the case against the organism found. Itis not easy at all times to be certain whether discovered spores belong to this or that organism, or group of organ- isms, although with certain groups as the anthracnoses, species of Fusarium, etc, the spore forms give somewhat clear evidence. The differences between parasitic and saprophytic fungi are not always simple matters admitting of ready determination; further, we must bear in mind that after a parasite has caused death or even minor lesions ina plant, the organisms of decay may be expected to appear to do their great work as the scavengers of the world. The fungi or bacteria found in a dying plant may be both saprophytic and para- sitic, or these may be only saprophytic. The methods of proof of parasitic cause in the bacterial diseases of animals including man have been extended to the study of bacterial diseases of plants and finally to the diseases caused by parasitic fungi. These methods consist of a group of rigorous exact rules which are stated by Dr. E. F. Smith in the following terms: (a) Constant association of the organism with the disease. (b) Isolation of the organism from the diseased tissues and careful study of the same in pure cultures on various media. (c) Production of the characteristic signs and lesions of the disease by inoc- ulations from pure cultures into healthy plants. (d) Discovery of the organism in the inoculated, diseased plants, re-isola- tion of the same, and growth on various media until it is determined beyond doubt that the bacteria in question are identical with the organism which was inoculated. Smith—Bacteria in Relation to Plant Diseases. Vol. 1, p. 9. DISEASES OF CULTIVATED PLANTS 331 While these methods and rules are stated with special reference to bacteria as the cause of disease, they apply with equal force to the proof of cause in the case of any given parasitic fungus. These methods require rigorous and exact work in the isolation and subse- quent culture of the parasite upon sterile media, followed by equally careful inoculation work using these pure cultures as a source of the organism. METHOD OF INOCULATION FROM CULTURES The methods of inoculation tried by the investigator are of great importance. ‘These determine, in fact, the success or non-success of his efforts. There must be adaptation of the method to the life history of the parasite and the developmental stages of the host plant, including the appearance of the parts more commonly attacked by it. Fig. 13. This shows method of infecting field plots by means of the hand spray pump, using the washings of samples of wheat and other grains. The washing of grain containing spores of disease such as anthracnose or scab may be used, Cultures may also be sprayed upon plants in this way or by means of blow-bottle in smallertests. (From Bul. 203, Ohio Experiment Station). Following the methods of earlier bacteriologists, needle pricks are often employed both in the inoculation of fungi and bacteria into plants. One seeking to pursue a special line of inoculations will need in all cases to study his conditions as wellas the methods of other investigators. ‘Thus, doubtless, inoculations like those of Phytoph- thora and Plasmopara may be best attained by using drops of sterile water to carry the spores. The same principle applies in field 332 OHIO EXPERIMENT STATION: BULLETIN 214 methods upon many crops. In the case of grain diseases, notably anthracnose and scab upon wheat, rye, oats and grasses, inocula- tions may be made by spraying the cultures upon the grain ata proper stage of its development. While some groups of fungi do not lend themselves readily to culture upon the usual media, it is the aim of plant pathology to make this possible witha constantly increasing number of these parasites. CULTURE PROOF NOT ALWAYS POSSIBLE While in all cases of bacterial diseases where the body of the organism is so little different from that of the bacteria of decay, fermentation, etc., these rigorous proofs are required before the disease is listed as of proven bacterial origin, we do not find it nec- essary in practice to reprove again the case as against frequently occurring species of fungi associated with particular plant diseases. This does not make it less necessary to prove all cases as to para- sitic cause, although the practicability in any single laboratory of pathology is admittedly one of narrow limits. ENZYMATIC DISEASES OF PLANTS: CHLOROSIS OR PANACHURE To this form of physiological breakdown, induced however, by specific causes recently determined, we attribute some very wide- spread and injurious diseases which belong under the head of chlorosis. Peach yellows, possibly peach rosette, frenching or mosaic disease in tobacco, and in general variegated or special yellow foliage types of plants as in Arundo, Acer and other genera of plants belong here. The yellows in peach has long been studied, as also the tobacco mosaic disease. In yellows the contagious character of the disease and its transmission in pruning by contact of parts of the harness of team and by or through the atmosphere has been recorded. A few years ago it was determined by Beierjink and by Hunger that this infection exists as a chemical compound or compounds of complex nature belonging to the oxidizing ferments ofa group called the oxidases. Oxidase, peroxidase and others of these ferments are known. They act by breaking down or oxidizing the plant leaf tissues and especially the chlorophyll or leaf-green of foliage and young tissues, converting it into xanthophyll. The tests for these © ferments are of some importance. Woods and others have shown their action with peroxid of hydrogen. From a practical point of view the transmission of the ferments, _ and, therefore, of the disease, by touching first diseased and then healthy foliage is: rather surprising. The work of Hunger in Java | DISEASES OF CULTIVATED PLANTS 333 upon the transmission of the tobacco mosaic disease. makes the risk of transmission from diseased to healthy plants by such handling, stand out clearly. ‘This line of transmission was verified on tobacco by the writer’s assistant in 1903 (See Bulletin 156, of this Station). While the same class of proof for peach yellows is very difficult, owing to the latent nature of the disease for some months after first infection, the actual results of infection from nearby diseased trees make clear the danger of such exposure and the necessity for the destruction of diseased trees. Chemical examination of variegated or chlorose tissues shows the same compounds, the oxidases, etc., to be present and to account for the transformation of the leaf-green or chlorophyll, into xanthophyll, or leaf yellow. ‘Thus by degrees apparent plant disease mysteries are solved. ‘The weakness of variegated plants and their ready susceptibiity to attacks of para- sitic fungi are now explained by thisimpaired condition of the leaf parts. PLANT DISEASES TRANSMITTED IN THE SEED The public in general little realizes how many diseases of plants are transmitted in the seed, although as the years pass the general dissemination of knowledge concerning infection by spores and by germs has partly prepared the way. The public mind does not longer expect something to grow from nothing. The treatment of seed grain, as wheat, oats, barley, etc., todestroy adhering spores of the smut fungi, and thus prevent these smuts in the crop, has been known for many years. In the early days of the Agricultural Experiment Stations, these doctrines and practices in this regard were widely disseminated, new impetus being given by the success- ful use of hot water following’ the methods of Jensen in Denmark; but despite the conquest of the practical control over the order Ustiligineae, the smuts, we have only really begun to study the matter of seed infecting diseases produced by seed infesting fungi. These seed infesting fungi are of two types, viz, first, those whose spores adhere to the seed grain as in the case of the smuts of grains generally, and second, and more exactly, those fungi which develop upon or within the seed largely by their threads or mycel- ium, and may, or may not, prevent the germination of the infested seed grain. Our knowledge of these strictly seed infesting fungi is quite recent; we may point to the work of Prof. Bolley and his assistants at the North Dakota Experiment Station, especially upon the matter of flax diseases; to the work of Dr. Halsted in New Jersey and to Bulletin 173 of this Station by Van Hook. With the tendency to continuous growing of flax, in the west there was developed m 334 OHIO EXPERIMENT STATION: BULLETIN 214 that new area specific seed and soil troubles which have been proved to be perpetuated in the infected seed. An anthracnose of flax and a Fusarium attacking flax seed are examples. No less conspicuous is the case of the blight fungus of peas, Ascochyti pist, which is also an anthracnose, and the allied anthrac- nose of beans, Colletotrichum lagenarium. Investigations made at this Station by Van Hook show the source of the trouble with peas to be the infected seed employed and show also that seed treatment will not destroy these internal fungi without destroying the vitality of the seed. It was further shown that the source of relief zs in growing healthy seed through the use of fungicides upon thé pea vines from which seed is gathered; likewise that infection way remain in the soil, (See Bulletin 173). Fig. 14. Showing physician’s centrifuge and other apparatus used in making examinations of grain washings for smut spores and spores of other diseases adher- ing to the exterior of seed. The flasks at the right show samples of washed grain. Those at the left show amounts of grain and water used. The glass tubes in con- tainer are used in the metal holders of the centrifuge. The precipitates in bottoms of tubes were obtained from washing of oats and wheat samples in flasks. (From Bul. 203), More recent work at this Station has shown the presence of seed infesting and seed infecting diseases in wheat. (See Bulletin 203). The illustration, Fig. 15, exhibits the germinating seeds of wheat with the outgrowth of the parasitic fungus (Fusarium) which we find associated with wheat scab. Thisis upon seed grains (kernels) that are not destroyed by the fungus; many of the kernels of scabby. DISEASES OF CULTIVATED PLANTS 335 heads will not germinate. It was also found in continuous wheat land as much as 6 percent of the young wheat plants were destroyed in the fall by this same parasite which appears to survive in the soil under continuous ‘wheat growing as well as to be propagated in the seed grain. Fig. 15. This shows results from germinating ten wheat kernels in Petridish containing agar. Both the agar and the kernels were sterilized. After five days it was found that five kernels had produced healthy plantlets, and four kernels had germinated but were attacked by the scab fungus, Fusarium, and two by anotherfungus. One kernel in the center did not grow and gave only growth of the scab fungus, Fusarzum, (From Bul. 203). HOW TO EXAMINE SEEDS FOR INFECTION Recently good success has been obtained in the laboratory of this Department in determining the presence of certain seed infesting fungi in seed wheat, oats, rye, etc. In regard tothe matter of adhering spores this is accomplished by making washings of the seed in distilled water and separating the spores from the washings by means ofa physician’s centrifuge. (Fig. 14). The spores and similar particles washed from the seeds are thus collected in the bottom of the tubes of the centrifuge and may be identified by microscopic examination. (Fig. 16). 336 OHIO EXPERIMENT STATION: BULLETIN 214 Examples may be multiplied to illustrate the range of seed infection both by adhering spores and by internal development of the mycelium of the invading fungus. Many of these are treated under the particular diseases of the crops. ‘The bean, pea, barley, broom- corn, flax, millet, potato, sorghum, rye, sweet-po- tato, and wheat will all furnish examples. Not only have we to test the actual survival ofthe par- asites thus found but we must discover the behav-_ ior of the disease with respect to the germina- tion and seedling plants which grow from such infected seeds or tubers. Examination for infection of seed bulbs and tubers may be made either with or without the growth of Fig. 16. Microscopic photographs from centrifuge precipitates of wheat washings. 2, from wheat washings, narrow, slightly curved anthracnose spores, small spherical, loose smut spores, large spore of stinking smut and portions of the setae of anthrac- nose. 4, from wheat washings, small, loose smut spores, large stinking smut spores and curved scab spores, All magnified about 180 times. plants from them. With potato scab and rosette, the external scab effects or the sclerotia of Ahz- zoctonia are not difficult tosee. With the latter the moistened tubers show _ > marked color contrasts and make the work easier. These diseases are reached by seed treatment. Where the infection is internal by the threads or mycelium of the fungus, the seeds may be germinated in Petri dishes where the kernels are surrounded bya moisture retaining, sterile medium DISEASES OF CULTIVATED PLANTS 337 such as agar or gelatin. This method has been worked out in Bulletin 203 and may often be applicable. The illustrations above will show the results in these cases as before referred to. With internal tuber infesting diseases as in the bacterial wilt disease of the potato, the Fusarium wilt or dry- rot fungus of the potato and the soil-rot of the sweet potato, we must go further than mere external examination. For the two named wilt diseases of potato, infection usually shows earliest at the stem end. Thin slices across this stem end of the tubers will show wheth- er or not there is discoloration in the vessels. In the absence of infec- tion there will be no discoloration with bacterial infection by Bacillus solanacearum, black areas or rings will be seen in these tissues while tubers infected with Fusarium oxysporum will show local areas of browned or blackened tissues. This infection applies usually to “harvest time. As the infect.on advances, one-half the length of the tuber or even more may become infected. In all cases sections from sterilized tubers may be used asa source of cultures in Petri dishes. The same applies to soil rot of sweet potato. These diseases are not reached by seed treatment. THE LIMITS IN SEED TREATMENT — Fig. 17. Pea stem showing lesion from It will be apparent that serious _ blight fungus, Ascociyta pisi, near surface eee iN of ground. Thisfungus came from the seed limits hold in regard to seed treat- pea. (Natural size). From Bul. 173. ments. Where the spores are external and simply adhering to the seed grain, treatment will destroy these spores if rightly adapted to the seed in question and the germination need not be much, ifany impaired. On the other hand where the seed infection is internal rather than external, grave doubts arise as to the possibility of successful seed treatment. 338 OHIO EXPERIMENT STATION: BULLETIN 214 It has not been found possible in thé cases of seed peas when infected with the blight fungus, or of seed wheat, rye, etc.,infected with the scab and other fungi to apply any seed treatment which would destroy the infecting fungus without destroying the vitality of the seed grain. In genera] we may say that where the seed infection or fungus spores, etc., are external to the visible or germinable grain, seed disinfection through treatment is possible, but for the internal fungiitis rarely possible. The loose smut of wheat may be amen- able to special seed treatment with only partial loss of vitality in the seed wheat. Fig. 18. Potato tubers attacked by Dry Rot Fusarium, showing sections near the stem-end of infected potatotubers. This infection may be easily discovered by cross sections made with a sharp knife, and sections from sterilized tubers gives culturesin Petri disnes. At times the discolorations extend to the middle of the tuber. (From a photograph by T. F, Manns). : METHODS OF SEED TREATMENT The methods of seed treatment heretofore employed are set forth in the spray calendar and consist in an immersion of the seed in hot water of definite temperatures or in solutions of formaldehyde of different strengths. The formaldehyde solutions may also be employed to sprinkle piles of seed grain and in this manner less handling of the grain is required. More recently it has been pro- posed to disinfect seed potatoes, onions, forage, etc., through fumi- gation with formaldehyde gas liberated by boiling the solution, or better by mixing formaldehyde or formalin solutions with pulver-' ized potassium permanganate by which the gas is liberated. With seeds, tubers, roots, bulbs, etc., the limitations of the treatment are not so narrow and these may be immersed for longer or shorter periods in solutions of corrosive sublimate, formalde- hyde, etc., or they may be exposed to fumigation with gaseous formaldehyde as has just been stated. (See Seed and Soil Treat- ment, pages 344 and 345 following. DISEASES OF CULTIVATED PLANTS 339 SOIL INFESTING PARASITES IN FIELD AND FORCING HOUSE The cultivated soil is a medium in which many species of bacteria and fungi survive from year to year. The public is famil- iar with the doctrine of bacterial infection or inoculation of the soil in its relation tothe nodules or tubercles of clover, alfalfa, soy beans, cowpeas and other cultivated plants of the Family of Leguminosae. One form of bacterium is not sufficient for both clover and alfalfa. This flora of the soil both in relation to bacteria and fungi of consid- erable range of species, is enriched by the applications of manure and by the practices of culture; by this is meant that the growing ofa given crop a second time ora third time consecutively in the soil increases the probability that the plant roots remaining in the soil are carried over from one crop to the next together with root parasites which cause dis- ease in the plants of this crop. Manifestly, like- wise, ifin preparation for a given crop to be grown for the first time upon the Jand, rather liberal appli- cations are made of fresh stable manure containing spores or mycelium, more especially the resting forms of mycelium called sclerotia, the soil will be- come infected by this manurial application. While this source of. in- fection is rather rare in field culture we bave spe- tific examples as in the ; isease of potatoe Fig. 19. Bases of potato stems (Carman No. 3) collected scab dise s a om oes June 7, 1902, Cheshire Ohio, showing injuries by Rhizoctonia, transmitted in this Way; The shaded areas are darker lesions occupied by an abund- ugar beets ence of Rhizoctonia hyphae; the tops showed conspicuous the scab of - & : in Rosette effects. Reduced from Bulletin No, 139. may be carried in like manner. But in forcinghouse culture where heavy applications of manure are made, the chances are greatly increased that soil infec- ion will be produced from the manure. It is of value to remember that seed infesting or seed infecting organisms are also very largely capable of survival in the soil nidus of cultivated soils, thus our troubles multiply adequately if our care be inadequate to avoid them. 340 OHIO EXPERIMENT STATION: BULLETIN 214 Fig. 20. This shows root portions of seedling lettuce plants with dark spots, lesions caused by attacks of the rosette fungus, /th/zoctonia, With the younger plants these attacks cause large mortality and in very small seedlings the stem of plantlet may early collapse after the manner shown in rotting specimens, (From Circular No. 57). THE AVOIDANCE AND PREVENTION OF SOIL INFESTING DISEASES We, perhaps, may assert that the law of nature is that of a diver- sified plant covering; at any rate the law of successful culture will permit of statement in termsofcrop rotation. And it is true that as culture ages the number and seriousness of plant diseases increase almost in geometric ratio. Itis further conspicuously true with respect to those areas devoted largely to continuous culture ina single crop or ina group of closely related crops such as the growing of wheat in Western United States and Canada, also in the growing of flax and other crops. Potato growing in San Joaquin county, California, illustrates this danger. Muck lands devoted to vegetable culture, tempt the grower to continue his crops of celery, onions, etc. Here we haveasatrue result the accumulation of diseases which attack these plants; thus for field culture we are sao ca DISEASES OF CULTIVATED PLANTS 341 a serious decrease in the return from the crop on the special type of soil. While for general field culture avoidance of conditions may be successful, this is by no meansa simple matter. Rotation is often absolutely necessary, but this same rotation will not rid the soil of the onion smut fungus, nor of some other parasites such as in the case of the club-root fungus of cabbage and related plants. In these cases some soil treatment must be applied to field areas. In the case of the onion smut it is sufficient to apply a formalin. drip which will fall with the seed and disinfect the soil layer in proximity to the seed. This is effective because the smut fungus can penetrate the germ- inating onion plant only in the earlier stages of growth. On the other hand with cabbage club-root, where plants are transplanted from the beds in which they are grown, some general method of soil treatment which involves the soil mass is more effective. In this case it is the application of stone lime or caustic lime in liberal quant- ities. These examples are only illustrative of general conditions to be met with. In the case of potato scab, itis found necessary both to disinfect the seed where scab is present, even toa limited extent, and to plant upon new soil not infested with the scab organism. Potato rosette is certainly an acid loving parasite. | ee Fig. 21. Lower portion of two celery plants Showing effects of root rot, RAzzoctonia, It will be observed that nearly all the roots of the plants have rotted off in consequence of the attack. (From Cir- cular No. 72). 342 OHIO EXPERIMENT STATION: BULLETIN 214 Under the diseases described for each plant, methods of avoid- ance and treatment are indicated and the diligent student will find other instances of similar character mentioned therein. SOIL TREATMENT IN THE FORCING HOUSE About our great centers of population have grown up conspicu- ous developments of the forcing house industry; large areas are covered with glass and these hothouses are maintained at the necessary temperatures for the production of the special green crops found profitable, At the same time the soil of the hothouse beds is very liberally manured and enriched further by applications of commercial fertilizers. Not only do these applications of manure tend to enrich the flora of the soil and to introduce particular root parasites, but the decay of the organic matter of the manure also tends to produce humus acids in considerable quantities. Along with these are brought parasitic eelworms which are peculiarly fatal to curcurbitaceous plants, to violets and to some other hothouse crops. _ Next to the eelworms the most conspicuous organism in our area is the sterile fungus Rhizoctonia, whose resting formsor sclerotia may be readily introduced in manure. To meet this indoor soil infection, special methods of soil sterilization had to be developed, since soil rotation is practically impossible in the forcing house. These methods of treatment are in brief, steaming oi the soil to render it practically sterile, so far as these parasites are concerned, and a method of formalin drenching. This method of steaming is ideal, or almost so, in its results upon sandy or loamy soils, but often entails unfavorable changes in heavy silty or clay soils. Essentially, it consists in burying a series of perforated pipes in the soil at inter- vals, covering the surface of the beds and passing live steam in suffic- ient volume into the pipes. These pipes are prepared insets with cross heads and boiler connections and are perforated at desired distances. The surface of the bed is covered with canvas and the steam passed into the system for such period as is required to heat the soil toa temperature from 180° to 212° Fahz., to be maintained for one hour or more. ‘This applies to high pressure boilers; for low pressure or hot water heating systems where the steam is applied in subdrains, four to five hours steaming will be required with a pressure of six to seven pounds. ‘This treatment is effective for destruction of the eelworms or nematodes of cucumbers, violets and | lettuce, the rot or drop fungus of lettuce, the rosette fungus, and in general of all soil infesting parasites. DISEASES OF CULTIVATED PLANTS 343 Another method, the formaldehyde or formalin drench, has proved successful for the control of AAzzoctonza in greenhouse soils. This consists in a solution of 40 percent formaldehyde in water, say at the rate of two to four pounds in 50gallons of water. ‘This is applied at the rate of one gallon per square foot of area and will involve a very severe wetting down of the bed and prevent immediate resetting of the soil owing to the persistence of the formaldehyde. One secondary effect of formaldehyde drench and lime in sandy soils in the forcing house was an increased yield of lettuce amounting from 60 to 90 percent over the ordinary crop. This was explained on the assumption that the parasitic fungus was destroyed and certain other inhibiting organisms at the same time. THE BEST FORCING HOUSE PRACTICE The best forcing house practice will contemplate a recurrent disinfection or sterilization of the soil during the idle period; it should be preceded by whatever applications of limestone and manure that are to be made tothe soil, then after thorough working and appli- cation of water to correct unevenness of moisture the soil may be sterilized by steam, or the formaldehyde drench be applied with assurance of results: obviously also this treatment must extend most vigorously to the plant beds and bring healthy seedlings to soil in which the soil parasites have been destroyed. ‘The following tables of seed and soil treatments taken from the spray calendar will be of more use than extended description or discussion: OHIO EXPERIMENT STATION: BULLETIN 214 344 “(TET UIETING seg) “10S OUT ITs pu ‘puL] pamord ATYSeIj UO Butpsas 0} snorAcid qsnf eloe Jed sjeysnq eay-4jUeM} pUe peIpUNyY uO 03 aAY-AjUAAIS JO 9} eI BY} 7e OUTITYIIND punois Ssdde iQ “pass 34} YUM Sulyes uorpNjos ay} ‘yUeWYIEIIe dip pur [TIP YIM Mos 19}}aq Jo ‘aou0 38 12409 PUL [LOS [JIM 308 UD UI! pass UO papYUlIds JozeA Jo “s[eS YE OF “QI [ UOIFN[OS epAYapyeussoy as—) *sdoi9 101730 YUM SUOIUO 932701 $}NUIS UOTUO Joy Se apATOSP]VUIIOJ YITM Peas MOG) ‘AIP Way} ‘BUIIIS YIM SAnoyY FZ JO} Py[Ns unissejod uorynyos yueo1ed yf Ul psas HOS *£1p 0} auU0 ye pealds ‘-1yey soo1gap Z-OF] 7B SONUIUT BAY 10j 10 *-1Yeyy Se01Bap Oey 3B SazyNULUL UWaAas JO] **IGe SeiFap ¢-ZET 72 19}eM JOY Ul SazNUIW Ud} JO} JasseA Usdo U! 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UO !quds 10} SY apAYAP|eUIIO] UL Pas YLOS]----+-- qeos ro] se ops ¢ ‘ -OpleulLIoy ul pass yEOG) *pasn oq ALU SBS SPAYSP[EWAIOJ IOS BoTf | ++ eee eee ayeullqns -qeos Uo JuRId pue AIp Uat]} ‘a; BULI[qns VAISO1IOD Ul InoY suo JO epAYap[ewW.Ao} Ul SINOY Om} 10} peas Yeo ‘ “epuayed Aeidg seg “syue{d Jo Buldeids YBno1q} pass jo uoreyuI UMOp daoy ‘(SeploIsuNy Japun poyzour jo Uor}dl1osep 93g) “SaNoY gF 0} Fz Jo POliod B IO S9}zV1D LIS Jo Safid pasojIue UI ses apAYap[eUsio} yRIM ‘SUOIUO AIP 9} JOsUISIP 0} eye sUIN . cs a "++ T1B94S UPI [IOS Jeayy ttrss-q1OS UL OLITTyIING vette JaqzeM-qns JO qorny wre8}S YHA [10s ozt1103¢ press urvoys HM azit4e3s Jo apAyepreur -10y YEA spaq) youarq dase ecenes ‘apAyepleuti0og “rere sss -apsyapreuni9 gy WIB9}S YIM TIOS ezITL1971g PATSO1I09 10 apsYapyeur -10} UL pees ynoun yeog rie cibletere's xnveplog qqIM doid BUIMOIs ay} ABIdg ++ SB OpAY “OPTEUIOY YM Joystiq) uleil# pai0}s Ul s}IesSUT vrrotts anus BuLUns saraeletous vee JnUISs Bs00'T asoudeIyyuy asnoyjzoy Ul sepoyewany ++ gsnoy}oY Ul oOI-yUIOg lasnoyyoy ul sepozeweyT “*}01-Pag PUB 701-JO0y asouseIyyOy lasnoyzoY UI SepoyeUayy| “+ (ByUOyDOZIYY) 9779S0y, “qeog srertsees=* SIOI-93RIOIS tresses ess) *qoor-qntg| + yor-uIa}g pure }O1-yORTg]- “*(W3Nq) esousvayquy| J9TOLA ers Giang sieereysieew Barer oyeUIOL, osseqoy, “oOJe}0g J2aMg opr alahs niosenans aussas esate asy sasoy oye10g wag uwolugO INAWLVAUL JO dOHLAW INGDALVANL GaLvanl LVHM 10a INV1d 40 aaas Pepnpu0d SLNANLVaaL TIOS GNV daas 346 OHIO EXPERIMENT STATION: BULLETIN 214 ROOT DISEASES AND ROOT-ROT FUNGI OF ORCHARDS Diseases upon the roots of herbaceous plants are very commonly due to soil infesting parasites. As explained under that topic, the soil conditions may be favorable to certain parasitic organisms or without being especially suited to them soils become infested with fungi which tend to remain indefinitely and become a source of loss in crops and effort. See lettuce rosette, tobacco root-rot, potato dry-rot and root-rot of violet. The root-rots of woody growths are commonly more or less truly wood invading fungi of the semi-parasitic type and become of interest to foresters as wellas orchardists. A partial exception to this wood-invading character of these root-rot fungi is found in a recently discovered development of the tobacco and violet root-rot fungus, Thielavia basicola Zopf., upon catalpa seedlings in nursery. However, since even tree seedlings in their early stages have not developed their woody tissues to any great extent, they are susceptible to the same root parasites as are found on herbs. ‘This will likely explain cases like that cited on catalpa and the trouble may pass as the seedlings become older. Yet it must be confessed that this still raises a ques- tion as to the effect of the general parasitism of even Thielavia upon the rootlets of trees like catalpa. In forest woodlots, root-rots are likely to become of increasing effect. Wherever these tend to limit the reproduction of certain species in the woodlot, they will be injurious. In this respect they may prove an added reason for the rotation of conifers with decid- uous growths. In coppice or cut-over lands such as prevail in the charcoal furnace districts of Ohio, the roots and stumps of the parent stem must be an eventual menace to the'new growths which spring up about them. The exposure to the wood fungi which become timber or heart-rot sorts will be very great in all such cases. The gradual invasion of the new growths must often occur when these approach a size that gives a considerable heartwood cylinder. These are the great sources of trouble in coppice reproduction of timber trees. Root-rots in orchard plantings are known more especially when these are made following oak and other species somewhat after the manner of coppice conditions. The rhizomorphic development of these root-rots is difficult to determine but is usually referred to Agaricus melleus (Armillaria mellea). See root-rots of apple, peach, etc. An especial feature to be noted in root-rots of all sorts are the soil conditions as toexcess moisture and aeration of the soil. In silty or clay soils of close texture and coagulable nature, with excess moisture, serious conditions arise. Any traces of root-rot fungus DISEASES OF CULTIVATED PLANTS 347 under such conditions will involve increased risks. The necessity for drainage will usually be apparent and due consideration of the limits of certain orchard trees needs also to be given. Cherry trees and even peach and apple trees will not survive under moisture con- ditions wherein plum and pear trees may grow with profit. Orchard replants in “‘clinker’”’ locations wherein failures have been numerous, will raise these questions of root-rots and relative adaptability of different orchard trees. Rotation planting, as pear or plum after apple, plum after cherry, etc., may at times succeed and replace unsightly gaps in the orchard by flourishing trees of another sort. At present, drainage and aeration are our known methods of restricting root-rots under out-door conditions. PARASITIC FOLIAGE DISEASES Foliage diseases of every sort are caused by oxygen loving or aerobic species of parasites, and very often this development on the leaves consists of the imperfect forms of the fungus life history. These forms are none the less aggressive and injurious for this reason, but the exact manner of survival from year to year becomes important wherever not known.. ‘The application of this to prevent- ive measures in the control of these diseases upon foliage and fruit is seen in the case of apple scab, the monilia rot of plums, peaches, cherries, etc., and in apple bitter-rot. These last two rot troubles survive in the “mummy” or dry rotted fruits and this explains the reason for the oft repeated injunction to destroy all “‘mummies” in addition to spraying operations. ‘The bitter-rot of apples is propa- gated by means of summer branch cankers on the tree, as brought out-in recent years. Other leaf forms survive on the fallen leaves or possibly in bud scales as with the leaf curl and “‘bladders”’ of the Exoascae. A large number must live over on the branches. Parasites upon foliage soon become apparent from the spots on the leaves and dropping of fruit resulting. This dropping may come asaresult of impaired vigor by reason of disease—‘hen it is later, but is more often the direct result of parasitic attack by the disease upon the young fruits. Herein as elsewhere the philoso- phy. of fungicides comes to our relief. A good foliage fungicide is a relatively insoluble compound which will not greatly injure the leaves with which it is in contact. The remedies for foliage troubles are applied in anticipation of attack and for the purpose of checking the fungus when it may appear. ‘The relative efficiencies of various fungicides in early summer will possibly depend upon the sticking qualities of the sprays. 348 OHIO EXPERIMENT STATION: BULLETIN 214 Foliage diseases, moreover, are liable to recur each year and - tnis is an added reason for anticipatory treatments to ward them off. - Foliage diseases may not be neglected with impunity since the leaf ts the plant's vital working organ and the plant must suffer from its im- pairment. BITING AND SUCKING INSECTS AND LEAF DISEASES The part played by insects which wound the leaf epidermis, in the spread of leaf diseases, is often very important. Such wounding Fig. 22. Showing sections of blades of oats attacked by green lice (aphides), The right hand specimen shows type of injury re- sulting from the sucking of the aphis. In case these lice are carry- ing the organisms of oat blade blight, this sucking will lead to infection by the disease. (From a photograph by T. F. Manns), of the leaf or green stem whether by insects such as flea beetles, foliage eat- ing worms, or by sucking insects such as mites, leaf hopp- ers and plant lice, opens the way for the spores of para- sitic fungi or of bac- teria or mere molds, any one of which may be injurious to the leaf. The eariy blight disease of ’ potatoes is a good example. In seasons when there aremany of these little black flea-beetles to punc- ture the leaves, the thorough control of both these insects and the early blight, Alternaria fungus, is called for. ' Many fungi of doubtful penetrating powers are truly injurious when they follow in- sect puncturesof the leaves. Fortunately both these are se- cured by Bordeaux sprays. The reasons for such applications are of double character since they are to combat both the insect and the fungus to follow it. DISEASES OF CULTIVATED PLANTS 349 With shade trees the leaf hoppers and mites may be so numer- ous that tip-burn and various leaf dying results from the injuries or punctures they inflict. A more startling relation is that of the blade blight of oats, a recently investigated bacterial trouble. This bacterium is distributed and inoculated very obviously by the aphids or green flies (plant lice), and other sucking insects which prevailed during the seasons of 1907 and 1908 upon oats almost throughout Ohio. For fuller details see Blade Blight under Oats, and Bulletin 210. WOUNDS AND WOUND INFECTION With woody growths, especially in trees which attain considerable size, we have the various phenomena of disease infection through wounds; this infection later becoming evident by reason of decays set upin the woody tissues. Of course, in instances such as the bark disease of the chestnut, Diaporthe parasitica Murr., the dis- ease may penetrate the living tissues. Not so, generally, in wounds of woody plants. Any large woody growth, as in forest or shade trees and in larger fruit trees, shows the combination of an external or living sapwood layer Wiget Reel atone and an internal dead or _ heart- — with large branch cut off. Below this wood cylinder. The commoner forms jytinay nas ja yen Ani of wound infections are attributable cap is nine inches across. This shade to those species of fungi which cause are ns Ome eee decay of this dead heartwood. Among pole. (From a photograph by J. M. these are the long list of saprophytic, S"*e"” agarics, polypores and stereums. Because of the fact that this heartwood cylinder is dead, these saprophytic species of fungi, once they gain entrance into it, flourish there and invade the wood toa very great extent, even by adaptation to parasitic habit extending their work into the living parts causing death. The removal ofa large branch of a shade tree or a fruit tree, unless the wound thus 350 OHIO EXPERIMENT STATION: BULLETIN 214 formed is properly protected by dressing, opens the way for spores of these fungi which cause timber decay to obtaina start and thus eventually to invade the heartwood of the interior. For dressing cut. off branches, asphaltum is admirable; in its lack gas tar is good, and either is better than ordinary paints. Fig. 24. A wound parasite (Pleurotus ulmarius) on the trunk of a maple tree. (After Freeman, Minnesota Plant Diseases). There is always to be borne in mind that the protection of the woody cylinder of trees depends on its being covered by the living layer of sap wood. Every branch of considerable size connects directly with the extensive heart cylinder; we thus see that the wound fungi which attack the heart wood are the timber decays and their presence emphasizes the need for care in providing protection for all wounds, especially those caused by pruning. i Any decay becoming established inthe dead heartwood may extend for long distances through this dead wood and in the end so destroy it as to be ina position to invade the external or sapwood _ layer. DISEASES OF CULTIVATED PLANTS 351 In addition to the exposure of the internal woody cylinder to these decays, we have sap-rots due to various species of fungi which belong on the border line between the parasitic and saprophytic sorts: Among them are species of Fomes, Polyporus, Lenzites, etc. Any wound of the sapwood even though it does not reach to the dark heart- wood, exposes to the danger of this infection, and with infection, to all the consequences of sapwood decay and premature death of the tree.- These decays and those of heart- wood are inline with those of the rots of structural timbers, but we are at this time interested only in their effects on the parts of the living plants. TIMBER ROTS AND TIMBER PRESERVATION The decay of dead logs, wood- en frameworks, or other structural timbers is caused by the attacks of .saprophytic fungi belonging to the gill and pore fungi mentioned under wounds; these are of the great class of basidium bearing fungi, to which the fleshy forms, everywhere more or less abundant, belong. The most of them are included in the “‘mushrooms,’ which there is a Fig. 25. Another wound parasite (a strong impulse now to study and frets oir The fungus ond illustrate by photogra phs. ‘These tree (an oak), and as shown by the fungus timbers are dead and are subjected [uting tates, is efadully, rorresing to invasion by timber infecting the photograph was taken, (After Free- species wherever the conditionsas _™"" toair and moisture are suchas to favor their development. Dry timbers are not subject to such attack because lacking the requisite moisture for the organism. Floors and other timbers of houses adjacent to the earth or to unheated cellars are often attacked by rot-causing species. The timbers of trestles, railway ties and the bases of fence, telephone and telegraph posts, where inserted into the earth or in contact withit, are kept sufficiently moist to invite attacks of this sort. 352 OHIO EXPERIMENT STATION: BULLETIN 214 Wood that has been invaded by such fungi is reduced to the state called punk: that is, the wood fibers and arrangement in vessels to which the timber owes its strength, are broken down by the invasion of the fungus which flourishes at the expense of this woody tissue. There is no help for timber after it has once been attacked by rot fungi. Whatever preventive measures are taken must precede the attack. The most effective means of timber preservation is to cause it to be injected or permeat- ed with creosote or other antiseptics. ‘This is done by plac- ing the timbers in vats containing the solution and extract- ing the air from the timbers so far as possible. The per- manence of the effects of such tim- ber treatment de- pends upon the resistance offered by the material used to t| gradual solution by water. In the case of creosote the re- sults are quite satis- factory; with chlorid ° of zinc, subsequent Fig. 26. Fruiting bodies of the fatty Pholiota (Pholiota adipose, solution takes place in a wound of an oak tree trunk. (After Freeman). ; : : \ too readily, while with crude petroleum there is a tendency toward the evaporation of this substance when injected. The increasing cost of timber will stimulate timber treatments by making treatment profitable. One drawback at present is the necessity toimport creosote for use in such work; possibly refiinery by-products from petroleum of a character analogous to asphaltum may find application in timber treatment. DISEASES OF CULTIVATED PLANTS 353 ATMOSPHERIC CONDITIONS AS AFFECTING PLANT DISEASES The relation between weather and the prevalence of certain | plant diseases has been often recorded. The diseases which prevail are none the less parasitic, the difference exists solely in the temperature and moisture conditions of the atmosphere. Here we must distinguish clearly between the cause of the diseases and the conditions which favor the given diseases. Certain parasitic fungi develop more rapidly under cooler con- ditions than the normal or average while others are favored by higher temperatures; all fungi are favored by large amounts of moisture when these stop short of water immersion and shutting out the needed air. In temperature we have an optimum which usually lies within certain maximum and minimum limits for any given species, but this temperature optimum varies with the organism; it is a matter which admits of exact determination for any organism. As to moisture, an abundant supply of water is the optimum for most fungi with which we deal in plant disease investigations. In these atmospheric conditions of temperature and moisture the seasons of the year, in our climate, vary one with another. The seasons of heavy rainfall are commonly those of low temperatures by reason of the check on temperatures exerted by evaporation. Further, our weather service records show a tendency for our seasons to come in groups of cooler alternating with groups of warmer seasons; that is, we may have several years as with 1904 to 1907 (excepting parts of 1906) in which the mean monthly tempera- tures of those months which affect crops were decidedly below the normal or average. Evidently this normal lying as it does between the extremes, is surpassed by the warmer seasons which are said to be above normal. We have likewise, other alternating groups of years in which the season’s temperatures are decidedly above the normal. The effects of these cool seasons upon diseases are most clearly shown in outbreaks of leaf-curl of the peach and plum bladders in early season, and of potato late blight and rot, Phytophthora infes- tans, upon the potato crop. It is understood that plenty of moisture is the usual accompaniment ofa cool season; from the combined effect of this supply of moisture and cool weather we have outbreaks of the potato disease even in northern Ohio where it does not appear certainly to survive from year to year. Such groups of cool seasons culminate as arule in particularly injurious outbreaks of the potato Phytophthora with us;in more northerly situations, the disease is present nearly every season, but the outbreaks culminate with favorable weather conditions of excessive rains and lowered temperatures. 354 OHIO EXPERIMENT STATION: BULLETIN 214 Stress has been laid upon the downy mildew of potato and cucumber respectively. It must not be inferred that other diseases do not offer like contrasts between dry, hot seasons and those of heavy precipitation and low temperatures accompanied by relatively high atmospheric humidity. Mention has already been made of the greater prevalence of the shot-hole disease of the plum and leaf-spot of cherry, Cylindrosporium padi, in rainy seasons over drier ones. The same facts will apply with respect to practically all external parasites of plants as in the scab fungus on the apple, the rot of plum, cherry and peach, and to the countless number of foliage diseases with which we deal from year to year. Contrasting with the potato Phytophthora is the allied disease of curcurbits, the downy mildew, lasmopara cubensis, which appears to flourish during our hot seasons and to disappear during the cool ones where grouped as above described. The writer has suggested that this Plasmopara does not survive in our climate but is carried northward each year by its conidia alone; the extent of spread will thus be limited by the length of period favorable to it. This period must be one of relatively high temperatures since this parasite is more widely distributed near the tropics. All these instances only make more clear the intricacies of the mutual adaptations of parasite and host which have resulted from the long periods in which these dwell together. MEAN SUMMER TEMPERATURES AND RAINFALL IN OHIO, 1883-1909 Mean temperatures degrees Fahr. Mean Rainfall, inches Year Three 3 Mos. May June_| July -|August|/months}| May June | July |August] Total Mean inches 69. 72.1 68.2 69.8 5.72 4.25 -16 .88 | 10.29 1.1 71. 70.8 71.1 3.87 2.96 83 45 8.24 7.1 vin 68.9 70.4 3.97 34 20 .33 | 13.87 7. 72. 70.9 70.1 23 53 88 .62 | 10.03 L. 7. 77.9 75.6 87 85 16 .39 8.40 0.4 72. 70.4 71.0 77 41 4.40 16 | 12.97 66. 72. 69.1 69.4 i 4.13 25 .50 9.88 3. 2B. 68.8 VL? 5.52 4.50