/• s/'o- Bulletin No. 23. v. p. p. 76. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. GALLONA^AY, Chief. SPOT DISEASE OF THE VIOLET. Al 1 1 fitii ml lUdld' 11. sp. »v P. H. DORSETT, A-ssoeiate, Divi>sioii < .f Venetahle Physiolocjy and Pathology. ISSIIEP NOVEMBKR 2S, 1900. WASHINGTON: () <) \' !•; R N M I', NT 1' K I \ r I N ( ; ( ) K l' UK. I 9 (J (J. DIVISION OF VEGETABLE PHYSIOLOGY AN1> rATHOLOGY, SCIENTIFIC STAFF. B. T. G\\.}.oyfKy, Chief of Dmsnon. Albert F. Woods, Amstant Chief. associates. Erwin F. Smith, Oscar Loew, Mertox B. Waite, W.\i. a. Orton, Newton B. Pierce, P'rxst A. Bessey, Herbert J. AVebber, Flora W. Patterson, M. A. Carleton, Hermann von Schrenk,* P. H. DoRSETT, Marcus L. Floyd.''' IN charge of laboratories. Albert F. Woods, Plant Physiology. Erwin F. Smith, Phmt Pathology. Newton B. Pierce, Pacific Coaxt. Herbert J. Webber, Plant Breeding. Oscar Loew,* Plant Nutrition and Fermeniaiion. 1 Special agent in charge of studies of forest-tree diseases, cooperating with tlic Divisinn of Forestry, United States Department of Agriculture, and the Henry Shaw School of Botany, St. Louis, Mo. 2 Detailed as tobacco expert, Division of Soils. •'In charge of the tobacco-fermentation investigations' of the Division of Soils. Bulletin No. 23. V. V. p. 7G. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. GALLOWAY, Chief. SPOT DISEASE OF THE VIOLET. (Alternaria violm n. sp. ) BV P. H. DORSETT, Associate, Division of Vegetable Physiology and Pathology. Issued November 28, 1900. WASHINGTON: O O V i; R N M E N T PRINTING O K K I C E . I 900. LETTER OF TRANSMITTAL. U. S. Department of Agriculture, Division of Vegetable Physiology and Pathology, ^Vaddngton, D. C, Amjicst 29, 1900. Sir: I respectfiill}' transmit herewith a paper b}- Mr. P. H. Dorsett, of this Division, giving the results of some investigations of a disease affecting cultivated violets and generally known as spot. There is not less than a million dollars' worth of violet flowers sold every year in the United States, and were it not for the disease in question the amount would doubtless be increased 20 per cent. The annual loss from the disease, therefore, represents probably a money value of fully 1200,000. In view of the general interest in violet culture and the importance of the knowledge of a means of preventing the disease, I respectfully recommend the publication of the paper as Bulletin No. 23 of this Division. Respectfully, B. T. Galloway, Chief of Division. Hon. James Wilson, Secretary of Agricultxire. 3 CONTENTS. Page, Introduction ' General appearance of the disease - 8 Theories of the cause and treatment of the disease 9 Weakness of the plants . .■ 9 Improper soil conditions 10 Improper conditions furnished the plants during the growing and flower- ing season 10 Fungous nature of the disease 10 Conditions favoring the development and spread of the disease 13 Susceptiliility of varieties - 1"* Preventive measures 1^ Explanation of plates 1^ 5 ILLUSTRATIONS. Page. Plate I. Healthy and diseased leaves of ]\larie Louise violets 16 II. Young plants of Marie Louise violet from the cutting bed showing spot on their leaves 16 III. A healthy and a naturally infected ^larie Louise plant 16 IV. A healthy (control) and an artificially infected Marie Louise plant.. 16 Y. Plate cultures of Alternaria violx and mycelium and spores from a diseased leaf 16 VI. Development and germination of spores of Alternaria violse and pure culture upon Lima bean 16 Vn. Diseased and healthy leaves of California violet 16 6 SPOT DISEASE OF THE VIOLET. INTRODUCTION. The subject of this paper is one of the most widespread and destruc- tive maladies known to attack the violet. The disease has been dis- cussed in the florists' journals under a variety of names, such as leaf spot, leaf rust, leaf blight, smallpox, etc. More commonly, however, the trouble is known as the " violet disease," growers not generally recognizing the fact that there is more than one malad}^ attacking the violet. The disease occurs throughout this countr}' wherever the violet is grown, and is probably of American origin. The cultivation of the violet has been abandoned in many sections of the country on account of its ravages, while in others it has become necessary to adopt new methods of handling the plants during the growing season. Five or six years ago, for example, 50,000 to 75,000 square feet of glass in the vicinity of Alexandria, Va., were devoted to the cultiva- tion of this crop, but on account of the disease the industry has been practically abandoned. A large grower near Boston, Mass., was forced, a few years ago, to abandon growing stock plants at his place on account of this trouble. He had to have them grown for him dur- ing the summer, at considerable expense, in localities that were free or comparatively free from the disease. After transferring these plants to his place in the fall and setting them in the houses he expe- rienced little or no difficulty in keeping them healthy during the remainder of the season. Many other instances of the destructive nature of this disease could be cited. The large amount of florists' litcn-ature relating to this subject when collected and condensed was found by the writer to contain only a con- fused mass of contradictory opinions regarding both the cause and treatment of the disease. This is not strange to one familiar with the violet. All growers know the violet to be variable, seldom if ever behaving any two seasons alike. Practical growers recognize the fact that methods of handling the plants followed with little disease and good results during one season may, though rigidly adhered to, result in disease and failure the next. It is also a well-known fact that gi'ow- 7 8 e.rs in the same section and in close proximity to one another often prac- tice widel}^ different methods in growing this crop, and 3'et the results obtained are practicall}^ the same. A novice in violet growing may have little or no difficulty the first few years in growing good flowers. After this, however, his troubles usually begin and failure more often than success crowns his efforts. Unless he is possessed of peculiar abilities and a determination to succeed a few years of reverses are suf- ficient to cause him to abandon the culture of violets and turn his attention to some other industry where the chances of success are at least equal to those of failure. GENERAL APPEARANCE OF THE DISEASE. Spot disease of the violet {AUernaria molce) attacks the plants at any stage of their growth from the small unrooted cutting in the cutting bed to the mature plant in full flower. (See Pis. II, III, and IV.) Plants that are making a vigorous, rapid, but soft or succulent growth are most subject to the disease. The disease may occur on any por- tion of the plant above ground, but causes the greatest amoimt of loss when present upon the foliage. Its first appearance upon the leaves is characterized by small, definite, usualh' circular, greenish or yel- lowish white spots, resembling very much the ))ite or sting of an insect. They var}' in size from dots scarcely perceptible to the unaided 63^6 to spots a thirty- second of an inch or more in diameter. The light- colored central portion or point of infection is surrounded by a narrow ring of discolored tissue, usually black or very dark brown at first, but changing to a lighter shade as the spots grow older. (Pis. II, III, IV.) As the spot develops the central portion remains unchanged in appearance, while the tissues immediately surrounding it, either to one side or more frequently in a circle, become diseased by the ramify- ing growth of the mycelium of the fungus through this portion of the leaf. This usuall}- takes place within a few hours after infection. The freshly diseased portion of the leaf at first presents a water- logged appearance, frequently being semi-transparent, and is lighter in color than the adjacent healthy tissue. The diseased portion around the central point of infection in a few days fades or bleaches to a yel- lowish or grayish white, sometimes to a pure white, the time depend- ing somewhat upon the conditions of the weather. The development of the disease may stop at this point and the plants apparentlj^ entirely recover from its effects; in which event the diseased portions of the leaves after a few days separate from the healthy tissue and fall out, leaving the leaves full of holes. More frequently, however, the disease continues to develop in the parts of the leaf adjoining or sur- rounding those alreadj^ diseased. These freshly diseased areas in turn pass through the same changes as the parts previously attacked. Unless checked by some means the disease continues to spread in this 9 way until the entire leaf is destroyed. It is seldom, however, that a single spot upon a leaf develops to this extent. More frequentl}' the leaf is attacked at a number of different points (Pis. I, 11, III, IV), and as the disease progresses the spots become larger and one or more of them coalesce, forming large irregular areas or blotches upon the leaf. (Pis. I, II.) A well-developed spot of this disease therefore shows a light-colored central portion, the point of infection, partly or wholly surrounded liy alternate rings of dark and light colored tissue, the lighter colored portions as a rule being ver}' much broader and more conspicuous than the darker. (Pis. I, II.) The majority of these spots are usuall}'^ free from fungous spores exc(?pt under conditions peculiarly favorable to their development. Spores are produced, however, in great abundance upon most of them, especially upon the central or older portions of the spots, after the leaves have been placed in a saturated atmosphere for from twenty-four to forty-eight hours. It is frequently the case that spores are produced in sufficient numbers to be discernible by the unaided eye, but usually the aid of a hand lens or a microscope is necessarj^ to determine their presence. The spores are borne in chains on dark brownish hypha? that rise from the dis- eased surface. PI. V, fig. 2, shows a photomicrograph of some of the mycelium and spores of this fungus taken from a diseased spot in a living leaf. The spores break from their attachment and separate from each other easily, and being very small and light they are car- ried around liy currents of air and finally settle upon other leaves. THEORIES AS TO THE CAUSE AND TREATMENT OF THE DISEASE. Perhaps no subject relating to floriculture has received more atten- tion in the floricultural and horticultural journals during the past eight or ten years than the disease in question. The most varied opinions have been expressed in regard to it, and the explanations advanced as to its cause and the possible course of treatment are numerous. Some of the more important of these hypotheses are given here. WEAKNESS OF THE PLANTS. Some writers claim that the plants are of necessity weakened by being forced during the winter into heavy flower production, and that the taking of cuttings from such plants, and the rooting and forc- ing of them in the same way from year to year has resulted in pro- ducing a weak strain peculiarly susceptible to injury of all kinds. They reconnnond fall propagation to secure strong, vigorous, health}^ wood before the plants are weakened by flowering. The cuttings, after being rooted in clean, sharp sand, are transplanted into thumb-pots or into flats and carried through the winter in a house or in frames, where the temperature is kept as low as possible, not allowing the 10 plants to freeze, however. By this treatment the plants are given a rest, which is believed b}- many to be necessary to strong, vigorous growth. While growers generallj" admit that slightly better results are usuall}" obtained b}' this treatment than b}- the one generally prac- ticed, the}' are, as a rule, of the opinion that the benefits derived will not justify the expense necessary to carry the young plants through the winter in good condition for spring planting. This is an impor- tant problem, the practical solution of which would no doubt prove of great value to all interested in the cultivation of the violet. We have this work under way at the present time, and hope in a few years to obtain some interesting results. IMPROPER SOIL CONDITIONS. Other writers claim that the disease is due to improper soil condi- tions. The soil is either too heav}^ or too light in texture, and as a consequence holds, or gives up. too much or too little moisture, or con- tains too much or too little plant food. They advise selecting soil suited in every wa}^ to the best growth and development of the plants. Since good soil is one of the prime factors governing strong, vigor- ous, health}' plant growth, their advice is good, but extremely difficult to follow. The question of securing proper soil is one of the most perplexing with which the grower has to contend, requiring judgment that can be gained only by many years of practical experience. IMPROPER CONDITIONS FURNISHED THE PLANTS DURING THE GROWING AND FLOWERING SEASON. Still others attribute the disease to improper methods employed during the growth of the plants, such as growing them in the open field, where they are exposed to drought, rains, dews, and the direct rays of the sun during the summer, and lack of attention to properly heating, ventilating, and fumigating the houses and to cultivating, watering, and cleaning the plants. As a remedy they propose fur- nishing the necessary conditions for vigorous, healthy plant growth at all times. This is a good doctrine, but begs the question. FUNGOUS NATURE OF THE DISEASE. Over four years ago the writer succeeded in producing upon violet leaves spots that were in every way identical with those above described by spraying the leaves with distilled water to which spores of the fun- gus Alternaria vioim had been added. Since that time he has proved by nmuerous laboratory and greenhouse experiments (details of which will appear in the following pages) that the so-called ''spot disease" of the violet is unquestionably due to the attacks of this fungus. Other fungi, Cercosjxjra viohe Sacc, F/iyUostictavwhv Besm.. Sepforia violce Wesid., etc., are known to attack the violet, producing upon the 11 leaves spots very similar in outline and appearance to those caused by xllternaria vlolm (with which they are often confused), but in the writer's experience in the study of the violet and its diseases he does not recall a single instance where these fungi have come to his atten- tion as causing any serious trouble. It is possible, however, for them to do considerable damage under conditions peculiarly favorable to their development. Ninety-live per cent of all the specimens of the .so-called violet disease received at the Division laboratory during the past four or five years were found, upon careful microscopical examination, to contain spores of the particular fungus mentioned.' The fungus was isolated by agar poured cultures in Petri dishes, and comparatively little difficulty was experienced in securing pure cultures for inoculation experiments. The growth and development of the fun- gus on artificial media is, as a rule, quite rapid, normally producing spores in from four to six days after the sowing of the spores or the transferring of a single germinating spore from one plate culture to another. The growth of the fungus in agar is normally in concentric rings, each ring marking the amount of growth made in twenty-four houra (PI. V, fig. 1). The color of the fungus growing on agar before spore formation is grayish white (PI. V, fig. 3). Spore production begins at the center on the older growth, and gradually extends outward, until the entire surface of the colony is covered with a dense mass of olivaceous spores. The fungus grows well on other culture media, especially young lima bean pods (PI. VI, fig. 18). The first inoculation experiment with Alternaria molce was made February 12, 1896. Two plants of Marie Louise violet, in 4-inch pots, were removed from the Department greenhouse to the labora- tory. They were quite uniform in size and, as far as could be ascer- tained by observation, entirely free from disease. Plant No. 1 was sprayed with sterile distilled water and placed under a bell jar in a sat- ^ SCIENTIFIC DESCRIPTION OF THE FUNGUS. Alternaria violse Galloway and Dorsett. Amphigenous, but especially epiphyllous, olivaceous, velutinous, on light-colored subcircular definitely limited spots 2-4 mm. in diameter, extending into arid patches 10-12 mm. in diameter, which show one or more dark concentric lines; spore-bearing hyph?e fasciculate, erect, pale olivaceous, septate, simple, 4 l)y 25-30 a; conidia borne at or near tips of the hyphte, catenulate, clavately Hask-shapcd, muriform, strongly conHtricted at the septa, which are variable in number, oliva- ceous, 10-17 l)y 40-60 /.i exclusive of istlnnus, which is 3-5 by 3-25 //. Dr. Gino Pollacci has described and figured (Atti del K. Inst. Bot. dell'Univ. di Pavia (Laboratorio crittogamico) Ser. II, Vol. V, pp. 1-2, PI. VII, figs. 1-5, 1897) a spot disease of violet occurring in Italy which is due to a fungus which he names }f(icnjspnriiiiri rlolir. If his description and (h-awingsare correct the two diseases are quite distinct. 12 urated atmosphere, where it was kept. Plant No. 2 was sprayed with sterile distilled water in which spores from a pure culture of Alter- naria molce had been sown, and was then placed under the same con- ditions as plant No. 1. The temperature of the laboratory at the beginning of the experiment — 3.30 p. m. — was about 80- F. The fol- lowing- notes, made during the progress of the experiment, are descrip- tive of the results obtained: February 14, 1896, 9.30 a. m. Plant 1 apparently in a perfectly healthy condi- tion, leaves covered with moisture, but showing no ill effects from the spraying or from being kept in a saturated atmosphere. Plant 2 liadly diseased, nearly every leaf showing one or more spots of infection, which are in every particular identical with the first stages of the disease as naturally produced. February 15, 1896, 9.30 a. m. Plant 1 still remains healthy and apparently unin- jured by the treatment. On plant 2 the disease is progressing rapidly. There is a peculiarly disagreeable odor present when the bell jar is removed that is not noticeable under the same conditions with plant 1. This odor, so far as I am able to judge, is identical with that noticed with plants suffering from an attack of the disease under normal conditions. This odor is one of the characteristics of the dis- ease, and its presence in the house, frame, or field is usually the first intimation the grower has of the presence of the disease among his plants. February 19, 1896. Plant 1 still healthy and apparently in good condition. The spots on plant 2 are a little further developed and resemble more closely those produced under natural conditions. A striking example of the results obtained b}' artificial inoculations of violets with this fungus is shown in PI. IV. The two plants shown in this photograph received the same treatment as that given the two plants in the experiment described. The plants were sprayed at 3.30 p. m., August 26, 1896, and examined first at 2.30 p. m., August 27, 1896, just twenty-three hours after treatment. At this time plant No. 1 appeared free from disease, and showed no ill effects whatever from the treatment. On the contrary each leaf of plant No. 2, with one or two exceptions, showed from 1 to 30 or more spots of the dis- ease, which were in every way identical with those produced on plant No. 2 in the previous experiment. The plants were photographed August 31, 1896. At this date plant No. 1 was apparently free from disease, while the disease on plant No. 2 had made considerable progress and the spots w^ere gradually assuming the normal colorings which are characteristic of this disease. In these spots thus produced a careful microscopic examination demonstrated the presence of the mycelium of the fungus, and subse- quent observation showed that the fungus pushed through to the surface of the spots and fruited, whenever the leaf was put under a bell jar in moist air, exactly as it did on spots occurring naturally. The disease was again produced in health}- plants by inoculation with the spores- thus formed. That the spots produced upon violet leaves by artificial inoculations with spores of Alternaria violce closely resemble those occurring natu- 13 rally can be readily determined by comparing PL III, which is a photo- graph of a naturally diseased and a healthy plant from greenhouses at Garrett Park, Md., December 18, 1897, with PI. IV, which is a photograph of a healthy and an artificially infected plant. The simi- larity of the spots is, however, more strikingly shown in PI. I. Fig. 1 is a healthy leaf; fig. 2 is a diseased leaf from a plant naturally infected with the disease, while leaves 3 and 1 were taken from the diseased plant shown in PI. IV, which was artificially infected with the disease. Spores of the fungus Alternaria violw sown in water in a van Tieg- hem cell and kept at a temperature of 65° to 80° F. germinate readily in from one and a half to three hours. Fig. 9, PI. VI, is a camera lucida drawing of a group of spores that were sown in distilled water and placed in a van Tieghem cell at 10 a. m., January 15, 1898. The dotted lines at right angles to several of the germ tubes mark the amount of growth made by them between the time of sowing and the time noted; all subsequent growth and the production of all germ tubes not marked with a dotted line occurred between 11.55 a. m. and 2.10 p. m. the same day. Figs. 1 to 6 show a camera lucida drawing of a group of Alternaria spores in distilled water just previous to being- placed in a van Tieghem cell, and figs. 7 and 8, two spores in the same sowing, nineteen hours later. Figs. 11, 13, and 11: show spores that were sown in distilled water in a van Tieghem cell at 10.20 a. m., August 19, 1898, at which time they showed no signs of germinating. Four hours later, however, at 2.20 p. m., the time at which the draw- ings were made, the number of germ tubes had developed as indicated. Fig. 12 is a second drawing of fig. 11, made twenty minutes later. Fig. 10 is a camera lucida drawing of two spores from four to six hours after being placed in distilled water. Figs. 15, 16, and 17 show the chain formation of spores and their attachment to the mycelium. This drawing was made from a pure plate culture of the fungus. Numerous greenhouse and laboratory experiments under strict con- trol conditions have confirmed these results, and show that spot disease of the violet is due directly to the attack of the parasitic fungus Altermiria violce, and not to any of the other causes suggested. Indi- rectly, however, other conditions may have their efi'ect. Any one or a combination of all of the conditions included in the various theories advanced may cause the plants to become susceptible to the attacks of the fungus. CONDITIONS FAVORING THE DEVELOPMENT AND SPREAD OF THE DISEASE. The conditions favoring the development and spread of the fungus may be considered under two heads, viz, natural conditions and arti- ficial conditions. Among natural conditions those of the damp, warm, cloudy weather of the summer season are the most diflicult to modify or control. 14 Conditions of this nature are almost invariably present during the months of August and September. The da^^s are long and usually hot and dry, followed, as a rule, by cool, moist nights. The plants at this time are subjected to extreme changes, viz, from the hot, dry atmosphere during the da}", which frequently causes them to become wilted and remain so for several hours, to the cool, moist atmosphere of the night, which causes them to become excessiv^ely turgid. Conditions of this kind induce a I'apid, weak, soft, or succulent growth of the plants which is particularlj' subject to disease and at the same time favors the germination and development of the spores of the fungus. It is at this season of the year, as a rule, that the spot disease is most abun- dant and destructive. This is the time for great vigilance, and every condition influencing plant growth must be made as favorable as pos- sible to a hardy, health}^ growth which will be able to withstand the attacks of disease. The grower who is able to accomplish this and tide his plants over this critical period of their growth in a compara- tivel}' healthy condition is fortunate, and, as a rule, has little to fear from the disease during the remainder of the season. Artificial conditions include those wholly or in part under the con- trol of the grower. They are too often neglected, resulting as a rule in disease and consequent loss and discouragement. They may be enumerated as follows: (1) Not keeping the houses or frames clean, fresh, and sweet by frequently repairing and painting them and b}^ removing and destroj'- ing rubbish of all kinds as soon as it appears. (2) Not keeping the plants clean and in the best possible growing condition at all times. (3) Not selecting stock from strong, vigorovis plants that have been entireh' free from disease. (4) Not being careful to select only strong, vigorous, healthy stock from the cutting bed for planting in the spring. (5) Not giving the proper attention to the selection and preparation of the soil, to the date and method of planting, and to the care and cultivation of the plants during the growing season. (6) Not giving due consideration to the several varieties and their adaptability to the soil and location in which they are to be grown. SUSCEPTIBILITY OF VARIETIES. While the susceptibilit}" of the plant to disease depends largel}' upon the way in which it has been grown, still, as a whole, some varieties are more susceptible than others; Marie Louise, for example, even under conditions most favorable to growth, is more subject to injuiy from spot than is Lady Hume Campbell. The former variety can be grown to perfection only under the most favorable conditions, but when thus grown it has no equal for size, color, and excellency of flower. The hardier, more resistant, and more prolific variety Campbell stands 15 next to Marie Louise in qualit}" of flowers, lacking onl}' the deep rich color of the latter. The single varieties are as a rule more resistant than the double, though occasionally they are seriousl}" afi'ected. (Plate VII.) PREVENTIVE MEASURES. So far as we are aware there is at present no effective remedy for this disease when it has gained a foothold. The principal fungicides in common use for the prevention and check of plant diseases have frequently been tried for this trouble, but with varying results. The experiments of the Division in spraying violets with some of the more important of these, among them Bordeaux mixture and ammoni- acal solution of copper carbonate, seem to show that they possess little or no value in preventing the disease, while on the other hand they render the foliage worthless for bunching with the flowers, and thus occasion considerable loss and inconvenience. From the writer's experience and that of many others it would seem that the solution of this problem of controlling the disease lies in preventing it by giving careful attention to the production of vigorous, healthy, plant growth rather than in attempting to check the trouble after it has once gained a foothold. The successful growing of violets free from disease and the pro- duction of flowers of the best quality are governed by a number of factors which must be kept in mind. The principal rules which should govern the grower are the following: (1) Study carefully the behavior of the plants under the varying conditions surrounding them. Endeavor by modifying these condi- tions, when necessar}", to secure plants of ideal development. Set the standard of excellence high and be satisfied with nothing short of its attainment. (2) Grow the plants during the entire season where the}" can be given the conditions necessary for making a vigorous, healthy growth, and where they can be protected at all times from conditions likely to induce disease. (3) Keep the houses or frames clean, sweet, and in perfect condition for growing healthy plants, by repairing and painting them when nec- essary, and by removing and destro3dng all rubbish likely to harbor vermin or disease. (4) Propagate only from healthy, vigorous stock (;f known parent- age at the season most favorable to the plants. (5) Select each spring none })ut perfectly healthy, vigorous plants from the rooted cuttings for planting into the houses or frames. Old plants are sometimes carried over, and occasionally yield a large crop of flowers. They are not as reliable as the young plants, however, and are much more liable to all kinds of disease. The best growei's rarely use them if it is possible to secur(> strong, health}' young plants for spring or early summer planting. 16 (6) Keep the plants clean of yellow, dead, or dying leaves, being careful to destroy them after removing them from the plants, (7) Keep the plants free from insects and other animal pests. (8) Give careful attention to ventilating, heating, and shading the houses or frames and to watering, cleaning, and cultivating the plants. (9) Renew the soil in the beds each season before setting in the yovmg plants by removing from eight to twelve inches of the surface soil and replacing it with that freshh^ prepared, (10) Set the young plants early in the spring in the beds where the}^ are to remain during the season, so that they may get well estal^lished before the hot. dry weather of summer makes its appearance. Careful attention given to the above directions for a number of years will, it is believed, result in the production of a strain of plants that are not only practicalh' disease resistant, but are also ideal as regards regularity and symmetry of growth, length, and strength of flower stems, and 3'ield, size, substance, and qualit}' of flowers pro- duced. EXPLAXATIOX OF PLATES. Plate I. Healthy and diseased leaves of Marie Louise violet. (Natural size. ) Fig. 1, healthy leaf. Fig. 2, naturally infected leaf. Figs. 3 and 4, artificially infected leaves from the diseased plant shown in Plate lY. II. Young plants of 3Iarie Louise violet from the cutting bed, showing spot on the leaves. III. Fig. 1, healthy plant of ^larie Louise violet for comparison. Fig. 2, dis- eased plant (natural infection). IV. Fig. 1, healthy plant of Marie Louise violet (control). Fig. 2, diseased plant (artificial infection). Y. Fig. 1, growth of the fungus in eleven days from a single spore <>n an agar plate. Fig. 2, photomicrograph of mycelium and spores of Alter- narki violee from violet leaves. Fig. 3, pure plate culture of Alfemaria riola: YI. Figs. 1 to 6, inclusive, show spores as they appear when brushed from a diseased spot. Figs. 7 and 8, some of the same spores after nineteen hours in distilled water in a van Tieghem cell. Fig. 9, a group of ger- minating spores. The length of the germ tubes at the time of the first examination is indicated by the dotted lines, and time marked; all subsequent growth of these tubes occurred, and all unmarked tubes developed, between the time marked and 2.10 p. m. Fig. 10, two con- nected spores germinating at several points after l)eing about four hours in distilled water. Fig. 11, germinating spore. Fig. 12, the same spore twenty minutes later. Figs. 13 and 14, germinating spores. Figs. 15, 16, and 17, spores produced on agar, showing manner of attachment to mycelium and chain formation of spores. Fig. 18, pure culture of Alter- naria rioliv on sterilized lima bean. The darker part of the culture is thickly covered with spores; the white marginal portions are young grow- ing mycelium. VII. Fig. 1, healthy leaf < CO Bui. 23, Div. Veg. Phys. 8i Path., U. S. Dept. of Agriculture. Plate 11 Bui. 23, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. Plate IV. Bui. 23, Div, Veg. Phys. & Path., U. S. Dept. of Agriculture. Plate V. Plate Culture of Alternaria viol^, and Mycelium and Spores from a Diseased Leaf. Bui. 23, Div Veg. Phys. & Path., U. S. DeDt. of Agriculture. Plate VI. Development and Germination of Spores of Alternaria viol/E and Pure Culture UPON Lima Bean. Bui. 23, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. Plate VII Diseased and Healthy Leaves of California Violet. Bulletin No. 24. v p. p.— 77. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. GALLOWAY, Chief. THE BASIS FOR THE IMPROVEMENT OF AMERICAN WHEATS. BY MARK ALFRED CARLETON, Cerealist, Division of Vegetable Physiology and Pathology. Issued December 10, 1900. WASHINGTON : GOVERNMENT PRINTING OFFICE. I 900. DITISION OF TEGETABLE PHTSIOLOGT AND PATHOLOGY. SCIEXTIFIC STAFF. B. T. Galloway, Chief of Division. Albert F. Woods, Assistant Chief. associates. Erwin F. Smith, Oscar Loew, Merton B. Waite, Wm. A. Ortox, Newton B. Pierce, Ernst A. Bessey, Herbert J. Webber, Flora W. Patterson, M. A. Carleton, Hermann von Schrenk,^ P. H. DoRSETT, . Marcus L. Floyd.'* ^ IN CHARGE OF LABORATORIES. Albert F. Woods, Plant Physiology. Erwin F. Smith, Plant Pathology. Newton B. Pierce, Pacific Coast. Herbert J. Webber, Plant Breeding. Oscar Loew,^ Plant Nutrition and Fermentation. 1 Special agent in charge of studies of forest tree diseases, cooperating with the Division of Forestry, United States Department of Agriculture, and the Henry Shaw School of Botany, St. Louis, Mo. 2 Detailed as tobacco expert, Division of Soils. s In charge of the tobacco fermentation investigations of the Division of Soils. BUL. 24. PIV. VEG. PHYS. & PATH., U. S. DEPT. OF AGR. FRONTISPIECE. AHCEMftCO e-AUIMORH Bulletin No. 24. V. P. P.-77. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. OALLOWAV, Chief. THE BASIS FOR THE IMPROVEMENT OF AMERICAiN WHEATS. BY MARK ALFRED CARLETON, Cerealist, Division of Vegetable Physiology and Pathology. Issued December io, 1900. WASHINGTON: GOVERNMENT PRINTING OFFICE. I 900. LETTER OF TRANSMITTAL. U. S. Department of Agriculture, Division of Vegetable Physiology and Pathology, Washington, D. C, July 19, 1900. Sir: I have the honor to transmit herewith, and to recommend for publication as Bulletin No. 24 of this Division, the manuscript of a paper by Mr. M. A. Carleton, on The Basis for the Improvement of American Wheats. During the past ten years this Division has had under investigation a number of problems connected with cereal pro- duction, and in order to carry on this work intelligently it has been necessary to make a careful study of the wheat industry generally. To this end a thorough survey of the field has been made, and the results are brought together here. The bulletin will prove especially valuable as showing the lines along which further work must be carried on. Part of this work is already under way, and other lines will be taken up as rapidly as the means at hand will permit. Respectfully, B. T. Galloway, Chief of Division. Hon. James Wilson, SeOi etary of Agriculture. 8 CONTENTS. Page. Introduction 7 Personal exijlorations 9 Characteristics and needs of the several wheat districts of the United States .. .9 General needs of all the districts - 10 Yielding power 10 Early maturity 11 Soft wheat district - 12 Semihard winter wheat district 13 Southern wheat district 14 Hard spring wheat district 15 Hard winter wheat district 17 Durum wheat district - 18 Irrigated wheat district 20 White wheat district 22 Sources for desirable qualities - - 25 Characteristics of botanic groups of wheat 26 Common bread wheats ( Triticum valgare) 26 Club or square head wheats ( T. compactum) 28 Poulard wheats ( T. turgidum ) 29 Durum wheats ( T. durum) 30 Polish wheats ( T. polonicum) - - - 32 Spelt (T. spelta) ---- 33 Emmer ( T. dicoccum) 34 Einkorn ( T. monococcum ) 35 Geographic groups of wheats - - - 36 Improvements accomplished - 37 Introduction of new varieties 38 Work of the Department 40 Wheat breeding 63 Improvement by selection 64 Improvement by hybridization 69 Summary 77 5 ILLUSTRATIOXS. PLATES. Pasre. Fkontispiece. Map showing the distriljution by districts of the different nat- ural groups of ^Yheat varieties in the United States. Plate I. Wheat field near jNIount Vernon, Va 12 II. Fig. 1, Wheat fields of the Red River Valley near Grand Forks, N. Dak. Fig. 2, Self-binders at ^\'oi-k near Grand Forks, N. Dak 16 III. Fig. 1, Field of wheat on "Tule" lands near Stockton, Cal. Fig. 2, Steam combined harvester-thresher harvesting on "Tule" lands near Stockton, Cal 22 IV. Fig. 1, Bags of wheat just harvested on the Bidwell estate, Chico, Cal. Fig. 2, Wheat field near Tehama, Cal 22 V. Fig. 1, Harvesting with the combined harvester-thresher near Walla Walla, AVash. Fig. 2, Wheat fields before and after harvesting, near Walla AValla, Wash 24 VI. Fig. 1, Combined harvester-thresher at work near Walla Walla, Wash. Fig. 2, Harvesting with the wide-cut binder near Colfax, Wash 28 VII. Spelts and Emkorns in exi^erimental plats at Garrett Park, Md. 34 VIII. Fig. 1, Group of Russian wheats in experimental plats at Garrett Park, Md. Fig. 2, Experimental wheat plats at Garrett Park, Md. , showing earliness of King's Jubilee 62 IX. Hybrid wheats. Early Arcadian and Diamond Grit, shown by the side of the parent varieties 72 X. A comjjosite cross by the Gartons, showing samples of the prog- eny of the last cross 74 TEXT FIGURES. Fig. 1. Diagram showing pedigree of Early Genesee Giant Wheat 71 2. Diagram showing pedigree of one of the Gartons' hybrid wheats 74 3. Diagram showing pedigree of one of the Gartons' hybrid wheats 75 4. Diagram showing pedigree of one of Farrer's hybrid wheats 76 5. Diagram showing hypothetical cross of wheat and spelt 76 6 THE BASIS FOR THE IMPROVEMENT OF AMERICAN WHEATS. INTRODUCTION. In 1894 the Division of Vegetable Physiology and Pathology began experiments on an extensive scale to test the comparative rust resist- ance of different varieties of cereals, especially wheat. This work was carried on for three seasons at Garrett Park, Md., Salina, Kans., and Manhattan, Kans., respectively. An account of the results of this work has already been published/ so that it is unnecessary to refer to them in detail here. Suffice it to say that in the course of the work it became apparent that constant rust resistance is not to be obtained among the ordinary bread wheats known at present, though on an average a few such varieties are fairly resistant during a long period of years. By the results obtained it was rendered highly probable that this quality must be bred into a variety either by rigid selection of the most resistant individuals of that variety or by crossing with resistant varieties of other wheat groups and selecting from the result- ant progeny such types as combine in the highest degree the usual qualities of the bread-wheat group with that of rust resistance. It was found, moreover, that in regard to other qualities than rust resistance it is not possible to obtain varieties which even approximate perfection, and especially is it rarely, if ever, true that many desara])le qualities are found in the same variety. However rust resistant a cer- tain variety may be, it will usually l)c found lacking in some other ^sential quality, and manifestly the most perfect rust resistance is of no consequence if other essential qualities are absent. As a rule the wheats that are most highly resistant to orange leaf I'ust^ are not varieties of the common bread-wheat group {Trlttcxiii vi(lg((r<') at all. though it by no means follows that they can not be used in bread making. At the same time some of the most valuable sorts for bread Hour, including a number of Russian varieties, rust very badly in certain seasons. Occasionally good qualities may neutralize bad ones __^ ^ ^ • — ■ ■ 'Cereal Rusts of the United States, Bnl. No. 16, Division of Vegetalile Pliysioiogy and Tathology, U. S. Department of Agr., lS!)i>, by M. A. Carleton. "■"For descriptions of the two wheat rusts of this country and illustrations of their differences see Bui. No. Ki "I" this Division, a))ove referred to. 8 in the same variety. For example, a variety may be very susceptible to rust when attacked, but usually be able to escape it by virtue of its quality of early maturity. Consideration of such facts linally led to the determination to study thoroughly wheat varieties themselves in all their relations, and not simply wheat diseases. Such a study of course naturally presupposes the investigation of all associated problems, such as drought resistance, early maturity, yielding power, and other matters of great economic interest. The different phases of the subject of wheat culture in its broadest sense are so intimately connected that no one of them can be intelligently studied separate and apart from the others. During the iirst season (1895) of the investigations above mentioned about one hundred crosses were attempted with wheat varieties (besides a number with varieties of oats), mainly to determine the facility with which hybrids might be produced by crossing varieties of quite differ- ent groups. One-third of these crosses resulted successfully, unless a few of them ma}' possibly have resulted from accidental pollination. Some of them were readily effected between varieties of common wheat {Triticum vidgare) and the durums {T, duTimi)^ as well as between varieties of each of these groups and the poulards {T. turgi- dum). All the resulting hybrids were planted, but, the weather con- ditions of the following season being unusuallj^ severe, these and many of the other experimental varieties failed to survive. This work was continued, but in the meantime careful studies were being made in the several wheat districts with a view of determining the particular needs of each. In some districts greater hardiness of winter sorts is required; in others, varieties with a particularly tena*- cious chaff; in others, stiff er straw; in others, drought resistance, and so on. Varieties bred for North Dakota and Minnesota are of no value for California, and the best varieties for Texas would be useless in Montana. But aside from these considerations a knowledge of the different botanical groups of wheats is necessar}', in order to have at command all the sources from which ma}' be drawn the qualities reciuired for different districts. After five vears investigations it can bv no means be assumed that a full knowledge of the conditions of wheat culture and the demands of the country has been attained by this Division. Nevertheless, it is now possible to establish a reasonably complete basis upon which intelli- gent and systematic work may be accomplished — work that either could not be accomplished ut all from a narrower standpoint, or would require much more additional time than has been given to the acquire- ment of this foundation, and could not even then be as thoroughly done. PERSONAL EXPLORATIONS. At various times during the years 1894 to 1897 all the wheat States except New York, Pennsylvania, and the Pacific Coast States were pretty thoroughly explored by the writer, the conditions of soil and climate being noted and a careful study made of the nature and distri- bution of the wheat varieties. Finally, during the past season (1899), it became possible to make a similar investigation of such conditions in the Pacific coast and North Mountain States, special attention being given in this case to the region usually known as the Palouse Country, and also to wheat culture under irrigation. Naturally very valuable information was obtained through these personal observations, which will be of great use in future work in wheat improvement. During the summer and autumn of 1898, under the direction of the Section of Seed and Plant Introduction of this Department, an exploration was made of the greater part of European Russia, including the Caucasus, and of a small portion of the Kirghiz Steppes, as well as of Hungary and Roumania, in search of additional cereals for this country. A general report of this work has been published.^ In Huno-arv and Roumania no varieties better than our own were found that had not already been obtained from those countries. In Russia some very valuable sorts were secured, which together with four or five others yet to be received," give this country now practically ever3'thing of importance in the line of wheats from that, the second greatest wheat country of the world. All these explorations have been of great value in furnishing a long-desired opportunity for a comparative study of wheat varieties and the conditions of wheat environment in different countries. CHARACTERISTICS AND NEEDS OF THE SEVERAL WHEAT DIS- TRICTS OF THE UNITED STATES. From the standpoint of investigations so far made concerning the conditions of wheat environment and the adaptations of varieties in the United States, the country may be considered as divided into eight wheat districts, each possessing characteristics quite different from those of the others. In fact, in some cases they are as different from each other as though they lay in different continents. They are as follows: (1) The Soft Wheat district, including mainly the New Eng- land and Middle States; (2) the Semihard Winter Wheat district, including the North Central States; (3) the Southern Wheat district, including the northern part of the Southern States; (4) the Hard Spring Wheat district, including the Northern States of the Plains; ^ Russian Cereals adapted for Cultivation in the United States, Bui. No. 23, Division of Botany, U. S. Department of Agriculture, 1900, by M. A. Carleton. ■■'Since this was written these- varieties have all been obtained. 10 (5) the Hard AYinter Wheat district, including- the Middle States of the Plains; (6) the Durum Wheat district, including a part of the Southern States of the Plains; (7) the Irrigated Wheat district, includ- ing* in general the scattered portions of wheat area in the Rocky Mountain and Basin States; and (8) the White Wheat district, includ- ing the larger part of the Pacific Coast States. Just as these districts differ from each other in their characteristics, so do the particular needs of the wheat grower in each dijfJer widely from those of other districts. (See colored map, frontispiece of this bulletin.^) GENERAL NEEDS OF ALL, THE DISTRICTS. Before describing these districts separately, it will be well to note briefl}^ two general needs common to all of them. These are (1) greater yielding power and (2) earlier maturity. In the writer's experience these are found to be ever present needs, not onl}^ in all our own States but in all wheat countries. YIELDING POWER. This quality is of course always desirable, simply from the stand- point of obtaining the greatest possible profit from the same area. Nevertheless, on account of peculiar local conditions the demand for a large yield is given much more emphasis in some localities than in others. Besides, the need of a large yield does not always arise from the same cause, and in many cases it is not real, but only appears so because of defects in other regards. To illustrate, the Palouse country of Washington and Idaho may ])e taken as an example in contrast with that of the Southern States. In the Palouse country the regular aver- age yield is already probably near 25 bushels per acre, while 35 or 40 bushels per acre is a common crop in certain seasons, and 60 bushels not particularly rare. Yet from no part of the country has the writer had more requests for information concerning larger-\^ielding varieties. As a matter of fact prices of wheat are proportional^ so low on account of the great distance from good mai'kets, and the method of summer fallowing, which allows a crop only ever}^ second j^ear, is so ' It has been a most difficult matter to prej^are this map, and it is not claimed that it is accurate. Indeed it would he impossible at present to prepare an accurate map of this nature. But it represents ajjproximately the different wheat districts charac- terized mainly by the cultivation of certain natural groups of wheats. Of course the different groups will lap over more or less from one district to another. In all that part of the United States approximately east of the one hundred and fourth meridian the uncolored portions represent territory either from which we have no statistics, such as the Indian Territory, or in which the wheat itroduction averages less than 1 bushel to the square mile. West of this line the Mhite portion represents territory in which there is practically no wheat grown at all. The reports of the census of 1890 and those of the Irrigation Division of the Geological Survey have been of much help in the preparation of the map. 11 much practiced that to overcome losses in these directions exceeding'ly laro-e yields are considered necessarj' in order that much prplit may be gained in the end. On the other hand, in the Southern States the problem of increasing the yield is entirel}^ independent of deficiencies in other regards, for the home demand alone is sufficient to make prices good as a rule; but the average yield is extremely low, being under 10 bushels per acre. It would add one-half to the profit in these States if the yield could be increased even to the average of the entire country (slightly over 13 bushels per acre). In the South manuring the land must also be practiced in order to obtain the best results, which is an item not at present considered in the West. In the States of the Plains the actual average yield is also rather low (a little over 12 bushels), so that here, too, the reason for a demand for an increased yield is evident and is usually independent of other deficiencies. The average yield for the United States is far lower than it ought to be. The yield for the semiarid districts, which is much less, can and should be as high as that for the entire country at present. EARLY MATURITY. There is no part of the United States where early maturing wheats ar(^ not desirable for one reason or another. The reasons are various in difi'erent localities. As before stated, early ripening varieties are, in most seasons, more likely tp escape damage by rust. In a large por- tion of the country this is a very important matter for consideration, but especially so in the Southern States and the States east of the Mis- sissippi River, where the whole wheat crop is occasionally entirely destroyed by this parasite. But the need of early maturity is most ui-gent in the Palouse country, as the shriveling efi'ects of the annual dijought in that region which sets in just before harvest may be avoided l)y the use of early varieties. In the North Central States and the Great Plains region early maturing and winter varieties are less liable to the ravages of chinch bugs than are late maturing and spring varie- ties. In all the Northern States early maturity also allows the variety a l)etter chance to escape early autumn frosts. There are instances in which late maturity is apparently an advan- tage, but such cases are rare. Finally it should be noted that there is quite a distinction between early wheats and early-sown wheats. A late-maturing wheat will ripen earlier than usual if sown earlier, or will ripen still later than usual if sown later. In the case of winter wheats early seeding allows the wheat plant to accunmlate more reserve force in th(^ roots during the autunni, tluis enabling it to begin growth with greater vigor in the spi'ing and get the start of the later-sown crops. In the case of spring sorts earlier seeding, of course, simply I'naljles the crop to get an earlier start and 12 thereby to ripen earlier. By early sowing and the constant selection of the earliest ripening heads for seed a naturally late wheat may bj gradually transformed into an early variety. SOFT WHEAT DISTRICT. In this district are included approximately New York, Pennsylvaj nia, New Jersey, Maryland, Delaware, and portions of Virginia (Plate I), West Virginia, and eastern Kentucky; also such portions of New England as produce wheat to any considerable extent. The region is characterized on the whole by the production of rather soft wheats, containing a large amount proportionally of starch, though occasion- ally they incline to semihard. The color of the grain is usually yel- lowish white or amber, but sometimes quite reddish. The soil, especially if not heavily fertilized, does not possess the necessary amount of alkali, phosphate, and humitied organic matter required for the production of hard, glutinous wheats. Moreover the climate is against their production, being too moist and cool in summer. Nevertheless in New York and Pennsylvania, by means of the plenti- ful application of fertilizers and the unusual attention paid to seed selection practiced in this region, a large amount of good wheat is annually grown in proportion to the entire area. Twenty-five or thirt}^ years ago, when the area given to wheat culture in this country was much more limited than at present, and when the hard red wheats were not so popular. New York had a deservedly great reputation both for her wheat production and flour'industry. And even at pres- ent, if there is a diminution of this reputation, it is not because of any actual decrease in wheat and flour production, but because of the over- shadowing increase in districts more favorably conditioned or situated, though we should add to this the fact that there has been a corre- sponding change in the kind of wheat used for bread making. The fact that so high a standard is maintained in the wheats of this region in the face of adverse natural conditions, is strong proof of the importance of intelligent wheat culture, particularly in respect to seed selection and the proper treatment of the soil. In some localities of this dis- trict the standard is considerably above what one would expect, while in some other districts it is far below what it should be. In the most northern portions of this district spring sowing is almost entirely practiced, and there is a need for hardy winter sorts which will be able to extend the winter-wheat area farther northward. In some localities rust is occasionally very injurious, the black stem rust sometimes completely destroying the crop. Early maturing and rust resistant sorts are therefore desirable for escaping or overcoming the attacks of this parasite. Bui. 24, Div. Veg, Phys. & Path., U. S. D-pt. of Agriculture. PLATiI I. 13 SUMMARY OF CONDITIONS AND NEEDS OF THE DISTRICT. (1) Chief varieties now grown: Fultz, Fulcaster, Early Genesee Giant, Longberry, Jones's Winter Fife, Mediterranean, Eed Wonder, Early Red Clawson, Gold Coin, ' Blue Stem. (2) Average yield per acre, about 14| bushels.' (3) Needs of the grower: (a) Harder-grained, more glutinous varieties. (b) Hardier winter varieties for the most northern portions. (c) Early maturity. (d) Rust resistance. SEMIHARD WINTER WHEAT DISTRICT. Ill this district we may include Ohio, Indiana, Illinois, Michigan, and a small part of Wisconsin. It produces a wheat of medium qualit}^, and on the whole is one of the most important cereal regions of the United States. The wheats grown are generally semihard, rather reddish in color, and either bald or bearded. Throughout this district, as well as over a large portion of the country, there has been a decided tendency during the last twenty years or more toward the use of harder red wheats and also of a larger proportion of winter compared with spring varieties. The increasing use of the harder wheats has been coincident with the advent of the roller-milling proc- ess, but not necessarily a forced result of the latter, as some have inferred. The two have worked together. The proportion of such wheats now grown in this region is nuich larger than ten years ago. Especially is this true in Michigan, where special impetus has been given to such improvements through the efforts of Prof. R. C. Kedzie, assisted by the millers of the State. Similarly the area in which it is considered possible to grow winter wheats has been extended much farther noi'thward, now including practically all of Michigan, nearly all of Illinois, and even a small portion of AVisconsin. Thus this group of States may now be properly called the semihard winter wheat district. These changes have been accomplished by the grad- ual introduction of hardier winter sorts, which are at the same time usually harder and red grained. Nevertheless there has been little more than a beginning in these improvements, and there is still a demand for hard red wheats, and in the northern portion of the roo-jon for hni'dicr winter varieties. The black stem rust is sometimes very d(>structive in these States, particularly in the lower, moist, and timbered portions of Ohio, Indiana, and Michigan. Hence there is great demand also for rust resistant sorts. 'Calculated ai^ accurately as possible from data collected by the Division of Statia- tics of this L)ei)artmeut covering the perioil IS'.IO-lHUy. 14 SUM>rARY OF CONDITIONS AND NEEDS OF THE DISIRICT. (1) Chief varieties now grown: Fultz, Poole, Eudy, Valley, Early Red Clawson, Nigger, Dawson's Golden Chaff. (2) Present average ji.eld per acre, about 14 bushels. (3) Present needs of tlie di«trict: (a) Hardness of grain. (b) Eust resistance. (c) Hardy winter varieties. SOUTHERN ^VHEAT DISTRICT. In area this district includes the larger portion of Kentucky, Vir- ginia, West Virginia, and North Carolina, all of Tennessee, and portions of South Carolina, Georgia. Alabama. Arkansas, and Missouri. The annual production of wheat is comparativeh' small, and is furnished principally by Kentucky, Missouri, Tennessee, and Virginia. In the greater portion of the region the combination of great rainfall with mild temperature is not conducive to the greatest success in wheat growing. The soil is also generally not of the best for such purposes. Rust is always ver}^ bad, because of the constantly damp, warm climate. In spite of these difficulties there is no doubt that with sufficient effort the wheat industry might be very materiall^^ improv-ed. Just recently there has been much interest awakened in the possibilities of success- ful wheat culture, particularly in Georgia and South Carolina. This increasing interest in the matter finalh' resulted in the calling together of a convention at Macon, Ga.. in July, 1899, when it was unanimously decided that Georgia can veiy easily and should supply her own demands for wheat for bread making. Man}^ members of the conven- tion gave very favorable testimon}" regarding their own experiences in wheat growing during the past year. Probably one of the greatest obstacles in the way of profitable wheat raising in portions of the South is the lack of good flouring mills, much of the grinding being at pres- ent performed by the most primitive of gristmills. With a continued increase in wheat acreage there will perhaps be a corresponding increase in the number of iii'st-class mills constructed. On account of the severe rust attacks which occur in this district it is highl}' desirable to grow early ripening and rust resistant sorts. But there are really not many early matui'ing wheats grown in this country, and of the early foreign varieties already tested none have yet proved to be sufficiently hardy. Canning Downs, an early Austra- lian sort, winterkilled even in so mild a region as Mississippi.^ How- ^See Tracy, S. M. T^Tieat. Sixth Annual Eeport Mississippi Agricultural Expert iment Station, 1893, pp. 23-25; also Eighth Annual Eepoi't, 1895, pp. 44-46. 15 ever, there has not been a sufficient number of trials of such varieties, and the different experiments have not been often enough repeated to give reliable results. As to the matter of rust resistance, experiments made in Louisiana^ showed that hard red wheats, including a number of Russian origin, resisted rust the best. In Mississippi two Austra- lian varieties, Beloturka and Defiance, were quite rust resistant, while varieties obtained from England rusted very badly. ^ Occasionally wheat is much injured in the noi'thern portion of this region l)y late spring frosts. It is on such occasions that late-maturing wheats and late-sown crops may have the advantage, since those ripen- ing early are likely to be caught by the frost just at blooming time and be prevented from "filling out," while the later ripening crops, blooming after the frost, escape such injury. It seems possible, how- ever, to p-row varieties that will resist the action of these frosts, and therefore varieties hard}' in this respect are desirable. The wheats at present grown in the Southern Wheat district are either soft or semihard, and usually amber or reddish in color. They are either bearded, as in the case of the Fulcaster, or beardless, of which the Fultz and May wheats are examples. In Arkansas and the Carolinas, Nicaragua wheat, a durum variety, is grown somewhat, but to no great extent as yet. Wheat from the Southern States is always more likely to be infested with weevil than that from other districts, and occasionally much annoyance as well as injury to the grain results from this cause. Nicaragua and the hard red wheats are more resist- ant to weevil than are the soft wheats. SUMMARY OF CONDITIONS AND NEEDS OP DISTRICT. (1) Principal varieties at present grown: Fultz, Rice, Fulcaster, Everett's High Grade, Red May, Bough ton, Currell's Prolific, Purple Straw. (2) Present average yield per acre, about 9| bushels. (3) Needs of the grower: («) Rust resistance. (6) Early maturity. (c) Resistance to late spring frosts. (d) Stiffness of straw. HARD SPRING WHEAT DISTRICT. The hard spring wheat area comprises the States of Minnesota, North Dakota, South Dakota, the larger part of Wisconsin, portions of Iowa and Nebraska, and small portions of Montana and Colorado. 'SeeStub1)s, W. C. Experiments in wheat. Louisiana Agricultural Experiment Station Bulletin No. 19, 1892, 2(1 scries, pp. 555-.562. ■•'See Tracy, S. M., in Mississippi Agricultural Exiieriment Station reports above cited. 16 In this district, because of the rich, black soil and dr}-, hot suminoi's, there is grown the highest grade of spring wheat in the world, except- ing the spring varieties of the middle Volga region in Russia, which are very siniihir. Two general types of wheat prevail throughout this district — the Velvet Blue Stem' and the Fife. A large proportion of the farmers in this region know no wheat which does not belong to one of these types. The chaff of the Velvet Blue Stem is covered rather closely with small hairs, and the plants are bluish gray near harvest time. In both types the heads are beardless and the grains are medium or small, hard, and red. There are several strains or varieties of each type. The gluten content of these wheats is comparatively very large, and especially of that quality which gives great lightness in bread making. The average annual wheat production of this district is larger than that of any other siuiilar area in the WT)rld, and is about 30 per cent of the entire production of the United States, The average yield per acre, however, is not very large — certainly far below what it might be. Almost everywhere the self-I)inder is used in harvesting the grain, and in some localities the farms given entirelv to wheat culture cover many thousand acres. (See Plate II.) On these bonanza farms 50 to 100 self -binding liarvesters are sometimes at work at the same time. The large size of the farms is one of the worst features connected with wheat growing in the Northwest. From this cause not enough atten- tion is given to details of the work. Operations delegated to the best of foremen and other emplo3^ees are never so carefull}' performed as when done under the direct scrutiny of the man who owns the farm, and wliose interests are therefore at stake. Little things that are of importance when summed up are overlooked. The tillage is not thor- oughl}^ accomplished, weeds are not kept down, there is more or less waste of land, and the grain is allowed to degenerate in quality. The needs of the grower in this district are not so great as in some others, though there is much to l)e desired. In the northern portion earliness of maturity is needed to enable the wheat to escape the early autumn frosts which sometimes catch the crop before harvest, while in the southern portion chinch-bug depredations and rust attacks might often be avoided through possession of the same quality. A combina- tion of earliness and rust resistance in the same variety would be espe- cially desirable. The average yield could be made ver}^ much larger, as already stated, but this is a matter depending fully as much on methods of culture as on the improvement of varieties. Proper seed selection, ' There are apparently four distinct varieties of so-called Blue Stem in the United States. The name Velvet Blue Stem is adopted here to designate the spruig variety grown in this district. The one grown in the Palouse country will 1)e called Palouse Blue Stem. Bui ?4 Div. Veg. Phys. & Patn.. U. S. Dept, of Agriculture. Plate II. Fig. 1.— Wheat Fields of the Red River Valley, near Grand Forks, in. Dak. (Original.) ■ !'■ ' » ». Fig. 2.— Self-binders at work near Grand Forks, N. Dak. (Original. 17 however, .should be rigidly practiced. The (\stiil)li.shuu'nt of hardy winter varieties in place of the spring -varieties now grown would no doubt be an improvement of the utmost value in Iowa, Nebraska, and portions of Wisconsin, and perhaps a small part of Minnesota. This border is now the battle ground between winter and spring varieties, and it should be the constant aim to carry the line farther to the north, thus increasing more and more the winter-wheat area. Such purpose can be accomplished either (1) by the introduction of winter varieties, of similar quality to the spring sorts now grown, from the Crimea, north Caucasus, and southern Volga region of Russia, or (2) by the actual origination of hardier winter varieties of good quality through hybridization and selection. As an example of the effectiveness of the former method, we have only to point out the work already accom- plished by Turkey wheat — a Crimean variety — in extending the Avinter- wheat area in Nebraska and Iowa. SUMMARY OF CONDITIONS AXli NEEDS OF THE DISTRICT. (1) Principal varieties at present grown: Saskatchewan Fife, Hayne's Blue Stem, Scotch Fife, Bolton's Blue Stem, Powers Fife, Wellman's Fife. (2) Average yield per acre, about 13 busliels. (3) Needs of the grower: {a) Early maturity. {})) Rust resistance. (c) Hardy winter varieties. (d) Drought resistance. HARD WINTER W^HEAT DISTRICT. In this district is comprised approximately the middle States of the plains, including Kansas, a large part of Missouri, portions of Iowa and Nebraska, and the larger part of Oklahoma. As the name implies, it is characterized b}- the production of hard winter wheats, such wheats as are rareh' found, but which are of the veiy best ({ualit}'. The onl}^ other wheat region in all the world tliat is exactly com[)ar- able to this one, so far as known, is that including northern Crimea and the country directly between the Sea of Azov and the Caspian Sea. The latter region, however, at present produces better wheats than are produced in this district, and therefore should ])e drawn upon for all improvements that are attempted through introduced sorts. The wheats of this district luue slender, stiff stems, narrow com- pact heads, usually bearded, and medium or small, hard, red grains. In this region there is tiie most interesting exampU* of the changes that may take place for the better in the development of the wheat industry. Twenty-five years ago the softer wheats (often white- grained) were chiefly grown oxer a large portion of this district, and .1871)— No. 2i '2 18 the cases of winter wheat sowing- as against spring wheat sowing were much fewer than at present. Now the hard red-grained varieties are principal!}' used, and only in Iowa and Nebraska are spring varieties grown to any extent. The introduction of these hard-grained winter sorts has added remarkably to the certainty and value of the wheat crop, and has greatly decreased the ravages from rust and chinch bugs. Such improvements are after all l:)ut fairly begun, and there is yet great demand for hai'd-grained sorts and varieties that will resist the winters of Iowa and Nebraska. As the wheat area extends farther westward — to the one hundredth meridian and beyond — there is also a special need for drought-resistant sorts. In fact, in this and the dis- trict just described there is the most exacting demand of the entire country for hardy varieties. The extreme severit}^ of the drought and winter cold combined forms a greater obstacle to winter wheat cul- ture than exists in any other district. The average j^ield per acre is alwa3's low, but the problem in a considerable portion of the region is not so much to increase the yielding power per acre as to make sure of a crop every year, since there are so man}^ complete failures from drought. A constant average of even 12 to 15 bushels per acre from year to year would be considered good.^ Early maturity is of importance in this district in order to allow an escape from the worst eflects of the drought in the western portion and from the rust in the eastern portion. Rust resistance is also important, but not so much so as in States east of the Mississippi River. SUMMARY OF CONDITIONS AND NEEDS OF THE DISTRICT. (1) Chief varieties at present grown: Turkey, -^J^ay, Fulcaster, Zimmerman, Fultz. (2) Average yield per acre, alj(jut 12| bushels. (3) Needs of the grower: (a) Hardy winter varieties. (b) Drought resistance. (c) Early maturity. DURUM WHEAT DISTRICT. The area contained in this district is comparativelj^ small and includes a large part of north-central Texas, the southwestern portion of Oklahoma, and a small portion of the southwest corner of Kansas. It also properly includes a portion of Colorado, but can not be so indicated on the map, as the particular portion is not yet definitely outlined. Some of this region (southwestern Oklahoma) has only ^ The problem of successful wheat growing in arid regions is receiving .earnest consideration and will be discussed in a later publication. 1^) recently been opened to .settlement, but wheat culture ha« developed rapidly in the new lands. The soil is generally black and rich in humus, just as in the district last described, and produces wheats with a large gluten content, which quality is further increased in the west- ern portion by the dry. hot summer weather. The general demand is for hard-grained, drought-resistant varieties, and such sorts are already grown to a considerable extent. In recent years there has been an increasing tendency toward the cultivation of the durum or macaroni wheats, the chief variety grown so far being Nicaragua which has become quite popular. This variety is very hardy, yields well, and the grain is extremely hard and glutinous. It is quite simi- lar to Ku))anka. Arnautka, and other macaroni wheats grown in southern Russia, and for which there is so much demand in France and Italy. Notwithstanding the usual notion concerning such wheats, Nicaragua has been very successfulh" ground into flour by a well- known milling company at lort Worth, Tex. By mixing slightly with other wheats an excellent bread flour is made. However, the chief profit to ])e gained frc^ni the cultivation of this variety in futui'e will no douljt arise from its use in manufacturing macaroni, just so soon as the possibility of furnishing a sufficient supply becomes cer- tain. Though its distribution is not yet very wide, Nicaragua is, nevertheless, grown over a lar^-e portion of Texas and also sparingly in Oklahoma and Colorado. For this reason, and because of the evi- dent adaptation of such wheats to this region, it seems proper to call it the durum Avheat district. These durum wheats grow rapidly, are tall, and have wide leaves with a harsh surface, and large heavy-bearded heads, compactly formed. The grains are very larg(> and long, and yellowish-white in color, becoming darker the blacker the soil in which the crop is grown. It being once proved that durum wheats succeed well, there is bound to be a still greater demand for them, so that the further introduction of such varieties l)ecomes at once one of the needs of the district. Aside from macaroni varieties, the red-grained winter wheats, similar to those described for the Hard Winter Wheat district, are best adapted for the larger part of this region. The best example is the Mediter- ranean, which is very conmionly grown. In central and southwestern Texas rust is very destructive, so much so that wheat cultuie has been completely abandoned in many places on account of it. There is, therefore, a great demand for rust resist- ant varieties. The durum wheats have the advantage of being highly resistant to orange leaf rust, but succumb to black stem rust. In the western poi-tion of the district the oft-recurring droughts are very detrimental, and therefore in that ])<)rti()n drought resistance and (>arly maturity are important (lualities. 20 SUMMARY OF CONDITIONS AND NEEDS OF THE DISTRICT. (1) Chief varieties at present grown: Mediterranean, Fulcaster, Nicaragua, Turkey. (2) Average yield per acre, II5 bushels. (3) Needs of the grower: (a) Macaroni varieties. (6) Drought resistance. (c) Rust resistance. (d) Early maturity. IRRIGATED WHEAT DISTRICT. In this region is included all those scattered portions of the Rocky Mountain and Basin States in which wheat is grown at all. The States thus included are Wyoming, a part of Montana, southern Idaho, Utah, Nevada, Arizona, New Mexico, and the greater part of Colorado. In this district we find conditions remarkably difi'erent from those exist- ing anywhere east of the Rocky Mountains. Three striking charac- teristics not present to so great a degree in any other district are (1) the extreme aridity, necessitating the application of water by irriga- tion, (2) the very low humus content of the soil, and (3) the superabun- dance of alkali usually present. These conditions are closely inter- related and mutually dependent upon one another. The absence of humus is a natural result of the absence of rainfall, upon which depends the existence of plant life. Rainfall also tends to equalize the distribution of the alkaline matters of the soil, which in this district, however, are concentrated, in places, in high percentages. The prac- tice of irrigation is often allowed to make conditions worse by grad- ually carrying and depositing in certain localities or on certain farms an excess of alkali largely above that which was already present. These features of extreme aridity, lack of humus, and excess of alkali are so particularly characteristic that they go far beyond any matters of temperature dependent upon latitude or elevation in their effects upon the nature of wheat varieties grown in this district. That is, wheats so far north as southern Idaho are very like those of southern New Mexico or Arizona, and in all parts of the district show uni- formly a great lack of gluten content, which is dependent mainly upon the presence of soil humus. Wheat does best in soil that is alkaline rather than acid in reaction, but an excess of alkali becomes very injurious. Different cereals are able to withstand different amounts proportionally of alkali in the soil. Barley and rye seem to tolerate a larger proportion than wheat, and the latter will usually tolerate a larger amount than oats. Of all the cereals barley will withstand the largest amount. The wheats of this district are almost always white-grained, soft, and 21 extremely starehy, and lack greatly in gluten content. The straw is so white and clean and glistening- that it is dazzling to the eyes in the hot sunshine. Rust on wheat is seldom injurious, and in some locali- ties is entirely unknown. Smut, howeyer, is often present to a con- siderable extent. The stiffness of the straw and the absence of rain preyent the grain from eyer lodging, so that haryesting may be delayed for weeks with little or no injury to the grain. Manifestly the greatest need of this district is an increase in the gluten content of the grain. While the introduction of hard-grained nitrot'-enous sorts from other sections is at first an improyement, the gluten content can not thus be materially and permanently increased. No wheat yariety, whatever its nature, can abstract from the soil ele- ments that are not present there. Wheats l)rought f rom the black prairie soils of other sections to this district show the most striking illustration of the radical changes that may be caused in a yariety by a simple transference to a new locality, and, eyen when grown under the best of care, quite etfectually disprove a notion prevalent eyen among scientists that yarieties will not deteriorate. The hardest red Fifes from North Dakota, Turkey wheat from Kansas, or Diamond Grit from New York become rapidly more starchy and of a lighter color on being grown in Utah or New Mexico. The first requisite, therefore, for wheat improvement in irrigated sections is the complete amelioration of the soil by (1) dispersing the excessive accunuilations of alkali and (2) increasing the humus content through the application of nitrogenous fertilizers and the growth of leguminous crops in alternation with wdieat. At the same time it will aid greatly to gradually introduce the harder red-p-rained wheats. In many portions of this district, at high elevations in the moun- tains, wheat is often seriously damaged by early autunm frosts. It is therefore important to obtain for these localities the earliest maturing varieties possible, or varieties that may perhaps resist the action of the frost. For example, in the San Luis Valley of Colorado wheat is trrown at an elevation of over 7,500 feet, where frost is likely to occur in any month of the year, but is especially liable to injure the crop in August. SUMMARY OF CONDITIONS AND NEEDS OF THE DISTRICT. ( 1 ) Chief varieties now grown : Sonora, Little Club, Taos, Defiance, Felspar, Amethyst. (2) Averajje yield per acre, about 21 bushels. (3) Needs of the grower: (a) Increase of the gluten content, (i) Pearly maturity. 22 whitp: wheat district. This district covers, in a general way, the Pacific Coast region, in- cluding California, Oregon, Washington, and northern Idaho. All varieties that have become at all acclimated are characteristically white- grained, soft, and starchy. Usually the factor which is probably most influential in producing a grain of such nature is the lack of humus in the soil, as is true in the irrigated district. The generally cool sum- mers, however, no doubt give aid to the same end. Hard red-grained varieties, when Ijrought to this district, deteriorate in a few years time. Nevertheless such introductions have in a number of instances proved beneficial. A majorit}^ of the more common varieties strictly characteristic of the district are of the group usually called club wheats and belong to the species Trlticmit cmn].)actum. Sonora, Defiance, and Australian of California, Red Chaff of Oregon (distinct from the Palouse Red Chaff' of the Palouse country), and Palouse Blue Stem of Washington are not, howev^er, club wheats. As the botanical name of the clul) group implies, these wheats have their spikelets (meshes) so compactly arranged in the heads that they stand out nearly at right angles with the rachis (or stem of the head). The head thus becomes squarely formed (hence the name square head applied to many of the varieties), and, ])eing usually a little larger at the apex than at the base, Appears club shaped. Thus, although the heads are usually rather short, each contains comparatively a large number of grains, which partially accounts, probably, for the large yields per acre in this district. Heads of Chili Club are occasionally found that contain over 160 grains each. A very valuable characteristic of the club wheats is their ability to hold the grain in the chaff so that there is little danger of shattering, even durinof the driest season, if there should be much delav in the harvest. In some localities the grain, though ripening in July, is sometimes left standing till September before harvesting, a habit which, however, has no good excuse for its practice. For the purpose of clearer discussion, the district may be considered as subdivided into three sections — California, Oregon, and the Palouse country of Washington and northern Idaho. In southern California the varieties Sonora and Defiance are much grown, the latter particularly for its rust resistance, which is an im- portant need in this part of the State. Sonora wheat has a reddish velvet chaff, is beardless, and is white-grained as seen in this district. The grain is a little harder than that of the club wheats and is used for export, while the grain of the latter is used for home consumption. From the latitude of Fresno to the Oregon State line Austndian and the various strains of club wheats are principally cultivated. The best known varieties that are given special names at all are Golden Gate Club, Salt Lake Club, and Chili Clul). The variety Propo is also Bui. 24, Div. Veg. Phys &. Path., U. S. Dept. of Agriculture Plate III. Fig. 1.— Field of Wheat on "Tule" Lands near Stockton, Cal. (Original.) Fig. 2.— bTEAM Combinld Harvester-thresher harvesting on Tule Lands near Stockton, Cal. i Original. - Bul^ 2A Div. Veg. Phys. & Path.. U. S Dept. of Agriculture. Plate IV. Fig. 8.— Bags of Wheat just harvested on the Bidwell Estate, Chico, Cal. (■Original. .v •,■ •v;:v'?' Fig. 2.— Wheat Field near Tehama, Cal. < Original. 23 grown to some extent. Other sorts from the East, such as Rudy, are occasionalh^ introduced, l)ut these do not seem to yield so well, and besides shatter so badly that they soon have to be given up. Nonshat- tering varieties are in great demand. In all portions of the State the increase of the gluten content is probably the greatest need. All varie- ties grown in the State arc winter wheats. One of the most interesting sections of California devoted to wheat culture is that of the "Tule" lands, near Stockton. (See Plate III, fig. 1.) The great grain fields there show strikingly the possibilities in a reclamation of immense marshes. They were once vast flats cov- ered with water, mud, and a growth of bulrushes {Sc/'rj)u.s lacmtris)^ called Tule in Spanish. By means of pumping, dredging, and throwing up levees these lands have been reclaimed, and now after many years they are among the most fertile of the State. Wheat yields from 50 to 80 bushels per acre here, and barley sometimes as much as a hun- dred ])ushels or more per acre. This remarkable fertility is a result, in part at least, of the deep deposits of organic matter. There is still apparently a lack of certain mineral ingredients, such as lime and pot- ash, which are needed to make the quality of the grain as good as the quantity. As in the case of the Hard Spring Wheat district the chief difiiculty in the way of successful wheat culture in California, so far as agricul- tural practice is concerned, is the enormous size of man}^ of the farms or ranches. They are even hirger than in the Dakotas and Minnesota, containing often from 20,000 to 30,000 acres. On this account it is impossible to give the attention to details in farming that are necessary for the best results. The lack of attention to nitrogenous manuring, and especially to the alternation of wheat with leguminous crops, is particularly noticeable. The combined harvester-thresher (Plate III, fig. 2) is used in har- vesting pretty generally throughout the State. This machine is either drawn with an engine or with 28 to 40 horses. By its use the grain is thrashed directly from the field, and left piled in bags. (See Plate IV, fig. 1.) Inmiense ricks of these bags of grain remain in the field sometimes for weeks umnolested and undamaged b}' the weather. All grain throughout the State is handled in this form and calculations are made in bags and not in bushels. There is therefore no use for the grain elevator, in the ordinary sense of the term. Each ])ag contains 2i Itushels or about 150 pounds. West of the ('ascades, in Oregon, conditions are somewhat similar to those in California. In a large portion of the State a consid(M-al)le amount of spring wheat is grown. In addition to the ordinary club wheats some other varieties, such as Oregon Red Chafi" and Foise, are also well represented. The midsununci-c limatcMs much cooler than in (Cali- fornia, and therefore harvesting is performed nuich later. On account 2-1 of the greater dampness of the atmosphere and the smaller size of the farms combined harvester-threshers are not used, but self-binders instead. There is great need of early maturing varieties, as the cool autumn weather begins so early. The nitrogen content of the grain is exceedingly small. In eastern Oregon climatic and other conditions are quite diflerent from those west of the Cascades, and a description of that section is more properly included in the discussion of the Palouse country. In western Washington the general conditions and the cjuality of the wheat are very similar to those of western Oregon, but in southeastern Washington and adjacent portions of Idaho and Oregon is a large sec- tion known as the Palouse country, which possesses peculiarities of soil and climate that are distinctively characteristic and radically different from those of the Pacific Coast region proper. Strictly speaking, the Palouse country is con.'5idered to be rather limited in extent, compris- ing approximately Latah County, Idaho, and Whitman Count}-, and very small adjoining portions of Adams and Franklin counties, in Wash- ington. Recently, however, the term has come to be applied practically to nearly all of these last-named counties, as well as to Garfield, Colum- bia, and Walla Walla counties (Plate V), and may even include the northern portion of Umatilla County, Oreg. The two features which most distinguish this region from the Pacific Coast proper are the dry- ness of the climate and very finely divided condition of the soil. The particles are so very tine that when dry the soil is practicall}^ mere dust. On windy daj's this dust fills the air, forming vast clouds that are very disagreeable to the traveler. At the same time, with very little rain the soil becomes quite sticky and diflicult to manage. The capacity of the soil to al)Sorb and retain moisture is remarkable. It is pretty generall3Mjelieyed that a rainfall of 12 inches in this district is sufficient to make a crop of wheat, while in the States of the Plains 18 inches is considered to be rather low for successful wheat growing. Wheat is the chief crop of the region, though barley and oats are grown to some extent. The principal wheat varieties (except Palouse Blue Stem) are of the cluli-wheat group. They are usually soft grained and starchy, and generally white, similar to those of the coast region, but a little better in qualit}. The three standard varieties commonly grown are Palouse Blue Stem. Palouse Red Chaff', and Little Club. As regards the comparative distribution of these varieties, if the region be con- sidered as divided into three parallel north and south belts, it will be found that Palouse Blue Stem prevails in the western belt, extending as far westward as North Yakima; Palouse Red Chaff' in the middle belt, passing through the heart of the region, and Little Club in the eastern belt, reaching the foothills of the mountains. The most serious obstacle to successful wheat culture in the Palouse country is the annually recurring drought which occurs about two weeks before harvest time, particularly in the western and southern Bui 24, Div Veg. Phys. & Path , U. S. Dept. of Agriculture. Plate V, Fig- 1.— Harvesting with the Combined Harvester-thresher near Walla Walla, Wash, i Photographed by A. B. Leckenby. > ^^^ -r • ^ 'r'm'^ ,V,. Fig. uJ;.-;uiiik\ .k .: »; w'Jtu ■ 'j;iif^^^ ■ '^ Vi>a: ■ '^^I^ISi -Wheat Fields before and after harvesting, near Walla Walla, Wash (Photographed by A. B. Leckenby. ' 25 portions. From this cause the wheat is often badly shriveled, and l>oth the yield and quality there1)y much att'ected. A slight compensation for this loss lies in the fact that shiiveled wheat in this district is more in demand for making macaroni than plump wheat, because of the greater proportional amount of gluten in the former. In order to escape the severe eti'ects of the drought, early maturing sorts are exceedingly desirable. It would probably be no exaggeration to say that a variety ripening ten to fifteen days earlier than the varieties now used, and as good in other respects, would add from one to three million dollars a year to the wealth of this region. In the central and southern portions of the region fall sowing is chiefly practiced, but in the northern and eastern portions, near the mountains, there is a larger proportion of spring varieties, and there a good, hardy winter sort is needed. In the drier western and southern portions, especially in the vicinity of Walla Walla, nonshattering varieties are necessary. There the combined harvester-thresher (Plate VI, fig.l) is used in harvesting, as in California. In the north and east, and in the more hilly portions, as in the vicinity of Colfax, the self- binder is more commonly employed. In a few places a comparatively new^ sort of machine has recently come into use. (Plate VI, tig. 2.) It makes a 10 or 12 foot cut, and is driven in front of the horses, as in the case of a header, but unlike the latter possesses a self-])inding attachment as well. SUMMARY OF CONDITIONS AND NEEDS OF THE DISTRICT. (1) Principal varieties at present grown: Australian, Palouse Blue Stem, California Club, Palouse Red Chaff, Sonora, ■» Little Club, Oregon Red Chaff, White Winter, Foise. (2) Average yield per acre, about 14f bushels. (3) Needs of the grower: (a) Early maturity. (6) Nonshattering varieties. (c) Hardy winter varieties in the colder portions. SOURCES FOR DESIRABLE dUALITIES. Having descriVjed the characteristic features of the different wheat districts of the country, and having noted the most pressing needs of the grower in each one, respectively, it will now l)e api)r()priate to discuss the sources from which the desii-able qualities may be ()l)tained for satisfying these needs. This subject may be considered from two different' standpoints, (1) the botanical subdivisions of th(> cultivated varieties of wheat (Triticum) in the broadest sense, and (2) the geo- graphic groups of varieties characteristic of difl'crent regions of the 2(> world. Manifestly a complete treatment of the subject can not be presented in the present state of om' knowledge, since wheat varieties and their adaptations have not been thoroughlj' studied in all parts of the world. Nevertheless, considerable investigation has been made in this line, and the future promises still more. Such studies are exceed- ing-ly interesting, and form an absolutely necessary part of the basis for rational wheat improvement. CHARACTERISTICS OF BOTANIC GROUPS OF WHEAT. The cultivated varieties of Triticum., according to Kornicke and Werner,^ whose classification will in the main be followed in this bul- letin, may be grouped into eight species and subspecies, as follows: Triticuin mdgare, T. conqmctitm^ T. durum, T. turgldimi, T. poloni- cimi., T. Kpdta., T. dicoccnim^ and T. monococciiiii. Only T. vulgare^ T. polonicum, and T. monococcum are considered to be good species- in all classifications. The other five are generally considered as subspe- cies of T. vidgare^ though T. coriipacturu is sometimes not even ele- vated to that rank. In this bulletin they will all be referred to as though they were distinct species. The chief characters of these groups of wheats will now be described, with special reference to their importance in wheat improvement. COMMON BREAD WHEATS [TrUicum vulgare). This is of course the most valuable and widely distributed group of wheats in the world, and is represented l)y a greater number of varie- ties than all other species taken together. Nevertheless a number of veiy important qualities can be found only among varieties of the other species. The characters of this group, both l^otanical and agricultural, are well known. The heads are long in proportion to thickness, as com- pared with those of some other groups. The}^ are broader in the plane of the rows of spikelets, as a rule, and narrower on the sides of the fui-row between the rows; taper toward the apex, but may be very blunt or even thicker above; are usually looseh^ formed comparatively^ bearded or bald, and usually possess smooth chaff, but may Ix^ velvety. The spikelets, or meshes, as they are popularly called, generally con- tain three grains, but sometimes two and rarely four. The empty glumes or outer chaff of the spikelets are slightly keeled above and merely arched below. The -stem of th(^ plant is usually hollow, l)ut occasionally somewhat pithy within and varies greatly in strength and height in different varieties. The leaves also vary in character, but are rarel}^ as wide as those of the durum and poulard groups, and are velvety in only a few varieties. 1 Kornicke, Fr., and Werner, H. Handbuch des Getreidebaues, 1885. 27 The species is usually divided into a number of botanical subspecies and varieties, based upon the presence or absenc-e of beards, nature and color of the chatf, color and qualit}' of the grain, etc. For our present purpose, however, 'it will be more useful to consider that there are live great suVjdivisions of the species, based not upon botanical characters, but upon characteristics induced by influences of environ- ment, as follows; (1) Soft Winter wheats, (2) Hard Winter wheats, (3) Hard Spring wheats, (-i) White wheats, and (5) Early wheats. The location of these groups in the United States has already been pretty well stated in the descriptions of our wheat districts. Their distribution throughout the world is approximately as follows: (1) The soft winter wheats, varying in color of grain from aml)er to white, are produced under the influences of considerable moisture and mild, even temperatures, and are distributed in the Eastern United States, west- ern and northern Europe, Japan, and in portions of China, India, Australia, and Argentina. (2) The hard winter wheats are red-grained, usuall}^ bearded, possess a relatively high gluten content, and are more limited in their distribution. They are grown usually on black soils and under the influences of a climate characterized by extremes of temperature and moisture, but especially by dry, hot summers. They are found chiefly in the States of Kansas, Nebraska, Iowa, Mis- souri, and Oklahoma in this country, in Hungary and Roumania, in southern and southwestern Russia, and to some extent in northern India, Asiatic Turkey, and Persia. (3) The hard spring wheats are also red-grained and rich in gluten content, and are adapted to con- ditions of soil and climate identical with those just mentioned for hard winter wheats, with the exception that the growing season is shorter and the winters too severe for winter varieties. They are found in central and western Canada, our Northern States of the plains, east Russia, and western and southern Siberia. (4) The white wheats are soft and very starchy, l)ut possess grains a little harder and nuu-h drier than those of the soft winter wheats. They are either fall or spring sown, and are sometimes sown at both seasons in the same localit}'. They are grown chiefly in the Paciflc coast and Rocky Mountain States of this CQuntry, in Australia, and in Chile, Turkestan, and the Caucasus. (5) The early wheats are soft or semihard and generally amber to red in color of grain, but are distinguished from oth(n' groups chiefly in their ability to ripen early. They are found in Australia and India, ai-e represented by a very few varieties in the Southern States of this country, and include some of the dwarf wheats of Ja[)an. The varieties of this species naturally include the most diverse char- acters, because of their cultivation under so many diverse conditions. Their greatest characteristic as a whole, how(>vcr, is. of course, the well-known and long-esta))lishe(l (juality of their grain for the produc- 28 tion of bread flour, for which reason the term ''bread wheat" is usually applied to them. Nevertheless, it should be noted that the difference between the best and poorest sorts of this group for bread making is full}' as great and sometimes greater than between the for- mer and some v^arieties of other groups. The hard, red-grained varieties are by far the best both in food content and for our present system of roller milling. They include the Fifes, Velvet Blue Stem, Turkey, Mediterranean, and Fulcaster, of this country and Canada; the Ghirkas, Ulka, Crimean, and Buivola, of Russia; and the Theiss and Banat, of Hungary and Roumania. On the other hand, the white wheats and soft winter wheats give the best success in the manufac- ture of crackers. Several of the most popular breakfast foods are also made from white wheats. In a few instances macaroni is made from the hard spring wheats and the white wheats, but not exten- sively. No varieties of the bread-wheat group are well adapted for this purpose. The special qualities that are found in varieties of this group may be summarized as follows: (1) Excellence of gluten content for bread making. (2) Excellence of certain varieties for cracker making. (3) Yielding power of certain sorts. (4) Rust resistance in some varieties. (5) Hardy winter varieties. (6) Resistance to drought (in some varieties). (7) Early maturity (in some varieties). CLUB OR SQUARE HEAD WHEATS {T. COmpactum). By most writers this is not even ranked as a subspecies, but the different varieties certainly form an isolated group which is quite complete in itself and distinct from all other wheats, and which will therefore be considered here as a distinct species. The various varie- ties are commonly known under the names "club" or "square head". In this species the plant is verv erect, with stiff", usually rather short, culm, attaining an average height of probably little more than 2 feet. The heads are extremely short as a rule, and often squarely formed, in some varieties much broader and flattened on the furrow side, usually thicker at the apex than at the base, commonly white but sometimes red, bearded or bald, the bearded varieties usuall}^ being native in hot countries. The spikelets are set extremely close together, often standing almost at right angles to the rachis (stem), three or four- grained, sometimes with four grains nearly throughout the entire head. The outer and inner chaff' are much the same as in the bread wheats. The grains are usually short and rather small, white or red, often boat-shaped, and occasionally appear much like those of naked barley. The peculiar structure of the head in this species allows the varie- ties to be comparativelv large yielders, which is naturalh' their most Bui. 24, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. Plate VI. Fig. 1.— Combined Harvester-thresher at work near Walla Walla, Wash. (Photographed by A. B. Leckenby. ' Flo. 2. -Harvesting with the Wide-cut Binder near Colfax, Wash. 'Original. 29 important quality. They are very deceptive in tliis regard, the short- ness of the head leading one to suppose at hrst that it can not contain so many grains as are present in reality. The chati' is usually very tenacious, so that these wheats may be harvested long after ripening without loss from shattering. This is especially true of varieties grown in California and Washington. Having short, stiff straw, these wheats also usually stand up well, any damage from lodging being quite rare among them. Besides producing the class of tlours desired in certain localities, club varieties are very good for cracker making and for the more starchy kinds of breakfast foods. They are grown either as spring or winter varieties except in Turkestan, where the winters are too cold for fall sowing. Being grown in dry, hot regions, they are usually rather drought resistant. Club wheats are at present cultivated chiefly in the Pacific Coast and Rocky Mountain States of this country, in Chile, Turkestan, and Abyssinia, and to a slight extent in Switzerland, Russia, and a few other districts of Europe. The special qualities of the group are as follows: ( 1 ) Great yielding power. (2) Stiffness of straw. (3) Freedom from shattering. (4) Early maturity (in some varieties). (5) Drought resistance (in some varieties). (6) Excellence of certain varieties for cracker making and breakfast foods. POULARD WHEATS {T. turgidum). This group of wheats is usually classed as being quite distinct from the durum (T. durum) group, the two ranking as subspecies of T. vulgare. But as a matter of fact there are intergrading varieties which bring them as close together as are the club wheats and common bread wheats. They will both be considered here, like T. co7nj?actum, as distinct species. The poulard wheats are usually rather tall, with broad, in most varie ties velvety, hairy, or often glaucous leaves. The stems are thick and stiff, and sometimes pithy within. Heads long, often squarely shaped, with long beards, that are white, red, or bluish red in color, or sometimes black. Spikelets two to four-grained, and arranged rather compactly. Outer chaff' strongly and sharply keeled. Grains large, proportionally shoit and rounded, sometimes ahnost semicircular in middle cross section, rather hard and glutinous, light yellowish red in color, sometimes nearly white, and becoming glassy in varieties allied to the durum group, or on growing in certain soils. The name poulard is most commonly applied to these wheats. In Europe they are sometimes called English wheats, a very misleading name, as they are really little grown in England. On th(> other hand the few varieties that have been grown there are known as rivet 30 wheats. A name often used in Germany is hauchigerWeizen^ ^ndi a French name of corresponding meaning occasionally used is hU Ijetanielle. The wheats of this group are used sometimes in the manufacture of macaroni and other pastes. They are also occasionally used in bread making, but are more often employed for mixing with common bread wheats in grinding in order to give the quality of flour that is desired in the French markets. To a small section of this species, having compound or branched heads, some have given the separate name of composite w^heats {T.com- 2?ositum). Some well-known varieties of this section are Seven-headed, Wonder Wheat, Hundred Fold, and Miracle. It should be noted, however, that the group of emmers {T. dicoccum) includes several varieties with compound heads similar to these. Many facts known in connection with the existence of these closely allied forms, together with that of the intergrading sorts between the poulards and durums, afford strong evidence of the occurrence of natural hybrids among the varieties of these three groups. The poulard wheats are native usually in hot, dry regions, and are therefore often rather drought resistant, but not so much so probably as the durums. Many of the varieties are also very resistant to orange leaf rust. These wheats are grown chiefly in France, Egypt, Italy, Turkey, Greece, southern Russia, and other districts bordering the Mediterranean and Black seas. In this country they are only rarely grown; so far, in an experimental way. Special qualities of value to be found in this group are: (1) Excellence of certain varieties for making macaroni. (2) Resistance to orange leaf rust. (3) Resistance to drought. (4) Stiffness of straw. DURUM WHEATS ( T. clurum) . As already stated, this group of wheats is rather similar to the poulard group. As a rule, however, the heads are not so thick and the grains are longer and much harder. The plants are rather tall, w ith stems either pithy within, or hollow with an inner wall of pith, or in a few varieties simply hollow as in the common bread wheats. The leaves are usually smooth, but have a hard cuticle, and are almost always resistant to orange leaf rust. The heads are rather slender, compactly formed, occasionally very short, and always bearded, with the longest beards known among wheats; spikelets two to four- grained. The outer chaff is prominently and sharply keeled, and the inner chaff somewhat compressed and narrowly arched in the back. The grains are usually very hard and glassy, sometimes rather trans- parent, 3^ellowish white in color, occasionally inclining to reddish, and 51 l)ioportionally rather long-. In the variety Arnautka the grains are ahnost or fully as large as those of Polish wheat, and are sometimes actually mistaken for the latter. The varieties of this group are generally best known as the durums. In Europe they are often called, and correctly so, simply hard wheats. They are the hardest-grained wheats that are known. The Fifes, Velvet Blue Stem, Turkey, and others of that class usually called hard wdieats in this country are not, strictly speaking, hard wheats at all when compared with these. On account of the resemblance of the heads of these wheats to those of barley they are sometimes called barley wheats or Gerstenimizen. Durum wheats are particularly sensitive to changes of environment, and quickly deteriorate when grown in a soil or climate to which they are not well adapted. Such a change of conditions may be encountered, too, within the distance of a few miles. For example, it is well under- stood in south Russia that the excellent variety Arnautka gives the best results only when grown within a very limited area bordering the Sea of Azov. So also the best Kubanka is found east of the Volga on the border of the Kirghiz Steppes. In the Caucasus this variety has actually developed into a red winter wheat, though the original is a yellowish-white spring wheat. The durum group furnishes the great bulk of the world's supply of macaroni wheat, though a considerable amount of these pastes is made from poulard and Polish varieties and a still smaller proportion from the common bread wheats. There is now not the least doubt that some if not all these durum sorts used for macaroni can be successfully grown in this country, thus adding immensely to the profits of our wheat industry. The success that has attended the trials of the variety Nicaragua in Texas has already conclusively proved the point. At the same time the idea that these wheats can not be successfully used for bread has never yet been shown to be more than mere assumption. Sev- eral mills in this country have successfully ground them, and i n southern Russia, where milling has developed to a high degree of perfection, it is no longer an experiment. In that i-egion durum wheat has become actually the most popular for bread making, though it is usually mixed with a small per cent of ordinary red wheat before grinding. In France there is an increasing demand for durum wheats for all ])urposes. Durum wheats are adapted for soils rather rich in nitrogenous matter but somewhat alkaline, and give the best results in a very hot, dry climate. They are, theri>fore, quite drought resistant. Almost all varieties are adapted only for spring growing except in mild lati- tudes. Tii<> young plants both of this group and the poulard group arc very light green in color at tirst and grow up rapidly. They are grown in Spain (where they predominate over all other groups) and other Mediterranean countries, in south and east Russia, Asia Minor, 32 and to some extent in Mexico, Chile, and Arjj^entina. In this countiy one variety, Nicaragua, is grown to a limited extent, chiefly in Texas. The special qualities to be obtained in this group are briefly: (1) Excellence of gluten content for making macaroni and other pastes. (2) Resistance to drought. (3) Resistance to orange leaf rust. POLISH wHE.\Ts ( T. polonicum). This group is considered by all writers to belong to a distinct species. Though there are several subspecies and varieties, apparently only one variety. White Polish, is very widely known. The plant is usually rather tall, with stems smooth and more or less pithv witJiin. It does not stool extensively. The heads are extremely large and loosely formed, and before ripening are bluish-green in color. A special pecu- liarity of this species is the rather long, narrow, outer chafi'. papery in structure, and standing out slightly from the head, instead of being rigid and closely applied to the spikelets, as in other wheats. The grains are of great size when normal, proportionately quite long, yel- lowish-white in color, and very hard. The name Polish wheat is univer- sally applied to this species, though for what reason it is not clear. There is no evidence at all that it originated in Poland, and in fact it has been very little grown in that region. It is more probable that its native home is some portion of the Mediterranean region. A red win- ter wheat ofrown rather extensivelv in Poland and southwest Russia and also called Polish wheat, should not be confused with this group, as it is radically dilierent, being one of the bread wheats. Other names have been given to this species but they are quite local in their use; such are Giant rye, Astrakhan wheat, Jerusalem rj^e, etc. In almost all of the few cases where Polish wheat has been tried in this country it has proved a success from both the standpoint of yield and quality of the grain. But it seems never to have occurred to anyone to make use of the wheat for the production of Auierican macaroni, though no doubt it would be excellent for that purpose, and a great demand for its increased production could thus be created. As it is, there is not sufficient incentive to the farmer for growing this wheat, since it is not well adapted for bread making if used alone. Though requiring considerable moisture at seedtime. Polish wheat is admirably adapted for cultivation in arid districts; in fact, it produces the best quality of grain when grown under arid conditions. It is also somewhat resistant to orange leaf rust, but not so valuable in this respect as the durum wheats. Varieties of this species are grown chiefly in Italy. Spain, and other portions of the Mediterranean region, and in southern Russia and Turkestan. They are also said to be culti- vated to some extent in Brazil. 33 The special qualities of value belonging to Polish wheat are similar to those of the durum group, and are as follows: (1) Quality of gluten content for making macaroni. (2) Resistance to drought. (3) Eesistance to orange leaf rust. SPELT (T. spelta). This and the two following species are in several respects very dif- ferent from any of the preceding groups. They are also not widely cultivated, although more commonly grown than Polish wheat, and are used only to a very limited extent for human food. Nevertheless, in the intercrossing of wheat groups for the improvement of our bread wheats some very valuable qualities may be obtained from varieties of these species. The varieties of this group are called spelt in English, Sj)eh in Ger- man, and ejpeautre in French. In Germany the old name Dinkel is also sometimes applied. The varieties often called spelt in this coun- try and Russia are not spelt, but emmer (T. dicoccurii). The spelt plant (Plate VII) grows to the average height of wheat, or perhaps a little higher, and possesses a hollow stem. The leaves are of ordinary size, usuall}'^ smooth, but sometimes with scattering hairs; heads loose, narrow, and rather long, bearded or bald, espe- cially characterized by a very brittle rachis, allowing them to be easily broken in pieces in thrashing. The spikelets are usually far apart in the head, arched on the inner side, and contain usually two grains; outer chaflf oval, four-angled, boat-shaped, and only slightly keeled; grains light red in color, somewhat compressed at the sides, with a narrow furrow, the walls of the furrow flattened, and with sharp edges. The grain is always held tightly within the chatf, and can not be hulled in thrashing. Spelt is used very little for human food, but is generally fed to stock. It is very important, however, for certain portions of our country, at least, to obtain for the bread wheats the particular quality of this group of holding the grain tenacioush'. This can readil}' be done, as the Garton Brothers have amply demonstrated in England, by inter- crossing varieties of the two groups. For certain varieties that would otherwise be of great value in the Pacific Coast and Rocky Mountain States such an improvement of preventing shattering at harvest is the most important that can be made. The few varieties possessing this (jualit}^ that are now grown in these districts are sometimes not desira- ble in other respects. At the same time complaint is oft(Mi made that c(M-tain introduced varieties which are most excellent from the stand- l)()int of yielding capacity and hardiness are rendered worthless because of the irreat loss from shatterino-. It has also been observed bv certain experimenters that the (juality of constant fertility, or of producing 487'J— No. 24 3 34 "well-filled" heads, is greatly increased by the introduction of the spelt element. No doubt we very little realize the loss in yield that is simply the result of the inability of the variety to fill out its heads. There are both winter and spring varieties of spelt, and some of the former are very hardy. Certain varieties are also rather drought resistant, but nearly all sorts are more or less susceptible to rust attacks. It is in just such cases as that of the use of spelt varieties in intercrossing with bread wheats that the greatest of judgment must be exercised because of the presence of undesirable as well as desira- ble qualities. While the experimenter is endeavoring to secure the qualities of nonshattering, drought resistance, etc. , it is equally impor- tant to select from the progeny of such crosses in such a way as to eliminate the characteristics of rust liability and brittleness of the head. Here also is shown emphatically the advantage of the practice of composite crossing (to be discussed further on), as in such case the variation induced is so great that there are almost certain to be indi- viduals present among the sporting ofispring which possess the desired qualities without having preserved the undesirable ones. Spelt is chiefly grown in Germany, Italy, Spain, France, and Swit- zerland, and perhaps to a small extent in Brazil. It is not grown in this country except mainly in an experimental way. Summarized, the desirable qualities found in the spelt group are: (1) Power of holding the grain in the head. (2) Constancy in fertility. (3) Hardiness of certain winter varieties. The undesirable ones are: (1) Brittleness of the head. (2) Rust liability. EMMER {T. dicoccuni). This species has no English name, but is often incorrectly called spelt in this country. The German name is Emmer and the French amldonnier. As the German name is best known and easily pro- nounced, it should be at once adopted with us, and the name spelt applied where it properly belongs. In Russia, where the group is well represented, it goes by the name of polha, which name is invari- ably translated spelt. But either the Russians wrongly apply the name polba or this is an incorrect translation. As a matter of fact, very little if any true spelt is grown in Russia, though a rather large quantity of emmer is produced each year. The plants of this species are pithy or hollow, with an inner wall of pith; leaves sometimes rather broad, and usually velvety hairy; heads almost always bearded, very compact, and much flattened on the two- rowed sides. The appearance in the field is therefore quite difierent from that of spelt. The spikelets (that is, the unhulled grains as they Bui. 24, Div. Vtg. Phys & Path., U. S Dept. of Agriculture. Plate Vll. 35 come from the thresher), however, look considerably like those of spelt, but difi'er principally in the presence always of a short pointed pedicel. This pedicel, which is really a portion of the rachis (stem) of the head, if attached at all to the spelt spikelets, is always very blunt and much thicker. Besides, the emmer spikelets are flattened on the inner side, and not arched as in spelt, so that they do not stand out from the rachis as the spelt spikelets do, but lie close to it and to each other, forming a solidly compact head. The spikelets are usually two-grained, one grain being located a little higher than the other. The outer chafi' is boat-shaped, keeled, and toothed at the apex. The grain is somewhat similar to that of spelt, but is usually harder, more compressed at the sides, and redder in color. ' For the production of new varieties by hybridization emmer has qualities similar to those of spelt, but still more valuable. At the same time emmer, besides possessing harder grain, is more resistant to drought, and usually rather resistant to orange leaf ru.'^t. It is well adapted for cultivation in the northern States of the Plains and has already proved very valuable as a hardy forage plant in that region, besides giving a good yield of grain per acre. Almost all varieties are spring grown. Of other countries emmer is chiefly cultivated in Russia, Germany, Spain, Italy, and Servia, and to some extent in France. The emmer of this country is descended from seed originally obtained chiefly from Russia, where a considerable portion of the food of the peasants of the Volga region is a sort of gruel ("kasha") made from hulled and cracked emmer. The desirable qualities furnished by this group of wheats are: (1) Power of holding the grain in the head. (2) Drought resistance. (3) Resistance to orange leaf rust. The undesirable qualities are: (1) Brittleness of the head. (2) Adaptability only for spring sowing. EiNKORN (T. monococcum). This species of wheat is very distinct from any of the others, though the heads resemble those of emmer somewhat. It has no F^nglish name, but is called Einkorn in German and erigrain in French. The German name is adopted here. Einkorn (Plate VII) is a short, thin, and narrow-leaved plant, which presents a peculiar appearance in the Held. It seldom reaches a height of more than 3 feet. The stem is iiollow, thin, and very stitt". The leaves are usually quite narrow, soinetimes hairy. Those of the young plant are sometimes bluish-green, and after flowering the plant becomes yellowish-green. Portions of the stem may also be brown. The heads are slender, narrow, very compact, bearded, and much flattened 36 on the two-rowed sides, and always stand erect, even Avhen ripe, but break in pieces easily. The spikelets are flat on the inner side, or form a concave surface with the projecting edges of the outer chaff. They are arranged very compactly in the head and are usually one- grained, except in the variety Engrain double (Plate VII), where they possess two grains. The outer chaff is deeply boat-shaped and rather sharply keeled, the keel terminating in a stiff* tooth. The grains, which are tightl}' inclosed in the spikelet, are light red and extremeh' flattened, becoming thus bluntly two-edged and possessing an exceedingly narrow furrow. This species is at present but little improved over the original wild form, and only a few varieties have been developed. Nevertheless some of the most valuable qualities maj^ be expected from these varie- ties if they can be successfully employed in hyl)ridization experiments. They are among the hardiest of all cereals and seem to be constant in fertility, and in the writer's experience are absolutel}^ proof against orange leaf rust. Einkorn is entirely unknown in this country, except among a few experimenters, but is grown to a limited extent in Spain, France, Germany, Switzerland, and Italy. The two chief varieties are common Einkorn and Engrain double. The valuable qualities to be obtained in this species may be summa- rized as follows: ( 1 ) Power of holding the grain in the head. (2) Resistance to orange leaf rust. (3) Hardiness. (4) Resistance to drought. (5) Stiffness of straw. An undesirable quality is: (1) Brittleness of the head. GEOGRAPHIC GROUPS OF WHEATS. From the description of the different natural groups just given and the statements concerning their geographic distribution, it may be inferred that the localities as well as the natural groups might also be given from which particular qualities in wheat can be obtained. This can be done, but not with the completeness that could be desired, as it is not yet accurately known what kinds of wheat grow in all regions of the world. However, the matter may be stated approxi- mately and briefly as follows: (1) White Avheats containing much starch are grown in the Paciflc Coast and Kocky Mountain States of this country, in Chile, in Turkestan, and to some extent in Australia and India. (2) Amber or reddish-grained wheats, also starchy, are to be found chiefly in the Eastern States of this country, in western and northern Europe, and to some extent in India, Japan, and Australia. ( 3 ) Large proportions of gluten content of the quality considered to 37 be necessary for making the best bread are found in the red wheats of Canada and our Northern and Middle States of the Plains, in eastern and southern Russia, in Hung-ary and Roumania, and in southern Argentina, (-i) Resistance to orange leaf rust is to be secured in the bread wheats of southern Russia (particularly in the Crimea and Stavropol government), in the poulards, emmers, and einkorn of the countries bordering the Mediterranean and Black seas, and in a few varieties in Australia. (5) Large gluten content of the quality necessary for making the best macaroni is furnished by the durums, poulards, and Polish wheat of Algeria, Italy, Spain, and especially of the northern shores of the Black and Azov seas in Russia, and to a limited extent in the State of Texas in this country. (6) Stiffness of straw, preventing the lodging of the grain, is found in the einkorn and some of the spelts, durums, and poulards of the Mediterranean countries, and in the dwarf bread wheats of Japan, and some of the club wheats of our Pacific Coast States, Turkestan, and Australia. (7) Great yielding power, at least in proportion to the length of the head, is obtained in the club wheats of the Pacific Coast States of this country and Chile, and Turkestan. (8) The quality of holding the grain, or nonshattering, is found in the club wheats of the Pacific Coast States, Chile, and Turkestan, and in all the spelts, emmers, and einkorn of east Russia, Germany, and the Mediterranean countries, and to a limited extent in the emmers of our northern States of the Plains. (9) Constant fertilit}^, so far as known at present, is probably best obtained in the spelts of Germany and Southern Europe. (10) Early maturity is found to a limited extent in some of the bread wheat varieties of Australia and India, and in the dwarf wheats of Japan. (11) Resistance to drought and heat is best secured in the conunon red wheats and durums of south and east Russia, and the Kirghiz Steppes, the durums of the south Mediterrean shore, and both the bread wheats and durums of Turkestan. (12) Resistance to drought and cold is found to the greatest degree in the red winter wheats of East Russia. IMPROVEMENTS ACCOMPLISHED. Looking to the future, the possibilities for wheat improvement appear to be unlimited, and it is with these that we are of course more directly concerned at present. It will be of interest, however, to con- sider briefiv some of the areat cliano(>s for the better that have already been made in the wheat industry of this country during its short his- tory. Some of tliese changes have been accomplished in line with similar ones in othei- countries, and have been coincident with improve- ments in the milling process or with the demands of consumers for greater variety in food, but in th(^ main they have followed as a nat- ural result of the development of the country. As wheat is not native 38 in the United States, necessarily all seed was originally brought from other regions. At first, the territor}^ being limited and the demands of the people comparatively simple, ver}?^ few varieties were sufficient; but as the country rapidly developed and new territory was from time to time added and thrown open to settlement, new and varied condi- tions of soil and climate were encountered, and to meet the require- ments of these new conditions other new and different varieties became necessary in order for the best success. INTRODUCTION OF NEW VARIETIES. A full history of the introduction of new varieties of wheat into this country, and from one section of the country to another, Avould be a matter of much interest but can not be attempted here. Only a few of the most important instances will be mentioned — those that mark real epochs in the development of our wheat industiy, and have in cer- tain localities entirely revolutionized wheat culture. By far the most important among the earliest varieties introduced is the Mediterranean wheat, obtained first in 1819 from the islands of the Mediterranean Sea. At various times after that date this Department secured seed of the same variety and distributed it to all parts of the country. It soon met with favor everywhere. It is a hard}^, bearded variet}'^, productive, and producing a large red grain of good milling quality. But more than all this it was found to be quite resistant to rust and to damage by the Hessian fly, two enemies of the wheat crop which had already begun to be very much dreaded. This wheat has maintained its excellence through all decades since, and is to-day one of the most popular varieties in certain States, particularh" Texas. It has also been used as a parent of several very valuable hybrids. A most interesting example of improvements that are possible in the adoption of varieties best adapted to a particular region is found in the Fife wheats of Canada and the Northern States of the Plains. These varieties, which have become the basis of the great wheat and flour production of the Northwest, originated, according to the Cana- dian Agriculturist of 1891, in the following manner: Mr. David Fife, of the township of Otonabee, Canada West, now Ontario, pro- cured through a friend in ( ilasgow, Scotland, a quantity of wheat which had been obtained from a cargo direct from Dantzic. As it came to liand just before spring seed time, and not knowing whether it was a fall or spring variety, Mr. Fife con- cluded to sow a part of it that spring and wait for the result. It jjroved to be a fall wheat as it never ripened, except three ears, which grew apparently from a single grain. These were preserved, and although sown the next year under very unfavor- able circumstances, being (juite late and in a shady place, it proved at harvest to be entirely free from rust when all wheat in the neighborhood was badly rusted. The produce of this was carefully preserved, and from it sprang the variety of wheat known over Canada and the Northern States by the different names of Fife, Scotch, and Glasgow. If the above is an accurate statement of the introduction of Fife wheats, indications are rather strong that the}' are of Russian origin, 39 judging from the description of the grain and source of the cargo, in connection with the present similarity of these wheats to Russian varieties. Their subsequent history in the Northwest and the impetus given to the wheat industry of that region through their cidtivation are well known to agriculturists generally. Various strains have been developed till there are now a dozen or more so-called varieties in use. They are red, hard-grained wheats (as we use the term) similar to the Ghirkas of the Volga region, yield fairly well, and produce flour of excellent quality. In Michigan there has been an energetic movement for a decade or longer to obtain hardy winter sorts, which has resulted in a great improvement not only for that State but for adjoining territory. The millers of the State have especially been active in this movement and the matter has been frequently a prominent topic of discussion at the meetings of the State Millers' Association. Budapest from Hungary and Dawson's Golden Chafl' from Canada have been introduced and become favorite varieties. Another variety, Theiss, introduced from Hungary, has obtained a well-merited reputation as a hardy, red winter sort in the North Central States and as far west as Kansas. It has, however, not even yet received the attention that it should have. Perhaps the most remarkable development in wheat culture in this country has been made in the Middle States of the Plains, in what we may now call the Hard Winter Wheat district, all brought about through the introduction of the hardy, red-grained wheats. Twenty- five years ago very little hard wheat was grown in this region, the seed being brought by the early settlers from States farther east, where soft wheats were chiefly cultivated. Also, spring varieties formed the basis of a large proportion of the wheat production. But the spring wheats were severely rusted, injured by drought because of late maturity, and in some seasons almost wholly destroyed by chinch Inigs, while the soft winter sorts, such as White Michigan and Poole, also rusted badly and were not able always to stand the winters. For some time these defects were overcome in great measure by the use of the variety Odessa, popularly called "-Grass'' wheat in some localities, which is probably equivalent to the variety LTlka of south- ern Russia. It is hardy, red-grained, rather rust resistant, and has the additional advantage of being adapted for cither autunm or spring sowing. A little later, the well-known vai-iety Fultz also became quite popular in the West, as it is still in the greater portion of the United States. But th(>, variety which more than all others tinally completely changed the status of wheat culture in this district, is that which is commoidy but unfortunately known as Turkey. It is a bearded, hard red wheat of the highest class, coming originally from the Crimea and other portions of Taurida in southern Russia, and not from Turkey as the name would imply. Within a very small area in Kansas, Turkey wheat has been grown al)out twenty-five years, but its merits have 40 become generally known only during the last twelve or fifteen years. It is now a favorite variety in the middle Great Plains. B3' the use of this variet}^ autumn sowing is now made practicable in most seasons to the northern limit of the district, and the winter-wheat flour from this region has obtained a reputation for quality of the very best, and distinctively its own, in the foreign markets. At the same time there is no longer so much damage resulting from the attacks of rust and chinch bugs. As it is also one of the most drought-resistant sorts, it has made it possible to extend the winter- wheat area farther westward as well as northward. In a large part of the Pacific coast region, including the Palouse country, the improvements which have resulted in such large yields and great profit in certain localities were made chiefly through the introduction of club wheats, which are very productive, hold the grain in the head, and are in other regards well adapted to the conditions of the region. One or more of these wheats came originally from Chile, and others probably from Australia and France, but the origin of many of them is not accurately known. Two other varieties, not club wheats, namely. Australian and the Palouse Blue Stem, are also two of the most valuable wheats of this district and probably belong to the Purple Straw group of Australia. In southern California and the Irrigated Wheat district the variety Sonora has had the greatest influence in the development of wheat culture. It is a white-grained sort with reddish, velvet chafl', but the grains are a little harder than those of the club type and better adapted for export. It came originally from the State of Sonora in Mexico. Among later introductions is the variety called Nicaragua, a durum wheat, alread}^ discussed in another part of this bulletin, which is likely to take a considerable part in the future wheat production of this country, both because of its adaptation — as is true of all durum varieties — to the hot, dry summers of the southwestern Great Plains, and because of its suitability for the manufacture of macaroni. No facts concerning the origin of this variety are at present known to the writer, though one would infer from the name that it came from Nicaragua, and it is true that varieties of the same group are known in that country. It has been known in Texas for many years, and its use has made it possible to grow wheat in portions of that State not before successful in wheat growing. A variety similar to this one, called Wild Goose, is grown to a very limited extent in North Dakota, and probably came originally from southern Russia. It is also likely to be of value for the production of macaroni, though it seems to be somewhat inferior to Nicaragua. WORK OF THE DEPARTMENT. In connection with the discussion of the introduction of varieties, it is hoped that it will he of interest to give an account of the experi- ments made by this Department with wheats from all parts of the world. Though the aim in beginning these experiments, as already 41 stated, was primarily to test rust resistance, the work naturally soon developed into a study of the characteristics in general of the varieties of different natural groups of wheats, and of groups considered geo- graphically, and some most interesting facts were thus obtained which will be of great value in the work of wheat improvement. In fact, many statements made in the foregoing discussion of the " Sources for desiral)le qualities" are based upon the results of these experiments. Varieties were obtained from every wheat country of the woi'ld, aggregating about 1,000 rather distinct sorts in all. The manner of securing these wheats, and the time and labor thus involved, together with the difficulties of nomenclature arising from the confusion of varietal names which prevails generally, have all been discussed in a former bulletin on Cereal Rusts of the United States, and need not be referred to here. The varieties were grown one year in Maryland and most of them one year in Kansas, while about 300 of them were grown two years in Kansas, or three years in all; that is, during 1895, 1896, and 1897, During the same time a number of the varieties, especially from Russia, Siberia, Japan, and Argentina, were tested by other parties in cooperation with the Department, in other localities, viz, in Michigan, Wisconsin, Indiana, Tennessee, and Colorado; and in the case of a few of these, the experiments have since then been repeated in Colorado, Kansas, and Nebraska, In conducting these experiments all the principal characteristics of the wheat plant, as shown in its different stages from that of the young plant to harvest time, were studied, though complete notes can not be given for every variety. These features include in a general way (1) the character of the young plant; (2) hardiness, including resistance to rust, drought, and cold; (3) character of the head; (1) character of the grain; and (5) time of maturity. Field experiments alone do not show those qualities of the grain which indicate the value of the variety for different uses, and which are after all more important than any others; though it must be remembered that certain characteristics of the growing plant often indicate quite correctly what these qualities will be. Therefore chemical analyses have been made of a large number of representative varieties, and for many of them the absolute weight and specific gravity of the grain have also ])ecn determined. As would be expected, a large number of the varieties either proved to be entirely unsuited to the conditions in this countrj^ or were found to be in other respects undesirable sorts. It was the purpose from the start to discard gi'aduallv all varieties that seemed to be uhjible to adapt themselv(!s to their new environment. During the tirst year oi the experiments at (iarrett Park, Md,, many of them were planted so late, on account of their late arrival in this country, that much allow- ance must be mad(^ for their behavior in comparison with others which were sown in good season. In other respects the trial for that year was very satisfactory, and afforded an excellent opportunity of c()m])iiring the behavior and <|ualities of a large number of varieties under average conditions. 42 But the larger portion of our area most important in wheat pro- duction lies much farther westward than Maryland and possesses a very different soil and a climate characterized by great extremes of drought, cold, and heat. At the same time it is manifestly desirable to search particularly for varieties adapted to such extreme condi- tions for two reasons: (1) It is found that as a general rule sorts which are able to withstand the most rigorous extremes of climate are also of the class which makes the best quality of bread and macaroni, the two principal purposes for which wheat is used.^ (2) It is com- paratively easy to obtain varieties suitable for mild conditions, as those which are resistant to climatic extremes are more easily grown in a milder climate than the reverse. It was therefore decided to test the varieties during the following years in the Great Plains. Accord- ingly, in 1896 the field experiments were carried on at Salina, Kans., and in 1897 at Manhattan, Kans. At Salina Mr. B. B. Stimmel kindly donated the use of about two acres of land for the experiments. At Manhattan, by courtesy of the board of regents of the Kansas Agri- cultural College, the farm department of the experiment station was authorized to cooperate with this Department in the experiments con- ducted at that place, by furnishing land and other facilities for the work. During the years of the experiments in Kansas the seasons were unusually severe even for that region. As a result it was found desir- able to discard a large number of sorts from year to year. Only about 300 varieties were grown at Manhattan in 1897 out of the 1,000 origin- ally obtained, and of these, less than 200 were selected as being worthy of continued trial. Over 100 of the varieties finally remaining have been made the basis of a large part of the series of field experiments now in progress at Halstead, Kans., while a number of the same vari- eties are still being tested at the Nebraska Experiment Station Farm, Lincoln, Nebr., and at the Arkansas Valley Experiment Station, Rockyford, Colo. Through this process of rigid elimination, which is a good example of the practical application of the law of the " .survival of the fittest" in agriculture, about 100 varieties have been determined upon as being fairly representative sorts of the world as regards hardi- ness and quality of gluten content. There are many varieties, how- ever, which can not be classed with these hardy, glutinous sorts, ))ut which, nevertheless, because of their early maturity or particular adaptation in other respects to certain localities where hardiness is not necessary, and where these sorts would fail because of the lack of other qualities, must be considered as equally important. Palouse ^ For the present, for the want of space, a full discussion of this proposition in detail can not be given, although the experimental proof concerning the qualities of varieties originating in regions of different climatic conditions will be brought out in connection with the table presented in the following pages. The whole matter is one of much interest and may be discussed in detail in another publication, The Relations of Soi) and Climate to Wheat Production. 43 Blue Stem, Australian, Little Club, Early May, AUora Spring-, Yemide, King's Jul)ilee, Early Genesee Giant, etc., are examples of this class. Adding still to these a number of other sorts, belonging to the Spelt, Emmer, and Einkorn groups, which are also hardy, but are especially valuable for certain qualities they may furnish in the work of hybrid- ization, and then still a few others, mentioned favorably by other experimenters, we have in all 2-45 wheats which may be regarded as leading varieties of the world. For comparative study the principal qualities of these 245 varieties are briefly stated in the following table, which is based mainly upon investigations of this Department, but to some extent also upon the work of others. As regards the field work, it represents a summary of the combined results of the three years' experiments, so that each column shows as nearly as can be given an average of that quality for each variety for the three years. ^ The table is made up of twenty-five columns, and the information given in each is in most cases clearly explained by the heading, but in a few cases a little further explanation is perhaps needed. In column 2, the following abbreviations are used: C. for common or bread wheat; CI. for club; D. for durum; P. for poulard; Pol. for Polish; S. for spelt; Em. for emuier; and Ein. for einkorn. In column 8, "stand" refers to the degree of completeness with which the plants fill the drill row, and of course often measures, though not always, the stooling quality of the variety. In column 9, under "spring condition," each number expresses in a scale of 1 to 100 the general condition of the variety in all respects about May 1. The figures in column 10 are percentages showing the comparative amount of leaf rust on the plant at the date when this rust was most abundant. This column is in the main a reproduction of the column of averages in Bui. No. 16 of this Division, Cereal Rusts of the United States, pages 26-32, table 3. Of course the smallest number represents the greatest degree of rust resistance. In column 11, the abbreviations C. and D. indi- cate that the variety corresponding is resistant to cold or drought and the figure shows on a scale of 1-5 how great is the degree of hardiness, 5 meaning extremely hardy. In column 19, the word " vit- reous" refers to grain which is not oidy very hard but is somewhat transpariMit and presents a glassy surface in fracture. Wheats so characterized are usually durums. In column 25 is shown the partic- ular wheat district of the United States to which the variety is best adapted. The districts are indicated by roman numerals having the following signiHcations: I, Soft Wheat district; TI, Semi-Hard Winter Wheat district; III, Southern Wheat district; IV, Hard Spring Wheat district; V, Hard Wint(u- Wheat district; VI, Durum Wheat district- VII, Irrigated Wheat district, and VIII, White Wheat district. 'There is one exception. In column 9, "spring condition," tlie data given refer only to the e\iieriinents of 1894-95 in Marvlaiid. 44 o s e a. £ a. 'S' ^1^ o I s S- a, ►o s , CO 'MM 1 5 ii i ^ 2^ a 'X gg a SO..S ^ n 3 s = 3 H^ ^ •-: ■-: "75 '-5'-5 1 •c i t» ! ! 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' • o3 of va Fife, an... i "S : '; : : a) ;j V. X 5 s ^ te • • • a S 'gs : ~s 5 £S« 5a a o3 X 1 a B , ^ r :i " 01 ; ^ a 5 § .£ ?:.^ 3 > l :a ■ 2a 1 ^^ ^^ & ^?">-NN : ?: ' ?: ^ ? * < ^^ c^ 3S 59 In comparing- the value of different Aarieties it is very desirable to knoAV both the absolute weig-ht and specific gravity of the grain, as these phj^sical qualities bear some relation to the chemical composition of the grain and to the nature of the plant in general. All the infor- mation concerning specific gravity, as well as a number of determina- tions of absolute weight given in the table, are the results of a series of interesting investigations made in the Seed Laboratory of the Division of Botany of this Department by the late Mr. J. C. Dabney, then a mem- ber of that Division. Almost all the data of the table concerning nitro- gen content of the grain are the result of chemical analyses made under the direction of Dr. H. W. Wiley, chief of the Division of Chemistry. The greater part of these analyses were made at the request of the chief of this Division with samples furnished b}^ the Division. The remainder, together with a number of determinations of absolute weight, are taken from reports of work formerly done by the Division of Chemistry.^ A few anal3"ses are also given on the authority of F. B. Guthrie' and Emerich Pekar.^ As the value of the grain for making bread and macaroni is meas- ured almost wholly by its quality and quantity of gluten content, only the percentage of moist and diy gluten and the total per cent of albuminoids are given. Jt will be noted that no column for yields is given in the table. For this omission, which under other circumstances would be a serious one, there are three good reasons: (1) In experiments of this kind each variety is given so small a space (an average space of one drill row 35 feet long) that it is not practicable to obtain accurate estimates of yield. (2) Many of the varieties tested are already well-known Ameri- can wheats, whose yields have often been reported by various experi- ment stations, while as to the foreign sorts it is most important, first of all, to know Avhether they will prove to be suited to our conditions at all or not, before we an^, ready to test their yielding capacity. (3) The 3'ield of a variety, whih^ it is of course directly the biggest thing to the practical man, is, after all, not a distinct constant qualit^Mii itself, but is the combined result of a luimber of qualities acting indirectly, and often not thought of at all. For example, such sorts as Clawson and Pool(> are fairlv ji'ood wheats, and under mild conditions would probably yield better than Turkey; but in west Kansas or. southern 'See "Analyses of Cureals collected at the World'w Colnmbiaii Kx|)osition," Bnl. No. 4o, Div. CheiH., T^ S. Dept. Agric, pp. 39-5.'5, 1895. '^ "Noten oil the inilling qualities of different varieties of wheat," Dept. Agric. N. H. W., misc. jml). No. 189, p. 47, 1898; "Milling Jiotes on the Lainl)rigg liarvest of 1897-98," Agric. (laz. N. S. W., Vol. X, Pt. 9, pp. 90()-9iri, Sept., 1S99, and "Absorp- tion of water by the gluten of different wheats," Dept. Agrir. X. S. W., misc. pub. No. 104, p. 7, 1S9<>. ^"AVei/.cii mid :\h'hl nnsercr Krde," ini Auftragedes Kig. Ing. .Ministeriunis fur Ackerbaiiiiidustric luid Handel, pji. 277, lUniapest, 1882. 60 Nebraska they would fail entirely in certain seasons because of drought or cold, while Turkey, being very hardy, would produce a much larger yield on an average than either of the former, though its absolute yield in a good season might not be so great. So, also, it is found in the Palouse country that there are certain varieties which have absolute yields in that region greater than those of the Little Club or Palouse Blue Stem, but thej^ shatter so badly that the net yield of the latter is greater. As regards the field trial experiments upon which is based the larger part of the results given in the table, it must be said that many of those sorts whose behavior indicated that they would not be well adapted for our use should be further tested before adverse judgment is pronounced upon them, especially so if their qualities in other respects are good. Nevertheless, the table as a whole shows pretty accurateh' which are the best varieties for different districts of the country. Nothing can be more interesting than the constant observation from year to year of the efforts being made by varieties from every country in the world, struggling with new" conditions of soil and climate, to obtain a footing in a strange land. The gradual elimination of the less-adapted sorts by the severity of winter, drought, etc. , soon shows unmistakably which are the varieties that will be most valuable. Of course it may truthfully be objected that mere hardiness is not of value by itself if other qualities are not also present. But, on the other hand, it is a further matter of interest how different qualities are often so closely associated in the same varieties that if a variety is adapted to a certain district with respect to one quality, it is apt to be so with respect to at least one or two other equally valuable qualities, though, of course, there are serious exceptions. It is also quite worth 3' of note that some apparently insignificant characteristics bear an important relation to the presence of qualities of direct economic importance. As an example of these we may note especially the characteristics of the young plant in its autumn stages in connection with the presence of certain economic qualities. Hardy winter varieties are rather slow starting in the fall, but produce good roots and soon spread out flat on the ground in preparation for the cold and snow of winter. The leaves are narrow and usually dark green or purplish at first, especially near the roots. Spring varieties and most durums and poulards, as well as some of the weaker winter sorts, on the other hand, germinate quickh' and make a large growth in the autumn, but are cut short or entirely killed b}^ the severity of the winter. They produce coarse, light-green leaves, but poor roots. In regions of mild winters durum and poulard wheats make excellent pasturage because of their rapid autumn growth. There is really very little, if any. check to the growth from seeding till harvest in localities well adapted for these varieties. 61 One well acquainted with wheat varieties i.s usually able to determine largely their general classification simply from their appearance in the autumn. It will be seen also by a stud}^ of the table that there is a very close constant relation between hardiness, color, size, weight, and hardness of grain, and chemical composition in varieties of the common group. Varieties very resistant to cold and drought have small, hard, red grains, possessing a large proportion of albuminoids and a relatively high specitic gravity, though the absolute weight is likely to be low. It is also a general rule that bearded varieties are less susceptible to leaf rust, l)ut there are a number of important exceptions to this rule. Varieties with harsh, hairy, or glaucous leaves are also usually rather resistant to this rust. Varieties early in ripening are often dwarfed, and come from warm regions nearly always. Hard-grained winter varieties are bearded, as a rule. Drought resistant sorts, whether bald or bearded, white or red-grained, possess a larger proportion of nitrogen than those which succumb to drought. The effect of a change of environment upon the wheat plant has already been referred to. That marked changes are effected in this way is proved, with respect to chemical composition, by the facts given in the table. In a number of instances duplicate analyses are given of samples of the same variety obtained from different localities. Almost invariably the samples from hot and more or less arid dis- tricts show a larger per cent of gluten content. In some instances the difference is considerable. Alsace wheat from Ekaterinoslav (Russia) furnishes 13.. 58 per cent of dry gluten, while the same variety from Poltava, farther northwest in a moister region, shows only 9.30 per cent. Improved Fife, though a much liked variety in Australia, produces but 11.20 per cent of gluten there in comparison with 16.16 per cent in Colorado. Kubanka from Kursk (Russia) possesses 37.79 per cent moist gluten and 13.63 per cent of dry gluten, while as grown in Germany it furnishes only 20.93 per cent of moist gluten and 8.50 per cent of dry gluten. At the same time a sample from the Caucasus shows 41.65 per cent of moist gluten. A remarkable difference is shown in the ease of Scotch Fife from Nel)raska and Oregon. The former sample contained 14.65 per cent of dry gluten, while the latter contained onh^ 5.13 per cent, slightly over one-third as much. There is a striking example in the case of Palouse Blue Stem of a differ- ence in gluten content between two samples from the same Stiite, Washington. These samples, however, were no doubt from diffiMcnt localities, and no two regions are likely to be much more different from each other than are western Washington and the Palouse country of eastern Washington. A most interesting example of correspondence between climate and chemical composition of the grain is exhibited in the case of samples 62 from Kursk. As will ))e seen in the table, all samples from this locality not only show a very large per cent of gluten, >)ut also a per cent always far above that of other samples of the same varieties from other localities. The Kursk samples are uniformly so superior in this respect that one naturally looks about for an explanation. The matter is no doul)t to be explained in this way: It is a fact already discussed by the writer in another i)ublication' of this Department and referred to before in this })ulletin that the nitroo-en content of the gi'am is greatest in regions having black soils, extremes of temperature, and very low rainfall. In Russia extreme heat and aridity increase eastward and southward as a rule. The government of Kursk, how- ever, presents a remarkable exception to this rule, especialh' as regards rainfall. The normal yearly rainfall is 16.9 inches, while in Woronetz, Tambov, and Ekaterinoslav, east and south of it, the normal is 21 and 22 inches. It is apparently situated in an arid area, with greater rain- fall all around it. At the same time the extremes of temperature are great and the soil is of the best in the ''Chernozem"" (black earth) region. As before stated, all the experiments and ()l)servations which form the basis of this table have been made with the view of obtaining some reliable foundation for future wheat improvement. The general con- clusions of immediate value to the wheat growers that are to be drawn from this work of the Department, and which are of rather wide application, maj' be stated as follows: (1) Considering all qualities, the best wheats in the world are of Russian origin, coming particularly from eastern and southern Russia. They are resistant to cold and drought, are more or less resistant to leaf rust, and have the best (pialit}- of grain. They are fairly earh^ in ripening and are good fielders. Under the head of remarks the jnelds per acre of several newly introduced Russian sorts in Kansas and Colorado are given in the table. For varieties not yet acclimated it will l)e seen that these 3nelds are very good. The yields and weights per bushel in Colorado are furnished bj' W. F. Crawle}^, superintendent of the Arkansas Valley Experiment Station at Rocky ford in 1897. The following may be considered as the best Russian varieties so far known: Arnautka, Kul)anka. Kubanka Red Winter, Crimean. Sando- mir. ITka. Chernokoloska. Buivola, Red Winter, Bearded Winter, Yx, Odessa, Sarui-bug-dai, Ghirka Spring, Ghirka Winter. Russian, Belo- turka. Mennonite, and Turkey. (See Plate VIII, Fig. 1.) (2) The earliest ripening wheats are often dwarfed and come princi- pally from India, Australia, and Japan, though a few are from the ^Mediterranean region. They are usually soft white wheats, but those from Japan are red, rather hardy, and possess a fair gluten content. ' Russian cereals adapted for cultivation in the United States, Bui. No. 23, Div. Bot., pp. 8-11. Bui 24, Div, Veg. Phys. & PaTh . U. S. Dept. of Agriculture. Plate VIII Fig. 1.— Group of Russian Wheats in Experimental Plats at Garrett Park, Md. (Original. < Fig. 2.— Experimental Wheat Plats at Garrett Park, Md., showing earliness of KING'S Jubilee; 1 , Leaks ; 2, King's Jubilee; 3, Tuscan ; 4, Purple. (Original.! 63 The ])ost varieties so fur known for our use from these regions are: Early Japanese, Yemide, Kintaina, Japanese No. 2, Onigara, Daruma, Japanese No. 1, Japanese No. 4, Shiro-yemidashi. AUora Spring, Stein- vvedel Early Baart, King's Jubilee (Plate VIll, Fig. 2), Roseworthy, Canning Downs, Kathia, and Nashi. (3) Though varieties of Russian origin are, on the whole, the best, there are certain sorts from other eounti'ies which behave much like them. These are P\ilcaster. Lancaster, Tasmanian Red, Fultz, Chu- but, Frolifero, Rieti, Nashi, Mediterranean, Tangarotto, and Valley. (4) Durum, Polish, and poulard wheats, besides being admirably adapted for making macaroni, are all rather resistant to leaf rust. The best known varieties are: Arnautka, Kubanka, Beloturka, Medeah, El Safra, Galland's Hybrid, Petanielle noire de Nice, Chernokoloska, Sarui-bug-dai, Volo, Missogen, Atalanti, Cretan, Wild Goose, Polish, and Nicaragua. (5) Common bread wheats can not be depended upon to resist rust, but the best in this regard are: Turkev, Crimean, Prino-le\s Defiance, Rieti, Oregon Club. Fulraster, Odessa, Pringle's No. 5, Mennonite, Velvet Blue Stem, Saskatchewan Fife, Mediterranean, Alsace, Nashi, Ghirka Sj^ring, Frolifero. Bellevue Talavera, Ghirka Winter, Red Winter, Bearded Winter, Theiss, Deitz Longberry, Arnold's Hybrid, Sonora, and Banat. (()) Einkorn resists leaf rust completel}^, and emmers resists it to a high degree at least. (7) The very hardiest winter varieties are Turkey, Crimean, Red Winter, Ghirka Winter, Yx, and Bearded Winter. During the unusu- ally severe winter at Manhattan, Kans., in 1896-97, these varieties fared very well when nearly all the experimental varieties of the regu- lar experiment station plats at that place winterkilled, though well acclimated. (8) Club wheats ai'e usually soft grained and tender sorts and adapted oidy to mild climates, like that of California. They are excel- lent yielders. Among the best of them are: Little Club, California Club, Palouse Red Chaff, Sicilian Red S(juare-head, Herissoa barbu, Herisson sans barbes, and Chili Club. WHEAT BREEDING. If we wish to continue our improvements in wheat culture, it is evi- dent that we nuist soon depend upon other means than simply the introduction of vai'ieties new to the country. During the earlier his- tory of the country it was a (juestion even whether wheat could be grown at all in many of the new regions open to settlement, and prac- tically every variety had to be tested. Their introduction, therefore, naturally played the greater part in wheat impro\'ement, and has continued to do so, in less measure of course, almost to the present 64 time. But the time will soon arrive when there will be no further varieties to introduce better than we already have. The work now being done by the Section of Seed and Plant Introduction of the Divi- sion of Botany of this Department is especially hastening- the approach of this period. So far as our knowledge goes at present, there are now but two regions in the world which produce varieties likely to be of particular value to this country from which we have not already secured seed for trial in considerable amounts. These regions are (1) the north- ern portions of India and China, including Tibet, and (2) Abyssinia. There ai"e still some of the very best varieties to be obtained, however, from regions alread}" drawn upon, such as southeast Russia, Turkestan, and Japan. No more important work could be done at present than that of securing all these new sorts from different regions, for of course it is a great waste of^ time and labor to the wheat breeder to spend years in the production of varieties having special qualities if other sorts alread}^ possessing these qualities can be readily obtained from other countries. But, as stated at the beginning of this report, although many valu- able improvements have resulted and are likely still to result from introduction, there are often certain combinations of qualities found to be extremely desirable for a particular region which, so far as we yet know, do not exist in an}" one variety, native or introduced. Such ideal sorts are therefore to be acquired by improvements of the vari- eties now in use, which must be accomplished through hybridization and selection. Besides, in certain varieties ideal in other respects, such qualities as rust resistance, yielding capacity, etc., may exist already, but not to a sufficient degree. In such cases these qualities must be increased by selection of seed from individuals which exhibit them to the greatest degree. But manifestl}" the greater number of varieties one has at hand, either native or introduced, especially if these have been chosen with great care, the greater are the number of chances offered him for selecting and improving these qualities. The trial of introduced sorts, therefore, in comparison with native ones simply gives one a practical knowledge of the facts herein discussed under the heading '"Sources for desirable qualities." With these facts in mind, together with those concerning characteristics and needs of the different wheat districts, one is prepared for effective work in wheat improvement. IMPROVEMENT BY SELECTION. During the last thirty or forty j'ears considerable work has been done in wheat breeding through selection, though it is only a begin- ning in comparison with the great amount that ma}' be done. It may be of interest to note a few of the most important instances of the actual production of new sorts in this wa}'. 65 In 1862, in Mifflin County, Pa. , Abraham Fultz, while passino- through a field of Lancaster wheat, which i.s a bearded variety, found three heads of bald wheat. He sowed the seed from these heads the same year, and continued sowing a larger amount each year, until he ol)tained sufficient seed to distribute it pretty well over the country. It soon became a well-marked and popular variety, called Fultz from the name of the breeder, and is now the best known of American wheats. In 1871 this Department distributed 200 bushels of the wheat for seed. This variety is rather early in ripening, fairl}^ hardy, and possesses a semihard, red grain of good quality. It comes nearest being a general purpose wheat of all our varieties, being grown with good success in nearly all parts of the country and in several foreign countries. Next to Fultz, one of the best known of our native wheats is White Clawson, or >;imply Clawson. This variet}^ originated in Seneca County, N. Y., in 1865, through the selection of certain superior heads from a field of Fultz by Garrett Clawson. On planting the grain from these heads, both a white and red-grained sort resulted the following season. The white wheat was considered the best, and the pint of seed obtained of this sort was sown, producing 3!J pounds the following season. The third year after this 254 bushels were harvested, and that season the variety was distributed to other farmers. In 1871 this variety took first premium at the Seneca County fair, and in 1874 seed was distributed by this Department. Though judged inferior by millers at times, this variety has become a very popular one. It must not be confused with Earl}^ Red Clawson, a very distinct variety. It is a bald wheat, rather hardy, with soft, white, or light amber grains. Early Red Clawson, because of its earliness, has taken the place of this variet}^ to a great extent in recent years. One of the best of the more recently produced varieties is the Rudy, which was originated at Troy, Ohio, in 1871, b}' M. Rudy, through a careful propagation of the seed from a superior and distinct stool of wheat found in a large field. It is a semihard or soft reddish-grained wheat, bearded and with white chatf. It is widely grown in Ohio, Indiana, and adjoining States. A number of tiie different varieties of Fife and Velvet Blue Stem of the spring- wheat States were also produced by simple selection. Wellman's Fife is a good example. In 1878 D. L. Wellman, of Frazee City, Minn., received a sample package of Scotch Fife wheat from the Saskatchewan Valle}', in Manitoba. This was sown in the sprmgof the following year, and as a result it was found that the seed was badly mixed. Removing all plants but those of the true Fife and propagating carefully from year to year, Mr. Wellman gradually bred upward a ver}^ pure strain of the Fife, which became known as the Saskatchewan Fife. From the crop of 1881 were seliM-ted some uinisually large heads, and from the seed of these as a beginning he finally produced a rather 4879— No. 24 5 66 distinct sort, now known as Wellman's Fife. In a similar manner Powers's Fif e, Hayne's Blue Stem, Bolton\sBlue Stem, and other sorts have been produced b}' the men whose names they bear. By the process of selection an unusually good variety of white wheat for the Eastern States, usually called Gold Coin, has very recently been produced by Ira W. Green at Avon, N. Y. Several years ago he grew a field of Diehl Mediterranean, a bearded, red-grained wheat, and while passing through this field one day found a bald head possess- ing white grains. Planting every grain of this head, he found as a result next season that he had heads with very long beards, some with short beards, and others with none at all. The grain also was mixed, some red and some white. He desired a bald wheat, since the beards interfered with his success in woolgrowing, hence only the grains from the bald heads were again planted. From this as a beginning, a practically new variety resulted, which he called '"No. 6." It has proved to be of considerable value for certain localities, and is already pretty well known. Various names have been given to it by difierent seedsmen, but it is best known by the name Gold Coin. In instances like those just related the change has been so great as to produce really a new variety. But, of course, the majority of improvements made by selection do not represent such marked changes, though there is a great tendency among breeders to establish new varieties on the basis of very slight improvements. In a majority of the instances above described the circumstances too are such that one can not escape the thought that the abnormal heads found in the fields were the result of natural crosses. In fact in the cases of Clawson and Gold Coin wheats this is almost certain, since the seed from the first heads continued to produce sporting progeny, the following year. Or it is possible in the case of Gold Coin that the sporting was simply a later cropping out of this phenomena in the Diehl Mediterranean, which is itself a hybrid. Besides these cases, there are also instances mentioned by other writers which pretty well establish the fact of the occurrence of natural crosses among wheat varieties,^ though, of course, such occurrences are rather rare. On the other hand, in the work of hybridization the selection of parent forms and the after .selection of the best individuals from the sporting ofispring are by far the most critical operations to be performed. Hence selection is both the most important part of all the work of wheat breeding, and is also to be considered from two rather difierent standpoints: (1) that of its operations in connection with hybridization (natural or artificial), and (2) in making the ordinary less striking improvements in the same »See especially Rimpau's statements in his article on " Kreuzungsprodukte land- wirthschaftlicher Kulturpflanzen," in Landwirthschaftliche Jahrbiicher, Bd. xx, S. 347-350, 1891. 67 variet}'. The former phase will be best discussed under the subject of h^'bridization. In eases like those of the different varieties of Fife and Velvet Blue Stem, such as Wellman's Fife, Haj^ne's Blue Stem, etc., above men- tioned, as well as many others that might be described, the new sort, if it is rightly called such, has been produced b}^ ver}^ gradual improve- ments during many years. It is not a selection of varieties, nor of offspring showing combinations of elements from different varieties as a result of crossing, but is simply a selection of individuals. The process is slower and the changes effected are not so great at an}^ one time, but in the end important results ma}^ be reached. Selection of this kind is, of course, the most common, and occurs constantl}' in nature, especiall}^ in connection with the qualities of rust resistance, hardiness against cold, etc. Farmers prett}^ generally practice a sort of selection of seed corn, and often too of potatoes, for seed. Comparatively little attention, however, is paid to the selec- tion of Av heat for seed, although the wheat plant is ver}" susceptible to its environment, furnishing therefore many variations as a basis for excellent results in this line. It is through this kind of w^ork, but carried on thoroughly and svs- tematically, that Prof. W. M. Hays, of the Minnesota Experiment Station, has attained some ver}^ interesting and practical results with the Fife and Velvet Blue Stem varieties of that region. He has prac- ticed rigid selection with these varieties for a number of 3'ears, giving special attention to yield and quality of grain as shown by the baker's test. Certain new strains capable of giving to the farmer substantial gains over others have already been produced in this wa}'. He has also developed a method of keeping records which is worthy of the attention of other experimenters. In the preceding pages the special needs of different wheat districts have been discussed, and also the groups of wheats from w^hich, in crossbreeding, the qualities for satisfying these needs may be secured. One must not forget, however, how much such qualities may ))e increased in the varieties already grown in the district, and should remember too, that even after great improvements have been secured through hybridization, very careful selection must be practiced in order to maintain the standard of excellence reached, especially if the variety is to be grow^n under conditions adverse to the production of the particular qualit}' accjuired. Some of the most important qualities of the wheat plant that may readily be increased on any farm simply by selecting seed from those plants which exhibit these qualities to the greatest degree, are yield, drought resistance, winter hardiness, rust resistance, earliness in rip- ening, quality of the grain in any respect, and nonshattering. If in passing through a field certain i)lants jire noticed which are almost or ('.8 (juitc I'rrc From rust, \vhil<*- thc.othors aro (;()nsi(l(M"iil)lv rust(Hl, nud tlio lociility should 1i!1|)))(mi to l)(^ one in whicli rust is usually vory hud, such heads should hy all means he sclecti'd, sown separately, and from th(^ ))roj^-eny the most resistant individuals a<4ain selected. It imist of (U)urse he noted thai -ly to he I'ust i-esistaid. in miiiiy instances. Great yieldi no- power and nonshattei'ino- ni:iy idso occui' in the same individual, while gluten content iind drought resistance may (^xist together in ceitain othei's. Ill Mil ai'ticie \)\ the writer on " Impi"o\cmeiits in wheat culture" ' a simple metluxl is suggest(Ml which, if practiced, would enable any farmer to constantly and cIlectiNcly im])ro\-e th(^ yield and (piality of grain with little lioiihle. hut with grejit profit in the <^w\. As this iiK^thod niiiy he employed e(|ually well tor the improvement of any othei- (piality of the plant, tiid'e is jjiohahly no mor(^ fitting way of closing the discussion of this topi<- than to I'cproduce here the desci'ip- tion ot" that method with such modilications as are necessary to make- it api)licahle for any improv«>ni(Mit desired. It is as follows: Hegin pr:icli<'ing the constant us(> of a wheat-hreeding ])lat of 1 acre or more from which to select sccmI each year. Locate this plat at ditter- i'nt paits of the farm every two or three years, preferably in alterna- tion with cloN'ei' or other leguminous ci'ops, and gix'e it the best of care, flust hefore liar\'est go through a tield of a good, hardy, standai'd variety that has gi\'en the best results in the locality, and mark ])lants that exhihit to the higlu\st degree the special (luality which it is desired to incn^ase, su, and which are at. the same time at least as good as the ax'cragii in other respects. At harvest tim<> cut with a sickle enough of these marked j)lants for sowing the plat and, after thrashing them, select the largest and most \igorous seed foi' this pur])os(\ by means of a scr(MMi or even hy hand picking. Sow the plat early, drilling it at theavei'age rate of ai)out \.\ bushels ]M>r acre. Next s(»ason use none of the Held crop for seed, but select in the same mannei' enough of the l)(>st plants from this brecMling ])lat for r«>seeding the plat and use all the remainder for sowing the general cioj). In the following season and eai'h succeeding season practice exactly the same method. In this way seed is never taken from the gre is no extra lahoi" involved exc(^pt the small amount iHMpured for seed selection each year. Of cours(> the hreediiig plat should he k(>pt constantly free from rye or other foreign heads and weeds. ' YciirlxKiU I'nittMl Stiilcs i)cii;iil mciit <>l' Au'ricultiirc, ISiXl, ])a^es 489-498; also ivpiiiito'i. IINU'UON KMKN r IH 1 1 V UKl I H/A'l'K )N. Ill inanv iiistjuu-os (luiilitics tliiit arc xcry (l('siral)lo or vyvu lu'ccssarv for a partifuliir districl aic ciitircly lackinu'. or at least not ])ivs('iit in any a])pr('cial)l(' dcurcc. in Niirictics which arc in all other respects adinirahly adapted to the district, in such instances th(> iinproN eincnt of the vaiMety must be accomplished hy l>r(M>din>i- into it the desired quality from som(> other sort ]M)ssessiny- it to m hiu'li deorcc. Thoiinh not so simi)lc a process as that just descrihed. and Irauolit with iinich more uncertainty in its operations, hyhridi/ation is 5)rten absolutely necessary for ])ro(lucinj>- radical chanycsoi' oi-cat moment, or, in ciisesof emerj^ency, for satisfyin«i' an inipeiati\-e need, wIumi the ordinary i)roc- ess of s(dection alone would either he too slow or f:iil entirely. 'The possi))ilities for im])rovement through hyhridi/ation, accomi)anied l»y discriminating- sidection, in the hands of skillful hnMulers, s(>em to he practically unlimited, especially in the <-asc of a [Anui so closely s(>lf- fertilized as wheat. Nevertheless, comi)aratively little work of this kind has vet been done with the cereals, and particularly so in this country. Also the j^reater part of what has Ix-eii accomplished, thoiiuh productive of important residts, has been of rather an elementary natui'e. It may l)e advisable^ before continuino- the discussion to ^ive lirst a l)rief account of some of the principal wlu'at hybrids produced in this country. Nearly all of these new sorts have i)roved to be of more or less value in wheat improxcment, while a few of them have become well-known factors in dev(dopin<;- the industry. 'VUo pioneer in the production of wheat hybrids in this country is ('. (i. Pi-in, Felspar, Ruby, and (Jranite. (Jypsum (Blount's Lambrigg), Iloi-nblende. Quartz, and Improved Fib' are the mo.st 70 popular in Australia. In New Mexico, where field tests of all his hybrids were last made, Ruby and Felspar are now most extensiyely grown. Blount's No. 10 is much prized in the northern portion of the Pacific coast district, where the yariety Oregon No. 10 is probably identical with it. An important characteristic of several of Blount's hybrids is that they are rather rust resistant and it is partly for this reason that the}' are so much used in Australia. Improved Fife, how- ever, has also an excellent quality of grain. One of the very best varieties of this country, standing probably next to Fultz in popularity, is Fulcaster. It was produced in 1886 by S. M. Schindel, of Hagerstown, Md., and is a hybrid between Fultz and Lancaster. This variety is a bearded, semihard, red-grained wheat, considerably resistant to leaf rust and drought. It is grown pretty generally throught the country, but especially in the region from Pennsylvania to Oldahoma, including Tennessee and North Carolina to the southward. Recently Professor Saunders, of Canada, has produced a number of new sorts adapted for growing in the Northern States and Canada. Perhaps a half dozen of these — such as Preston, Percy, Dawn, Alpha, Progress, and Countess — are now pretty well known. All the hyl)rids just described have been produced, as a rule, in the most simple way; that is, they were the direct result usualh" of crosses between varieties comparatively closely allied. That they have met with so much success, therefore, is convincing evidence that most remarkable results must follow extensive hybridization experiments with this cereal when composite methods are employed with parents selected from wideh^ different varieties. No experiments completely of this nature have been made in this country. Composite crossing, however, is practiced by A. N. Jones, of New- ark, N. Y., but always with parents comparatively closely allied. He has without doubt done the most important work in wheat hybridiza- tion in this country. Of all American wheat hj^brids recently pro- duced, Jones's varieties are to-day most widely used. In composite crossing, after one or more regular simple crosses have been made, one hybrid is either crossed with a fixed variety or with another hybrid, and the offspring of this last cross may be again crossed with another fixed yariety or hybrid, and so on. In this way the variations that are always induced eyen in ordinary simple crosses are of course multiplied many fold, giving practically an unlimited chance of select- ing from sporting progeny. The results ol^tained from composite crossing, therefore, even with varieties closely allied, are not to be compared with those from simple crosses. Aside from the practice of composite methods, another feature which characterises Jones's work is the tendency he has shown to adhere to a particular aim in all his operations. The Avheats grown in New York and other Eastern States are inclined, on account of the nature of the 71 soil and climate, to be soft and starchy. Recognizing that the best bread flour is made from varieties containing a large proportion of gluten, Jones has given much attention to raising the standard of Eastern varieties in this regard, and has in a large measure succeeded. Of his tirst varieties the two most popular are his Winter Fife and Early Red Clawson. The former is descended! from Fultz, Mediter- ranean, and Russian Velvet, and is a bald, velvet chaff wheat with amber grains, soft or semihard. It is grown chiefly in the Eastern and North Central States, and would be of great value in the Palouse country were it not for its shattering. Early Red Clawson is a hyln-id Mediterrane an Russian Velvet Jjancastev JTuhrid Earh^ White Leader )WinterFife ^Troih Straw Go Ide n Cro s s, Jn Hiibrid jITyhrid Iron Straw arlxf Genesee Giant Fii;. 1.— I>i!iKrani shcuving ix'rtigree of Early Genesee Girtiit. of Clawson and Golden Cross, the last named being a hybrid of Medi- terranean and Chuvson. Though in some respects similar to Clawson, it matures earlier and has a stifler straw. It has a reddish grain. It is a bald, red-chaflV X H O Z CO CO I O S CO > m CO I m TJ XI O m z H I m r > CO H O 3J o CO CO 75 Hybrid = ; [(Medeah X Gypsum) X Hornblende] X [Hornblende X Ward's White] | X Improved Fife. (See fig-. 4.) Medeah is a North African durum wheat. The others are common bread wheats. This new hybrid has been tested by the writer in the field experiments of this department, and was found to be a vigorous sort. Among Continental breeders probably the most important work with cereals has been done by W. Rimpau, of Schlanstedt, German3\ Though not characterized l>y the use of composite methods, Rimpau's work shows a number of important examples of the results obtained by crossing with parents from different wheat groups. Some of the most interesting of the crosses showing various forms similar to the parents and intergrading as to form, color, etc., are the following: Rivett's Bearded Spelt (poulard) X Red German Bearded. Rivett's Bearded X Square-head (club group), and Mainstay X Square-head.^ Black Spelt ffardcastle White TiTiite ChidcLcLm^ Hunqarian Hungctrian ^HedL Mainstcm Wrvite 'Hybrids, fijbrid < 'NubridS Huhridj 7 Fig. 3. — Diagrnin shuwing pudigree of one of the (iartons' hybrid wiipats. As already shown in the earlier part of this bulletin, wheat is, of all the principal cultivated crops, probably the most intluenced by its environment. Connect with this the fact also of its close self-fertiliza- tion, and it is readily explained why there are so many different varie- ties, each best adapted to its particular district. The same variety taken to localities chai-acterized by widely different conditions will gradually change to suit the new conditions, thus giving origin to dif- ferent strains. At the same time new hybrids, when well Hxed. are not likely to be broken up b}' subsequent natural crosses, as in the case 'For an interesting account of ponio of Riinpau's work, written l)y himself, see "Kreuzungsprodukte landwirthschaftlicher Knlturpflanzen." Lanthvirthsehaftliclie Jahrbiicher, Ltl. XX, S. o;>5-o71 (lUus.), 1891. 76 of some other species. It is important, therefore, that all hybrids intended for a particular district should either be produced in that dis- trict or transferred there before they have become fixed, in order that the careful selection necessary may be continued in accordance with the tendencies developed under the influence of the new conditions. Ward's White ^Gypsunh C Jiomblenae ^edeah Hybrid 1 Huhrid 2 KTjbrid4 Improved Fife SyhridJ Fig. 4.— Diagram showing pedigree of one of Farrer's hybrid wheats. Another matter of importance should be noted before leaving this topic. It was supposed for a time, and is still supposed by some, that varieties from different wheat groups will not cross with each other. Often this is true if it is attempted to cross them directly; but it shows another great advantage of composite crossing that if these same varie- ties are first crossed with others of the same group, or with those of groups more nearly allied, the resulting progeny will cross more readily with that of a widely different group. For example, instead of attempting to cross a common wheat with a spelt, the desired result Common Common Club Spelt Spelt Hiihrid 2 HvhricL Svhrid ^ Fig. 5.— Diagram showing hypothetical cross of wheat and spelt. would be more certainly and easily attained 1>y means of a composite cros.s .similar to that shown in the accompanying diagram (fig. 5), and at the same time there is a much better chance offered for selection because of the increased amount of variation thereby induced. 77 Through long, natural "in-and-in breeding'' the qualities of the variety have become specialized, as it were, in harmony with the con- ditions of the environment, and do not readily amalgamate with those of a widely different sort. But once produce variation among these qualities by means of crosses with allied sorts, and it becomes easier to blend them with those of very different sorts. SUMMARY. ■ 1. As a foundation for rational wheat improvement, a knowledge is required of (1) the characteristics and needs of different wheat districts, and (2) the characteristic qualities of the natural groups of wheats. 2. On the basis of conditions of soil and climate and the nature of the varieties adapted to these conditions, the United States may be con- sidered to be divided into eight wheat districts as follows: (1) Soft Wheat district, including mainly the Middle and New England States; (2) Semihard Winter Wheat district, including Ohio, Indiana, Illinois, Michigan, and a small part of Wisconsin; (3) Southern Wheat district, including approximately the Southern States; (4) Hard Spring Wheat district, covering the northern portion of the States of the Plains; (5) Hard Winter Wheat district, covering the central portion of the States of the Plains; (6) Durum Wheat district, covering the southern por- tion of the States of the Plains; (7) Irrigated Wheat district, including approximately the Rocky Mountain and Basin States, and (8) White Wheat district, including the Pacilic Coast States. 3. Certain general needs, such as earl}^ maturity and greater yielding power, are common to all these districts and must be kept constantly in mind in connection with all efforts made to improve varieties. Other characteristics and needs are more special and are stated here- with under headings of the different districts. 4. Soft Wheat district: (a) Present average yield per acre, about 14f bushels. (b) Chief varieties now grown: Fultz, Longberry, Fulcaster, Jones's Winter Fife, Early Genesee Giant, Red Wonder, Mediterranean, Gold Coin, Early Red Clawson, Blue Stem. (c) Needs of the grower: Harder-grained, more glutinous varieties. Hardier winter varieties for the most northern portions. Early maturity. Rust resistance. 5. Semihard Winter Wheat district: (rt) Present average yield per acre, about 14 bushels. {b) Chief varieties now grown: Fultz, Valley, TVii.le, Nigger, Rudy, Dawson's Golden Chaff, Earlv Red Clawson. 78 (c) Needs of the grower: Hardness of grain. Rust resistance. Hardy winter varieties. 6. Southern Wheat district: (a) Present average yield per acre, about Of bushels. (6) Chief varieties at present grown: Fultz, Everett's High Grade, Fulcaster, Boughton, Red May, Currell's Prolific, Rice, Purple Straw, (c) Needs of the grower: Rust resistance. Early maturity. Resistance to late spring frosts. Stiffness of straw. 7. Hard Spring Wheat district: («) Present average yield per acre, about 1.3 liushels. (b) Chief varieties at present grown: Saskatchewan Fife, Wellman's Fife, Scotch Fife, Hayne's Blue Stem, Powers Fife, Bolton's Blue Stem. (c) Needs of the grower: Early maturity. Rust resistance. Drought resistance. Hardy winter varieties. 8. Hard Winter Wheat district: (a) Present average yield per acre, about 121 bushels, (ft) Chief varieties at present grown: Turkey, May, Fulcaster, Zimmerman, Fultz. (c) Needs of the grower: Drought resistance. Hardy winter varieties. Early maturity. 9. Durum Wheat district: («) Present average yield per acre, 11 5 bushels. {b) Chief varieties at present grown: Mediterranean, Fulcaster, Nicaragua, Turkey, (c) Needs of the grower: Macaroni varieties. Drought resistance. Rust resistance. Early maturity. 79 10. Irrigated Wheat district: (a) Present average yield per acre aljout 21 bushels. (6) Chief varieties at present grown: Sonora, Little Club, Taos, Defiance, Felspar, Amethyst, (c) Needs of the grower: Increase of the gluten content. Early maturity. 11. Wliite Wheat district: (a) Present average yield per acre about 141- bushels. {b) Chief varieties at present grown: Australian, Foise, California Club, Palouse Blue Stem, Sonora, Palouse Red Chaff, Oregon Red Chaff, White Winter, Little Club. (c) Needs of the grower: Early maturity. Nonshattering varieties. Hardy winter varieties in the colder portions. 12. The cultivated varieties of wheat are naturally divided into eight rather distinct groups, corresponding to eight botanic species, as fol- lows: (1) Common Bread Wheat {Triticumvulgare)^ (2) Club or Square- head {T. compactum)^ (3) Poulard {T. turgidum)^ (4) Durum {T. durwn), (5) Polish Wheat {T. polonimm), (6) Spelt {T. spelta), (7) Emmer {T. dicoecu7it)^ and (8) Einkorn {T. monococcum). The special character- istics of. these groups of wheats that are of prime importance in the work of wheat breeding are her^ given: (1) Common Bread Wheat group: («) Excellence of gluten content for bread making. {h) Excellence of certain varieties for cracker making, (c) Yielding power of certain sorts. {d) Rust resistance (in some varieties), (e) Winter hardiness of certain varieties. (/) Resistance to drought of certain varieties. {g) Early maturity (in some varieties) . (2) C^lub or Square-head group: {a) Great yielding power. {h) Stiffness of straw. (c) Freedom from shattering. {d) Early maturity (in some varieties). (e) Drought resistance (in some varieties). (/) Excellence of certain sorts for making crackers and breakfast foods. (3) Poulard group: (f/) Excellence of certain varieties for making macaroni. {h) Resistance to orange leaf rust, (c) Resistance to drought. {d) Stiffness of straw. 80 (4) Durum group: {(() Excellence of gluten content for making macaroni and other pastes. ( b) Resistance to drought. (r) Resistance to orange leaf rust. (5) Polish Wheat group: («) Quality of gluten content for making macaroni. {b) Resistance to drought. (c) Resistance to orange leaf rust. (6) Spelt group: Desirable qualities — (o) Ability to hold the grain in the head. (b) Constancy in fertility. (c) Hardiness of certain winter sorts. Undesirable qualities — (d) Brittleness of head. (e) Ru.st liability. (7) Emmer group: Desirable qualities — (a) Ability to hold the grain in the head. (6) Drought resistance. (c) Resistance to orange leaf rust. Undesirable qualities — (rf) Brittleness of the head. (e) Adaptability only for spring sowing, as a rule. (8) Einkorn group: Desirable qualities — (a) Ability to hold the grain in the head. (6) Resistance to orange leaf rust, (o) Hardiness. (d) Resistance to drought. (e) Stiffness of straw. Undesirable quality — (/) Brittleness of the head. 13. Wheats may also be grouped geographically. On this basis groups of varieties having certain special qualities are located approx- imatel}^ as follows: (a) Starchy white wheats: Pacific Coast and Rocky Mountain States, Chile, Turkestan, Australia, and India. {b) Amber or reddish grained wheats, also starchy: Eastern States, western and northern Europe, India, Japan, and Australia. (c) Excellence of gluten content for making the best bread: Northern and Cen- tral States of the Plains, Canada, eastern and southern Russia, Hungary, Rou- mania, and southern Argentina. (d) Resistance to orange leaf rust: Southern Russia, Mediterranean and Black Sea regions, and Australia. (e) Excellence of gluten content for making macaroni: Southern Russia, Algeria, and the Mediterranean region in general. (/) Stiffness of straw preventing lodging: Pacific Coast States, Japan, Turkestan, Mediterranean region, and Australia. ig) Yielding jiower (at least in proportion to size of head): Pacific Coast States, Chile, and Turkestan. (h) Nonshattering varieties: Pacific Coast States, Chile, Turkestan, Germany (spelts), and East Russia (emmers. ) 81 (i) Constancy in fertility: Germany (spelts) and southern Europe. (j) Early maturity: Japan, Australia, and India. (k) Resistance to drought and heat: East and South Russia, Kirghiz Steppes, Turkestan, and southern Mediterranean region. (0 Resistance to drought and cold: East Russia. 14. Of the many wheat introductions made into this country in the past, the following are among* those of the greatest moment, and which have completely revolutionized the wheat industry in places: (a) Mediterranean, introduced first in 1819. (b) Fife wheats, introduced first into Canada and then into the northern States of the Plains. (c) Turkey wheat, first introduced into Kansas about twenty-five years ago from Taurida, in Russia. (d) The California Club, Australian, and Sonora, introduced into the Pacific coast States. 15. Field experiments of the Department have shown that in the common bread-wheat group there is a very close constant relation between the autumn condition of the young plant on the one hand and winter hardiness and quality of grain on the other. 16. Wheat is very susceptible to changes of environment, but espe- cially in regard to color, hardiness, and chemical composition of the grain. 17. In general, regions possessing black prairie soils and character- ized by violent climatic extremes, especially extremes of heat and drought, produce wheats that are hardiest, have the hardest grains, and are the best in quantity and quality of gluten content. 18. Considering all qualities, the best wheats of the world are of Russian origin, coming particularly from eastern and .southern Russia, the Kirghiz steppes, and Turkestan. Of Russian varieties so far known, the following are among the best, if not the very best: Arnautka, Turkey, Kubanka, Ghirka Spring, Ghirka Winter, Russian, Crimean, Buivola, Sarui-bug-dai, Kubanka Red Winter, Mennonite, Yx, Chernokoloska, Beloturka. 19. The earliest ripening wheats are often dwarfed. The following varieties are among the best of this class: Yemide, Early Baart, Onigara, Early Japanese, Shiro-Yemidashi, Japanese No. 2, Kinta,nia, Nashi, Kathia, AUora Spring, Roseworthy, Stein wedel, King's Julnlee. 4879— No. 24 «) 82 '20. The following varieties are among the best known of the durum and poulard groups: Arnautka, Galland's Hybrid, Kubaiika, El Safra, Beloturka, Petanielle noire de Nice, Chernokoloska, Volo, Medeah, Missogen, Sarui-bug-dai, Atalanti, Cretan, Nicaragua. 21. Common bread wheats can not be depended upon to resist rust, but the best in this regard are probably the following: Turkey, Crimean, Pringle's Defiance, Oregon Club, Rieti, Odessa, Pringle's No. 5, Mennonite, jfashi. Velvet Blue Stem, Saskatchewan Fife, Sonora, Theiss, Prolifero, Bellevue Talavera, Mediterranean, Arnold's Hybrid, Deitz Longberry. 22. Einkorns resist leaf rust completely, and emmers resist it to a high degree. 23. Some of the very hardiest winter varieties so far tried in this country are: Turkey, Crimean, Yx, Ghirka Winter, Bearded Winter. 24. Club wheats are usually soft-grained and tender sorts, adapted only to mild climates like that of California. Among the best of this group are: Little Club, Palouse Red Chaff, California Club, Chili Club, Herisson barbu, Sicilian Red Square-head, Herisson sans barbes. 25. Some of the most popular and valuable wheats of our country have been produced by simple selection, though in some cases the indi- cations are strong that they were originally the result of natural crossing. The best known of such varieties are: Fultz, Rudy, Clawson, Wellman's Fife, Gold Coin, Currell's Prolific. 26. Selection plays far the most important part in wheat breeding, and necessitates on the part of the experimenter a thorough knowl- edge of varieties and their relations to each other and to their envi- ronment. 83 27. Simple selection of individuals, however, for the improvement of the same variety can and should be practiced on every farm. Very little extra time or trouble is required, but the gain is great. 28. Among the most valuable wheats of the United States that have been produced through hybridization are the following: Fulcaster, Pringle's Defiance, Gypsum, Pringle's No.5, Improved Fife, Hornblende, Quartz, Felspar, Ruby, Blount's No. 10, Jones's Winter Fife, Diamond Grit, Early Genesee Giant, Early Red Clawson, Early Arcadian, Early White Leader. 29. For the most complete success in wheat improvement through hybridization it is necessary to practice composite crossing with parents selected from widely different wheat groups. 30. The wheat plant is so closely self-fertilized in nature that the practice of composite crossing produces some most interesting and remarkable results. There is apparently no end to the variations exhibited by the sporting progeny in such cases, and, accompanied by discriminating selection, the possibilities of wheat improvement in this way are practically unlimited. INDEX OF VARIETIES. Page. Algerian 44, 4.5 Allora Spring 43, 44, 4.5, 63, 81 Alpha VO Alsace 44,4.5,61,63 American 44, 45 American Bronze 44, 45 Ames 44, 45 Amethyst . , 21, 44, 45, 69, 79 Amidonnier. (^'eeEmme^.) Arnautka 19, 31, 44, 45, 62, 63, 81, 82 Arnold's Hybrid 44, 45, 63, 82 A six rangs 44, 45 Assiniboine Fife 44, 45 Astrakhan 32 Atalanti 44,45,63,82 Australian 22, 25, 40, 43, 44, 45, 79, 81 Au.stralian Indian 44, 45 Au.stralian Purple Straw 44, 45 Baggi 44, 45 Banat 28, 44, 45, 63 Barletta 44, 45 Barley wheats. {See Durum wheats.) Basalt 44, 45 Bauchiger Weizen. {See Poulard wheats. ) Bearded Winter 44, 45, 62, 63, 82 Bearded Winter Fife 72 Bellevue Talavera 44, 45, 63, 82 Belokoloska 44, 45 Beloturka 15, 44, 45, 62, 63, 74, 81, 82 Berthoud 44,45 Bianchetta 44, 45 Big English 44,45 Big Frame 44, 45 Black Spelt 74, 75 Black Velvet 44, 45 Bl(5 petanielle. (See Poulard wheats.) Blount's Fife 44, 45, 74 Blount's No. 10 44, 45, 69, 70, 83 Blue Stem 13, 16, 44, 45, 77 Bolton's Blue Stem 17, 44, 45, 66, 78 Boughton 15, 78 Bread wheats 7, 26, 28, 30, 31, 32, 33, 37, 43, 63, 71, 74, 75, 76, 79 Buca Nera 44. 45 Bucke ve 44, 45 Budapest 39, 46, 47 Buivola 28, 62, 81 California Club 2.5, 63, 79, 81, 82 Candeal Redondo 46, 47 Canning Downs 14, 46, 47, 63 Cape 46, 47 Cartagena 46, 47 Chernokoloska 46, 47, 62, 63, 81, 82 Chcrnouska 46, 47 Chiddam de Mars rouge 46, 47 Chili 46, 47 (;hili Club 22, 46, 47, 63, 82 China Red 46,47 China Tea 46, 47 China White 46, 47 Chinese 46, 47 Clhubut 46, 47, 63 Clawson 46, 47, 59, 65, 66, 71, 82 Club wheats 22, 23, 24, 28, 29, 37, 40, 43, 63, 73, 74, 75, 76, 79, 82 Common wheats. {See Bread wheats. ) Composite wheats 30 Countess 70 Cretan 46, 47, 63, 82 (Crimean 28, 46, 47, 62, («, 81 , 82 Currell's Prolific 15, 46, 47, 78, 82 Page. Dallas 46, 47 D'Arblay's Hungarian 46,47,74 Daruma" 46, 47, 63 Dattel 46,47 Dawn 70 Dawson's Golden ChaflE 14, 39, 46, 47, 77 Defiance 15,21,22,46,47,79 Deitz 46,47 Deitz Longberry 63, 82 De la Basse 46, 47 Diamond Grit 21, 46, 47, 72, 83 Diehl Mediterranean 46,47,66 Dinkel. (See Spelt.) Dividenden 46, 47 Duro di Apulia 46,47 Durum wheats 8, 15, 19, 2(i, 29, 30, 31, 32, 33, 37, 40, 43, 60, 63, 72, 73, 74, 75, 79, 80 Earlv Arcadian 46, 47, 72, 83 Early Baart 46, 47, 63, 81 Early Genesee Giant 13, 43, 46, 47, 71, 72, 77, 83 Early Japanese 48, 49, 63, 74, 81 Earlv May 43,48,49 Earl V Red Clawson .... 13, 14, 48, 49, 65, 71, 72, 77, 83 Earlv Rice 48,49 Earlv White Leader 71,83 Einkorn 35, 36, 37, 43, 48, 49, 63, 73, 79, 80, 82 El Safra 48, 49, 63, 82 Emmer 30, 33, 34, 35, 37, 43, 72, 73, 79, 80, 82 English wheats, (.sve Poulard wheats.) Engrain. (See Einkorn.) Engrain double 36, 48, 49 Entre Rios 48,49 Epeautre 33 Essex Red 74 Everett's High Grade 15, 48, 49, 78 Farquhar 48, 49 Farrer's Durum 48, 49 Felspar 21, 48, 49, 69, 70, 79, 83 Fern.. 48.49 Fife 16,21,28,31,38,65,67,81 Flourelle 48,49 Fluorspar 48,49 Foise 23,2.5,79 Frampton 48,49 Frances 48,49 Frankenstein 48, 49 Fulcaster 13, 15, 18, 20, 28, 48, 49, 63, 70, 77, 78, 83 Fultz 13, 14, 15, 18, 39, 48, 49, 63, 65, 70, 71, 74, 77, 78, 82 Galland's Hybrid 48,49,63,82 German Amber 48, 49 German Emperor 48, 49 Gerstenweizen. {See Durum wheats.) Gharn( )vka 48, 49 Ghirka 28,39 Ghirkii Spring .56, 57, 62, 63, 81 Ghirka Winter 58,62,63,81,82 Giant Rye. {See Polish wheats.) Glasgow 38 Glvndon 673 48,49 Glvndon Sll 48,49 Gold ( -oin 13, 16, 48, 49, 66, 77, 82 Golden (Toss 48,49,71 Golden Cross Jr 71 Goldene Aue 48, 49 GoliU'u ( iate Club '-'2 Graf Walderdorff's Regenerated 48, 49 Granite ^9 Grass ( .sV'C Odessa. ) G viisum 48, 49, 69, 74, 75. 76, 83 Hairkani 48, 49 85 86 Page. Hallett'f? Pedigree 48, 49 Hardcastlc White 74, 75 Hayne's Blue Stem 17, 48, 49, 66, 67, 78 Herisson barbii 50, 51, 63, 82 Herisson sans barbes 50, 51, 63, 82 Hickling 50, 51 Hopetowu 50, 51 Hornblende 50, 51, 69, 74, 75, 76, 83 Hudson's Early Purple Straw 50, 51 Hundred Fold 30 Hungarian Red 74, 75 Hungarian White 74 Hunter's White 74 Igel mit Grannen 50, 51 Igel ohne Grannen 50, 51 Imperial 50, 51 Improved Fife 50, 51, 61, 69, 70, 74, 75, 76, 83 Iron Straw 71 Japanese No. 1 50,51,63 Japanese No. 2 50, 51, 63, 81 Japanese No. 4 50,51,63 Jejar de Valencia 50, 51 Jerusalem Rye. (See Polish wheats.) Jones's Square-head 50, 51 Jones's Winter Fife 13, 50, 51, 71, 72, 77, 83 Kastamuni 50, 51 Kathia 60,51,63,81 Khel 50,61 King's Jubilee 43, 50, 51, 63, 81 Kinney 50, 51 Kintama 50,51,63,81 Krasnokoloska .60, 51 Kubanka 19, 31, .60, 51, 61, 62, 63, 81, 82 Kubanka Red Winter 62, 81 Kubb 50, 51 Ladoga 50, 51 Lai 50,61 Lamed 50, 51 Lancaster .50, 51, 63, 66, 70, 71 Lehigh 50, .51 Linaza 50, 61 Little Club 21, '24, 25, 43, 50, 51, 60, 63, 79, 82 Longberrv 13, 77 Lost Nation 50,61 Macaroni wheat. {See Durum, Poulard, and Polish wheats.) Mainstay 74, 75 May 15,18,78 McKissick's Fife 50, 61 Mealy 52, 53 Medeah 52,53,63,74,75,76,82 Mediterranean 13, 19, 20, 28, 38, .52, 63, 63, 71, 77, 78, 81, 82 Meekins 52, 63 Melka 52, 53 Mennonite 52,53,62,63,81,82 Minnesota Fife 52, .53 Miracle 30 Mirado 62, 63 Missogen 52,63,63,82 Moscow 52, 53 Mundia 52, 53 Murcia 52, .53 Muzaffarnagar .52, 53 Nab-el-bel 52,53 Nashi 52, ,63, 63, 81, 82 Nicaragua 15, 19, 20, 31, 32, 40, 63, 78, 82 Nigger 14, 52, 63, 77 Noe 52,63 Nonetto de Lausanne 52, 53 Nonpareil 52,53 No. 6. ( See Gold Coin. ) Odessa 39,52,63,62,63,82 Onigara 52,53,63,8] Oregon Club 52, .53, 63, 82 Oregon No. 10 70 Oregon Red Chaff 23,25,79 Palouse Blue Stem 16, 22, 24, 26, 40, 42, 52, 63, 60, 61 , 79 Palouse Red Chaff 22, 25, 63, 79, 82 Pedigree Red 74, 76 Pedigree White 74 Penquite's Velvet Chaff 52, 63 Percv 70 Petanielle noire de Nice 62, 53, 63, 82 Pilli 52, 53 Pissi Hydrabadi 52, 53 Page. Polba. {See Emmcr.) Polish wheat.... 31,32,33,37,43,52,53,63,73,79,80 Poole 14, 39, 52, 63, 59, 77 Poulard wheats 8, 26, 29, 30, 31, 37, 43, 60, 63, 73, 74, 75, 79 Power's Fife 17, 52, 53, 66, 78 Preston 70 Pringle's Best 69 Pringle's Defiance 62, 53, 63, 69, 82, 83 Pringle's No. 4 69 Pringle's No. 5 52, 53, 63, 69, 82, 83 Pringle's No. 6 69 Probsteier 52, 63 Progress 70 Prolifero 52, 63, 63, 82 Prophet 52, 53 Propo 22, 54, 55 Pulavka 54, 55 Purple Straw 15, 40, 54, 55, 78 Quartz 54, 55, 69, 74, 83 Rattling Jack 54, 55 Red Bearded 54,55 Red Chaff. {See Oregon Red Chaff and Pa- louse Red Chaff.) Red Chaff Club. {See Palouse Red Chaff.) Reddish White Bearded 54,55 Red Fife 54, 55 Red German Bearded 75 Red May 16, 78 Red Provence 54, 55 Red Spring 54, .55 Red Tyrol 54,55 Red Winter 54, 55, 62, 63 Red Wonder 13, 77 Rice 16,64,56,78 Rieti 54, 55, 63, 82 Rio Grande 64,55 Rivett's Bearded Spelt 75 Rivet wheats. (iSee Poulard wheats.) Roseworthy .- 54, 55, 63, 81 Ruby 69,70,83 Rudy 14, 23, 54, 65, 66, 77, 82 Russian 62, 81 Russian Hard 54, 55 Russian Spring 54,55 Russian Velvet 71 Rve Wheat -. 54,55 Safeed 54,. 55 Saida 54, 55 Saldom6 54,66 Salt Lake Club 22 Samara 64, 65 Sandomir 54, .55, 62 Saratov 54, 55 Sarui-bug-dai 54, 55, 62, 63, 81, 82 Saskatchewan Fife. . . . : 17, 54, 55, 63, 65, 78, 82 Saumur Winter 64, 55 Scotch 38 Scotch Fife 17,54,66,61,65,78 Seneca Chief 54, 55 Seven-headed ■. 30,54,55 Shirosawa 54, 55 Shiro-yemidashi 54, 55, 63, 81 Sicilian Red Square-head 54, 55, 63, 74, 82 Sindhi 56,.57 Sonora 21, 22, 25, 40, 66, 67, 63, 79, 81 , 82 Soules 56,67 Spelt 33, 34, 35, 37, 43, 72, 73, 76, 79, 80, 81 Spelz. {.See Spelt.) Spring Ghirka (.see also Ghirka Spring) .56, 57 Square-head 75 Square-head wheats. (See Club wheats.) Steinwedel 66, 67, 63, 81 Swamp 56, .67 Taganrog ,. 56, 57 Talavera 56, 57, 74 Tangarotto 56, 57, 63 Taos 21, 66, 57, 79 Tasmanian Red 56, 57, 63 Theiss 28, 39, 56, 57, 63, 82 Tourmaline 56, 57, 74 Touzelle 56, 57 Trimenia 66, 57 Tritieu m 26, 26 Triticum compact am 22, 26, 28, 29, 79 Triticum cowpo.srtWHi 30 Triticuvi dicoccum 26, 30, 33, 34, 79 87 Page. Triticim durum 8, 26, 29, 30, 79 Triticum inonoco( cum 26, 35, 79 Triticu m polonicum 26, 32, 79 Triticum spelta 26,33,79 Triticum turgidum 8, 26, 29, 79 Triticum vuigare 7, 8, 26, 29, 79 Turkey 17, '18, 20, 21, 28, 31, 39, 56, 57, 59, 60, 62, 63, 78, 81, 82 Tuscan 56, 57 Ulka 28,39,62 Urtoba 56, 57 Valley 14,56,57,63,77 Varesotto 56, 57 Velvet Blue Stem 16, 28, 31, 56, 57, 63, 65, 67, 82 Velvet Chaff 56, 57 Victoria d' Aiitomne 56, 57 Victorian Defiance 56, 57 Volo 56, 57, 63, 82 Vyssoko-Litovsk 56, 57 Walla Walla - - 56,57 Page. Walker 56, 57 Ward's Prolific 56,57,74 Ward's White 74, 75, 76 Wellman's Fife 17, 58, 65, 66, 67, 78, 82 White Chiddam 74, 75 White Clawson. (See Clawson.) White Michigan 39 White Polish 32 White Tuscan .58 White Winter 25,58,79 Wild Goose 40, 58, 63 Winter Fife. (See Jones's Winter Fife.) Winter Ghirka (see, also, Ghirka Winter)... 58 Wonder 30 Wyandotte Red 58 Yemide 43, 58, 63, 81 Yx 58, 62, 63, 81, 82 Zaruta 58 Zimmerman 18, 58, 74, 78 o Bulletin No, 25. V. p. p.— 78. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. GALL-bwAY, Chief. SOME DISEASES OF NEW ENGLAND CONIFERS: A PRELIMINARY REPORT. BY HERMANN VON SCHRENK, Instructor in Botany^ Henry Shaw Sr/iool of Botany, Special Ayent, Division of Vegetable Physiolocfy and Pathology. WASHINGTON: GOVERNMKNT PRINTING OFFICE. I y oo. DIVISION OF VEGETABLE PHYSIOLOfn' AND PATHOLOGY. SCIENTIFIC STAFF. B. T. Galloway, Chief of Division, Albekt F. Woods, Assistant Chief. ASSOCIATES. EmviN F. Smith, Mertox B. Waitk, Newton B. Piekce, Herbert J. Webber, M. A. Garleton, r. TI. DoKSETT, OsCAli LoEW, AVm. a. Orton, Ernst A. Bessey, Flora W. Patterson, Hermann von Sciirenk,' Marcts L. Fj.oyo.^ IN charoe of laboratories. A-lbert F. Woods, Plant PhysioJogij . Erwin F. Smith, Plant Pathology. Newton B. Pierce, Pacific Con. y the action of the mycelium of this or that fungus, and passes on before it, up the heart- wood of old branches and out through them, forming resinous lumps, which harden in contact with the air. These lumps occur at all heights on the trunk and increase in size from year to year. The accumulation of these resinous masses prevents the normal healing of the wound or healing over of the stub of the branch, and results in the formation of a marked protuberance at that point, commonly called a knot, with its various modifications. The turpentine often drips from such a spot or runs down the bark in small streams. It may be many years before the decomposition within the tree advances sufficiently to enable sporo- phores to form, and a system of prophylactic treatment must take into account phenomena such as these to aid in detecting diseases in their early stages. What has been said with reference to these resin accu- mulations applies particularly to fungi like Trametes jnni {^vot.)FY. and its form ahletis Karsten, to Polyporm schweinitzU, and Pohjjyorm sulfurcm^ and one or two others not yet definitely identified. RELATION TO INSECT ATTACKS. The nature of the fungus injury is often very obscure, and there are so many factors which have to be considered in tracing the nature of any one disease that the results of the present paper are but fragmen- tary, and it is very probable that they will be modified largely by future discoveries. The intimate relationship which exists between the attacks of insects on the one hand and fungi on the other nuist be pointed out. There are without doul)t many fungi which find their way into the wood of trees through the holes which l)oring insects have made in the bai-k. The injury which the insect makes may be very slight, but it has opened the way for the action of the fungus, which may be very destructive. An example of this kind is to be found in that most curious of all the Poh/porel^ Poh/ponis volvatm Peck. This grows on the trunks of spruces which have been attacked by various spe(•ie^^ of boring beetles, notably species of Dcndroctonus. These be(»tles bore throuirh the bark into the cambium laver. The fungus enters through 14 these holes and grows in the sapwood of the tree, destroying it in a few mouths. Whether it grows there while the tree is still alive, and what its possible relations may be to the Dendroctouus, are problems yet to be solved. In many parts of the Maine woods every tree where the beetles had been or were still active was covered with the rounded fruiting organs of this Polyporus. (See PI. I, fig. 2.) Their association with this Polyporus offers a promising field for study. The holes made by the beetles allow the spores of several other fungi to enter, notably those of Poly poTUH pi ni cola. These germinate and grow throughout the heartwood, rendering it worthless in a very short period. The possible role which beetles and boring larvaB may play in carry- ing the spores of a fungus from one tree to another will be referred to below. These few instances will serve to show that it is all important that a study of the insect and fungus enemies of a tree should be made hand in hand. There are grave inherent difficulties in deteimining the exact cause of death of a large tree, for there are many factors which may influence its growth so that the tree becomes weakened. There is a widespread opinion that insects or fungi will not attack an absolutely healthy tree, but that the latter must be more or less weakened before such an attack takes place. That this is not always the case need hardly be said, but the mere fact that a fungus is growing in the tree or an insect is at hand upon it is no positive proof that one or the other is the active agent in bringing about its death. Such evidence, particularly if oft repeated, will become very valuable when taken in conjunction with other proofs. SCOPE OF THIS REPORT. In the following a number of fungi will be described, together with the characteristic changes which their mycelia induce in the wood of the trees in which they grow. These fungi were found again and again, always associated with the forms of decay ascribed to them, and never was such decay found without the fungus in question, or with- out a mycelium from which the fruiting portion of the fungus devel- oped. These fungi occurred on all coniferous forest trees, with few exceptions. Some of them started in the living trees and caused the heartwood to decay. They were found in large numbers destroying trees injured by insects, and on some tracts where fire had swept through the woods and had injured the bases of the tree trunks sev- eral of them had gained a foothold and had destroyed every tree thus injured. The principal ones met with were: Pohjporus sclnoeinitzii Fr.; Polypmnis pnnicola (Swartz) Fr., Trametes pini (Brot.) Fr. forma alietk Karsten; Pohjporm mlfnrew (Bull.) Fr.; and Polyjwrus suh- acidns Peck. A number of doubtful forms will be mentioned near the end of this report. 15 NEW ENGLAND FORESTS. VEGETATIVE CONDITIONS. The original forests of most of the New England States are gone. The White Pine, which at the advent of the white settler formed such a large part of the forests, is present in any large quantity only in the most inaccessible places and elsewhere as ripe timber only in isolated spots. The chief forest trees from the lumberman's standpoint are the Red Spruce and the WhitQ Spruce. Millions of feet of Red Spruce lumber are now being cut year after year in the States of Vermont, New Hampshire, and Maine. The time is not far distant, however, when the stand of spruce timber will be in a similar condition to that in which the White Pine is now. The conditions which prevail in the forests of Maine and New Hamp- shire can be touched upon only in so far as they relate to the presence of and probable influence on the diseases which form the basis of this report. The forests are usually moist. The forest floor is covered with a large variety of mosses, which hold water very readily. Sphag- num covers many square miles. Springs and brooks are abundant everywhere. The annual rainfall, often very heavy during the spring and summer months, accounts for the general humidity of the air. Near the coast the fogs keep the woods moist for a large portion of the growing season. The summer season is usually comparatively short, but while it lasts very warm days are not uncommon. Warmth and humidity, chiefly the latter, are very influential in promoting the growth of many saprophytic as well as parasitic fungi. Before describing the various fungi and their efl'ects, it may be well to say something of the trees which are affected by these fungi. RED SPRUCE. Foremost among the coniferous trees of New England at the pres- ent time is the Red Spruce, Picea ruhens Sarg. (P. mariana (Mill.) B. S. P., P. nigra Link). It is a tall, stately tree, which grows to be 70-80 feet (21-2^ meters) high and 2-3 feet (0.6-1 meter) in diameter. It occurs all over northern New England, together with the Balsam Fir and White Pine. Sargent says of this tree:^ Picea ru})ens, which is the principal timber spruce of the northeastern United States, and, with the exception of the White Pine, the most valuable coniferous timber tree of the region which it inhabits, produces light, soft, close-grained wood, which is not strong nor durable when exposed to the weather. It is pale, slightly tinged witli red, with paler sapwood about two inches thick, and a satiny surface * * *. Now that the most valuable White Pine has been exhausted in the forests of the Northeastern States, the Red i^pruce is their most important timber tree, and immense (piantities of its hun1)er are manufactured every year from trees cut in Maine, New Hampshire, Vermont, and northern New York. * * * ' Sargent, G. S. Silva of North America. 12:35. 1898. 16 The wood of the Red Spruce is used for construction, and thousands of trees of all sizes also find their way to the pulp mills for the manu- facture of paper. During the summer of 1899 several large new mills were building in central Maine, one of which was expected to con- sume 300 tons of spruce wood daily. In a recent article in The Forester, Mr. Lyman, of the International Paper Company, discusses at length the use of this spruce for making pulp. The tree is one of moderately slow growth. It reproduces itself well from seed, and grows up readily to replace the original stand of timber. In the forest, when growing in close stands, the lower branches die gradually and break off, leaving dead stubs which, in the case of larger branches, offer inviting spots for the entrance of fungus spores for several years after the fall of the dead branch. Attention has already been called b}^ the writer to the manner in which diflferent trees heal the wounds caused by dead branches.^ There are resin channels scattered through the summer wood. Their number varies considerably in the individual tree. In some trees there are but one or two in a given ring, while six or eight years later there may be two or three dozen. WHITE SPRUCE. The White Spruce, Plcea ccmadensis (Mill.) B. S. P. (P. alha Link), a very much more stately tree than the Red Spruce, grows to a height of 150 feet (about 16 meters), with a trunk 3 to 1 feet (0.9 to 1.2 meters) in diameter. In the Northeastern States it is found in abundance, especially along the coast, and on some of the islands it is the only tree. It is widely distributed to the north and northwest, extending into Alaska. In the New England States it is not as abundant as the Red Spruce and is not used for lumber purposes to the same extent as its near relative. In the eastern provinces of Canada, where it is prol)a])ly the only Sprnce cut in large quantities, it is used in construction and for the interior tinish of buildings and for paper pulp. * * * White Spruce lumber is also occasionally -manufactured in Dakota and Montana, etc.^ * * * The wood of the White Spruce is straw yellow, very light, and not strong. Resin passages occur now and then in the very narrow band of summer cells. As an ornamental tree it is more extensively used than the Red Spruce. ^Von Schrenk, H., Two Diseases of Red Cedar. Bui. No. 21, U. S. Dept. Agr., Div. Veg. Phys. and Path. 2 Sargent, C. S. Silva of North America. 12:37. 1898. 17 BALSAM FIR. The Balsam Fir/ Ahies halsamea (L.) Miller, is a tree common all over New England, .springing- up wherever the White Pine or Spruce are cut away. It produces great quantities of seed, which germinate readily the succeeding year. The trees are usually smaller than the Spruces, growing to be 50 feet (15 meters) in height and 6 inches to 1 foot (15 to 30 cm.) in diameter. Its wood is used for a cheap grade of lumber, for it is ver^ light and does not have any resisting power. In central Maine it is often cut with the Spruce and sent to the pulp mills. The trees are very subject to the attacks of insects and fungi. The large black ants^ annually destroy hundreds of trees. HEMLOCK. The Hemlock, Tsuga canadensis (L. ) Carriere, is a stately tree, usually 60 feet (18 meters) in height, having a trunk 2 to 4 feet (0.6 to 1.2 meters) in diameter. It is an important element of the northern forest, and has long been valued for its l)ark, which is extensivelj^ used in the tanning of leather. As an ornamental tree it has few equals among our native trees. In stately grace it has no rival among the inhabitants of the gardens of the northern United States, when, with its long lower branches sweeping the lawn, it rises into a great pyramid, dark and somber in winter and light in early summer, with the tender yellow tones of its drooping branchlets and vernal foliage.'^ It is one of the most valuable trees of the Eastern forests. It is estimated that in the year 1887, 1,200,000 tons of bark of this tree were harvested, and although a large part of the timber of the trees cut and stripped of their bark is allowed to rot on the ground it is believed that the average annual value of the material of all kinds obtained from this hemlock is not less than $30,000,000. The tree is one which grows ver}^ slowly. The seedlings are very sensitive to exposui'e and do not recover I'cadily Avhen inju)-ed. The wood is very coarse and ])rittle and is worked with ditKeulty. It is, however, used considerably in various localities for a cheap grade of lumber, and at times, when other wood is not to be had, for railway ties, fence posts, and railing, but its resisting powers to weathering inlBluences are very slight. ARBOR VITiE. The Arbor Vita\ Thuja, occidentalk^ L.,* is a tree found throughout the northern parts of New England, pai'ticularly in wet, boggy lands, where it forms dense forests, the individual members of which grow 'Sargent, C. S. Silva of North America. 12:107,108. 18<)S. ■"Tlopkins, A. I). Preliminary Report on the Jiisect Knemies of the Forests of the Northwest. Bui. No. 21, U. S. Dept. Agr., Div. Entomology. 1899. •''Sargent, loc. cit. ()(>. * Sargent, C. S. Silva of Nortii America. 10: 120. 1896. 5776— No. 25 2 18 to be 50 feet (15 meters) in height, with trunks 6 inches to 1 foot (15 to 30 cm.) in diameter. The wood is very durable and is on that account prized for fence posts and railway ties, for foundation walls, and for making shingles. The wood itself is rather coarse, yellow brown, and is free from resin ducts. The trees are grown as orna- mental trees, particularly in the form of hedges. WHITE PINE. The White Pine, Pinus 8trohxis L., once so large a factor in the lum- ber industry of the New England States, is now comparativeh' rare as mature timber. It is subject to a number of diseases which will be treated of in a special paper. It is left out of consideration on that account in the present report. TAMARACK. The Larch, or Tamarack, Larix laricina (Du Roi) Koch (Z. ameri- cana Michaux),^ is a tall, stateh^ forest tree which is found growing with the White Pine and Spruce and in some sections forms extensive forests, especially in low swampy lands. It grows throughout the Northern States, ranging from Maine westward to the western slopes of the Rocky Mountains, and southward to northern Pennsylvania, Indi- ana, and Illinois, and to central Minnesota. As an ornamental tree it is highly prized because of its graceful habit and thrifty growth. The wood of the Tamarack is extensivel}' used in shipbuilding, for railway ties, and telegraph poles. It is veiy durable and hard. Compared with the AVhite Oak, it has a crushing strength of 1.38. Dudle}'" sa3^s of it: The quality of the wood of this tree is such that it deserves to be widely known and more extensively used for ties than it has been. * * * The wood is easily treated with antisejitics to prevent decay, especially with sulphate or acetate of iron, and ties so treated have lasted over thirty years under heavy traffic. POIiYPORUS SCHWEINITZII Fr. Polyporus schweinitzii Fr., Syst. 1 :351. Polyporus schweinitzii Fr. , Epic. 433. Boletus sistotremoides Alb. and Schw., 243. [Figured in Fries's Icon. Hyni. No. 179.] OCCURRENCE. This fungus is one which is very common throughout the Northern forests on the Spruce and Fir, and, as Dr. Farlow remarks,'' appears to be very much more prevalent in this country than in Europe.* It certainly stands near the top of the list in point of destructiveness. 1 Sargent, C. S. Silva of North America. 12:7. 1898. ^Dudley, P. H. Bui. No. 1, Division of Forestry. Ajjpendix I. 51. "Sargent, C. S. Silva of North America. 11:11. 1897. ''Hartig (Lehrbuch der Pflanzenkrankheiten. 177. 1900) says it occurs only on Pines. 19 It attacks young trees as well as older ones, entering the tree through the root system and growing up into the trunk for sometimes 40 and 50 feet (12 to 15 meters) from the ground. The mycelium makes the wood of the Spruce very brittle. Diseased wood is of a yellowish color; it has a cheesy consistency so that it can be cut across the grain with a knife quite readily and without much resistance. When dry, it is readily powdered. The brilliantly colored. fruiting bodies are to be found in July and August growing about the base of the affected trees, more rarely on the trunks. (See text fig. 1; also PI. I, fig 1.) It was found more frequently in places where the air was laden with moisture — for instance, along the coast and near lakes. On many of the islands which lie off the Maine coast the fungus was found to be very plentiful, even to a distance of 5 miles (8 kilometers) from the mainland, showing that the spores must be carried for a considerable distance. One small island had some 12 trees on it, all White Spruces, of which 7 had old fruiting organs of this fungus growing about the bases of the trunks. STRUCTURE OF DISEASED WOOD. The wood of the Red Spruce or the Fir when first invaded b}^ the mycelium turns yellow, and after a time cracks here and there as if dried rapidly. A cross section of the trunk of a young Fir, made about 6 feet (about 2 meters) above the ground, is shown in PI. II. The large crack at the side was made in chopping down the tree; the other cracks in the heartwood show plainl}^ how the wood has shrunken. The structural changes which take place are as follows: Soon after the myce- lium enters the wood of the Spruce the color changes and the wood becomes more or less brittle. This is due to the fact that at various points in the summer wood cracks appear in the walls of the tracheids and extend in the spiral direction around each tracheid. The break deepens gradually until it extends entirel}^ through the secondary lamella up to the primary lamella. The latter remains unbroken. The spiral breaks increase in number and at last the tracheid has the appearance shown in PI. IX, fig. 1. There appear to be two series of cracks, one extending upward from left to right, the other from right to left. This appearance is due to the fact, as Hartig has shown, that one sees the breaks in the walls of two tracheids at the same time. Hartig mentions that these cracks all extend in a spiral direction, none paralh'l to the walls. This is certainly a striking fact, and seems to distinguish wood attacked by this fungus from that injured by many others. It will be shown that some other fungi, Polyporus sul- fureiis and a form of Polyporus destructor possibly, attack the wood of the Spruce and the Yellow Pine, respectively, in a similar way. The spring wood has few cracks. These aic inaiidy in the pits, where four radiating cracks appear in the secondary lamella. 20 Wherever a hypha has passed through a wall is to be found the peculiar double spiral crack. (PL IX, fig. 1.) The diseased wood of the Balsam Fir differs little from similarly affected Spruce wood. The summer tracheids as a rule are not so wide as those of the Spruce, hence the spiral cracks are not as evident as in that tree. They appear to extend more or less parallel with the walls. They are likewise present in greater numbers, so that there is very little left of the wall. Wood which is ir. the last stages of deca}^ is exceedingly brittle. It does not partake of the character of brown charcoal as much as does Pine wood similarly diseased, but is much firmer. It absorbs water very rapidly, and when boiled in water for a few minutes becomes soft and putty like and can be kneaded like bread dough. When dry it can not be cut with a knife without crumbling, but when soaked in water it can be readily cut into the thinnest sections. These have no elasticity, however. The walls of the wood cells are very thin and swell to several times their size on addition of dilute potash. Here and there are found masses of resin, more frequently in the Balsam Fir than in the Spruce. As the wood grows older the action of the mycelium seems to stop. The wood changes no further except that it cracks more or less. It appears to be very resistant to change brought about by weathering. FRinTING ORGAN. The first fruiting bodies observed began to appear toward the begin- ning of July. Small rounded masses grew out from the l)ark and very soon became flattened horizontally. The specimen shown in the photo- graph (fig. 1) was watched closely and measured daily from the time of its first appearance until it had reached its full size. By means of wires stuck at the edge of the growing shelves, it was easy to measure accurately the daily increase in diameter. The hyphaj rapidly grew around the wire so that it became embedded in the mass of the sporo- phore. One of the wires is visible at the right side of the middle shelf of fig. 1. The measurements show that for the first two weeks the larger shelves grew about one-fifth of an inch (one-half centimeter) a day in all directions; on warm days, however, the increase was more than that and on other days not so much. The youngest portion of the sporophore was yellow-brown, which in three or four days deepened to a red brown. The unequal development of the mass caused concen- tric rings to ap[)ear on the top of the pileus, showing b}^ the low ridges and shallow furrows, respectively, where any particularh^ rapid growth had set in and where it had stopped. On August 15 the growth in width suddenly stopped. When full grown, the largest of the three shelves was 16 inches (40*"") across at the widest point and 8 to 1-1 inches (20 to 35*^°^) from front to back. 21 The, sporophores grow either on the roots of an affected tree or on the trunk, the former being the usual position. When growing on the ground the pile us is supported on a ver}' short stalk; it is sessile when growing from the trunk. There are usually several shelves which are grown together at the center in the ground form, or grown one above the other in the trunk form (see text figure 1 and PI. I, fig. 1). The whole body varies greatl}" in size. The smallest specimens collected during the past summer were 4 inches (lO^"") in diameter; the largest about 14 inches (35""^). The hjnnenial layer begins to form some three days after the body of the pileus is com- Fiu. 1. — Folypuras ncliuxiiiitr.ii Fr. growing on u fallen Fir. plete, so that there is always a wide band of sterile h3'phj« on the under side of the pileus during the period that the pileus is growing in width. When this growth stops, the tubes gradually form close up to the edge. The h3'menium when f n^sh is rose colored; when touched or bruised it turns dark red very cpiickly. The bright colors of the 3'oung pileus gi'tiduallv give wa\^ to more su)>dued ones as th(^ fungus grows older. A few days after growth has come to a standstill, the spores ripen and begin t(^ bi; discharged. T'hey come off in clouds plainly visible to the naked eye. Slips of glass placed under the pileus and left overnight had so thick a layci- of spores deposited 22 on them that it was impossible to perceive anythino- throuo-h the glass. Attempts were made to grow the spores in the woods on bread cultures. These all failed, however, because of constant interference on the part of inquisitive squirrels. The spores came off at intervals as if they were being discharged by some force acting within the tubes. Pieces of the pileus were accordingly turned over in a jar so that the tubes of the hymenium pointed up. Glass slips were supported over the tubes overnight, and on the following morning a few spores were found on them, but the number was so small when compared with the large number discharged from a similar piece laid with the pores pointing down, that it did not seem probable that there was any very active discharge going on. The spores were borne far away from the spot where the^^ fell. Owing to their exceedingly small weight, ever}^ disturbance in the air carried them ofl'. It was surprising to see how slight a disturbance sufficed. The flame of a candle held near the sporophore remained perfectly motionless while clouds of spores were swaying to and fro under the hymenial layei's. The spores were sown in aqueous decoctions of humus, but did not germinate. The facilities for doing more careful work were not at hand in the woods, so experiments on the manner of germination had to be left for a future time. At the time of ripening of the spores it was noticed that hundreds of drops of a yellowish liquid were hanging from the hymenial sur- faces ever}' morning when the fungus in question was visited. Some of these drops were carefully collected and Avere examined. In them floated a number of spores and flocculent 3 ellowish brown masses, which stained j^ellow with nitric acid. These were present for several daj's. Thereafter the liquid was almost clear except for numberless spores which were in every drop. For three weeks the drops were collected with a pipette during the da}', and during the night a plate, carefully protected against dew and rain, was placed under the fun- gus. In this way about three-fifths of a pint (300"'') of liquid were collected. This was poured into an open dish and put in a cool place, where the water was allowed to evaporate. A thick brown sirup was left after some weeks, which had the odor of verj^ impure molasses. The sirup was transferred to a vial, which was corked and placed in a warm place. In a few days delicate needle-shaped cr3^stals shot out, which upon examination proved to be melezitose and mycose, sugars sometimes found in fungi.* At the same time that this secretion appeared on the h^'menium, or rather shortly afterwards, a number of small beetles began to devour the hymenium with great avidity. So active were they that within 'The writer is indebted to Dr. O, Lt)e\v for the deterinination of these sugars. 23 three weeks of their appearance the hymeniiim was entirely destroyed, and of course with it whatever spores had remained. It is suggested that the secretion of this sugar and the destruction of the hymenium by these beetles may have some meaning in connection with the dis- persal of the spores. It is a point worthy of further observations by local observers in future years. The rapid destruction of the hyme- nium is very marked. It is exceedingly difficult to get perfect speci- mens of the sporophores after the end of August. The upper surface, which is usually moist, becomes covered with a fine layer of fallen spruce needles, and before long a covering of mosses hides the brown sporophore completely. It is no unusual occurrence to find these old moss-covered sporophores several years after their formation, at the base of some old Spruce. The basidia and spores have nothing about them which is very dis- tinctive. Numerous peculiar hairs project from the hymenium, which are surrounded with a film or drop of clear liquid in which numerous spores are caught. When viewed by reflected light these glisten like dewdrops within the pores. The latter are exceedingly irregular, so irregular in fact that one can hardly call them pores. They partake more of the nature of pockets, which are divided by many much con- voluted walls into various chambers. The pores extend almost to the margin of the pileus and are usually about one-eighth of an inch (S™"") deep. EFFECT OF FUNGUS ON THE TREE. The fungus seems to spread through the ground, attacking the tree first at its root system, and growing thence up into the trunk. Wher- ever one tree is affected, others similarly diseased will usually be found close by. Infection may take place through the root on one side of a tree. The heartwood of that root will be destroyed and then the wood of the portion of the trunk nearest that root becomes affected. Many trees were cut down where but one-half of the trunk had been rotted by th<» fungus, and oftentimes only a small spot was visible where the fungus had just begun to grow. The tree continues to stand until either the roots or the trunk become weakened to such an extent that they can no longer hold the tree erect, and then the first wind storm overturns it. Fig. 1 shows a large Fir, the root system of which was almost entirely destroyed. In its fall the lower part of the trunk split, revealing decayed wood to a distance of 12 feet (al)()ut 3f meters). The tree was probably blown over in the spring of 1S!>!», and in the follow- ing July the sporophores formed on the trunk. A large tree thus dis- eased is a constant source of danger to all others about it. Not only may the disease be comnuuiicated to them, l)ut in its fall such a tree breaks down many a small tree, not to speak of the large numbers of very small second growth which it destroys. The sporophores form 24 on fallen trees for several years in succession, possibly omitting a j^ear now and then. As a rule but one set of sporophores is found on one tree. As has already been said, young trees are subject to the attacks of this fungus as well as older ones, although the latter are probably more so, because the points of infection are so much more numerous. Nothing is known as yet of the manner in which the fungus enters the tree, nor of the rate at which it grows within a tree after having obtained a foothold. TREES ATTACKED. Polyporus schweinitzii was found growing on the roots of the White and Red Spruces, Balsam Fir, and Arbor Vitce. It is likewise com- mon on the White Pine {Pinus strohus). METHODS OF COMBATING THIS FUNGUS. Because of its destructiveness Polyporus schweinitzii is perhaps the most to be feared, where living trees are concerned. As it spreads through the soil it is difficult to detect, and still more difficult to com- bat. In the European forests a deep trench is dug around an infected tree or group of trees; this trench prevents the spread of the mycelium through the ground to neighboring trees. Such a method can not be recommended for American forest tree conditions, at least not for the present. If a group of infected trees is met wnth in the forest while lumbering it may prove advantageous to cut all trees in the vicinity of the diseased ones. Some of these may produce a hollow sound when hit near the base, an indication that the decay has started. It may not have gone up into the tree very far as yet, so that one or more logs can be obtained from the top. It will not be profitable to hunt out diseased trees as is done in European forests. There is as j^et no evidence that the fungus can infect a tree above ground, consequently it need not be feared in burned-over regions, or such as have been attacked by bark beetles. POIiYPORUS PINICOIiA (Swartz) Fr. OCCURRENCE. This fungus occurs widely distributed over the world, growing on conifers and occasionall}^ on Birches and other deciduous trees. In the New England forests it is one of the most frequent fungi found on living or more often on dead trees of Spruce, Pine, Fir, and Hem- lock. From three to ten of its bright colored sporophores may grow on a single log for several, varying from three to five, years. At the end of that time the mycelium has used up the available food supply in the loo' and dies. The sporophores grow on living trees, but these alwa5's appear weakened or sickly. No vigorous, healthy trees were found on which this fungus flourished. It is essentialh^ a wound parasite, entering 25 the trunks and branches above ground. Old knot holes or branch wounds, wounds produced by fire, or wounds made by animals, are favorable spots for the entrance of the spores. Wherever a tree dies from any cause this fungus is sure to attack it before long. In the sections where the bark beetles had been active some years ago there were many trees the wood of which had been destroyed by this Polyporus. The large holes made by woodpeckers offer excellent opportunities for the entrance of spores. As the woodpeckers are very active in exterminating insects inha])iting the bark (presumably the bark beetles among others), we have here a case of their allowing one enemy to enter while destroying another. In old windfalls the dead trees were covered with sporophores, some of them many years old, showing that these trees had become infected very soon after the trees had been blown over. This fact is of importance, as it suggests that these trees could be saved by the lumberman if carried to the mills shortly after their downfall. This will be referred to again. Plate IV is from a photograph of a portion of a Spruce trunk. The small white spots in the bark are holes of a borer filled with the mycelium of the fungus. STRUCTURE OF DISEASED WOOD. Wood of the Spruce in which the m^'celium of this Polyporus has been growing for some time deserves the description "entirely rotted" par excellence. The wood has been changed to a 1)rittle red-brown mass, which has cracked in many directions. The individual pieces are barely held together by countless sheets of mycelium which have filled the spaces resulting from the cracking of the wood and form an intricate network of larger and smaller sheets. In PI. IV a portion of a log in the last stages of decay is shown. At one side a sporo- phorc one year old and another just beginning are visible. The sap- wood has mimerous tunnels of a borer filled with remnants of the borings. Such wood has lost all strength, and falls to pieces at the slightest touch. If the mycelium attacks a standing tree the decay goes on within it until the trunk becomes so weak that an ordinary wind blows it over. The shrinkage which takes place in the wood as it is being metamorphosed is very considerable, as is evidenced by the large number of wide cracks which fill it, passing both across the annual rings and parallel to them. The changes which result in the wood may be descril)ed as follows: In a tnra just attacked the wood a))out the point of entrance of the fun- gus turns darker jind finally becomes a decided red-bi'own. Before k)ng small whitish areas appear here and there scatt(n-ed irregularly through the wood. Some of these are mere lines, while others form white patches circular in shape, surrounding small areas of wood about the size of a pinhcMid, which are red-brown (PI. III). Others again have 26 the .shape of broad, irregular liancls. which extend across the rings of growth. The point.s at which these white areas appear and the direc- tion which they take do not seem to be controlled by any particular factor, for they are exceedingl}^ irregular. The areas are shown in PI. Ill, which represents a radial view of a spruce log in the earlj^ stages of attack ])y the mycelium. The very fine white lines which are visible near the center of the log, extending across the annual rings, are of a different character from the white areas spoken of. It will be noted that in the white areas the parallel lines which indicate the summer wood are ver}' distinct. A microscopic examination of a white area shows that at this point the cells of the wood are com- pletely filled with fine hypha\ which form a dense mass within that area. Mixed in with the mycelium are the granules of an amorphous substance, readily soluble in alcohol, which is evidently resin. This, together with the mycelium, gives the white appearance to the spots. As the summer tracheids have a very small lumen, they have compara- tively little mycelium, which accounts for their being visible as lines extending through the areas. The size of an ai'ea is thus dependent upon the distance to which the mycelium has grown, and probably varies from time to time. It is suggested that the smaller areas are also the ones most recentl}' invaded. At this stage of the decomposi- tion the mass of the wood is already very brittle. Here and there cracks have appeared in the walls of the wood cells wherever a hypha has passed through them. Some tracheids appear like sieves because of the numerous holes. The changes subsequent to this stage of decomposition consist essentially in a carbonizing of the wood and the formation of the sheets of mycelium. The former change is one proba- bly induced by some ferment, the nature of which it is the intention to discuss more fulh' in another report. The cells of the wood gradually show more and more cracks and fissures, and the diameter of the walls decreases about half. The main shrinkage takes place in the secondarj" lamella^. The fissures which appear in the spring wood usually emanate from the holes formed by hypha?. The outline of these holes is irregular, but approaches a circle in form. In the secondary wall a fissure is soon formed which extends diagonally from left to right across the cell. Viewed from the top there are apparently two fissures, but these can readily be shown to belong to the secondary lamella? of adjoining cells. The fissures never extend into the primary lamella?. Various stages of such fissures are shown at PI. X, fig. 4. In the bordered pits at first two and later four fissures are visible in the secondary ring, which, as Hartig has surmised,^ are probably brought about b}' drving. Here and there (PI. X, fig. -I, c) a hypha has passed directh' through a bordered pit and in its passage has ^ Hartig, Robert. Zersetzungserscheinungen des Holzes, etc. 27 dissolved more or less of the iiie]n])rancs. In the summer tracheids the iuim])er of fissures in the walls is ver^^ larg-e. They all extend diagonally across the tracheids as in the spring- cells, and wherever a hole occurs there two fissures seem to cross. The smaller fissures have no counterpai't in the secondary lamella of neighboring cells as a rule, showing the complete independence of the two halves of the cell wall. The margins of the fissures are ragged, and the fissures themselves are very irregular in shape, and look as if they had been formed suddenly. The wood substance itself has been changed com- pletely. With phloroglucin and hydrochloric acid it stains red, and when the wood, finel}" powdered, is extracted with absolute alcohol quantities of hadromal are obtained. No reaction for cellulose can be obtained, and it seems as if the latter has been completely destroyed. An aqueous solution yields several compounds — an amorphous sub- stance, possibly a humus compound; a faint trace of some sugar, as shown by the phenylhydrazine test; and small quantities of citric and succinic acids. These are doubtless all decomposition products. There is also some compound present which reduces Fehling's solu- tion vigorously. After the walls of the tracheids are filled with holes and fissures they have reached the last stage of decomposition. A touch will then cause them to fall into many pieces. When boiled in water, the walls swell somewhat, and a pasty mass results when squeezed. With dilute KOH the walls swell to three times their size, and portions of them dissolve completely. The formation of the sheets of mycelium seems to take place in one of two ways. In one case the hyphaj in the white areas mentioned above exert a solvent action on the walls. This first becomes evident in the cells of the medullary rays. Their walls disappear completely, and the spaces are rapidly filled by the growing mycelium. The walls of the wood cells adjoining the medullary ra3^s are attacked, possibly at the same time. The secondar}^ lamella* shrink and finally disappear altogether, leaving a fine framework of the primary lamella. This framework is usually broken in hundreds of places, and as a result only pieces of the walls remain em])edded in a web of h3'pha?. The })ordered pits are destroyed from within outward, the torus resisting longest (PI. X, fig. 5). The latter is often freed and can be seen lying free in the cell. The triangular areas (as seen in cross section) of the primary lamella, formed where several cells join, are the last to dis- appear. A hole is thus formed which is completely filled with myce- lium. The latter spreads from this point in several directions. The writer is of the opinion that very little actual solution of the wood takes place, resulting in the fornuition of holes or cavities. Hero and there it doul)tless does occur, but rather as the exception, and possibly in the early stages of the development of the mycelium. It seems very much more prol)ab1(* that the socond mode is the usual o!ie. As 28 the m^'celium spreads through the wood it brings about the chem- ical changes spoken of, extracting substances from the walls. This reduces the volume of the wood and causes the fissures in the cells. But before long the shrinkage becomes so great that larger masses of the wood suddenly break away from each other at many points throughout the wood. Many small fissures are thus formed, which extend in every direction, both across the rings and within them in a tangential direction. The fissures are very irregular. Sometimes the}" extend for a short distance within one ring, then cross over into another, and so on. They ap- pear both in the spring and sum- mer wood, and not infrequently start in one ring and extend radially through the summer wood of that ring into the spring wood of the next. Often the breaks follow the lines of the medullary rays, but just as often they do not (PI. X, fig. 1). The process is evidently one of drying, for the same result is seen when wood dries, resulting in the formation of fissures, the so-called ''checking" of wood. If the fissures are near points where mj^celium of the fungus flourishes, the latter grows into the spaces and fills them com- pletely. Several fissures may join, forming an irregular longer ^ one. In PI. X, fig. 1, a sketch ^p?(fi- aOoc gaSf ^^OQ O inr\ Fig. 2.— Cross section of Spruce wood showing masses of mycelium of Poli/porus pinicola (Swartz) Fr. is shown of the cross section of several annual rings, showing how and where the breaks have formed. As the wood dries more and more the fissures widen and the mycelium keeps step with them. In small fissures it is very evident that the fissure has formed as a break and not by the solvent action of the mycelium. Fig. 2 shows such a fissure filled with mycelium. (The same figure is shown at c, PI. X, fig. 1.) A glance at the rows of wood cells will show how they have been forced apart, breaking one row. The rows are inclined toward one another, as one would expect them to be. The figure also shows a medullary ray at the right, the walls of which have disappeared. In the cells surrounding the break the mycelium flourishes, and here and there some of the walls are destroyed, making a small hole. 29 In the last stages of decay the fissures are very numerous, each filled with a solid felt of white rayceliuui. The felts extending in radial lines join those extending- in tangential lines here and there, and they hold in place the wood which would otherwise have fallen to pieces long before. In a live tree the heartwood is attacked first, and grad- ually the decay spreads to the sapwood. In the latter the browning of the wood is more marked, owing to its lighter color. Nothing positive can be stated at present as to the rapidity with which the decay brought about by this fungus proceeds. It appears to be very rapid, for trees l>lown down some two years before were found in an advanced stage of decomposition, with sporophores forming on their trunks at various places. FRUITING ORGAN. The sporophores of this fungus are very large and conspicuous, and are formed on logs during spring and summer, often many together on the same log. The form of the pileus is exceedingly variable (PL V). It is entirely resupinate when growing on the lower side of an over- turned log (PI. IX, fig. 6). In such a case there is no upper surface for several years. After three or four years the edge extends out beyond the curved surface of the log, and a narrow surface is exposed. Usually the pileus forms a distinct bracket on the side of a standing- trunk or log. This bracket is sometimes hoof-shaped, then again very much extended. In size it varies from an inch to a foot (2.5 to SO""") in width, or even more in extreme cases. The average specimen is 4 to 6 inches (10 to 15°'") wide. The upper surface of the bracket slopes toward the margin, and is divided into a number of regular divisions or lobes, which corresjDond evidently to periods of growth (PI. V). The lobes are smooth and dark red-brown when old. The youngest lobe is bright red, shading into a pale yellow at the very edge of the pileus. In many specimens the upper surface is almost black, and some of the lobes shine as if varnished. The number of lobes varies with the age of the specimen; one of the oldest found had fourteen. It has not been determined whether these lobes represent annual increments of growth, so it is not possible to say how old any of these large sporophores may be. The mass of the pileus is extremely hard and woody, and shows division into a number of zones (PI. IX, figs. 5, 6, 7), which are always greater in numljer than the lobes showing at the top. The hymeniuni is a pale yellow, very smooth, and assumes a watery appearance when bruised. It is very rarely perfect, as many insects are constantly at work eating away the tissue. The outer edge of the lower surface of the pileus is raised, forming a distinct ridge. At the inner edge of this ridge the formation of the tubes of the hymeniuni begins. This ridirc is continuous around the whole lower surface and forms a char- acter which is very constant. In young individuals it is wider than 30 in older ones. It is composed of looseh' interwoven hyphse, which form a continuation of the main hyphal strands which compose the body of the pileus. The hyphse of the latter start from a central point on the bark and radiate out in several directions (PL IX, fig. 7), forming- a mesh which at first is very loose. The hyphai are almost colorless and have a decided lumen. As they grow older their walls become brown and very thick, so that the lumen is reduced to a very, small one. The peripheral growth of the hyphaj takes place in such a way as to form well-defined la^^ers. For .several years these laj^ers are added one outside of the other. The lowermost portion of each laj'er is usually less dense than the outer portion, and after the hyphse turn brown large masses of crystals of calcium oxalate are deposited in the meshes of the outer portion. The alternation of layers of less density with those of greater densit}" makes a differentiation of laj'ers possible. The laj^ers vary considerably in width (PI. IX, figs. 5-7), and it is suggested that this is probably due to varying conditions; probably the amount of food supplied and the amount of available moisture exert a marked influence. The pileus grows in width and length by the direct elongation of the hyphie of the last layer. After several 3'ears' growth the hypha- on the under side of the developing shelf grow down in a vertical direction and give rise to the pores. The pores. are ver}^ long and are continuous from year to year. After a time the}^ become plugged at the l)ottom by hj-ph^e which grow into them from all sides. Different sporophores differ in this respect. With some the pores are open through eight or ten of the recent layers; in others the growth of hyphai is so vigorous that the pores are closed almost as rapidly as they are formed. The h^^menium arises on the surfaces of the pores from hyphte of the trama which turn at right angles to the general direction of the tramal hyphse. The latter have very thick walls (PL IX, fig. 12) and extend longitudi- nally, forming a very loose network. The tips of those hypha? which form the hymenial laver are thin walled. The hymenial layer itself is composed of hyphre of almost equal width. The layer is a very narrow one. Cystidia are practically alisent. The basidia barely rise above the general surface and do not differ materially in form from the paraphyses. The four spores are colorless. Amid the tramal and hymenial hyphfe accumulations of calcium oxalate crystals, colored red-brown, occur in great numbers, likewise large quantities of an oil readil}' soluble in ether and becoming solid at about 59" F. (15'^ C). The growth of the hymenial la^^er is very irregular. At one and the same time pores ma}' be forming on one side, while at the opposite side the old pores are completely plugged. The h3'menium renews itself at frequent intervals. The vitality of its hyphsB is very great, for it is not at all rare that insects eat away a considerable por- tion of the lower side of the pileus. These parts die and turn brown. 31 The remaining- portions then form separate centers of growth, which gradual!}' spread over the dead portion and unite, after several years perhaps, completely covering the dead part. A view of such a pileus is shown on PI. IX, fig. 4; several areas have already joined, forming a larger one, and a number of small centers are evident. The spores begin to be discharged in July. Growth of the lower side of the pileus takes place at the same time. Black cloths were pinned to the under side in June and by the end of August large por- tions of them were found completely overgrown with hypha?, and pores were beginning to form on the under side of the cloth. While the growing season lasts drops of a glistening yellow liquid are con- stantly being discharged from the hymenium. It is of interest to note here that the secretion of these drops was noticed b}^ Fries in a description of this fungus.^ Several cubic centimeters of these were collected and were found to hold in solution melezitose, the same sugar discharged from the sporophores of Polyporus schweinitzii. As insects, particularly small boring beetles, eat the hymenium with great avidit}^, it is possible that the sugar ma}^ serve to attract these insects to the sporophores, causing them to carry the spores to unin- fected trees. TRAMETES PINI (Brot.) Fr. forma ABIETIS Karst. Pobjporus jjiceinus Peck. Pohjpoi'us abielis Karsten. OCCURBENCE. This fungus is very common in the forests of the New England States, and occurs northward into Canada and Newfoundland. The writer found it conmion oh the Spruces and Firs in the Adirondack forests. It grows on nearly all the conifers and has been found by the Writer on the White Pine {Pimis strohus), the Red Spruce {Plcea ruhens), the White Spruce {Plcea ccmadensls)^ the Hemlock {Tsuga canadensis)^ the Larch, or Tamarack {Larix larichia), and the Fir {Abies halsamea). It attacks living trees after they have reached such a size that they form heartwood, and honeycombs the wood in such a way that it appears filled with small holes, many of which are coated with a shining white lining. The changes which are brought about in the wood are difi'erent somewhat for the ditt'erent kinds of trees and will be described separatelv. Of the six trees the Tamarack seems to be the most readih' attacked. A greater per cent of the older trees of this species were fecund affected than of the other five. The Spruces came next, and the Balsam Fir last. The fungus enters the trees through the stubs of broken branches > Friea, Eliaa. Epicrisis Syst. Myc. 468. 1836-1838. 82 and through wounds. The ni^x-clium tlourishes in both hcartwood and sapwood of the Spruces, the Fir, and Tamarack, and is confined to the heartwood in the Pine. It grows up and down the trunk from the point of infection, reaching into the root system and extending into the larger branches of the top. Affected trees may remain standing in the forest for many years until some more violent storm breaks the trunk at a weak point. The wood of the trunk is never destroyed completely, as in the case of the two fungi described above. In the most advanced stages of decaj^ some fibers of unchanged wood are to be found. The extent of their presence varies with the tree. DESTRUCTION OF SPRUCE AVOOD. The first effect noticed when the mj^celium grows in the wood of either of the Spruces is a change in color from the light straw yellow of the normal wood to a light purplish gray closely approaching the color indicated on the Milton Bradley Color Scale as Neutral Gray No. 1. Ver}^ soon this gray deepens to a red brown, the gray remaining as an outer ring surrounding the portions of red-brown wood. Small black lines appear scattered here and there through the red wood. These lines are present throughout an annual ring and extend longitudinally in the direction of the wood fibers for a distance of ^V to ^V of an inch (0.5 to 1 millimeter). Gradualh^ the black lines disappear and here and there small white areas appear (Fl. VI, tig. 1). The central portion of each area is absorbed and small holes are formed, which have white linings of loose fibers. The holes are at some distance from one another and are generally arranged in rows corresponding to the annual rings. Where the latter are very wide there may be a row of holes in each ring. The holes generalh' have their centers within the summer wood of the annual ring, but as they increase in size portions of the spring wood of that particular ring, as well as the spring wood of the following- ring, are included. The holes have a more or less spherical shape, which soon changes to a more or less elongated form, the greatest diameter extending radially. Fl. X, fig. 2, shows a cross section of a piece of wood at an early stage of the destruction. Some of the holes at this period are filled with a mass of white fibers, so that there is practicall}' no hole. The outlines shown in fig. 2 of Fl. X represent the outer limiting line of the white fibers, and the dotted lines (where present) indicate where the actual cavity begins. As the growth of the mycelium progresses, the holes increase in size and their walls approach one another until only a narrow lamella is left (Fl. X, fig. 3). A large number of holes appear between the original ones, and in the final stages there is practically no wood left except the narrow walls separating two holes (Fl. X, fig. 3, and Fl. VI, fig. 2). Adjoin- ing cavities rarely, if ever, unite to form a larger one in a lateral direction. They often unite at their upper and lower ends, forming 33 a long-er hole. The holes are never sharply defined, for there is always more or less of a white mass of metamorphosed fibers which remain in position next to the unchanged wood, and in many cases the whole area is thus occupied, and one can recognize the change only by the white color. In older holes this lining- is often replaced by felts of brown mycelium (PI. X, fig. 3) which partially or completely fill the cavity. The lameihe of wood between the holes ultimatel}^ become of an ahnost uniform thickness (PI. X, fig. 3), and on cross section show one or more black lines which extend completely around each cavity at an equal distance from the walls of two adjoining cavities. These black lines begin to appear at a stage intermediate ])etween that shown in fig. 2 and fig. 3 of PI. X. They are of variable width and grow darker and more marked as the decomposition advances. A lon- gitudinal section shows that they extend around the holes in a vertical direction also; in other words, a thin hwer of dark-brown matter sur- rounds the individual cavities. A closer examination shows that the brown lines are due to masses of dark-brown hvpha3 which fill each separate wood cell so completely as to plug it entirely. The hyphte are closely matted together and are incrusted with a brown substance which dissolves in part in dilute KOH and entirely in warm nitric acid. These hyphal plugs occur in every tracheid surrounding a hole and fill it for a shorter or longer distance. The plugs of adjacent cells may be continuous, or may follow one another much as a series of steps. This is shown in PI. IX, figs. 10 and 13. The latter represents a radial view of a number of tracheids at one side of a hole. The parts of the tracheids toward the hole (t) are completely changed to white cellulose fibers, while the parts on the other side of the plug (l) give lignin reaction. The brown hypha? fill the wood between the holes rather loosely, and it is only when about half way between two cavi- ties that they become matted together so as to form the plugs. The brown incrusting substances occur in or on the cell walls in the imme- diate neigh))orhood of the holes, and the manner of occurrence leads one to suspect that they were deposited in liquid form, for they have difiused through the various cells in all directions from the wall of the cavities. The changes in the cell walls which result when the mycelium attacks them are practically those so fully described l)y Hartig.^ There is a gradual extraction of those elements which give the so-called lignin reaction, the hadromal of Czapek. This begins in the tertiary lamella tuid proceeds outward slowly through the secondary lamella. The primary lamella at this period splits in the middle and is shortly after dissolved, leaving the individual tracheids entirely free from one another, each composed of approximately pure cellulose. ' Ilartijr, Robert. Zersetzungsersclieinungen des Holzes, etc. 32. 577«j— No. 2.5 3 34 The parts of the primar}^ lamella which are situated between three or more cells resist longest (PI. IX. fig. d,j)) and can be found free between the white cellulose fibers. The change to cellulose apparently takes place simultaneously over a considerable area. The first evidence of this change is to be seen in the white spots which come after the black lines. The white spots are the points at which the change to cellulose has taken place. The cellulose fibers are absorbed later on, giving rise to the holes already mentioned. Preceding the change from wood fiber to cellulose the wood is full of hj^phae, which become massed in centers here and there and bring about the dissolution of the wood. It is as yet undetermined what causes influence this local initiation of the changes, wdiich is characteristic of several other wood-destroying fungi. The growth in size of the white spots or cavities takes place rapidly. The hyphffi grow out in all directions from the original center, and as they do so the products of decomposition pass outward likewise, passing along the tracheids faster than across them. After a period the advancing hyphal masses of two adjacent holes meet in the narrow lamella of unchanged Avood between the two. A quan- tity of brown substance, representing decomposition products, has by this time accumulated. It fills the tracheids and coats the hj^phje so that these turn very dark, almost ])lack. Warm nitric' acid removes these substances entirely, leaving the hypha' and wood almost color- less. It is the opinion of the writer that this accumulation of the products of decomposition ma}^ account for the fact that the destruc- tion of the wood stops at this point, thus preventing the total destruction of the wood substance. That this can not be true in all cases is shown by the fact that many of the cavities join in the direc- tion of the fibers, but in this instance it is probable that diffusion takes place to more remote places. The mass of cellulose within the affected areas consists of free fibers Avhich remain in place for a period and are then gradually dissolved here and there, leaving an actual hole with a lining of white fibers. In the newly invaded parts of a trunk the mycelium is colorless and fills the tracheids completely. The Individual hypha- are somewhat thick-walled and have numerous short branches which penetrate the cell walls in all directions, leaving the characteristic figure 8 holes described by Hartig and others. Here and there a second form of decomposition occurs in which there is no reduction to cellulose. The process, as found in the spruce, is essentially the same as described by Hartig. The secondary lamella? arc gradually absorbed, leaving the primary lamella intact. The wood araduallv changes into a mass of red-brown fibers which fall apart at the slightest touch. The destruction of the wood takes place throughout the trunk, including the heart and sapwood, and finally even the bark (see PI. 35 VI). A trunk like that from which the log shown in PI. VI. fig. 2, was taken decays no further, and ma}" stand in the forest for man}^ years. After a tree has once fallen the destruction seems to stop. Two trees under observation for more than a N^ear did not change at all. In both the decomposition had reached, in 1898, the stage shown in fig. 2, PL VI, and in September, 1899, no further change could be detected. Further observations in this connection are desirable. This point is perhaps not as important from the standpoint of the for- ester as the power of the fungus to form fruiting organs after the fall of a tree, and this assuredly takes place with this fungus for several years, as will be mentioned. DESTRUCTION OF FIR WOOD. The destruction of the wood of Balsam Fir, Ahies hahaniea^ does not differ materially from that of the Spruce. White spots appear in newly attacked wood, which soon grow into larger ones; the black lines surround the individual holes sooner or later and then the decay ceases. On PI. VII a radial view is shown of a log taken from a Fir which had been blown down during the past summer. DESTRUCTION OF TAMARACK WOOD. The process of destruction is very different in the Tamarack. This is proba))ly due to the different nature of the wood of this tree, which seems to be far less resistant than the others. In the Tamarack the decay goes much beyond that described for the Spruce and Fir. In the early stages (PI. VIII, fig. 1) small white spots appear, which usually occupy the entire width of an annual ring. Two or more of these spots soon join, at first in a longitudinal direction, then laterally also. In that way it happens that very early in the process of destruction long stretches of one or more rings of wood are transformed to cellu- lose. This is well shown in fig. 1 of PI. VIII. This brings about the separation of one or more rings from the adjoining ones, forming in that way a series of tangential plates which can readily be separated. In the figure each one of the plates visible at the upper end represents one amuial ring. The line of separation between the rings is always at the point where the summer wood stops and the spring wood of the following year })egins. As the decay continues, more and more of the sound wood fibers are attacked, leaving loose cellulose fibers. When most of the wood has disappeared, black lines similar to those described for the Spruce appear, but as there are no such centers of decay as in that tree the lines are scattered irregularly. It would seem as if there were few decomposition products formed in the Tam- arack, and then only at a very late date. Ultimately the tangential plates become extremely thin; they are then c<)nii)osed of the more resistant sunnner wood cells of this or that wood ring, which ar(> more or less infiltrated with resin. The whole body of the former wood is 36 a mass of separate fibers, which can be pulled out individualh'. This can be seen at the ends of the piece of wood shown in lig. 2 of PI. VIII. FKUITING ORGAN. The fruiting^ or^an of this fundus is exceeding-ly common on all the affected trees and has been collected in Maine, New Hampshire, Ver- mont, in the Adirondack forests of New York, and in the forests of Toronto, Quebec, and New Brunswick. It is readily distinguished from allied forms by the light red- brown color of the hy menial sur- face, the regular small round pores, and characters of the hj'menial layer shortly to be described. The form of the pileus A'aries exceedingly and is almost a distinct one for every host plant. Hartig, in describing what evidently cor- responds to this fungus, ascribes the difference in form of the pileus and position on the trees to the different amounts of resin or turpen- tine which the wood of the different trees contains. Trametoi ijini^ according to him, forms brackets around the stump of dead branches in the Pine, the Spruce, and the Larch, while on the Fir the sporo- phores may appear at any point on the bark. This is true only to a certain extent for the trees of the Northern woods. Travietes pini is a very common fungus on nearly all the pines so far seen, and on these trees it alwaj's forms very large brackets, which grow, as Hartig says, from old branches. On the Spruce, the Fir, and the Tamarack this does not hold, for on all three of these trees the sporophores form at the ends of old branch stubs and at scattered points on the bark. The resin content of the Spruce is somewhat higher than that of either Tamarack or Fir, and on that account, possibly, the sporophores are more common at the ends of branches. In PI. XII a number of the forms as the}^ are found on the White and Red Spruces are shown. The bark of these trees consists of corky scales which are constantly being peeled off by newer ones developing beneath. The mycelium of the fungus, after having penetrated through the sapwood of an affected tree, grows rapidly into the 3'ounger parts of the bark and ultimately appears as small cushions under several of the bark scales. These cushions arc bright red-brown and have a velvety margin com- posed of thick-walled hyphtv. which rapidly spread out over the adjoin- ing scales, forming a flat sheet (tig. 4). While' the growth in a lateral direction is going on, and when the flat sporophore is scarcely one- sixteenth of an inch (about 1.5™"') in width, some of the central hyphse elongate, leaving small pockets Ijetween them which form the pores of the hymenium. The lateral growth may go on for several years, while at the same time a downward growth of the hyphse which form the walls of the pores brings about an increase in thickness. It ought to be said that this tvpe of spoi'ophore was found only on the under sides of fallen logs or branches. When the sporophores form on a 37 living standing tree they take the form of extended sheets on the lower side of the uppermost branches or form as sessile brackets of varied shape around old stubs of branches or again as sessile brackets at scattered points on the side of the main trunk. Fig. 3 shows a sporophore growing on the under side of a branch. In such a case the mycelium grows out through the bark, forming a long vel- vety cushion oftentimes several feet in length. This cushion rapidly grows laterally, and on its lower surface the pores arise. The growth of such a sporophore may go on for many years. The under side of the branch shown on PI. XII, iig. 3 was covered for a distance of 10 feet with the brown sporophore. As the latter increases in width it sooner or later develops a free upper surface where the body of the sporo- phore projects beyond the curved surface of the branch. The cracks appearing in the wood are due to drying. Fig. shows the sporo- phore as it occurs on the vertical trunk of a living tree. Here a form results which approaches most closely to Trametes ])ini (Brot.) Fr. The mycelium growls out from between the bark scales, forming a small knol) or sometimes several beside or above one another. On the lower side the pores soon appear as shallow pits, which are increased in depth by downward growth of the hyph^ forming their walls. The upper surface of the cushions becomes brown and, because of alter- nate periods of growth and rest, concentric lines arise which are more or less obscured by the hairiness of the surface. In forms of this kind the directive influence of geotropic forces on the position of the pores is very marked. The pores alwa3^s extend vertically, and on that account when found on a perfectly horizontal surface their open- ings are almost round. As one passes on into the oblique portion of the lower surface the openings become more irregular and the lower end portions of the tubes are exposed until they appear as hollow grooves. Where for any reason the position of the trunk or branch upon which a sporophore grows is changed, the direction of the pores changes like- wise, and instances of this kind are very common. On old Spruces ends of broken branches are points where the l)rown sporophores of this fungus may be found almost without exception. Two cases of this kind are shown on PI. XII, figs. 5 and 7. The Spruce loses many of its bran(^hes during windstorms, far more so than the Fir or Tamarack. The })utt end of a broken branch keeps on growing after the death of the outer portion, and in that way large knobs are foiined which may in time cover the wound entirely. It is an exceed- ingly slow process, however, and where, as is frequently the cas(\ the branch ))reaks off jit a distance of a foot or more, as shown on PI. XII, fig. 5, it rarely if ever heals over.' Such branches form the places ' There ia apparently in the Spruces little of that most efficient natural pruning which takes place in the Pines, where a dead branch breaks off very close to the trunk. 38 where the spores of this fungus find a most suitable place for entrance into the trunk. The spore germinates and the m3'celium grows down through the dead heartwood of the branch. From there it spreads through the heartwood of the trunk, growing both up and down. The growth in these directions takes place more rapidly than the lateral growth. When the sapwood is reached, the progress is a slow one, owing to the resinous contents. At about this time the sporophore begins to form. The wood of the callus and the living sapwood of the knob become so thoroughly impregnated with turpentine that the mycelium does not grow in them, but grows out through the dead wood of the branch. At the first point where the hyphse can reach the air without haA'ing to go through the collar of sapwood they emero-e. Where the dead branch has broken off close to the callus the hj'pha? grow out from the stub and form a cushion on it. More frequently, however, the red-brown cushion is formed at the point where the living callus touches the dead wood (PI. XII, fig. 6). The cushion is at first very small and looks as if covered with velvet. The hyphse rapidly grow radially and form a sheet which adjusts itself to the shape of the callus and branch. At the edges this sheet projects from the bark and forms an irregular shelf, the top of which after a time becomes zonate and brown-hairy, as in the more strictly bracket-like forms. On many old Spruces there are deep clefts between the vari- ous bark scales, and in them sheets of the sporophores form whose folds fill the crevices completely, forming pores on the outer surface of the newer bark and the inner surface of the old scale. Growth takes place rapidly during the latter part of summer and early fall so far as could be noted. The hyphjB at the edge extend the area of the sheet, while those forming the walls of the pores grow vertically downward. Within the pores many hypha? grow into the holes, so that after a 3"ear or two these are completely plugged at the base. There are at present no means of judging how old one of the sporophores described may orow to be. The oldest one found was about four-fifths of an inch {2"") in thickness. Trametes pi7ii forma ahletis was found ])ut rarely on the Fir. Its sporophores assume on this tree a different haliit from those on the Spruces. On vertical surfaces a distinct sessile pileus is formed, resembling a ])racket, rather than a hoof, as do those on the Spruce. The mycelium, after having grown throughout the heartwood, grows into the sapwood, where it flourishes much more vigorousl}" than in the Spruce because of the absence of resin. From the sapwood the hyphie enter the bark and break through it all over the trunk. At the points where they emerge they form small cushions, light red-brown in color, which are at first the size of a pin head, but rapidly increase in size (PI. XII, fig. 1). When barely /g of an inch (2 '"■") in width, a differentiation into an upper and lower surface takes place. A band of 39 veiy loosely interwoven hypha? grows out at right angles to the bark. From the loAver side of this band some hyph* split oil' and grow down- ward, adhering closely to the surface of the bark. Other hyph^ also turn down, growing faster at several points than at others, thus giving rise to small pits, which form the beginning of the pores. The pits are very variable in size. When they are still scarcely recognizable the hymenial layer begins to form in them, as evinced by the black cystidia which can be seen projecting from the lower surface of the band first mentioned even before any sign of a ridge is evident to indicate where the next pore is to be. Growth in these directions goes on rapidly. The hyphffi of the original band grow on horizontally, forming a rounded edge of loose hyph^e, which give the hairy appearance to the margin. At intervals, where the growth of the sporophore ceases, some of these loose hyphee stop growing, and when growth is resumed are left, forming a brush-like pi-ojection on the upper surface. These hyphse give the concentric appearance noted above for the Spruce. The hypha? on the lower side of the band grow downward to form the pores, and those adhering to the bark grow in the same direction, thus increasing the thickness of the pileus in that direction. A large num- ber of small cushions usually start together on thel)ark, many of which join as their edg«^s approach one another, forming a series of more or less imbi-icated sporophores (see PI. XII, fig. 1). On horizontal sur- faces the plicated form is lost, and sheets much like those found in the Spruce are formed. The pores in all the specimens on the Fir are more irregular than those found on the Spruce, but in all other important characters they are identical. On the White Pine the pileus is sessile and occurs at old knot holes. On the Tamarack both brackets and sheets are formed. The largest bracket forms found grew on the Tamarack; they often grow singly, and then again together, one above the other. One individual meas- ured 4 inches (10"") in width laterally, 2.8 inches (7 '"") from front to back, and 2 inches (.5 ''") in thickness at the back along the bark (PI. XII, fig. 2). The pores in the Tamarack specimens are exceedingly regular, far more so than in those of any of the other sporophores. The sporophores of Tixmietes pini forma ahietis grow both on living and fallen trees. They were found on trees which had been cut down four years before, and new ones were constantly appearing. It is this faculty of fruiting on dead trees that nuist enable tiiis fungus to spiead through a forest in a very short time, and accounts for the fact that it does so. After a Spruce has reached a certain age the chances that it will become affected with this parasite are, in the Maine woods, the very greatest. Older trees, i. e., Spruces which have reached a diameter of 10 to 12 inches, are more often subject to attack than younger ones. The fungus enters through any wound, and ai)parently spreads rapidly. There is no ('vidcMicc at ])rcsent to 40 show how rapidly it spreads, nor whether the (•haracteristic form of decay which it induces continues in wood after it has been cut from a tree or not. The present view seems to indicate that it does not grow after the death of the tree. HYIVIENIUM. Hartig ^ has given a very full description and numerous drawings of the hymenial layer of this fungus, and his observations can simply be confirmed. The basidia arise as slender hyphae, which gradually become much smaller at the apex and form four slender, rather long sterigmata, bearing the spores. These are colorless at first, but turn brown later on, and not infrequentl}" contain an oil globule in the center. The most striking elements of the hvmenial laver are the C3"stidia, called hairs by Hartig. They arise from internal hyphfe, which approach the hymenial laj^er at an angle. Pushing between the basidia and paraphyses one finds these large, pointed, brown, spine- like bodies, which project for a considerable distance into the pore canal (PL IX, figs. 2 and 3). They are thick walled and persist for a long time after the disappearance of the basidia and spores. As the pores grow older they are filled with a network of hyphse which grow out from the liody of the sporophore, growing over the hymenial layer and completely plugging the hole. The exact period when this takes place was not determined. POLYPORUS SULFUREUS ( Bull. ) Fr. OCCURRENCE. This fungus, although more frequently found on the hardwood trees, occurs now and then on living Spruces and brings about a brown rot of the wood of trunk and branches. The trees found were attacked after the trunks were 9 inches (23*^™.) in diameter. Entrance is efi'ected through wounds and broken branches, much in the same wa}" as the other parasitic fungi which enter above the ground. The mj^celium spreads through the trunk of an afi'ected tree, growing up and down, and reaching the highest branches in one direction and the roots in the other. No evidence of a diseased condition is usually visible on the outside, except such as noted for the other diseases. STRUCTURE OF DISEASED WOOD. Diseased wood is red-brown in color and can readilv be distino-uished from wood changed by the other fungi described by the fact that it breaks into slabs or flat pieces, which correspond each to an annual ring of wood (PI. XIII). The brown rotted wood is hard, verj^ brittle, 1 Hartig, R. Wichtige Krankheiten der WaldbiiunK'. 50. pi S. 1874. 41 and }ireaks into more or less rectangular pieces. When in its final stages, it is exceedingly l)rittle and can be crushed to a tine powder in a mortar. It is always nuich firmer than wood destroyed by Polyporus schveinitzu and differs from the latter in the character of the cracks or breaks, which are most readih^ seen on a tangential view. The progressive changes which take place in the wood of a Spruce may be noted as follows: The wood at first turns slightl}^ red-brown in irregular patches, as seen when a trunk is split longitudinally. • These, patches grow larger, spreading from ring to ring and in a longi- tudinal direction along each ring. Small cracks next appear in these areas, extending part way through the thickness of each ring, both from the side of the spring and of the summer wood. These cracks are ver}- much more visi])le on the tangential view of an annual ring (PI. XI, fig. 1). At first but scattered cracks are to be seen extending longitudinally, which, however, soon elongate and pass both diagonally and directly across the direction of the fibers (PI. XI, fig. -I). At this stage the wood is still hard and has acquired a light-brown color. Immediately about the fissures it is more deeply colored than else- where. A microscopic examination shows that there has been great shrinkage in the volume of the cell walls and that the breaks and fissures occvir practically throughout the whole mass of the ])rown wood; though onh' the larger breaks are visible to the unaided eye. The shrinkage goes on rapidly, and after a time the tension becomes so great that the annual rings separate one from the other. A break usually occurs in a radial direction also, and as a result the free ends of the ring sw^ing outward. Breaks along the lines of the larger medidlary rays take place at the same time. This gives rise to long flat slabs of wood, each the width of an annual ring, Avhich hang together loosely at one end and at isolated points on their tangential walls (PI. XIII). Very ])adly decayed wood is so thoroughly traversed by larger and smaller breaks that it readily fails to pieces when struck. It nuist be noted, however, that the nature of the cracks is such that individual pieces of wood are, as it were, mortised into each other end to end, and this no doubt makes the wood as firm as it is. MINUTE CHANGES IN THE WOOD. The minute changes which the mycelium of Polyporus Hulfm^e\(i< indu- ces in the wood cells are such that the}' can not well be mistaken. It has been mentioned that the annual rings break into bands which curve inward as the ])rocess of drying goes on. A tangential view of several of these bands before they have broken will present an appearance such as is shown on PI. XI, fig. 4. A large munber of fissures have formed both across the wood fibers and parallel with them. The latter are more prominent — the cross fissures never occurring alone, but generally con- necting several longitudinal fissures. It will be noted that the Invaks are 42 characterized by sharp right angles, and in many places a stepladder arrangement is evident. In the early stages of attack the wood fibers turn red-brown and shrink. As a resvdt, tissures are formed in the walls of the tracheids, which extend diagonally across the wall at an angle of approximately 45 degrees (PI. XI, fig. 1). The medullary ray cells are at this period still intact, and hold together the more or less brittle wood libers. The next stage in the decomposition consists in the absorption of the medullary rays. This allows the wood fibers to con- tract more than up to that time, and as a result breaks occur. These breaks form at first so as to connect adjacent cavities left by the absorp- tion of the medullary rays. The wood fibers tend to curve in one direc- tion or another and break at the weakest point, namely, between two cavities, where the opportunity for curvature is greatest. What deter- mines the direction of curvature of the wood fibers has not yet been explained. In the illustration the curvature is toward the right. This curving has the efl'ect of bringing medullary rays which are in differ- ent longitudinal rows approximately into a line. Thus at •'«" two cav- ities are shown which are separated bj^ a curved fiber which sooner or later will break, uniting the two. At first two ray cavities are joined, then more, until long longitudinal holes are formed, such as are shown in fig. 4 of PI. XL The reason for the sharp angles' is now very apparent, likewise w^h}^ these fissure figures appear only on a tangen- tial view while on the radial view one simply sees the fissures as lines extending at right angles across a ring of wood (PI. XIII). The marking of the individual wood cells is a very regular one. The fissures extend through the secondary lamella, and at first sight remind one of those which the m3^celium of Polyporus schiceinitzii induces. The latter are very much steeper, however, and do not occur at such frequent intervals. The mycelium of Polyporm mlfureu.^ is colorless and is present only here and there in the wood cells, a fact to which Hartig calls attention. No spores, such as are so common when this fungus grows in Oak wood, were seen in the Spruce wood, although diligent search was made for them. FRUITING ORGAN. The sporophores of Polyporus sulfureus are among the commonest and best known of the largest fungi. The sulphur-yellow shelves of this fungus occur widely distributed throughout the United States, and are found in late August and September on many of the Oaks, Walnut, and other broad-leaf trees. A large number of sporophores usually appear together, one above the other, when growing from an upright trunk, or scattered here or there on a prostrate log. They grow on living trees and on the dead trunks also, for several years after the latter have fallen. A marked periodicity iu this respect 43 was noted for a particular tree during the past summer. This tree, a large White Spruce, had been blown down some years when first seen. The standing stump was 12 feet (3| meters) in height, and on its south side there developed in August of 1897 a large number of the sporo- phores. These dried and broke away during the following winter. During the summer of 1898 no sporophores appeared on either the standing stump or the fallen log, and it was not until August, 1899, that a new lot of the brackets appeared, and then in the greatest num- ber. Three large patches broke out on the north and northwest side of the trunk, and the lower side of the fallen log was literally covered with the yellow brackets. No mention of this periodical occurrence of the fruiting portion has been found, and it will be of considerable interest to see what will take place this year. Several other large Spruces in the immediate neighborhood were caused to decay by this fungus, but no sporophores have so far developed on their trunks. The shape of the pileus varies materially with the position which it happens to occupy. When on upright trunks several sessile sporo- phores usually occur one above the other, the upper surfaces of the lower ones touching and uniting here and there with the lower surfaces of those above. The individual parts are comparatively thin plates, which have radiating lines and depressions extending outward to the margin. The body of each is soft and fleshv when young and full of a clear yellowish liquid. The upper surface when 3'oung is verj^ moist, somewhat hairy, and when ])ruised turns brown. As the plant grows older it becomes verj^ much harder, and when completel}' formed is quite hard and brittle. Masses of the young plants have a peculiar fungous odor, which l)ecomes very intense as the parts grow older. The lower surface of the shelf is smooth and even. The pores are formed verj- early in its development, and almost as soon as thej' are completed the formation and discharge of spores begin. The sporo- phores are very short-lived. They l)egin to appear on the trunk as small I'ounded knobs, formed by thick-walled hypha?, which come out from between the bark scales. Their growth is very rapid, even more so than that noted for Polyporm schvjeinitzl i . The various small kno])s soon flatten into a number of plates, consisting of strands of hyphai, some of which grow out horizontally, increasing the width of the pileus, while others grow downward to form the pores. When the sporophores develop on the under side of a log they grow out in all directions from a central point, and sometimes forms with a distinct stipe are met with. Numerous drops of the clear liquid mentioned before were found hanirino- from the under surface of the shelves on some days.' The api)earance of the drops does not seem to stand in any relation to the amount of moisture in the air, for they weie found alike on very dry * Fries notes this fact — Epicrisis, etc. 450. 44 and very foggy days. The same sugar, melezitose, that was found in IWi/por>/s schweinitzil was obtained from the liquid in quantity. The fungus is attacked when barely mature l)y insects and small animals, and within a month after the ripening of the spores there is little of it left except the harder vipper surface of the shelves and the contracted basal portion. This may account for the fact that the spores ripen and are discharged so very rapidly. Cultures of spores made in water, in sugar water, and on bread showed no signs of germination. These experiments are to be repeated with better cultural facilities. The spores spread through the air and are carried to all parts of the forest. Wherever any wound or broken branch offers suitable condi- tions they germinate and induce the rot described. Polypmms, sulfureus was found only on trees growing along the coast of Maine. They were all older trees of the White Spruce. Further search will no doul)t show that it attacks the Red Spruce also, and possibly the other conifers. Its large, conspicuous sporophores make its recognition easy, and the fact that they are edible in their early stages ought to lead to their collection and destruction. POL.YPORTJS SUBACIDUS Peck. Poria subacida Peck, Thirty-eighth Report N. Y. State Museum. 92. OCCURRENCE. There are a number of fungi which attack standing trees and destroy their wood, of which it is not possible to tell, without continuous observation and experimentation, to what extent they are responsible for the death of trees, and whether they attack perfectly healthy trees. Among these belongs the fungus which for the present will be con- sidered as Poly2}(yrus subacidus Pk. It is one which is found on decay- ing logs of coniferous as well as other woods, ^ forming its pores in late summer and winter. It was found once on a living Hemlock^ twice on living White Spruce, and once within the trunk of a living White Pine. In many of the spruce forests hundreds of trees, particularly the younger ones, were found dead or dying. Man}^ of these trees were pulled up, and on their roots yellowish masses of mj^^elium were occasionally found. In one locality some thirty of these young trees, ranging from 2 to 10 inches (5 to 25 cm.) in diameter, had the wood of the trunk decayed by some fungus. The wood appeared 3^el- low, was very wet and spongy, and was easily pulled into shreds. No fruiting organs could be found. Several of the trunks were taken and sawed into pieces a foot (30 cm.) or more in length. These pieces were buried to the depth of a foot (30 cm.) in a sphagnum bank and were examined every week. Other trees were simply broken near the 1 See Exsiccati, E. & E., N. A. Fungi. 45 ground and left standing, while in still others wounds were made with an axe to permit the entrance of air, as it was thought that fructifica- tion might thus be induced. After two weeks the ends of the pieces buried in sphagnum were covered with a white film of hyphte, which gradually turned yellow, and after two months began to form shallow pores. The same took place in practically every one of the trees which were overturned or wounded. In all the localities visited where trees, both older and younger, had been overturned, this fungus was found again, and again, and associated with it the form of wood decay described below. (Pis. XIV and XV.) Masses of yellowish mycelium were sometimes found growing out from under the bark scales of the roots of many healthy spruces in a way which seemed to indicate that they were ])egiiming to enter the root itself. Hypha3 from these masses extend into the soil, V)inding together the particles so that dense clumps are formed, varying from the size of a pea to as large as two fists put together. The growth of the hyphie in the soil is a very rapid one; they can be grown with ease in moist soil and form the peculiar lumps in a few weeks. Pieces of diseased trunks were buried in soil in a greenhouse in September, and in four months the hyphffi had grown through the soil of the bench in all directions. It is thus very evident that this fungus grows in the ground rapidly and that this is probably one of the ways in which it enters standing trees. This is made more probable by the fact that one finds all of the trees in a certain area affected with this fungus, both younger and older ones. Each probably infected its neighbor much in the way in which Polyporus sclviDeinitzii does. The fruiting portion of the fungus has been found on living White Pine, Red and White Spruce, Fir, and Hemlock. A large Hemlock, almost 2 feet (0.6 meter) in diameter (near Houlton, Me.), had been blown over and the trunk had broken some 6 feet (2 meters) from the ground. The wood was very soft and showed numerous black spots surrounded l)y white areas. The fruiting organs were forming in the chinks and crevices of the trunk, and on the stump. The tree was alive at the time it was seen. STRUCTURE OF DISEASED WOOD. The decay which the mycelium of this fungus induces is. not to be confused with that caused by any other fungus. Spruce wood when very much decayed is moist, almost wet at times, and can be comi)ressed nuich like a sponge, when (piantities of water will drip from the mass. Larger and smaller cavities of very irregular shapes, lined with a tough felt of hypha^ yellow on the inner side, are found throughout the wood. Such a cavity is shown in part at the bottom of PI. XIV, fig. 2. The cavities are scatt«M-ed throughout the wood in most triMvs and are generally partially filled with a pale straw-colored liciuid. The wood 46 itself dift'er.s markedl}' in ditferent trees. Ttiis difference appears to be due somewhat to the rapidity with which the solution of the libers takes place. As a rule, the wood in the early stages of the attack has numerous black spots scattered throughout its mass (PI. XIV, fig. 1). These black spots are surrounded by a white circle before long, and somewhat later disappear entireh', leaving very nuich larger white spots. The wood around the spots is now straw-\'ellow in color and begins to look somewhat frayed, as if groups of wood fillers were sepa- rating readily from the rest. A tendency for the different annual rings to separate now becomes very marked (PI. XIV, fig. 1, at the right), and a log of spruce wood at this stage can be split into concentric rings by mere pounding. Gradually the number of white spots increases. In one form of decay the white spots are confined almost entirely to the summer Avood. The newly formed spots are also in the summer wood, and l)efore very long all the summer wood of every ring, including also some of the adjacent spring wood of that ring, has turned white. This stage of decomposition is shown very well in PI. XIV, fig. 2, a longitudinal section of a spruce log, and in PI. XV, fig. 1, a cross section of the same log. It will be noted that the change to the white masses nowhere passes from the summer w^ood of one ring to the spring wood of the adjoining ring. There is evidenth' ^ome agent, presumal)ly of a chemical nature, which confines the solvent action of the fungus mycelium to the summer wood and prevents it from attacking the spring wood. It may be recalled here that where a similar change takes place in the spruce wood, induced by the mycelium of Traincttxi jjini forma ohkth (PI. X, fig. 2) 1)oth summer and spring wood were changed. This localized action of the dissolving agent takes place with such regularity and in so many different ways, depending upon the kind of fungus attacking the wood, that it suggests the presence of specifically distinct dissolving agents, enzymes, perchance, for each fungus. In the second form of deca}" the appearance of the white spots is limited to the summer wood in the same wa}' as above described. The white spots do not increase in number so rapidly and consequently do not form the white bands spoken of. Changes take place within the wood cells of the spring wood, which give to them a ver^^ light and porous nature. A cubic inch (16. -l*"") of such wood completely decayed weighs but 1.3 grams (sound spruce wood weighs 5.52 grams). The mycelium of the fungus spreads through the individual tracheids after entering the tree, and collects in spots here and there. Solution of the wood cells begins around these centers, which at this time appear dark brown or black. They are the black spots referred to above. The change which takes place around these centers consists in a solution of the hadromal and the other lignin constituents of the cell walls, leaving the pure cellulose fibers free from one another. These con- 47 stitute the white spots and also the white ))ands spoken of. The various steps leading to the complete separation of the cellulose fibers are exactly those which have been described for a similar process caused by the hyphee of Trametes pini forma abietln. A xc^vj different change is going- on at the same time in the spring wood, and gradually spreads from this to the sum- mer wood. This change may be likened to the one which Hartig has described as taking place in pine wood attacked })y PoJyjyortis horealix..^ The hypha? of the fungus develop in the wood cells with great rapidity, filling them completely. Numerous hyph« pass through the walls in all directions, making large irregular holes many times the diameter of the hypha^ which pass through them. The secondary walls of the Avood cells are gradually dissolved; a faint granular appearance of the walls is seen at first, and little by little the walls become thin- ner. At last only the primary lamelhi is left, and in the bordered pits the torus (PI. XI, fig. 3). The whole wall finally disappears, leaving simply that part of the wall belonging to two or three cells, namely, the portion having a triangular cross section. This solution of the walls goes on sinudtaneously throughout large areas. The medullary rays disappear completely, long before the wood cells are entirely gone. The spaces left by the dissolved cells are rapidly filled with hypha^ and these hold poi'tions of the cell walls not yet destroyed in place, and give consistency to the mass, which thus retains th(> shape of the wood befoi-e the attack. The whole mass can be compressed by slight pr(\ssurc and will not return to its original size. This accounts for the extremely light weight of wood thus decayed. In PI. XI, fig. 2, a radial view of wood in an advanced stage of decay is shown. The straight black lines indicate groups of wood vessels, two or more; the hyphai between Fig. 8.— Baso of spruce brnnoli, sliowing its resistance to the attack of tlie my- celium of Polyporus subacidug Fk. ' Hartig, R. Zeraetzuiigserscheinungeii dcs Ilolzes, etc. 48 them have dissolved out the missing fibers and now fill the spaces. Plate XI, fig. 3, represents a cross section of similarly attacked spruce wood, showing several wood fibers of the spring wood at one side and the gradual dissohition of adjoining ones, leaving onl}^ the more resist- ant portions which lie free in the masses of hyphte. These remaining parts stain with phloroglucin and hydrochloric acid, showing that they are still lignified walls. Heartwood and sap wood of the spruce are destroyed with equal rapidity. All parts become spongy, with the exception of the resinous basal pieces of the branches, which resist the attack of the fungus even after the whole trunk has been destroyed. This resistance of the basal pieces of the branches is quite a common feature in diseased trees attacked by several other fungi, notal)ly Polyparus schwelnitzii^ but nowhere is it more striking than in this instance. Text figure 3 shows such a branch piece as it appeared immediately after pulling it from a dead standing tree. FRUITING OKOAN. After the mj'celium has invaded the sapwood it grows out over the bark, forming yellow felts. This takes place in the early part of the summer, generally about July. A few weeks later the small pores begin to form. Certain hypha? of the sheet turn at right angles to it and grow out at this angle, forming shallow pores. These are almost round and are separated by ver}- thin dissepiments. Fig. 2, PL XV, is from a photograph of a spruce log, about the middle of Sep- tember, almost natural size. As the season progresses the fungus dies and splits up into smaller areas and some of the tubes become inclined. No pores occur at the edge of the sheet, thus leaving a fringe of sterile hyphae. This distinguishes this fungus from many allied forms. The hymenial layer and the pores are generally straw yellow, sometimes even more decidedly yellow, the color deepening toward the latter part of the fall. The pores do not form until December or January, and as a completely fruited fungus was col- lected but once, its description will be deferred until more material has been seen. The fruiting organ frequenth^ develops in cracks and breaks formed when a diseased tree is blown over. Fructification was induced in many instances, as described above, b}^ allowing the air and moisture to have access to completely decayed wood. When PoJijporuH siihacidus grows in Northern forests on dead conif- erous wood as a saprophyte, its habit and action difi^ers somewhat from that described above. Inoculation experiments were made during the summer to test how rapidly this fungus destroys sound wood. Dis- eased wood from both dead and living trees was placed in holes bored in healthy spruces, and the latter were labeled so as to be readily iden- tified in later years. The amount of destruction which this fungus does 49 in the spruce forests is very large, but careful experiments will have to ])e made to determine its relation to trees weakened by other causes, also its progress through the soil from tree to tree. KEMEDIES. This fungus ma}^ be accounted most destructive to dead timber, and any remedies spoken of for Polyporus plnicola apply here. Dead trees should be utilized before the chance for infection becomes too great. No practical remedies can be suggested at present to prevent its spread through the soil. OTHER DISEASES. Besides the diseases described in the foregoing there are a num])er of others of which not enough was seen to enable a full description to be given. POLYPOEUS VAPOKARIUS (PERS.) FK. This is frequent on Spruces and Firs, and induces a brown rot of the sapwood. The fungus occurs widely spread over the United States and Canada on all coniferous woods. Its fruiting body is very variable, and there are probably many fungi included under this name which do not belong there. From observations made in the Maine woods it seems that this fungus attacks dead much more than living trees, destroying them for timber very rapidly. A fuller description of it will be given at a later date. POLYPORUS ANNOSUS FR. This fungus is a parasite of European trees much feared by the for- esters of the Continent. Diligent search was made for it, but fully formed fruiting ])odies were not found. A single Spruce seen on the top of Mount Kineo, Moosehead Lake, had its roots covered with tirm leathery sheets, such as PoJyjJoi'u.s annosus sometimes forms on the roots of the Southern Pines. Unfortunately there were no means at hand to cut down the tree, so that an inspection of its trunk was impossible. Other diseased trees of Spruce and of the Fir were seen north of the Kangeley Lakes. One of those Avas overturned, having grown in a dami) locality. Its roots were covered with the yellowish leatherv felts which extended into the surroundino- soil. The trunk of this tree was completely rotted in the center, th(> decay going up the trunk for 25 feet (almost S meters). At this point the wood was brown, showed some white areas, and smelled strongly of prussic acid. The stumps of many other Spruces were examiniHlfor evidences of this fun- gus. Some S})ruces were found which had small holes in the sunnner wood of many annual rings. The wood when cut longitudinally showed many of thes(> holes, which dUl'ered fioni those formed by Ti'dtitctea pln'i. 577r) No. 25 4 50 Thoy (x-cuiit'd chieHy in the .summer wood, and were lilled with a red- brown powder. There is no Avhite lining as in the wood attaeked by Trametes jy'mL Black spots appear here and there in the wood, and when they disappear the holes take their place. The holes increase in size and number, and in the last stages of decomposition the wood has become a shredded mass of yellow-brown fibers, which feel much like straw. It is completely honeycombed in ever}" direction. The annual rings of wood separate from one another, forming thin plates per- forated by thousands of small holes. The transformation of this fibrous material takes place from the root up into the trunk for from 3 to 20 feet (1 to 6 meters). In some trees the innermost rings of wood are afi'ected. As the wood becomes more and more rotted a hole is formed wdiich gradually increases in diameter, eventually sometimes becoming so large that the weakened trunk is blown over by the wind. On other trees one or the other side of the trunk may be affected. Two or more separate holes may be formed which join near the base of the tree. A more lengthy description of the changes in the wood just described is not deemed necessary, in view of the fact that the active agent which brings about the changes is as yet not fully determined. If it proves to be P(Ayp(>rui< nnno^im Fr. it would seem that the injuiy done in the Eastern forests by this fungus is not A'ery large, which may be con- sidered a fortunate circumstance, as this fungus is one naturally to be dreaded h\ the forester, as it is combated only with the greatest diffi- culty and expense. AGARICUS MELLEUS VAHL. Many trees were fovuid in which the well-known rhizomorph strands of this fungus grew under the bark. The summer of 1899 was exceed- ingly dry. and on that account the development of Agaricinese of all kinds was a very meager one. On the various excursions made through the Maine forests but one tree was found on which the yellow fruiting organ of this fungus w\as developing. The manner in which this fun- gus grows on the roots of the trees and brings about their death has been so fully described b}" Hartig and others that it seems hardly necessary to describe it here. The fungus grows within the living roots and cambium of a tree and speedily brings about a disturbance in its absorbing organs which results in ultimate death. The wood is rarely if ever affected to any extent, so that lumbermen use the dis- eased trees for lumbering purposes, making no distinction between them and live trees as long as the wood is entirelj^ sound. Diseased trees should be cut at once when recognized. 51 CONCLUSION. The conditions in the New England forests are very favorable to the growth and development of timber-destroying fungi, conditions which are made still more favoralile by an ever increasing supply of dead wood. Radical changes will be necessary in the present lumbering methods in certain localities before any betterment can be hoped for. During the summer of 1899 the wasteful cutting of timber was noticed in particular in the region north of the Moosehead Lake, where the old S3^stem of measuring logs by the top scale is still in vogue. The lum- berman cuts the logs on the stumpage plan, and in his endeavor to obtain as high a scale as possible he cuts the tree high up on the trunk and low in the top, leaving almost half the top in the woods. This is not only wasteful lumbering, but ofl'ers an excellent opportunity for the development of several of the fungi described in the foregoing pages. From the dead trunks and limbs their spores spread to stand- ino- trees which mio-ht otherwise remain sound. The same is true for the insects, as recently pointed out by Hopkins.^ In the foregoing it has been pointed out that as trees grow older they become more liable to insect and fungus attack. An old tree has many vulnerable points, such as old branches and wounds made by animals or by hail, where insects or fungi may gain entrance to begin their work of destruction. As a tree grows older the chances that it will be attacked become greater. This point ought to be taken into consideration in the harvesting of a timber crop. In certain sections of the Maine forests, particularly in the Rangeley Lake region, the trees have reached an age where it appears that the rate of annual accretion, and con- sequently the annual increase in value, is very small, while the danger of infection is increasing every year. It is recommended that such trees he cut immediately where practicable, as they are practically ripe and proba))ly at their point of greatest value. This may not alwaj^s ])e possil)le, owing to practical difhculties in reaching water courses, etc., but the principle should ])e established that it will prove more protitable in the long run to cut trees after they have reached a certain age, to prevent depreciation due to the attack of fungi or insects. Future investigation will have to determine what the exact age is at which it will be most profitable to do this cutting. It has also been pointed out that there are several fungi which attack trees after they have been killed by insects or other agents. This is of gi-eat practical significance, for it may often l)e possible to harvest such dead trees before the fungus in question has had time to l)egin its work. ' Hopkins. A. I). Ptvliininary Report on llic Insect p]neniies of the Forests of the Northwvst. I'.iil. No. 21, Div. of ImiLpiiiuIo^'v, V. S. Dcpt. Agr. 181)9. 52 In the Maine forests great areas of forest lands were killed hy bark beetles some years ago. If the dead trees had been cut shortly after their death, the timber might have been utilized, and it would have been as valuable as that from live trees, for the beetles do not mine in the heartwood. This was not done, however, and before long the whole forest of dead trees was rendered worthless by several fungi, notably Polypm^ics pinicola and Polyporus suhaoidm. What is true of larger areas holds for individual trees in the forest, and also in those sections where strong winds blow over many trees. Such an area, technically known as a windfall, offers opportunities for the action of destructive fungi, and the same recommendations just made for areas where trees are destroyed by insects hold good. A dead tree is as valuable as a live tree, provided its wood is sound, and it ought to be cut immediately. There is some prejudice among lumber bosses that such trees are of no account; nothing can be further from the truth, and this fact ought to be insisted on by those in charge of cutting operations. The trees, now in the forest, which are diseased are beyond help, and it is at present neither practical)le nor economical to practice the methods in use by the European foresters, which consist in the prompt removal and destruction of the diseased trees. The time will come when this may prove protital)le in the regenerated forests, but for the present the most hopeful method of combating fungi is by conservative lumbering. Men who are acquainted with the manner in which insects and fungi work and who can direct the cutting operations ought to be employed. It may not be out of place here to refer to the growing sentiment in favor of restricted cutting, which was very much in evidence in the localities visited. Much agitation is still going on decrying the lum- berman as the greatest enemy of the forest; but with the growing reali- zation that it is possible to utilize the timber of the forest and still leave a forest which will yield timber from year to year, this feeling is gradually lessening. The lumberman has not been slow in realizing that restricted cutting will be more economical in the long run than the indiscriminate destruction of the past years. It is gratifying to note that two of the largest lumber owners of western Maine are employ- ing trained foresters, under whose directions the cutting operations are carried on.^ These men will not only be able to make operations more prolitable, but can also aid in gathering information which may go to solve many of the problems still to be unraveled in connection with the enemies of forest trees. 1 See also Graves, Henry 8. The Practice of Forestry l)y Private Owners. Year- book, Dept. of Agr. 1899: 415. 1900. 53 EXPLANATION OF PLATES. platp: I. Fig. 1'. Sporophores of Polyporiij^ ^clavemitzii Fr. Fui. 2. A piece of the bark of Red Spruce with sporophores of Polyporun volvntun Peck growing from holes forinetl by a boring beetle, a species of Deiidroclonua. PLATE II. Cioss section (X|) of the trunk of a living young Balsam Fir [Abies halsamea (L. ) Mill.) at a point 4 feet (1.2 meter) from the ground. Decay, caused by Polyporus sclnreluitzii Fr., has shrunk the wood, jjroducing a number of cracks and giving it a rough appearance. It is so nonresistant that the saw tore the fibers instead of cut- ting them. The large crack at the top, extending through the sapwood, was formed when the tree was cut down. A small sporophore of the fungus grew at the base of this tree. PLATE III. Radial view (X j) of a log of White Spruce (Picea canndnisi.'i (L. ) B. S. P.) , showing an early stage of decay induced by the mycelium of Polyporus pinicola (Swartz) Fr. The fine parallel lines indicate the annual rings of wood. Here and there white spots with darker centers are seen; likewise long white lines parallel to the course of the wood fibers, and others near tlie center of the figure, which extend in an irregular manner across the direction of the libers. PLATE IV. Radial view (Xz) of a log of White Spruce {Picea canadensis B. S. P.) , showing an advanced stage of decay induced by mycelium of Poly poms pinicola (Swartz) Fr. The wood has cracked throughout. The white masses are sheets of mycelium. At the right of the figure two sporophores are shown — one just beginning to develop, the other about 1 year old. The sapwood has been partially destroyed by boring larvse, whose tunnels are filled with sawdust. PLATE V. Three sporophores (Xj) of Polyporus pinicola (Swartz) Fr. The uppermost one is a young one. The one on the right is growing on a stump, and its lower surface is much eaten by insects. The one on the left is a very old sporophore, in which the ridged ujtper surface is very marked. PLATE VI. Fig. 1. Radial view of a piece of wood (natural size) of the Red Spruce (Picea ruhens Sargent), showing an early stage of the decay induced by the mycelium of Trainrtes pini ( Brot. ) Fr. forma ahietix Karsten. The white spots indicate where the wood luus been (rhanged so as to leave cellulose fibers. Small black lines are visible here and there. Fig. 2. Radial view of Red Spruce log (natural size), showing advanced stage of the same decay. The mimber of white spots has increased. Tlu' decay rarely goes beyond this stage. PLATE VII. Radial view of a lug uf i'.alsain i'ir ( .lA/Vx Ixilsunird (!,. ) Mill.), shitwing advanced stagi' iif ilccay due ti> '/'ndiuiis />iiii { ilnil. | l''i-. f's jiiiii iormsi ahidis, showing how the fibers are grachially changed until only cellulo.se is left; " w," unchange>nl forma (thirds. Parallel lines of holes filled with cellulose fibers, here indicated Ijy dots, appear in the wood. The black lines bounding the cavities siniplj^ indicate the limit of change of cellulose, for in reality there is no such sharp line of demarcation. The short line at the right equals about 25 of an inch (1™"'). Fk;. 3. Later stage of the same form of decay. The wood is now simply a network of narrow wood lamelhe separating larger and smaller holes. In these lamelUe ]>lack lines are shown, which represent plugs of brown hyphfe incrusted with decomposition products. (See PI. IX, figs. 10 and 13. ) Cellulose fibers and mycelium fill some of the cavities. The short line at the base equals about ^V of an inch (1'"'"). Fig. 4. Longisection of wood (Spruce), showing effects of destruction by hyphje of Polyporus pinicola. Fig. 5. Cross section of several wood cells, showing changes which take ])lace in wood such as shown in fig. 4. PLATE XL Fk;. 1. Tangential view of Spruce avoo<1 destroyed by mycelium of PoIi/jkh'uk snJ- fureus (Bull) Fr. : "a" wood elements which have been curved, bringing two med- ullary rays into line; " e" part where a break occurred, uniting two medullary rays. Fk;. 2. Radial view 01 wood in last stage of decay, induced by mycelium of Poli/- jwnis Kiihitriilus Pk. The straight lilack lines represent one or more wood elements held in i)lace by the hyplue wliich are wound all around them. Remnants of medul- lary rays are to be seen here and there. Fk;. 3. Several cells from such a piece as is shown in fig. 2 (also PI. XIV, fig. 3). Normal wood cells of the spring wood are shown at the left, and going toward the right various stages in the solution of the cell walls. Fk;. 4. Tangential view of a piece of Spruce wood destroyed by mycelium of Poly- porus sulfureus, showing characteristic breaks in the wood, formed by the uniting of many medullary rays by cross breaks. (See fig. 1 of this plate.) The short line at the left is equal to l'""'. PLATE XII. Various forms of sporophores of Tixnnck's phil forma abietis. Fig. 1. On Balsam Fir. Fk;. 2. On Tamarack. Fig. 3. On horizontal branch of Spruce. Fig. 4. On bark (jf trunk of Spruce. Fk;. 5. At base of dead branch of Spruce. Fk;. t). Semipileate form on Spruce. Fig. 7. At base of dead branch of Spruce. PLATE XIII. Radial view 33 Ti 33 Bui. 25, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. PLATE VI. k'^^^itjiim Fig. 1. Red Spruce: Early stage of the decay caused by Trametes pini forma abietis Fig 2. Red Spruce: Advanced stage of the decay caused by Trametes pini forma abietis. Bui. 25, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. PLATE VII. Log of Balsam Fir showing decay caused by Trametes pini forma abietis. Bui. 25. Oiv. Ve|, Phys. & Path., U. S. Dept. of Agriculture. PLATE VIII. FIG. 1. FIG. 2. Fig. 1 EARLY STAGE AND FIG. 2 LATE STAGE OF DECAY OF LARCH CAUSED BY TRAMETES PINI FORMA ABIETIS Bull 25, Div, Veg. Phys. & Path,, U, S. Dept. of Agriculture. Plate IX. POLYPORUS SUBACIDUS PK., POLYPORUS PINICOLA 'SWARTZl FR., AND TRAMETES PINI (Brot.) Fr. forma abietis Karst. Bull. 25, Div. Veg. Phys. & Path., U, S Dept. of Agriculture. Plate X. Work of Polyporus pinicola iSwartzi Fr. and Trametes pini iBrotj Fr. forma ABIETIS KaRST Bull. 25, Dlv. Veg, Phys. & Path., U. S. Dept. of Agriculture. Plate XI. Stages of decay induced in Spruce by Polyporus subacidus Pk. and Polyporus suLFUREus I Bull. > Fr. Bull. 25, Div, Veg. Phys & Path , U, S. Dept. of Agricultuie. Plate XII, & ( 7 I^ Various i-ciHMh of sporophores of Trametes pini i Brot.) Fr. forma abietis Karst. Bui. 25, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. PLATE XIII. Block of White Spruce wood showing injury caused by Polyporus sulfureus. Bui. 25, Div. Veg. Phys. & Path., U. S. Dept. of Agriculture. PLATE XIV. FIG. 1. FIG. 3. Fig. 1 EARLY STAGE AND FIGS. 2 AND 3 SUCCESSIVELY LATER STAGES OF THE DECAY CAUSED IN White Spruce by Polyporus subacidus Peck. Bui. 25, Oiv. Veg. Phys. & Path., U. S. Oept. of Agriculture. PLATE XV. Fig. 1. Cross section of log of Spruce showing decay caused by POLYPORUS SUBACIDUS PECK. Fig. 2. Resupinate form of sporophore of Polyporus subacidus Peck on Spruce log. Bulletin No. 26. V. P. P. 79. U. S. DEPARTMENT OF AGRICULTURE, DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. ALBERT F. WOODS, CHIEF. WAKKER'S HYACINTH GERM. Pseudoinonas Jiyacinthi (Wakker). BY ERWIN R SMITH, BOTAAitt^ML IN CHARGE OF LABORATORY OF PLANT PATHOLOGY. Issued February 21, 1901. WASHINGTON: GOVKRNMENT PRINTING OFFICE. I 90 I . OFFICE OF PLANT INDUSTRY. B. T. Galloway, Director. AFFILIATED DIVISIONS. Gardens and Grounds, B. T. Galloway, Superintendent. Vegetable Physiology and Pathology, Albert F. Woods, Chief. Agrostology, F. Lamson-Scribxeb, Chief. Pomology, G. B. Brackett, Chief. Section of Seed and Plant Introduction, Jared G.- Smith, Chief. DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. SCIENTIFIC STAFF. Albert F. Woods, Chief of Division. Mertox B. Waite, Assistant Chief. associates. Erwin F. Smith, Wm. A. Orton, Newtox B. Pierce, Ernst A. Bessey, Herbert J. Webber, Flora W. Patterson, M. A. Carleton, Hermann von Schrenk,i P. H. Dorsett, Marcus L. Floyd.^ Thos. H. Kearney, jr. in charge of laboratories. Albert F. Woods, Plant Physiology. Erwin F. Smith, Plant Pathology. Neavton B. Pierce, Pacific Coast. Herbert J. Webber, Plant Breeding. 1 Special agent in charge of studies of forest-tree diseases, cooperating with the Division of Forestry, U. S. Department of Agriculture, and the Henry Shaw School of Botany, St. Louis, Mo. 2 Detailed as tobacco expert, Dirision of Soils. Bulletin No. 26. V. P. P. 79. U. S. DEPARTMENT OF AGRICULTURE, DIVISION '' VEGETABLE PHYSIOLOGY AND PATHOLOGY. ALBERT F. WOODS, CHIEF. WAKKER'S HYACINTH GERM, Pseudojiioiias hyacinthi (Wakker). BY ERWIN F. SMITH, IN CHAROE OF LABORATORY 6f PLANT PATHOLOGY. Issued February 21, 1901. WASHINGTON: GOVERNMENT PRINTING OFFICE. I 90 I. LETTER OF TRANSMITTAL. U. S. Department of Agriculture, Division of Vegetable Physiology and Pathology, Washington, D. C, October 6, 1900. Sir: I respectfully transmit herewith and recommend for publica- tion a report by Dr. Erwin F. Smith, of this Division, on a bacterial disease of hyacinths commonly known as "the yellow disease" or "Wakker's disease." The fact that large numbers of hyacinth bulbs are forced each year in the United States makes it desirable that their diseases be understood. The information gained regarding the biologj^ of the organism will also be of great value to those investigating the bacte- rial diseases of plants. The report confirms earlier work done in the Netherlands and adds much new and important information respecting the nature of the parasite. The latter belongs to a group of bacteria, hitherto but little studied, several members of which (also studied by Dr. Smith) cause diseases widely prevalent in the United States. The report while primarily for pathologists and bacteriologists will also be of interest to florists and any others who wish to detect this disease and to avoid its introduction into the United States. Respectfully, Albert F. Woods, Hon. James Wilson, Chief of Division. Secretary of Agriculture. PREFACE. This paper was prepared for publication in August, 1897, at which time I had secured characteristic infections and had worked out many of the cultural and other characters given in the following pages. The fact that I had not again produced the disease with germs isolated from mj^ first series of infected plants, the further fact that I could not satisfactorily explain the meager growth of the parasite in the host plant, and on steamed j)otato and the other culture media which I had used, and, finally, a shadow of doubt concerning the accuracy of two or three other observations, induced me to withhold tlie paper and repeat the experiments. In the time which has intervened I have gone over nearly or quite all of the experiments detailed in the origi- nal paper, without, however, discovering any serious errors. During this time reinfections have been secured, the reason for the feeble parasitism lias been discovered, and a number of other interesting facts have been brought to light, so that the long delay of publication has not ])een without its comiDensations. Throughout this study numerous comparisons have been made with two other yellow bacteria, Pseudomonas campestris and Ps. phaseoU, and occasional mention has been made of them in this paper, both being plant parasites. Occasional comparisons have also been made with other bacteria, especially with Ps. Stewarti. The leading cultural characters of the hyacinth organism are mentioned in the synopsis at the end of this paper, but it has been decided to relegate an account of the numerous experiments on which these conclusions rest to a second bulletin, which is now ready for publication and in which they will be discussed in connection with the cultural pecu- liarities of the other yellow species of Pseudomonas here mentioned. It is too much to hope that this bulletin is entirely free from mis- takes. Nevertheless great pains have been taken to make it trust- woi'thy, all of the experiments having been performed in duplicate, and iiearlj^ all of them having been repeated several times on differ- ent occasions to eliminate unsuspected sources of error. Some brief statements respecting the morphology and physiology of this organism, as determined by the writer, were made at the Detroit meeting of The American Association for the Advancement of Science in August, 1897, and were pul)lished in tlie Proceedings of the Associ- ation for that year (Vol. XLVI, 1897, Salem, June, 1898). Erwin V. Smith. 5 CONTENTS. Page Historical ... 9 Source of material _ 10 Inoculations of 1 897 ... 10 Series 1 (hyacinths) 10 Series 2 (onion) .. 18 Natural infection of a daughter bulb _ 18 Inoculations of 1898 19 Series 3 (hyacinths) 19 Series 4 (onions) .. ._ 20 Series 5 (hyacinths) 20 Series 6 (Roman hyacinths) /. 23 Series 7 (hyacinths) 28 Series 8 (hyacinths) 25 Series 9 (hyacinths) 26 Series 10 (hyacinths) 26 Series 11 (cabbage) .. 27 Series 12 (amaryllis). 28 Series 13 (hyacinths) 28 Series 14 (hyacinths) . 29 Series 15 (onions) ._ 30 Series 16 (hyacinths) 30 Series 17 (hyacinths) 31 Series 18 (plunge experiment) 82 Inoculations of 1899 ., 33 Series 19 (hyacinths) 33 Remarks on pathogenesis . 33 Morphology of the parasite _ 36 Size and shape 36 Motility 37 Zoogloeae 38 Spore formation 39 Involution forms 39 Behavior toward stains 40 Synopsis of characters 40 Remarks on relationship 42 Explanation of illustrations 44 I ILLUSTRATIONS Plate. Page. Plate I. Pseudomonas; hyacinthi ( Wakker ) Erw. Sm 46 Text figures. Fig. 1. Diseased scale of hyacintli ...,. 22 2. Inoculated leaf of hyacinth ... 23 3. Culture of Pseudomonas hyacinthi on slant 30 per cent cane-sugar agar, showing '• shagreen"' surface . ..._.. 38 4. Slightly magnified diagrammatic views of slime of Ps. hyacinthi on sweet potato, showing ' ' shagreen '" surface 38 5. Typical behavior of Ps. hijacinthi in fermentation tubes containing peptone water, or peptonized beef bouillon, with addition of vari- ous sugars and other carbohydrates 41 6. Ps. hyacinthi growing in strongly alkaline (0) gelatin with 10 per cent cane sugar. No liquefaction. The surface curves are due to the very gradual drying out of the gelatin 42 8 WAKKRR'S HYACINTH GERM, Pseudovionas hyacinthi (Wakker). HISTORICAL. Dr. J. H. Wakker published five papers on the hyacinth germ between 1883 and 1888. His studies attracted wide attention because he was one of the earliest investigators in the field of plant bacteriol- ogy in a time of general skepticism and uncertainty, and also because of the great care with which he seemed to have worked out his results. Since the conclusion of Dr. Wakker's studies, which were begun in 1880, no bacteriologist or plant pathologist seems to have given any personal attention to the disease. Several pathologists have written about it or referred to it,' but nothing of any value has been added, and some of the comments have served only to throw doubt on the original inquiry. In reading Dr. Wakker's papers for the purpose of making an abstract, I was at once struck with the need of a reinvestigation of the subject. This seemed necessary for two reasons: (1) Methods of isolation were not then as well understood as at present, and most of Wakker's successful infections seem to have been direct ones; (2) the germ is so imperfectly described that, excluding the test of patho- genesis, the identification of any particular organism as Bacterium hyacinthi Wakker would be altogether impossible. No disparage- ment of Dr. Wakker's beautiful studies is here intended. At the same time nothing perhaps better serves to illustrate the important advances ' De Bary: Vorlesungen iiber Bacterien, Leipzig, 1885, p. 137; also 8. Auflage, Leipzig, 1900, p. 173. Sorauer: Handbuch fler Pflanzenkrankh. . 2. Auflage. 2. Theil, Berlin, 1886, p. 99. Kramer: Die Bakteriologie in ihren Beziehungen zur Landwirtschaft, etc. Erster Theil. Wien. 1890. p. 145. Comes: Crittogamia Agrai-ia. Napoli, 1891, p. 510. Ludwig: Lehrbnch der niederen Kryptogamen. SUittgart, 1892, p. 90, Tubeuf : Pflaiizenkr. durcii kryptogame Parasiteii vernrsacht. Berlin. 1895, p. 550. Prillieux: Maladies des Plantes agricoles et des Arbres frui tiers et forc^stiers causees par des parasites vegetaux, Paris. 1895. Tome I, p. 22. Frank: Die Krankheiten der Pflanzen, 2. Auflage, 2. Band, Breslau, 1890, p. 23. Migulaj System dei- Baketrien, 1. Bd.. Jena, 1897, ]). 320. Hartig: Lehrbuch der Pflanzenkraiikheiten, 3. Auflage. Berlin. 1900, p. 209. 9 10 which have been made in the technique of bacteriology than a perusal of the best earh' papers. It is not unlikely that the additions which I shall make will also be insufficient, exclusive of the pathogenic test, to differentiate this germ ten or twenty years hence, but they will at least help toward definitely settling the group to which it belongs. Readers who wish merely a summary of Dr. Wakker's conclusions will find it in my critical review already cited, and those who wish to read the original papers will find the necessary references in the same paper. ^ Inas- much as that review is very full and readily accessible, I may be excused from going over the ground again in this place. SOURCE OF MATERIAL. The hyacinth bulbs from which the germ that I have studied was isolated were said to be in the first stages of the yellow disease, and were sent to me in October, 1896, by Messrs. Van Meerbeck & Co., growers of bulbs at Hillegom, near Haarlem, Netherlands. The bulbs were sound externally. They had been " visited," ^ and some of the vascular bundles of the inner scales were yellow, broken down, gummy, and full of bacteria. Penicillium was also present in places. No difficulty was experienced in isolating a yellow micro-organism from the broken down bundles of one of these bulbs, and subsequently the same germ was isolated from another bulb of the same lot. By plant- ing a third bulb the disease was also obtained tlie following year in a daughter bulb. I have now cultivated this organism over four years in hundreds of cultures on a great variety of media, and have also obtained very satisfactory infections— infections so exactly like those described by Dr. Wakker that there can be no doubt either as to the nature of the organism with which I have worked or as to the substantial accuracy of Dr. Wakker's conclusions respecting its patho- genic properties. INOCULATIONS OF 1897. SERIES 1 (HYACINTHS). The first set of inoculations was made February 16, 1897, from a pure beef-broth culture. Eight vigorous hyacinths were inoculated. They were all of one variety, a robust, single-flowered, deep-blue sort (name unknown). The plants were just coming into blossom and were the picture of health, six of the eight bulbs being large and well- stocked with food, and the other two smaller daughter bulbs. Part of the inoculations were by means of ordinary needle punctures and 1 The Bacterial Diseases of Plants: A Critical Review of the Present State of our Knowledge, Parts III and IV, The American Nati(ralist, October and Novem- ber, 1896. pp. 797, 912. 2 Removal of the top of the bulb with a sharp knife for purposes of inspection is called •• visiting." This is done after the bulbs are dug. 11 the rest by means of a hypodermic syringe, the results being the same, except that the symptoms appeared sooner when a large nnmber of germs were inserted. All of the inocnlations were made in the middle or terminal parts of healthy leaves, with one exception, in which case the germs were inserted into the upper part of a flower shaft before the buds opened. Much to my surprise, the progress of the disease was very slow, exactlj^ as described by Dr. Wakker, and the striping down of the disease was restricted in most cases to long, narrow areas, with healthy green tissue to either side. In case of the hypodermic injections, however, a width of three to eight or more vascular bundles was involved, i. e., as much breadth of tissue as appeared water-soaked after the injection, but not much more. Even when a great quantity of germs Avas injected (0.5 cc. or more of a fluid culture) the disease did not appear immediatel}^, develop rapidly, or cause widespread infection of the bulbs. To show how closely my results tally with those obtained by Dr. Wakker,^ I will here set down the course of the disease in each of the eight plants first inoculated. ' See Contributions k la pathologie vegetale, I, La maladie du jaune, ou maladie nouvelle des jacinthes, causes par le Bacterium Hyaciiithi, Archives neerlandaises des sci. ex. et naturelles. 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The plants were on a bench in a greenhouse, where there was an abundance of light and air and where they received water from time to time as required. The external symptoms were so slight and progressed so slowly that no record was kept after March 22. The plants were, however, under almost daily observation during April and May. The}^ made a vigorous growth for two months or more after flowering time, but as the warm weather of summer came on the leaves gradu- ally dried out and died from the top down, and, with the exception of the bases, were pretty uniformly shriveled by the middle of June. This shriveling was not, however, the result of the disease. In fact, so little increase of symptoms was observed in April and Maj^ that when the bulbs were cut open (June 23) it was with no expectation that any diseased places would be found. That some of the leaf inoculations did dry out and fail after starting was evident, but enoiigh succeeded to place the success of the experiment beyond doubt, one or more bundles in the bulb scales of each one of the 8 different plants being yellow, broken down, and full of bacteria (see PI. I, fig. 1). None of these plants became wet-rotten or bad-smelling as a result of the bacterial infection, the symptoms being wholly unlike those obtained by Dr. Heinz with his Bacillus hyacinthi-septiciis. So far as observed the diseased plants had no odor whatever; certainlj^ no pronounced odor. No mycelium was present in any of these yellow, broken-down bundles, or in any of the bulb scales, and in most cases no micro-organism of any sort was present except the one which had been introduced into the leaves (and scape) in the preceding Febru- ary; i. e., more than four months before and at a distance of from 15 to 25 cm. from the bulbs. No animal parasites were observed. Nearly all the scales of these eight bulbs were still entirely sound, but from the condition of the plateau, when the germs had penetrated that far, it was evident that a general infection of the scales and a more or less complete destruction of the bulbs would have been only a matter of time. Even in the attacked scales the greater part of the tissue was still sound. Previous to making these inoculations I was inclined to attribute the slow progr(!ss of the disease in Dr. Wakker's inoculated plants to the low temperature at which his plants were kejit, or else to his hav- ing used cultures containing very few living germs. ^ Having myself inoculated from a culture in prime condition for experimental pur- poses (i. e., swarming with motile rods), having in nearly one-half the cases inserted great numbers (that is, thousands) of the germs, hav- ' From my critical review, published in 1896, it will be seen that even then I was inclined to regard Dr. Wakker's statements respecting pathogenesis as substan- tially correct, and my subseciueut studies have fully contirnied this view. 8970— No. 20—00 2 18 ing kept the plants at considerably higher temperatnres (20° to 30° C), and yet having obtained the same results as Dr. Wakker, I am forced to the conclusion that the organism is a rather feeble parasite and that the slowness of its progress in the plant is due to natural causes, tne discussion of which I will undertake later on. SERIES 2 (onion). On December 13 six shoots of an onion (Allium cepa) were inocu- lated with bright yellow slime from a potato culture (tube 12, Decem- ber 4), by means of numerous needle punctures. Result: The plant developed no leaf symptoms, and when the four bulbs (all from one root) were dug and examined in June, 1898, there was no trace of yellow bundles or other indication of disease. NATURAL INFECTION OF A DAUGHTER BULB. In April, 1897, a diseased bulb was potted and placed in the hot- house. This was the last remaining bulb of those received from Holland the preceding fall, the rest having been cut for study or having fallen to pieces in the dry air of the laboratory, to which they had been exposed for six months. The planted bulb did not sprout for a long time, but finally developed some feeble leaves. No par- ticular attention was given to it during the summer and fall, but in midwinter I noticed that the leaves were dying at the top and were crooked — i. e., came up exclusively from one side of the bulb and curved over toward the other side. In February, 1898, the plant was knocked out of the pot and examined. The bulb which I had planted was completely decayed. All of the leaves were from a small daugh- ter bulb, which was not present, or at least not visible, when the mother bulb was planted. This bulb was one-sided, had only a few leaves, and these were dying at the top. There was no wet rot of the leaves or bulb and externally the bulb was sound. On cutting it open more than forty vascular bundles in the otherwise sound white scales were found to be bright j^ellow, and a careful microscopic examina- tion showed them to be full of the hyacinth germ. These yellow bundles were in eight different scales. That the daughter bulb had contracted the disease from the mother bulb which I planted was evident (1) from the fact that there was no other visible source of infection — i. e., this bulb was planted in good soil, in which hyacinths had never grown and was the only hyacinth in the gi-eenhouse; (2) from the fact that the plateau was the most badly affected part of the bulb; and (3) from the fact that the scales seemed to have been infected from below up, the yellow slime in more than two-thirds of the affected bundles being visible to the naked eye only in the lower half of the scales, whereas in bulbs which became diseased as the result of my leaf infections the upper half of the scales (so far as examined) was always the first to show the symptoms. Probably the 19 leaves curved toward the decayed mother bulb from whicli the infec- tion was received, as in case of one described by Dr. Wakker, but this I neglected to determine. INOCULATIONS OF 1898, The following year these experiments were rei>eated. All the plants were in the same greenhouse. The night temjperature of the house for a month or two, during which symptoms were slowly extending in the hyacinth leaves, was 10° to 18° C. ; the day temperature was 21° to 31° C. Subsequently, during May and June, the temperature fluctuated more, and some of the time it was considerably higher, especially in the daytime — that is, 10° to 20° C. by night and 30° to 46° C. by day. On quite a good many days during this period the air temperature for some hours ranged from 35° to 40° C. — i. e., too high for the growth of tliis organism, as shown by maximum temperature experimrotective state entered into by bacteria whenever the physical or chemical conditions of the substra- tum are unfavorable to growth, these conditions being either independent of the organism, as in this case, or brought about by its own metabolism? In beef In-oth and other fluid cultures the tiny aggregations of this organism showed a marked tendencj^ to gather into a ring or rim on the wall of the tube at the level of the liquid, and sometimes floating islands appeared, but the flocculent matter seldom united into any tough pellicle, being easily jarred apart and into the depths of the fluid. These zooglo^fe appear to the naked eye either as small whitish flecks or, Avhen on the rim at the surface of the liquid, as round, yellow, colony-like bodies, espe- cially when they have reached some age and density. These bodies also formed on substrata rich in assimilable sugars; here, perhaps, owing to the development of acids. On the solid, sugar- rich substrata, e. g., sugar-agar, potato with sugar, sugar beet, sweet potato, etc., they pro- duced a papillose, verrucose, or shagreen-like surface, the tiny rounded elements forming this surface being very smooth and distinct in their upper part, but fused below next to the substratum. This shagreen also appeared, on old cultures, on nutrient starch jelly containing 5 per cent glycerol. This appearance is shown in figs. 3 and 4. Flu. o.— Culture of Pseudomonas hya- cinthi on slant 30 per cent cane-sugar agar, showing "shagreen" surface. Fig. 4.— Slightly magnified diagrammatic views of slime of Ps. hyacinthi on sweet potato, showing "shagreen ■■ surfaoe. 39 SPORE FORMATION. No spores have been seen, and I am in considerable doubt as to whetlier the spores observed in some of his cultures and studied so carefidly by Dr. Wakker belonged to this species. He was never able to find any in the host plant, and those which appeared in his tubes may have been due to the fact that he was working at times with contaminated cultures. None of his successful infections with sporiferous material were made with spore masses entirel}^ free from vegetative rods, and the latter are long-lived. This supposition of mixed cultures is the more likely because his work was done at a time when it was impossible to decide with ease and certainty on the purity of any given culture — i. e., before the era of poured plates — and especially because some of his gelatin cultures were certainly contaminated, i. e., yielded gas bubbles. (See Fermentation tube experiments described in Bulletin No. 28 dealing with the cultural characters of this organism.) While, therefore, not wishing to deny absolutely the existence of spores in this species, it seems to me that further and more exact proof is necessary to demonstrate their occur- rence. A great many old cxdtures, grown on a variety of media at 18° to 26° C, have been examined without finding any spores. None were observed in the diseased bulbs, many of which were examined with care. Neither did any spores form in cultures exposed for fif- teen da3"s to air deprived of its oxygen by the potash-pyrogallic-acid method (test by microscopic examination and by exposure for ten minutes to 60° to 70° C. in alkaline beef broth). None developed in solid or fluid cultures exposed six weeks in the thermostat at 34° to 35° C. These cultures included alkaline and acid beef broth and cylinders of turnip and sugar beet standing in distilled water. Fur- thermore, this germ will not grow at all or grows onlj' very feebly at the temperature Avliich Dr. Wakker states to be most suitable for the formation of the spores viz. 35° C. (See Maximum temperature for growth, in Bulletin No. 2S.) Finally, no spores developed in cultures which were first grown foi' a week or two at room temperatures and then put into the thermostat at 34° to 35° C. Several different media were tried, but vigorous growth stopped immediately, and after two weeks all such cultures were dead. INVOLUTION FORMS. Some astonishing involution forms have been observed. They formed a whitish i-inx at the surface of the fluid in strongly (soda) alkaline beef-broth cultures to which 10 per cent cane sugar had been added. The color was so pale that at fii-st the tubes wei-e su])posed to be contaminated. Wheu examined mici-oscopically the cultures were five weeks old. These bodies wei-e so immensi-ly swollen, fused, twisted, and irregular in outline that seen on the slide no one to wliom 1 showed them liad any suspicion tliat they were bacteria. Involution forms were also seen on old tui-nip and banana cultures. 40 BEHAVIOR TOWARD STAINS. Beyond the fact that the flagelhini was stained with difficulty and that old growths, whether in the plant or out of it, took stains feebl}^ nothing- peculiar was observed, unless it be that the bacterial j^recipi- tate resulting from growth was not stained in Dunliam's solution con- taining methylene blue, and was stained in the same medium with rosolic acid. The following are transcripts from records scattered through my notes: Germs from an old culture in strongly alkaline (soda) beef broth stained slowly and rather feebly in a saturated alcoholic solution of gentian violet diluted with an equal bulk of distilled water and allowed to act for half an hour. This culture had lieen killed by heat in the thermostat. Germs from an old culture in acid beef broth which had become alkaline, stained feebly in Ziehl's carbol fuchsin with ten minutes' exposure. This, also, was undoubtedlj- a dead culture. Germs from a month-old culture on sugar beet were exposed for some time to a dilute watery solution of gentian violet, whereupon all the zoogkea^ stained deeply, but tlie loose rods rather feebly. On long exposure (over an hour) everything stained deeply. Germs from sweet potato cultures a month old (zoogloese, rods, doublets, and chains) stained feebly in a deep-colored Avatery solution of gentian violet, although exposed for one-half hour. Germs taken from one of the bright-yellow bundles of a diseased bulb (June 23, 1897) stained feebly in water made deep red with Griibler's basic fuchsin. Germs from the yellow bundles of another bulb (Feb. 3, 1808) showed a verj^ weak stain after five minutes' exj)osure to water saturated with Grii- bler's basic fuchsin. Exposed two minutes to water saturated with gentian violet, the stain was much better, but not deep enough. The rods from j'oung cultures stain readily. SYNOPSIS OF CHARACTERS. For convenient reference I have drawn up the following brief account of this organism : Pseudomonas hyacinth i ( Wakker) . A yellow, rod-shaped organism, multiplying by fission; ends rounded; single, in pairs, or 4's, more rarelj" in the form of chains or filaments; motile b}^ means of one polar flagellum. In the host plant, when the bundles are crowded full of the 3'ellow slime and broken down, it is, generally, 0.8 to 1.2 hj 0.4 to 0.6 /<. In alkaline beef broth or on agar it usually measures 1.0 to 2.0 b3^0.4 to 0,6 //. In old cultures rich in sugar it often grows out into long, slender chains, or into filaments (50 to 100 a^ long) in which there are no distinct septa. Nonsporiferous. Color distinctly yellow, but somewhat variable. Chrome yellow to pale cadmium in the host plant, i, e., bright yellow^ (Ridgway's Nomenclature of Colors). On ^ Saccardo's Jiavus and citrinus, but brighter (Chromotaxia) . The Standard Dic- tionary's i/pjlojc III. lemon . and cauari/. approximately ( under Spectrum ) . Prang's yellow. Plate I y. in the Prang Standard of Color. Popular Ed.. No. 1. 41 culture media, when not interfered with by the brown pigment, gen- erally gamboge, chrome j^ellow, or canary j^ellow, but sometimes paler. Old cultures on some media darken from the production of a soluble, pale-brown pigment. This feeble brown stain is best developed in hyacinth broth, in potato broth with peptone, on turnips, on radishes, and on banana rinds. It was not observed in acid or alkaline beef broth, on coconut flesh, on sugar beets, in nutrient starch jelly, in agar, or in gelatin, with or without sugar. This organism grows readily on potato cylinders standing in distilled water, but it never becomes copi- ous or fills the water with a solid yellow slime, owing to its feeble dia- static action. Potatoes on which it has grown, even for several months, always give a strong starch reaction with iodine. It behaves the same on nutrient starch jelly free from assimilable sugars. It liquefies nutrient gelatin and Loetfier's blood serum, but does so slowly, and will not liquefy gelatin at all if 10 per cent cane sugar is added (fig. 6). Growth on nutrient agar or nutrient starch jellj^ is inhibited (unless the inocula- tion be from a solid culture and very copious) by the addition of 10 per cent glycerol, and is greatly retarded bj" 5 per cent glj'cerol ; even 2i per cent of glycerol retarded growth. Growth in beef broth was much retarded by the addition of 1.5 per cent sodium chloride. Organism extremely sensitive to plant acids, including those of the hyacinth. Aerobic; doubtfully, if ever, facultative anaerobic; not a gas producer (see fig. 5). Does not redden litmus milk, but makes it bluer, and slowly separates the casein from the whey b}' means of a lab ferment. Produces under some cir- cumstances, and slowl}^ a small amount of nonvolatile acid (slime acid?) with various sugars (grape, cane, etc.), which acid is fre- quently obscured by the moderate production of alkali. In the presence of air produces an organic acid (probably acetic) from ethyl alcohol dissolved in milk or bouillon. Inverts cane sugar, but apparently without the intervention of any enzym. Will not grow on 30 per cent grape-sugar agar. Resists dry air very well, i. e., more than forty-eight days when spread on cover glasses in thin layers. In Dunham's solution with methylene blue the color is reduced in a few days, but reoxidizes quickly on shaking; final color (56 days) bright blue. In Dunham's solution with indigo carmine the color changes to a briglit blue, which persists for a long time; final color yellowish. In Dunham's solution with rosolic acid and enough IICI Fig. 5.— Typical behavior of Ps. hyacinthi in fermentation tubes containing peptone water, or peptonized beef bouillon, with addition of vari- ous sugars and other carbohy- drates. Fluid clear in closed end, clouded in U and open end. 42 to render the fluid yellowish, Ps. hyacinth i did not redden the fluid, but made it colorless, the bacterial precipitate becoming rosy or salmon- colored. Produces indol slowly in peptonized beef broth and in pep- tonized Uschinsky's solution; does not produce nitrites in these solutions. Does not reduce potassium nitrate to nitrite in peptonized beef bouillon. Not a strong-smelling germ. Not readily destroyed bj^ its own decomposition products except in media containing alcohol. Will not grow in the thermostat at 37° C, and grows verj- feebly on some media and not at all on others at 34° to 35° C. Optimum tem- perature 28° to 30° C. , or thereabouts. Minimum temperature approx- imately 4° C. Thermal death point (10 minutes' exposure) 47.50° C. ; nearly all the rods are killed at 47° and a great many at 46.50° C. Did not grow at room temperatures after 6 days exjDOSure in alkaline beef broth in the thermo- stat at 35° to 36.35°. Does not grow well in Uschinsky's solution. Grows much better in Uschinsky's solution when j)eptone is added to it. Grows well with a bright yellow color on cylinders of steamed coconut flesh, standing with one end in distilled water. Pathogenic to hyacinths. Enters the plant through wounds, through the blossoms, etc., and multiplies in the vascular system, filling the ves- sels, especially those of the bulb, with a bright yellow slime consisting of bacteria. The walls of the vessels are destroyed and extensive cavi- ties are formed in the bundles. The parenchyma around the bundles is also involved, but only very slowlj-, the organism being a feeble de- stroyer of cell walls. The host plant is not rapidly destroyed, a year or more being neces- sary. The cells are first separated by solution of the middle lamella, but the wall itself seems to finally disappear. The cavities contain innu- merable bacteria mingled with fragments of the dissolved bundles and of the surrounding parenchyma. First described by Dr. J. H. Wakker from the Netherlands, where it often causes serious losses in the hyacinth gardens. Not known to occur in any other part of the world. Feb. 10 JMar.l4 .Apr. 12 Fig. 6.— Ps. hi/acinthi grow- ing in strongly alkaline (0) gelatin with 10 per cent cane sugar. No lique- faction. The surface curves are due to the very- gradual drying out of the gelatin. REMARKS ON RELATIONSHIP. CloseU" related to Ps. campestris (parasitic on Cruciferous plants), Ps. phaseoU (parasitic on beans), and less so to Ps. stewarti (parasitic (?) on corn, especially sweet corn). Readily distinguished from the two organisms first named by (1) its brighter color; (2) its lower thermal death point; (3) its manner of growth on potato cylinders 43 standing in distilled water, i. e., by its feeble action on starch; and (4) its pathogenic properties. Other distinctions are given in Bulle- tin No. 28. Readily distinguished from Ps. siewarti by (1) its differ- ent, brighter color; (2) its feeble growth in Ilschinsky's solution; (3) its liquefaction of gelatin and Loeffler's blood serum; (4) its lower thermal death point; (5) its lab ferment; (6) its much greater sensi- tiveness to acids ; (7) its more luxuriant growth on turnip and rutabaga. From facts in possession of the writer it is certain that there are many yellow organisms moi-e or less closely related to the four men- tioned in this paper, i. e., nonsporiferous, rod-shaj)ed, micro-organ- isms, multiplying by fission, possessing one polar flagellura, and capable of living parasitically or semiparasitically upon various plants. All of these parasitic yellow organisms, at least all I have examined, are morphologically quite different from Bacillus coU, Bacillus amijlovorus, Bacillus tracheiphilus, or any other micro- organism having flagella distributed over its whole surface. They also differ in many cultural peculiarities. They are, however, related to each other in many ways, and appear to form a natural group. I have an idea also that in some species the production of the brown pigment, and in others the production of the yellow pigment, has been nearly or quite extinguished. The species in which both pig- ments come the nearest to being equally well developed is, perhaps, Fs. cainpestris. The yellow pigment appears to be a lipochrome. (See Bui. 28.) There are also, I believe, many morphologically similar yellow bacteria which are purely saprophytic. 44 EXPLANATION OF ILLUSTRATIONS. TEXT FIGURES. Fig. 1. Porfion of a bull) scale from plant No. 20. inoculated February 7, drawn June 14. showing four health}- and four diseased vascular bundles. The parenchyma between two of the latter has largely disappeared, its ■olace being occupied by a cavity full of bacteria. Smaller cavities in the paren- chyma, close to the vascular tissue, are visible in each one of the diseased bundles. The bundle in the middle of the scale also shows the bacterial occupation of anastomosing veinlets. The diseased portions w^ere bright yellow from the presence of enormous numbers of the parasite, which, how- ever, had not reached the surface of the scale. The infection of this scale was from below upward. (Page 22.) Fig. 2. Leaf of plant No. 25, inoculated February 7, drawn April 80. The figure shows shriveled apex and dead central stripe, na-row border of yellow, and beyond this to either side healthy green tissue (white in the figure). In the yellow border on the right side are some dotted areas intended to represent water-soaked tissue: i. e., spots recently invaded apparently by a slow side- wise movement of the bacteria from the central stripe. ( Page 23. ) Fig. 3. Slant. 30 per cent, cane-sugar agar showing the '• shagreen '" surface. Culture No. 9. June 30, 1898. Photographed August 2. (Page 38.) Fig. 4. Enlarged diagrammaticverticalandhorizontalviewof a similar shagreen surface from a sweet-potato culture twenty days old. (Page 38.) Fig. o. Fermentation tube showing behavior of Ps. hyacinthi in peptone water or peptonized beef broth with various carbohydrates, e. g., grape sugar, fruit sugar, cane sugar, milk sugar, galactose, mannit, glycerin, ethyl alco- hol, etc. Maltose is a possible exception, tubes with this sugar having finally clouded very feebly in th'^ closed end. None yielded any gas. ( Page 41. ) Fig. 6. Stab culture in gelatin + 10 jjer cent cane sugar inoculated Febrnary 10. On March 14 there was a well-developed stab and a good surface growth, but no liquefaction, the curved surface being due to the drj'ing out of the gelatin. On April 12 the gelatin had dried out as indicated by the dotted line, but there was no liquefaction. (Page 42.) PLATE FIGURES. Fig. 1. Cross section of the bulb of plant No. 8, inoculated in the upper part of the scape February 16. 1897. Photographed June 23, 1897. Six vascular bundl( s broken down and filled with the bright yellow bacterial slime. Fig. 2. Onion leaf, inoculated January 29. 1898. Painted by F. A. Walpole, March 5. The yellow color of the leaf in the vicinity of the inoculations was due to the slow and long-continued growth of the organisms: i. e.. it is the yellow color of the bacteria showing through. Fig. 3. Leaf of plant No. 51 (series 9). inoculated near the apex (at x) on Febru- ary 11. Painted by F. A. Walpole, .March 5. The water-soaked lines shown in the lower pai't of the stripe were conspicuous. This leaf was injected with 0.3 cc. of a clo