UNIVERSITY OF B.C. LIBRARY 3 9424 05047 460 7 U.B.C. LIBRARY THE LIBRARY THE UNIVERSITY OF BRITISH COLUMBIA Prof • A.H.' fiutchinson. Digitized by the Internet Archive in 2010 with funding from University of British Columbia Library http://www.archive.org/details/peachleafcurliOOpier X Bulletin No. 20. V. p. p.— 72. U. S. DEPARTMENT OF ACzRICULTURE. DIVISION OF VEGETABLE PHYSIOLOGY AND PATHOLOGY. B. T. GALL^OWAY, Chief. PEACH LEAF CURL: ITS NArURE AND TREATMENT BY TsTEAVTOlSr B. PIERCE, lu Charge of Pacific Coast Laboratory, Santa Ana, California. WASHINGTON: (iO VKKNMENT PRINTIN(; OFFICE. 1 <) 0 0 DIVISION OF VEGETABLE PHYSIOLOGY AM) PATHOLOGY. SCIENTIFIC STAFF. B. T. Galloway, Cliiif ()Ksktt, Mekton B. Waite, Oscak Loew, Newto.v B. Pierce, W.m. \. Outon, llEHIiEHT ,1. WeIJHEK, KkNST A. BeSSEY, M. .\. Cakleton, Fi.oKA W. Patterson, Maucis L. Fi.oyi).* I\ ('IIAKdlO OF l.AMOUA roIUES. >\m!EKT v. Woods, I'lmil riii/sia/iK/i/. Krwin 1*'. Smith, J'Imil I'tilliolni/i/. Newton i'>. I'iickci;, I'tn-ijii- CuukI Liiliiirri/. Herbert .). Weisbkr, J'lant Brccdimj. Oscar Loew, I /'l<(iii Nutrition and Ftrnicnidtion. *I)cluilcil as lohacco cxpi-Tt, Divisiim of Soils. till (liuitjL' of loliacco liiiiKiitalioii iiivtsligutions, Division of Soil.s. .EITER OF TRANSMHTAl U. 8. Dei'aktment of Acjkicultuke, Division of Vegetable Physiology and PAXiioLOciY, WasJungUm, D. C, February '20, IDOO. 8ik: I respectfull}' transmit herewith a report on peach leaf curl, prepared by Mr. Newton B. Pierce, who has charge of the work of this Division on the Pacific coast, and reconnncnd that it be published as Bulletin No. 20 of the Division. The report embodies the results of investigations and experiments carried on for a number of years, and shows conclusively that peach leaf curl can l)e controlled l)v com- paratively simple and inexpensi^'e treatment. Respectfully, B. T. Galloway, Chief of DivUloi). Hon. James Wilson, Secretary of Agriculture.- 3 HTTER OF SUBMITTAL Pacific Coast Laboratory, Santa Ana., Cal., Deconhcr 15., 1899. Sir: I herewith submit a report of investigations on the nature and treatment of peach leaf curl. The experiments described were con- ducted under the most varied conditions of soil, climate, etc., in all the leading peach centers of the United States, and it is believed that the recommendations for treatment here given are equalh' applicable wherever peaches are grown. Respectfully, Newton B, Pierce, lu ( '/i(fr(//; of Pacific ( ''oast Laljoratorij. Mr. B. T. CirALLOWAY, Chief., Division Thinning the fruit of sprayed trees ^ 108 (iathering fruit of sprayed and unsprayed trees 10(> (■omparative quantity, quality, and cash value of fruit fioiu sprayed and unsprayed trees 112 Comparative value of sprays in relation ti > fruit 115 Comparative size of fruit on sprayed and unsprayed trees 119 Color of sprayed and unspraj'ed fruit 1 20 Method of thinning and cost of picking peaches 121 Thinning by hand and by curl 121 Co.st of picking peaches 122 The local action of curl on foliage and fruit 122 Records of trees sprayed on one side ' 1 22 CiiAPfER VII. — Preventive Spray Work Conducted rv ()r( iiahdists 126 General consideration of the auxiliary work 126 Notes on the auxiliary experiments in Michigan 127 Notes on the auxiliary experiments in Oregon 135 Notes on tlie auxiliary experiments in California 140 Notes on the auxiliary experiments in New York, Indiana, and other peach-growing States 144 7 8 CONTENTS. Page. Chapter VIII. — Preparation, Composition, and General Characters of Tii E Sprays Used 1 46 Preparation of the copper sj^ra ys 140 Copper sulphate sohition 147 Bordeaux mixture 149 Eau celeste l'>2 Modified eau celeste l-ili Ammoniacal copper carbonate 1 •">:> Preparation of the sulphur sprays 154 Preparation of combined co])per and sulphur si)niys aii- The enduring qualities of the sprays - . . 1(>2 The corrosive action of the spi-ay s 1 (U Advantages of discernible and indiscernible sprays Kia Sprays adapted to use in wet and in dry localities 1 (>(> Chapter IX. — The Application of Sprays 1()7 ( ieneral accessories for winter spraying 1<)7 Nozzles suited to winter work 1 ()7 Hose and extension pipes 1 ()H Protection of the sprayer 1 70 Pumps for various sized orchards 171 Spraying tanks 1 7r) A])i)lying winter sprays for curl 17") The time for winter spraying 1 7() The manner of applying winter sprays 1 7() Spraying where other diseases are present with curl 177 I'rune rust on the peach {Puccinla ])rinii Pers.) 177 Mildew of the peach {I'orlo.'<]))i:rr(i ii.ii/dcinillur l)e P>. ) 1 78 Brown rot of the peach {Mojiilitt friiclifffiid Pers.) 1 7S Black spot of the jn-ach {('IddoxporiHtn (vrpoj)lilli(in Thiim.) 1 7S Winter blight of the peach and other spot and shot-hole diseases, such as I'liyUoxtirta nrcniniic'tstfd Berk., ('crronjxirK f/fYrwyy/.sr/.v.s-a Sacc., etc.. 170 Sooty mold of the peach 17il Animal parasites of the j)each tree ISO Chai-ter X. — Nati're and Soihck ok riiE Si'uavino Maiioui.m.s I'sed ISl Cop])er sulphate IS! Copper c-arbonate IS.! AnuMonia 1S5 Sodium carljoiiate IS7 Sulphur i!tO Cmai-i i:it \I. — Pi:a( II N'arietiis and Xiusery Stock in Uei.ation io Ci hi.. I;t4 ( 'oiiiiiari.son of peach varieties : 1!M Treatment of nursery stock L'tHi Summarv l-'()2 ILIllSTRATIONS. PLATES. I 'use. Plate I. Curl -infested peach slioot from Rifigs, Cal 11 II. Rlyccliuni of E.ronsais dcfmnnaii — the fungus causing peacli leaf curl . 85 III . Fruiting stages (»f Exonscus dej'ortiKiux 36 IV. Germination of the ascospores of E.rodsciix (h'/onnaiix 88 V. Terminal peach twigs infested with peach leaf curl 42 VI . Sprayed and unsprayed peach branches 42 VII. Sprayed and unsj)rayed Crawfords Late trees, Live Oak, Cal 77 VIII. Unsprayed trees in Lovell orchard, Biggs, Cal 89 IX. Lovell trees sprayed with Bordeaux mixture. Biggs, Cal 89 X. Fruit produced by row 15, experiment block. Biggs, Cal... 107 XL Lovell trees sprayed with sulphur, lime, and salt. Biggs, Cal 112 XII. Lovell trees sprayed with sulphur and lime. Biggs, Cal 112 XIII. Lovell trees sjjrayed with Bordeaux mixture, Biggs, Cal 115 XIV. Lovell trees sprayed with eau celeste. Biggs, Cal 115 XY. Lovell trees sprayed with modified eau celeste, Biggs, Cal 115 Y VI /^^e- ^' -^I^'mt^r of spraying trees on one side, Biggs, Cal "I ,^.j iFig. 2, Action of curl on trees sprayed on one side. Biggs, Cal' XVII. Condition of trees sprayed on one side at maturity of fruit 124 XVIII. Fruit gathered from sprayed and unsprayed halves of tree 124 XIX. Showing fruitfulness of sprayed half of tree 124 XX. Sprayed and unsprayed Crawfords Late trees, I.,ive Oak, Cal 142 XXI. Steam sprav-cooking appliances for small orchards 158 XXII. Steam spray-cooking appliances for large orchards 1(51 XXIII. Appearance of (jrchard at close of spray work, Biggs, Cal 17(i XXIV. Cnj)runed trees, too late for spraying 1 76 XXV. An orchard properly pruned, but too late for spraying 176 XXVI. An outfit for spraying small orchards 204 XXVII. An outfit for spraying medium-sized orcliards 204 XXVIII. Spraying eight trees at a time, Rio Bonito orchard. Biggs. ( al 204 XXIX. A power sprayer in use at Santa Barbara, Cal 204 XXX. A power sprayer, San Diego, Cal 204 FICUUES. Fk;. 1. Cyclone nozzle, w'ith direct discharge and degorger 168 2. Cyclone nozzle, with lateral discharge, for thin sprays 168 8. Heavy cyclone nozzle, with oblique discharge, I'or thick si)rays 168 4. Wire-extended suction hose 169 5. Bamboo extension pipe 169 6. Spray pump for use on barrel or tank 171 7. Spray pump for use on barrel or tank 172 8. Spray pum}) for general orchard work, upright lever 178 9. Spray i)ump for general orchard work, upright lever 174 10. Pneumatic pump for general spraying 175 9 Bull.20, Div. Veg. Phys & Path.. U. S. Dept. Agriculture. Leaf Curl. H-Ii-r DESCRIPTION OF PLATE T. Curl-infested peach .shoot from Biggs, Cal. Leaves of tliis character are badly infested with Exoascus deformans. The greatly broadened and distorted leaves, which are characteristic of this disease, are shown, and the whitened, spore-covered surface of some of the more elevated portions of the upper surface may be distinguished. The petioles of the affected leaves are greatly enlarged, the branch is much bent and distorted, and the internodes of the diseased portion of the branch are greatly shortened. A branch thus badly diseased is apt to die during the year unless con- ditions for growth are very favorable. It is in shoots of this character that the mycelium occurs in greatest abundance, but the hyphaj have been seen to spread only a short distance beyond the parts showing the hypertrophy. (Compare with Pis. V and VI.) PEACH LEAF CURL: ITS NATURE AND TREATMENT. By Newton li. Pierce. CHAPTER I. PRIMARY CONSIDERATIONS RELATIVE TO PEACH LEAF CURL. INTRODUCTION. This bulletin has been prepared to place before the peach growers of the United States the results of experiments conducted during- sev- eral years past for the prevention of peach leaf curl. The losses arising from this disease frequently' amount to several millions of dol- lars annually, and it is believed that a wide dissemination of the results obtained by the experiments here outlined will lead to a large saving to the peach industry. During the progress of the Department's work over one thousand six hundred peach growers in all peach-growing States have ))een requested to test the preventive measur(\s here rec- ommended. A large number have done so, and some of the more important results of their work are also given. From conservative data it has l)een estimated that the experimental work thus widely set on foot l)y the Department has saved to the coiuitry in a singU^ year the sum of three-fourths of a million dollars. This is but a fraction, however, of what may easily be saved in the future, when all growers have obtained a more thorough understanding of the disease and its prevention. The obscure views held by many growers in the past upon the true nature of peach leaf (uu'l, and the total lack of previMitive UK^asures up to a recent date, make it desirable to thoroughly consider the sub- ject at this time and to record the detailed work upon which the con- clusions reached arc l)ased. These conclusions are that peach leaf ciirl may be prevented Avith an ease, certainty, and cheapness rarely attained in the treatment of anj' serious disease of plants, and that there is no longer a necessity for the losses annually sustained from it in the United States. 11 12 PEACH lp:af cukl: its nature and tkeatment. GENERAL CHARACTERISTICS OF THE DISEASE. The disease of peach trees here considered is variousl}^ known in dilterent regions and languages. In the United States it is commonl}^ know as peach leaf curl, or curl leaf of the peach; in England and all Britisli possessions, as leaf blister, leaf curl, or curly leaf; in France, as cloque du pecher; in Germany, as Krauselkrankheit; in Italy, as Fillorisseraa, etc. Peach leaf curl is a disease which seriously affects the leaves, flowers, tender shoots, and fruit of the peach. Its action is most severe in the spring of the year, shortly after the hnifing of the trees, and the greatest injuries are caused in wet settsons and in humid localities. The leaves become enlarged, thickened, much curled, and distorted. As the dis- ease progresscvs the healthful green of the foliage is changed to a yel- lowish, sickly appearance. The leaves soon fall, and the newly formed fruit ceases to grow, yellows, wilts, and likewise falls. The total loss of foliage and crop is common in seasons favorable to the disease. A second growth of leaves develops more or- less rapidly, according to the severity of the disease and the favorable or unfavorable soil and atmospheric conditions prevailing at the time. If the soil and atmos- phere are dry and the temperature high, new foliage may appear slowly and nuich of the terminal growth may die throughout the orchard. In severe attacks young trees are frequently killed. The second crop of leaves, appearing on affected trees after the spring defoliation, usually remains c()m})aratively free from curl for the rest of the season. The amount of disease which will appear upon this later crop of foliage depends largelj'^ upon the humidity or dryness of the atmosphere, excessi\'e moisture favoring a continuance of the ti'ouble. The action of the disciise upon spi'ing branches causes them to enlarge, become curved iind distorted in various ways, and often to dry up and die.' GEOGRAPHIC DISTRIBUTION. IN'ucli Iciif cui'l exists in most peach-growing countries. Its distri- Itutioii in llic I'liitcd States extends from the Gulf of Mexico to Can- ada and Ironi tlie Atlantic to the Pacific. The cent(M-s of greatest prevalence, and where the greatest losses are sustained from tiiis cause, are in tiie leading ))each-growing districts bordering the Great Lakes, e.s|)ecijilly in Michigan and w(^stern New York; in the central, north- ern, and co:ist I'egions of California; and west of the C^ascade Moun- tiiins in Oiegori juid Washington. Tlu^ disease is less serious, or is of minor importance, in tlios«i peach-growing counties of New York most distant from the lakes, in Pennsylvania, Ohio, Indiana, Illinois, and in southern California. Still less injui'y is reported fiom New .Jersey, Dtdawaie, Connecticut, Rhode Island, Massachusetts, Mary- GEOGEAPHIC DISTRIBUTION. 13 land, Virginia, West Virginia, Kentucky, Tennessee, North Carolina South Carolina, Arkansas, Oklahoma, Louisiana, Mississippi, Ala- bama, and Florida, but in most of these regions occasional serious outbreaks are reported in seasons favorable to curl or in particular localities. It prevails rather more seriously in portions of Geor- gia, Kansas, and Missouri. In Texas, New Mexico, Arizona, and Colorado it has occasioned but little loss and is not widel}^ known. Reports from Utah and Nevada are meager, but it is probal^lc that the disease prevails to a limited extent in both States. The more northern States not mentioned here have either failed to report the prevalence of the disease or are properly included within that portion of the United States unsuited, by rigor of climate, to successful peach culture. In Canada both Ontario and British Columbia, which are the leading peach-growing provinces, are favoral)ly situated for the serious devel- opment of peach leaf curl in wet seasons. Mr. John Craig, horticul- turist of the Central Experimental Farm, Ottawa, writes that the disease "obtains in Canada in all the peach-growing districts, including British Columbia and the Province of Nova Scotia." It is known to cause considerable losses of fruit in some sections.* Peach leaf curl exists also in some if not all the peach-growing coun- tries of South America. In Chile the peach is widely grown, being planted from the snow line of the Andes to the Pacific Ocean, and from Copiapo south as far as Valdivia, a distance of 800 miles. Mr. C. T. Ward, Jr.,^ of the Hacienda Loreto, Department of Limache, says that the parasite of peach leaf curl "exists all over the country where the peach grows," but that no satisfactory method of control is A^et practiced there. In Europe Dr. K. Sadebeck'' records the disease from Denmark, Ger- man}^, Austria, Switzerland, and Ital}^ He states that in central Ger- man}^ it prevails more extensively than in the vicinity of Hamburg.* Among the many German scientists who have written upon this ' ^Ir. L. Woolverton, secretary of the Fruit Growers' Association of Ontario, said, in 1890, in a paper entitled, Points onPeach Growing in the Niagara District, pul)- lished in the Annual Report of the Society for that year, pj). 50 and 57: "The peach has its share of enemies and diseases, chief among which are the curl, curcnlio, the borer, and the yellows. For the curl I know no remedy. It is not often severe, but sometimes with the diseased leaves the fruit also drops." Mr. John Craig, in writing from Ottawa under date of October 7, 1897, says, relative' to the disease in Ontario: " It is only severely injurious here during years of unusually heavy rainfall. This year it was very l)ad." 2 Letter of March 22, 1890, to Mr. J. M. Dobbs, U. S. Consul at Valparaiso, Chile. ^Sadebeck, Dr. R., Die para.sitischen Exoasceen. Fine Monographic, Hamburg, 189:5, p. 94. 'Sadebe(;k, Dr. R., Untersuch. iilx-rdie Pilzgattung Exoiuscus, Hamburg, 1884, p. 115. 14 PEACH LEAF CURL: ITS NATURE AND TREATMENT. disease luid its cause are Sadebeck/ Winter,- De Ear\^,'' von Tavel,* Haitig/ Zopf.« Tubeuf/ Ludwig,* Sorauer,' Frank/" Kirchncr," Fuckel,'' and others. Winter says (1. e.) that the fungus of this niahid}' causes great damage by early defoliation of the trees, and that it even kills the diseased trees by its repeated occurrence. In Great Britain peach leaf curl has been common for a great many years. In 1821 it was accurate!}' described 1)}^ an English gai'dener under the name of "blight." He says:" "Under this denomination [])light] are frequently confounded two varieties of disease materially dillerent in their appearance, and which I shall distinguish ])y the appellation of Mister and curl. The former is generally confined to such peach trees as have glandular leaves, which are mostly subject to it in the months of April and May, and when attacked it is not until the latter part of the season, if at all, that the}' become healthy. The leaves so attacked are crisp, and assume a swollen, crumpled, and succulent appeai'ance; the shoots themselves are atfected l)y it in the same manner, and never produce either good blossom or healthy wood." Berkeley '* has described the fungus causing this disease, and it has been mentioned by Bennett and Murray^'' and many other English writers. (Consult a poi)ular article on Peach Blister, by W. G. Smith, Gardeners' Chronlde.^ Vol. IV, pp. 36, 37.) 'tSadebeck, Dr. K., see locations cited; also Einige neue Beobachtungeii uiul krit- ische Bemerkungen iiber die lilxoascacejc, Bot. Ges., 1895, Band XIII, Heft 0. ^Winter, Dr. Georg, Die durch Pilze verursachten KninkluMtcii d(>r Knltnrge- wiichse, Leipzig, 1878, p. 47; also Ral). Krypt. Flora, 18S5, II, p. (i. ^De Bary, Prof. A., Comparative Morjjhology and Biology of the Fungi, INIyceto- zoa, and Bacteria, English edition, Oxford, 1887, p. 265; see also in the same volume various other references to the arrangement and jxtsition of tiie Exoascus group. 'Tavel, Dr. V. von, Vergleicliende ^Morphologic der Pilze, Jena, 1892, pp. 55,5(5. •^Ilartig, Dr. Robert, Lehrbuch der Baumkrankheiten, Berlin, 1889, j). 118; also the English edition, Text-lx)ok of the Di.seases of Trees, London, 1894, j). WVl. "'Zopf, Dr. Wilhelm, Die Pilze in mori)hologischer, phy.siologisclici', biologisclier, und systematischer Beziehung, lireslau, 1890, pp. 2:^6,282. 'Tubeuf, Dr. Karl Ennheer von, rilauzenkrankheitcn durch Uryptogamc I'ara- siti-n verursacht, Berlin, 1895, pp. 167-188. ''Ludwig, Dr. Friedrich, Lehrbuch der Niedcrcn Krvptoganuii, Stuttgart, 1892, p. 205. '•'Soraucr, Dr. Paul, ilandl)U(li der i'llau/.cukranUlicilcu, /wcitcr Tlicil, Die para- sitiircri KranUhi-itcn, Ik-rlin, 188(), p. 278. '"Frank, Dr. \. B., Dit; Krankheiten der IMlauzcu, l>aiid 11, Die I'ilzpanisitarcu Krankhcilcu, I'.re.siau, 1896, pp. 24!», 250. Fdition of 1880-81, Vol. II, p. 52(). "Kirchncr, Dr. O.^kar, Di(t Krankheiten und Heschiidigungen unserer lamhvirl- wrhaftlichen Kuiturpflanzeh, Stuttgart, 1890, j.p. ;}24, -107. 'M''u(;kel, L., Synd)ohe niy(!ologica', 1869, j). 252. '•' See (juotation in li<'port of Miciiigan Pomological Society for 187:5, i>p. 16, 17. " l'.erkfley, M. .1., Introduction to Cryptogamic liotany, 1857, p. 284, and Outlines of r.rilisli Fungology, Lfindon, 1S60, pp. :)76, 444, tab. 1, lig. (i. '' IJcniifIt, .\. W., and Murray, < Icorgc, .\ Handbook of {"ryptogamic I'.otany, I.on PEACH LEAF CURL: ITS MATURE A>'D TREATMENT. the development of the fuiitru.s liegin.s. .so that in Juno all leaves at the ends of the young branches are curled and deformed and perhaps all the blossom buds fall off. If the more developed leaves at the l)a.se of the 3'oung shoots prove more resistant to the fungous action, then fresh shoots are formed in June, even if not in normal condition, but yet somewhat healthy, so that the tree remains intact. In the more pro- tected hill regions of the vineyards, at the foothills of the Carpathian Mountains, this disease is also troublesome, but less intense than in other parts of the country. Peach leaf curl exists in South Africa, and prolxibly also throughout Algeria and other peach-growing portions of 'the continent. Profes.sor ^lacOwan. of the department of agriculture of Cape Colony, has written of the disease in South Africa, giving his views as to the proper manner of treating the same. ^ He also writes that it is '"a great plague at the Cape." ^ A peach grower of Drysdale. Frere, Xatal. in writing to the Cape Colon}' agricultural department under date of October 81, 1893, .says that he has a good many peach trees of the yellow, white, and St. Helena varieties, and that they are all affected with the discolored and curled-up leaves characteristic of this disease; that several of his neigh- bors are complaining that their peach trees are suffering like his: and that the disea-se seems to be spreading. The young trees were simi- larly affected. ^ Perhaps no foreign country has suffered more from peach leaf curl than New Zealand. Mr. W. M. Maskell, of "Wellington, writes as follows : * ■■ The curly blight has ])een for many years prevalent in this country — so nuich so that whereas in the early days peaches were exceed- ingly luxuriant and fine, they have dwindled to comparatively very sn)all and poor trees and in many i)arts of the eolony almost di«'d out. In the last two or three years the people have been advised to emi)loy remedies, and have done so to sonic extent, so that thcic is a marked improvement in the j)each orchards. * * '^ I can myself recollect, early in the sixties, when the most sj)lendid peaches used to grow wild in the warm iiortheiii distiicts. wheic now scarcely a tree is seen; and thecui"ly blight has been a dicadful curse all over the colony." Australians r«'p(»rt peacii leaf curl among their serious plant dis- eases. In South Australia it "has been known (jiiite twenty years."'* and })rol>ably longer, and occasions consi(leral)le losses in seasons favoring it. The situation is much the same in New South ^\'ale>. ' MacOwan, Prof. P., Agricultural Journal, puhlishtMl by the department of a>;ri«'ultnn' of CajK- CVdony, 1H92, Vol. V, pp. 2.">2, 2n:i. - Ix'tterj;t<.ii, New Ziahiiid, I)c'ceniUT24, l.S«».'i. ^()l>s«TvationH of .Mr. \. Molineiix, jiencnil secretary for the atrrienltural l>iinaii of South .\uftralia, l«'tt«T sc calculations ga\-e a loss to the gi-owers averaging >:^1(>.1»5 per acre for the aci'cage r('port('r c(Mit of the full aresi. This IS (•(|iii\alcrit to about !i^l.t)(» per acre foi' the entire acreage", or about 1 cents \)vr tree. At first thought this may seem high, t)ut this is more ajjpaicnt than real. If one l(»-acr(M)rchar(l loses its cro}) from curl, valued at $150 ])cr aci-c. the loss amounts to ^1. .■"><»(). There may lie ;'>2 other oi'chards of K) acres each all about this orchai'd where not a [)each is lost, yet the average for such a district is the same as that Htat<' a clearer manner of putting the matter than LOSSES FROM THE DISEASE. 21 by placing an average loss for all orchards. The loss may ))e viewed in still anotlun* manner. If an orchardist has grown peaches for 82 years and lost only one crop during that time from leaf curl his loss for the third of a century will average as high as here calculated. There are large sections of the country where curl is scarcely known, as in portions of Texas. For such regions the preceding estimates may appear high. On the other hand, there are other prominent sections of the country devoted to peach culture where these estimated losses will certainly ])e far too low. If the preceding calculations and statements are accepted as fairly representing the situation throughout the country, the annual losses fi'om curl in the United States may be approximated. The Eleventh Census reports the orchards of peach trees in the United States at that time (1889-90) as 507,786 acres, and from replies to our circular we are led to believe that curl was present in 63 per cent of these orchards and that 12 per cent sustained some loss from the disease. Most of the orchards included in the 12 per cent sustained only a slight loss, but a very small percentage sustained a heav}^ loss, some- times amounting to the entire crop. The average loss for the 12 per cent of the orchards is found to amount to $10.95 per acre, or about 10 cents per tree, averaging the trees at 108 per acre. The total acreage of the country being 507,786, the loss should be calculated upon 12 per cent of this, or 213,219 acres, which gives a total esti- mated annual loss from peach leaf curl of $2,335,076. In this estimate no account has been taken of the groat injury to the growth of trees, the injury to nursery stock, the death of young orchard trees, nor the loss to succeeding crops from the reduced number or quality of fruit buds on affected trees. There is also the loss arising from the culti- vation and pruning of unproductive orchards, which, if it could be determined, would probably bring the entire annual loss to the country up to $3,000,000 or more. Since 1898, when the investigation of this disease was undertaken by the writer, a vovy large amount of correspondence has been con- duct(Kl with peach growers in all parts of the Union who have sus- taiiied losses from curl, and this correspondence has resulted in the accumulation of a large number of facts respecting these losses. These data, however, have not l)ecn drawir upon in the above esti- mates, as it might ])e claimed that they were from growers only who have suffered from the disease, and consequently would not fairly represent the industry as a whole — a claim which can not be made against tlu^ circular letter, the basis of tlu^ estimates, which was addressed to peach growers in general in all parts of the United States. In fact there* appears to have been a larger percentage of replies received from sections of th(^ country where curl is scarce than from the more affected portions. CHAPTER II. NATURE OF PEACH LEAF CURL. The study of the nature of plant diseases is intimately linked with the study of plant physiology, and the true science of vegetable pathology is largely, as Ward has defined it, the study of abnormal plnsiologv. (Introduction to Hartig's Text-book of the Diseases of Trees.) These facts become evident when studying the etiology of peach leaf curl and the conditions attendant upon its widespread development. The direct cause of peach leaf cui'l has long been known as a parasitic fungus, Exoascus (Jcforinanx (Berk.) Fuckel, but it is evident from a careful study of the disease that the injurious development of the fungus is distinctly correlated with special physi- ological phenomena of the peach tree itself. These conditions of the tree are in turn dependent upon such external influiMices as tem- perature, the humidity of the soil and atmosphere, and others. Such facts were foreshadowed by the theories advanced by p(>ach grow(M's as to the cause of the disease. Many growers have considered peach leaf curl as tiie direct result of excessive moisture and low tem- perature or sudden changes, and as these physical conditions cer- tainly have an important bearing upon the injurious de\ elopment of the disease, they are considered together with the direct relations of the parasite to its host. However, too nuich stress can not be placed upon the fact that the fungus alone is responsible for the injury to the tree.- Without the parasite not a leaf would curl nor a peach fall on account of this malady — in fact, no such disease would exist. This is shown by the work hereafter detailed. It is fortunate that the direct cause of peach leaf curl is a parasitic fungus rather than unfavorable atmospheric conditions^ for the latter could not be controlled, while the conti'ol of tin; fungus has been found practicable, simple, and inex- pcnsivi!. I'llVSICAI, CONDniONS INFLIiKNCIM! TIIK DISKASK. Tlx' influences of temi)erature, humidity, situation, soil, etc., upon leaf curl are often so well marked that they have frecpiently and in factase. Indeed a Nci'v large ])ercentage of peach growers have maintained, to within the past ten or fifteen years, that sudden changes of tiMuperature occuning in conjunction with west w«>ather are the solo cause of the curling and 22 PHYSICAL CONDITIONS INFLUENCING THE DISEASE. 28 loss of foliage. Notwithstaiidino- the nunil)er of known facts to the contrary, there are even now many growers who retain this idea to the utter and needless loss of their crops. The writer has met men who so firmly believe that leaf curl is due to uncontrollable climatic influences that they would not consider other explanations, bcMng unwilling to visit the orchard, though the crop was being lost through curl and by so doing future crops might have been saved. To gather the experience of peach growers in general respecting the conditions under which leaf curl develops most severely, a circular of inquiry was addressed to several hundred orchardists in November, 181);:>. The replies to some of the questions are presented. Among the inquiries the growers were requested to state if they had observed the disease to be more prevalent after a cold spell in the spring. To this question 97 replies were received, 89 affirmative, 6 negative, and 2 growers said they had observed no difference, which shows that the orchardists are almost unanimous in holding that a cold spell in the spring favors the development of curl. To the second question, as to whether the trees were most affected by curl in a wet or dry season, there w^ere 104 replies. Of these, 78 stated that peach trees were most affected in wet seasons, 8 that the}" were most affected in dr}^ seasons, and 18 that there was no difference. Here again is seen a marked agreement in the replies, a great majority of the growers recognizing that wet years favor the disease. The above-considered conditions — a cold spell in the spring and wet weather — may be explained by stating that such conditions favor, on the one hand, the serious development of the fungus causing the dis- ease, and, on the other, they result in a much greater susceptibility of the tissues of the peach leaves to the attacks of the parasite. Where both cold and rain occur together in the spring, about the time the leaves are pushing, the disease is liable to develop seriously and few varieties can then resist it. The action of wet, cold weather upon the tissues of the peach, making them much more subject to curl than they otherwise would be, has been considered in relation to other plants in a paper by Prof. H. Marshall Ward,^ who sa3'S that irJten the combined effects of tJie physical eiwironment are uiifavorahle to the host, hut not so or are even favorable to the parasite, 'wefind the dtseaxe axKuinhxj a more or less j^ronoimced epidemic chara,cter. He is not here speaking of curl, but the statement holds perfectly true for that disease. A cold, w^et spell succeeding warm spring weather, has a tendency to saturate and soften the tissues of the peach, as in the case of other plants. The sudden checking of active transpiration, due to lowered temperature and saturated atmosphere, soon results in the tissues of the plant being suffused Avith water. "'The stomata," as Ward puts tt, ''are nearly ^Wanl, Prof. W. Marshall, The Relations between Host and Parasite in Certain Diseases of Plants, Croonian Lecture, IVoc. Roy. Soc, Vol. XLVII, No. 290. 24 PEACH LEAF CURL: ITS NATURE AND TREATMENT. closed, the cell walls bounding the intercellular passages and the air in the passages themselves arc thoroughly saturated with water and aqueous vapor, respectiveh', and the movements of gases must be retarded accordingly; turgescence is promoted, and the water contents accumulate to a maximum, owing to the disturbance of equilibrium between the amounts absorljed by the active roots in the relatively warm soil and those passing off into the cold, damp air; much more water is absorbed by the roots in the relatively warm soil than passes off as vapor in equal periods of time." Further than this, Ward states that "'the low temperature, feeble light, and partialh'^ blocked ven- tilation system have for a consequence a depression of respiratory activity and the absorption of oxygen genei-alh'."' This must give a lowered vitality and an accumulation of organic acids. The reduced light also leads to a decided reduction in the assimilative power of the leaves. "The turgid condition of the cells, and the diminished inten sity of the light," Ward says, "will favor growth." If this takes place, "the tendency will be for the very watery cell walls to Ijecome relatively thinner than usual, as well as watery, because the ill-nourished protoplasm does not add to the substance of the walls in proport^pn. This being so. we have the case of thinner, more watery cell walls acting as the only mechanical protection between a possible fungus and the cell contents." It is generally known that the conditions of moisture and shade, which are above shown as making the tissues of a host plant more tender and watery (more subject to fungous attacks), are also the conditions most favorable to the development of fungi. This holds ec^ually as good for J'J.rod.scus defoi'man.'t as for other forms. In speak- ing of these conditions in relation to a fungus known as Botrijtis^ Pro- fessor Ward gives some generalizations equally applicable to Kiy>ai inches at Portland. From Albany to Portland the peach industry has been greatly injur<>d by curl, and on its account many growers in this region have considered peach culture a failure. Curl, it seems, was introduced into the central part of the Willamette Valley. Marion County, nearly half a century ago. Prior to that time the peach was successfully grown in that region in spit(> of the humidity of tin; climate. In the Patent Oftice Report for 1855, p. 298, there is a statement of the situation in Polk and ^Marion comities from 1852 to 1855. This statement was from Mr. Amos Harry, of FaiiM \'idlev, Polk County, Oreg., and is of special interest in this connection. Mr. Harry says: "The peach in this county has been attectcd with a disease known as the 'curled leaf,' which threatens to destroy the trees. It made its ai)p(>ai-ance at Mill Creek, in Marion County, in 1852, and extended considerably on that side of tli(> rixcr (Willamette River) in 1853, but had reached most parts of tho valley in 1S51-55. Some trees seem to escape it much more than others, but if tlu' malady irici-cascs for two years to come as it has foi* two ])ast, 1 fear we shall conif entirely short of this delicious t'l-uit. Some think It is owing to cold, wet w<'ather. and reconnnend shortening all the limbs as a remedy, and some experiments seem to taxor this id(>a. Others think it i> produced by an insect, and that no icniedy will save the tree- iird<» it can l»e applied to llie whole siiit'ace of the leaves." PHYSICAL CONDITIONS INFLUENCING THE DISEASE. 29 The rainfall at Portland, as already said, is 41> inches, and eurl is com- monly prevalent and severe. At Umatilla, east of the Cascade jMoiui- tains, l)ut a])out the same distance north as Portland, the rainfall is only 10 inches, while on that side of the mountains the peach industry is extensive and everywhere prosperous, leaf curl beino' nuich less prevalent and of secondary importance. This shows that it is not the distance north and the consecjuent lower temperature which makes curl more severe at Portland than at Los Angeles for instance, T)ut that it is the excess of rainfall, for at the east of the mountains, near Umatilla, the temperature goes equally as low or lower than at Portland, and curl is of little importance there. In the Puget Sound region peach culture has never developed extensively, the general prevalence of curl and its injurious action being one of the chief reasons. The rainfall is 50 inches at Seattle and 56 inches at Ol3"mpia. It is only 7 inches at Kennewick and 9 inches at Ellensburg, on the east side of the Cas- cade Range. The peach orchards of North Yakima and neighl)oring sections on the east side of the Cascades and near Ellensburg, where this rainfall is taken, are noted for their extent, thrift, and general health, and curl is not a serious trouble. This case is parallel with that of Portland, already considered. The rainfall at the west of the mountains is 60 to 56 inches or more, while at the east it is only 7 to 9 inches. In the former region peach growing is not listed ])v the Washington Board of Horticulture as one of the horticultural indus- tries, but in the latter region the peach is a leading fruit, ))eing extensiveh' and successfully grown. The winter temperature east of the mountains should range fully as low Avhere the peaches are grown as at the west of the range. The contrast in peach culture in the two situations results from the difference of rainfall, and the heavy rain- fall at the west of the Cascades results in a development of curl almost prohibitive to peach growing.^ In replying to a circular letter sent to the peach growers of Mary- land, Novem})er, 1893, Mr. T. C. Staj'ton, of Queen Anne, makes some statements which bear directly on the matter here considered and are of nuich interest as resulting from personal observation. After speak- ing of the conditions in Maryland, Mr. Stay ton sa3's: ''I was in Washington State during the months of April, May, June, etc., this year, and I find they can not grow peach trees west of the Cascade Mountains or in western Washington, as that part of the State is called, as that is a ver}' wet part of our country.'' He adds that this was especially true in 1893, and continues: "Ahout all the .young trees that had been planted in that part of the State died from curl leaf, or so nearly so that they were worthless, ])ut over in eastern Washington I did not notice any curl leaf, the climate being dry." 'For a full and accurate account of the rainfall conditions i)revailin,ries of hca\ \ dews, 7 (iiat it is W(^-si^ or fungi liear- ing their spores in cases or asci. The classification of the Eroaxcen', which now la^vs greatest claim to scientific permanence is that outlined in the recent writings of Sade- beck, who has given careful study to these forms. ^ Of the five genera which he recognizes, only the last directl}^ concerns us at this time, as it is to this genus {Exoascus) that the peach curl fungus belongs, as well as numerous other species injurious to horti- culture. In considering this genus Sadebeck ^ has grouped thirty of its species according to certain characters of development. He recog- nizes the following characters of the genus: ExoAscus Fuckel. A. The mycelium is perennial in the iimer tissues of the axial organs. a. The development of the hymenium occurs only in the floral leaves of the ' host i)lant. Eight species. h. The development of the hymenium occurs only in the foliage leaves of the host plant. Seven species, including E. dcfonnaui^.^ c. The development of the hymenium occurs upon the leaves as well as upon the fruits. One species. B. The mycelium is perennial in the l)uds of the liost plant and develops only subcuticularly in the leaves. 'Sadebeck, Dr. R., Die parasitischen Exoasceen, Hamburg, 1893, p. 43. Sadebeck recognizes five genera in the Exuasccrv, which he arranges and character- izes in the following manner: EXOASCEjE : Ascoinycetes whose asci are not xinited in a fruit body. A. The asci arise as swellings at the end of the branches of the mycelial threads. 1. Eiidomyrcs Tulasne. Four-spored asci, no conidia within the same; the sterile threads develop chlamydospores and oidia. 2. Maymmclld Sadebeck. Parasitic. Asci with more than four spores; usually conidia formations in the ascus. Oidia and chlamydospores wanting. B. The asci take their origin fi'om a more or less loose hymenium. 3. Ascocortk'inm Bref. Saprophytic on bark. The ascus layers are arranged in a loose hymenium upon the mycelium. 4. Taphrbia Fries. Parasitic. Without perennial mycelium. In the formation of the ascogenous cells differentiations of material occur. Forming leaf spots. 5. Exodscufi Fuckel. Parasitic. With i)erennial mycelium. In the formation of the asci no differentiations of material appear. The subcuticular mycelium changes directly to ascogenous (xdls. Causing sprout deformations. ■^Sade])eck, Dr. R., Einige neue Beol)achtungen und kritisclie IJemerkungen iil)er die Exoascaceaj, pp. 277,278, reprint from den Ber. d. deutsch. hot. Ges., 1895, Bd. XIII. "Dr. von Derschau has described the occasional fruiting of E.mnxrus tfrformans in the blossoms of the peach. The figures given by this author do not show the nor- mal development of ascogenous cells in the blossoms which are so conunon in the leaf Vjlade of the peach. His figures show the asci as arising from lateral l)ranches of a continuous mycelial hypiia, and this mycelium is situateil beneath the epidermal cells instead of between the cuticle and t-pideruiis (Landw. Jahrb., Berlin, 1897, pp. 897-901, and Table XLI). 32 PEACH LEAF CURL: ITS NATUEE AND TREATMENT. ti. The develoimu'iit of the hynienimu occurs only in the Horal leaves of the host plant. Three sjiecies. h. The develoiiiuent of the hynieniuni oix-nrs only upon tlie foHajre leaves. Ten species. c. The resting mycelium extends intercellularly in the deformations of the leaves. One siiecies. It may be seen under A 7) of this arrang-enient that Kromcus defor- m< is said to possess perennial mycelium, inhabiting the inner tis- sues of the axial organs, and that the development of the hymenium occurs only in the foliage leaves of the host plant. As will be seen in another part of this l)ulletin, it is perhaps a perennial nature of the mycelium of E. deformans which makes it difficult to thoroughly rid an orchard of curl by means of spray treatment, but this matter requires further careful consideration. The synonymy of Exoascus deformans (Berk.) Fuckel has been given b}' numerous writers. Sadebeck^ gives it as follows: Aacomyces deformarnf Berk. Intro, to Cryptogamic Botany, 1857, \i. 284. AscoKporiiDn (h'foDiiauH Berk. Outlines, 1860, p. 449. T(q}hrhm deformans Tul. Ann. Sci. Nat., 1866, V. Ser., t. 5., p. 128. Exoascus deformans Fuckel. («) Persicx Fuck. Symbola? Micolog., 1869, }>. 252. This fungus has been very commonly observed and frc(j[uently desci'ibed ])v ])otanists since Berkeley called attention to it in 1857. It has thus been known as the cause of curl for a little less than half a centur3\ The peculiar behavior of peach foliage under its action has })een observed by horticulturists, however, for a nnich longer time. The disease was well descri})ed in England in the early part of the present century. In spite of the very conmion appearance of Exoasous deformmis upon peach foliage in peach-growing countries, the descriptive litera- ture relating to its life history is not free from conflicting statements. Several species of Exoasceo} have been confounded with this species in some instances, and subseqiuMit writers have perpetiiat(>d the confusion. Some eai'liei' wi'itrrs believed this species inhabits a considerable lunnbci" of host plants, tluis resulting in tlu^ (lescrij)ti()ii and disti'ibu- tioii of sciveral distinct species as Exod.scu.'^ dcfornttiits. To avoid such confusion it would 1)c best to confine remarks upon this species to the fungus as it develo])s upon the peacli {Pnimi.s perslca L.). which if ii(»1 its only host, is certainly its most coiiunon one. At least two modes of infection of the peach 1 i'C(> by Exiklscu.^ dij'oriiKiiis ai'c said to exist (1) by means of perennial mycelium, and (2) by means of the s))<)rcs of the fungus. SadebecU ' is authority for the stat(Mncnt, that the niyceliiini winters o\er in the youngest poilionsof the one-yeaioid Inanchesof the host 'Sadeheck, i)r. II., Dif parasilisclicn I'lxoasceeii, llamlmig, ls;i;'>, p. 53. ^Jdeni, I.e. THE IlSrFECTION OF THE HOST. 33 plant, and may be seen in the primary cortex, in the medulla, and in the medullar}^ rays of the first shoots of each period of \egetation, but has not been o])served in the soft bast. With the beginning of the new season of growth the mycelium, according to Sadebeck, extends into the leaves of the young shoots, penetrates first the inner tissue of the leaves, and finally progresses to the development of the subcuticular hymenium. From what foundation of experimentation Sadebeck has arrived at these views respecting this particular species, I am unable to state, but he has given the outlines of his investigations upon other species.' The facts given b}^ De Bary ''' can not be cited here, for this work was done upon the Exoascus infesting the cherry tree, and which is now considered to be distinct from E. deformans. The general acceptance of the view that spring infection of the peach foliage is largely due to the extension of the internal perennial myce- lium into the new shoots and leav^es from the shoots of the previous summer, has probably considerabh^ retarded the progress of prevent- ive treatment. Pathologists have thought it improbable that any considerable amovmt of disease could be prevented after a tree Avas once generally aflected, as the perennial mycelium, being internal, could not be readily reached by external sprays. Prillieux,'' writing in 1872, advises the gathering of the diseased leaves and the cutting away and burning of the diseased branches. Frank* has made like recommendations in both editions of his work on plant diseases. Assmning the mycelium to be perennial, he says that the curing of the disease might be aimed at through cutting back of the diseased branches and the prevention through <|uick removal of the diseased leaves. Winter' suggests a somewhat similar line of treatment, with the additional recommendation that the trees be protected from rain during the unfolding of the leaves. Dr. Cobb," as late as 1892, after speaking of the perennial mycelium of this fungus, discusses pre- ventive and curative measures, such as the destruction of diseased leaves, prunings, etc., while in the more severe cases he says the sooner the trees are cut down and burned the better it will be for the peach industr3^ ^Sadebeck, Dr. R., Die parasitischen Exoasceen, Hamburg, 1893, pp. 24-28. — Das perennirende Mycel der Exoascus- Arteii. '■'De Bary, A., Com. Mor. and Biol, of the Fungi, Mycetozoa, and Bacteria, Eng- lish edition, 1887, p. 26(5. •'rrillieux, Ed., Bui. de la soc. hot. de France, 1872, T. XIX, p. 2m * Frank, Dr. A. B., Die Krankheiten der Pflanzen, Breslau, 1881, Part II, p. 526; second edition, 1896, Vol. II, p. 250. ^Winter, Dr. Georg, Die durch Pilze verursachten Kranklieiten der Kulturge- wilchse, Leipzig, 1878, ]>. 47. «Cobb, Dr. N. A., Tlie Agricultural (iazette, Sydney, New Soutli Wales, IS92, Vol. Ill, pp. 1001-1004. 19093— No. 20 3 34 PEACH LEAF CURL". ITS NATURE AND TREATMENT. Relative to the use of fungicides the same writer sa^'s: "These treatments are of doubtful value as far as the curl is concerned, and were it not that they are viseful in other ways I would not mention them." It is evident that these views are the result of Dr. Cobb's belief that the perennial uncelium is responsible for the major portion of the spring infection of the tree. The writings of others to the same effect could be cited, but the views of the workers already named are sufficient to show that their recommendations for treatment have been based upon the hypothesis that the spring infection could not be prevented by treatment with fungicides, as it arose mainly from in- ternal mycelium rather than from the germination of external spores. That this view has held back the preventive treatment of the disease, as already claimed, can not be doubted, and that a perennial mycelium is not responsible for more than a very small percentage of the spring infections seems evident from the results of the present experi- ments; in fact it may even be questioned if such infection takes place except under exceptionally favorable conditions. Our experiments have demonstrated that as high as 98 per cent of infections may be prevented b}^ a single thorough application of a suitable fungicide. This is as high a percentage of control as is often obtained in the treatment of fungous diseases where no pei'ennial mycelium exists, aJid it seeuis probable that the infections by this means luay not com- monly exceed 5 per cent of each spring's infections. Were this not the case we would be forced to assume that the spray has a direct effect upon the hi))ernating mycelium, which certainly would be unusual and scarceh' to be expected. The second mode of spring infection — that by means of spores — is probably much more general and important in this disease than has been supposed. That 90 to 98 per cent of the infections of tlie tree are i)revented by a single spraying suggests that at least such ])ei-cent- ag(? of the infections is by means of spores. The mycelium of Kroa.sciis difoi'iiKinx as found in the ])(>acii. shows great differences in the form and appearance of its hyplne. These differences depend ui)on the stage of development of the funglis and the various functions of the mycelium. Th(> writer recognizes three types of hyphte, which may he teinied vegetative, distributive, and fruiting. 'I'he vegetative hypha* are found most connnonly in the leaf ])ai(>n- chyma, but are also met with in the leaf stalk and cortical ])ai'eiichyma {*f l)adly diseased and distorted l)i'anches. These hyphic may l>e most distinctly seen, and ar«> most highly developed, in infested leaves which have not yet foi'ined the hyiueniuni of ascogenous cells, but in which the parasite has been present a sutKcient time to (Mitirely altei- the (•hara<-ter of the palisade tissue and caus<' the loss of the chlorophyll. In the leaf l)lade the palisade tissue first shows the serious action of BULL. 20, DIV. VEQ. PHYS. a PATH., U. S. OEPT. AGRICULTURE. MrCELIUM OF EXOASOUS DCPORMANS, THE FUNOUS 0AU8INQ PEACH LEAF CURL. A.ll<,niik I'll, Mil, ll,illli,>„i„ DESCRIPTION OF PLATE II. Mycelium uf Exua-scus deformans (600/1) . Figs. 1 and 2, normal vegetative hypha?, as ff)imd in the leaf parenchyma, showing characteristic septation, modes of branch- ing, etc. ; figs. 3, 4, and 5, usual type of distributive hypha^ found in swollen branches in the cortical parenchyma and medulla; figs. 6, 7, 8, and 9, fruiting hyphie, show^- ing successive stages in the development of ascogenous cells from the subcuticular mycelium (6) to the half-formed ascogenous cells (9). (See PL III for further stages in the development of the ascogenous cells and asci. ) THE MYCELIUM OF THE FUNGUS. 35 the vegetative hyphae, which are usually found somewhat later among the cells of the spongy parenchjMiia, below the vascular network. The loss of chlorophyll from the two classes of leaf parenchyma commonly preserves the order here given. The form of the vegetative hj^phae is ver}^ irregular, and their elements, or cell members, are often of dif- ferent size, length, and shape. The cells vary greatl}" in diameter from one end to the other, are frequently much curved and twisted, and oftentimes appear triangular in cross section. The branches may arise from greatly enlarged triangular bifurcations, or in other instances directly from the sides of the cells. These vegetative hj^phje are all intercellular so far as observed, but are commonly found adher- ing closely to the cell walls of the host, frequently wrapping about the parenchyma cells. The walls of the hyphte are semitransparent but firm, commonly having a slight yellowish cast. The septa present peculiar characters. Two adjoining cells of a hypha have the appear- ance of l)eing separately closed at the end and united with each other by means of an intervening plate, which if it should be dissolved or lost would leave the cells separated l)ut closed. These peculiar septa are remarkal)ly refractive and characteristic. They are well shown in the drawings of Sadebeck (Die parasitischen Exoasceen, Hamburg, 1893, Tab. II, figs. 7, 8). The predominating characters of the vegetative hyphre are shown in PI. II, tigs. 1 and 2, of this bulletin. The hypha? there shown were carefully separated from the leaf parenchyma and drawn under the camera. The vegetative hyphre of the l)ranch are much like those of the leaf, and have been seen most commonly among the looser parenchj^ma cells of the cortex just exterior to the bast fiber bundles. Thus far they have never l)een found by the writer in the caml)ial tissues. Sadebeck states that the mycelium has been found in the pith and medullary rays. The distributive hyphae are shown in PI. II, figs. 3. 4, and 5. They have been found by the writer in the tissue lying close beneath the epidermal cells of diseased peach twigs, and in great al)undance in the pith. They are occasionally found in groups of several hypha^ but slightly separated from each other and following a course parallel to the longitudinal axis of the shoot. The cells composing these hypha? are much longer than either the vegetative or the fruiting forms, while they are nearly straight and of more uniform diameter. The septa are characteristic of those found in the other forms of the mycelium of this fungus. Such distributive hypha^ have l)een followed for some little distance in the swollen portions of the peach twig, and the name has been given them from their apparent function of spreading the fungus in the branch. Such hypha- bi-anch ])y bifurcation, the branches commonly assuming a course parallel to the parent hvpha and the direction of the peach limb. 36 PEACH LEAF CUELt ITS NATUKE AND TREATMENT. The fruiting hyphse have been seen to arise in Exoasaus deforma/ns from the vegetativ^e hyphse after the latter have become well developed in the parenchyma of the leaf. Large, well-nourished vegetative hypha? commonly develop just below the epidermal cells of the upper leaf surface.^ From these hyphse arise branches which penetrate between the cells of the epidermis, and press themselves between the epidermis and the cuticle. Such hyphje may be seen both in section and surface view. These subcuticular hyphaj now branch freeh', and follow with more or less regularity the triangular space formed by the juncture of two adjoining and somewhat rounded epidermal cells with the cuticle. This is presumably the line of least resistance to the advance of the hj'phre. By opening and following these channels the mycelium assumes the outlines of a quite uniform network l)eneath the cuticle. While this manner of following the line of juncture of adjoin- ing epidermal cells with the cuticle is conmion, it is not invariably the practice of the fungus, cases occurring where apparently no such agreement exists. Series of straight and parallel hyphje, at regular distances apart, are sometimes met Avith beneath the cuticle as the precursors of the hj^menial layer. These send off lateral branches on either side, which ])y enlarging, branching, and curving eventually occupy most of the surface of the epidermis between the main hypha\ It is probable that the path followed by the first subcuticular hypha? depends largely upon the firmness with which the cuticle is attached to the epidermal cells, and which ma}' largch' depend upon the amount of water in the tissues and upon the age and rapidity of their growth. With the leaf tissues full of water and making a rapid groAvth, the hypha' could naturally pursue a more direct course beneath the cuti- cle than under contrar}' conditions. After the esta])lishment of a much-))raiiched filamentous ni^twork of subcuticular hyplnv. the cells of which are usually slender, of medium length, thin-walled, and of comparative!}' uniform diameter (PI. II, fig. 6), these cells begin to distend, and are shortened ])V the formation of new transverse septa (PI. II, fig. 7, and PI. Ill, fig. 22). About this time all septa become much more distinct. At a later stage the cells become still more distended and subspherical (PI. II, fig. 8). As these enlarged cells ' Miss E. L. Knowles (Bot. Gaz., Vol. XII, No. 9, p. 217) has called atten- tion to th(! fact that Winter's statement that "the asci break throujili tiie lower si, and Krank. Kultur-dewiichse, I^'ipzijr, 1878, p. 47). Winter is not alone in statinji that thea.«., Notes on the; (Jenus Taphrina, Ann. Bot., Nov., 1887, Vol. 1, No. 11, J). H)8). Atkinson also says: "The asci are developed on both surfaces of the leaf" (Atkinson, (Jeo. F., I>eaf Curl and I'luni PiK-kets, ('ornell Ajrr. Kxp. Sta. I?ull. No. 7.'{, 1894, p. .'{25). These and other like statements have probably arisen from a Htudy of other foliap- than that of the j)each, and of other species of Exoaaciis, and qave been i)er[>etMat<'ward. Fig. 21, the pushing of a forming ascus tlirough the leaf cuticle (()00/l). Figs. 22 to 27 (600/1), various stages in the fonnation of ascogenous cells from subcuticular mycelium. (P'or several early stages in this process see PI. II, figs, (i to 0). Figs. 28 to 30 (000/1) show fully developed asc(jgenous cells as seen from above. BULL. 20, DIV. VEQ. PHYS. & PATH., U. S. DEPT. AGRICULTURE. PLATE in. NVwt.in II Picni- ;«1 iwit H':l FRUITING STAGES OF EXOASCUS DEFORMANS. i.llurn< r».l.iil>. Ka THE FEtTlTlNG HABITS OF THE FUNGUS. 37 spread out between the epidermal cells of the leaf and the cuti- cle they are much distorted, curved, and lobed, the branches and lobes eventually tilling, in a quite uniform and continuous manner the entire space between the elevated cuticle and epidermis so that a more or less perfect and continuous hymenial layer of 'ascooonous cells is formed (PI. II, fig. 9; PI. Ill, %s. 23, 24, 25, 26, and 27). At this time the cells become well rounded and heavv-walled, and thev may or may not become loosened and separated^ from each otheT (PI. Ill, tigs. 28-30). These are now the fullv developed ascoge- nous cells of the hymenium, and they are fully stored with nutritive materials for the development of the asci. In their compact, continu- ous, and rounded condition they resemble, when viewed from the sur- face, the stones in the pavement of an old Roman highway. The various phases of the development of the hymenium of ascog- enous cells may often be observed at one time in a single infected leaf. The center of a swollen spot frequently shows the fully devel- oped hymenium, while at the margin of the spot the first filamentous hypha' are just spreading beneath the cuticle. In such instances nearly all stages in the development of the ascogenous cells may be studied in a single well-prepared specimen. The development of a subcuticular hymenium has been observed in the petiole as well as in the ])lade of the leaf. The formation of the asci from the fully developed ascogenous cells has been carefully followed in the study of a large number of prepara- tions. Thus far no sexual phenomena have been observed in connec- tion with the formation of the ascogenous cells or with the develop- ment of the asci. As already said, the walls of the ascogenous cells are heavy. The early steps in the development of the asci from these cells (the development of a papilla-like elevation on the upper surface of the cells) cause the rupture or dissolution of the heavy wall where the elevation occurs. The phenomenon is that of the germination of a heavy-walled spore, or, perhaps, more properly, the outgrowth or prolongation of an endospore through the rupture of the epispore (PI. Ill, figs. 17, 18, etc.). The fact to be noted is the perfect rest- ing condition into which the ascogenous cells may pass before the development of the ascus, as shown by the marked delimitation between the thin wall of the forming ascus and the heavy wall of the ascogenous cell. The entire isolation of single ascogenous cells or groups of cells from all sources of vegetative supply indicates that the ascus is entirely dependent for its nourishment upon the stored materials of the cell from which it arises. The pushing of the ascus after the com- plete development of the ascogenous cell instead of in direct con- tinuation of the development of the latter, also points to a probable cessation and renewal of the reproductive activity of the ascogenous cell. 38 PEACH LEAF CURL: ITS NATURE AND TREATMENT. In view of these facts, it seems possible that the asoogiMious cells may be capable of enduring, under especiall}^ favorable conditions, a resting period of considerable time. Such resting ascogenous cells have been sought for upon the swollen branches of the peach, how- ever, without success. Further research along this line is desirable. As the fungus is already known to fruit upon the ))lade and petiole of the leaf and upon the blossom, and a vegetative mycelium is found growing thriftily in the swollen branches, there seems to be no good reason why the parasite may not fi'uit upon the infected twigs. The perpendicular growth of the developing asci in the leaf soon rup- tures or pierces the cuticle, and where largo numbers of asci develop at the same time the cuticle is lifted, torn, and lost, the asci forming a more or less continuous plush-like surface growth. Isolated asci press through the cuticle so as to form separate perforations (PI. Ill, fig. 21). The contents of the forming ascus are finel}^ granular, and as the ascus elongates these contents crowd into the upper portion and a septum is formed across the basal part in such a manner as to cut off the now emptied ascogenous cell as a stalk cell for the ascus (PI. Ill, tig. 15). When fully developed the asci are usually broader at the top than at the base, and often somewhat clavate in form. A series of asci measured \aried in length from 3-1: to -i-i /<, the average being 38 M '■, the width of the asci ranged from 10 to 12 yw, and the height of the stalk cells varied from 8 to 13 ;^, the average l)eing slightly over 10 m (PI- III, tigs. 1-18). The formation of the ascospores in Exoascus deforrrums has not been carefully studied by th(^ writer. Sadebec^k has shown, however, for E. turg'iduH^ that mitotic nuclear division occurs in the ascus in connec- tion with spore foi'mation (Untersuch. iiber die Pilzgattuiig Exoas- cus, Hamburg, 1884, PI. Ill, tig. 20). The ascospores devek)ped in the asci of E. deformans varj^ in number from 3 to 8, the latter being the full and typical numl)er. When matures they are surrounded l)v a moderately tirm cellulose wall, which is rather inconspicuous, owing to its transparency. The spores are usually somewhat oval in form, being longer than broad, but occasionally some are seen which appear nearly or (^uite spherical. Fresh ascospores sometimes show distinct nuclear phenomena. This has been obscu'ved with spores still within the ascus, as well as in many which have escaped. The nucleated appearance seems less common in budding or germinating ascospores than in those in a resting condition (PI. IV, tigs. I, 2, 3, 4, and 10). Th(^ average; length of the ascospores measuivd was T/f /<, the length vaiying from <> to 9 a*, and the average width was (>,•',, /<. varying from 5 to 7 }i. The ascospores escape from the ascus through an ajjical I'up- tuic of the latter. (icrmiriation of the ascospores has been observiMl by the writci" to pi'oce(;d in two ways: (1) \\\ means of budding or conidia formation; (2) by nutans of stocky germ (iil»t's. often one branched and rcsiMnbling promycelia. DESCRIPTION OF PLATE IV. Germination of tiie ascosporet- anj>hi}t(>r(i hifextaiix. Thus far efforts to induce tilamentous gerniiiiation of the bud eonidia or of the ascospores of ExoancuH defonnmis in culture media have proved unsuccessful. Brefeld has worked with this problem for months, and the writer has frequentl}' attempted to ol)tain this form of germination.' Budding occurs, as alreadj^ indicated, quit(^ readily in various luitrient solutions, and short promycelia from the ascospores have been found in some cultures. In nearly if not all cases, however, the ascospores showing promycelia or short m^^celial germination have .shown that this germination occurred under natural conditions upon the peach leaf, the germinated spores l)eing transferred from the leaf to the culture^ in preparing the latter. It may be added here that the ))ud eonidia are also formed in vast numbers upon the surface of the infested leaf after the maturing of the ascospores. It i-; largely these eonidia which give the infested leaf the marked white appearance conmionly observed at the height of the disease. The leaf appears as if covered with flour or a heavy white bloom. RELATIONS OF THP: FITNGUS TO THE HOST. Under a previous heading in this chapter the physical conditions which influence the serious development of peach leaf curl liave been considered in accordance with the light which we now have relative to such influences, and there remain to l)e taken up at this time the more intimate and direct relations of the host and parasite. These relations includ(> the action of the fungus upon the cell contents, the cell walls, and the cellular tissues of the host; the pi'obal)le mode of infection and the spread of the parasite within the tissues; the wintering of the fungu.s upon the tree; etc. 'A very considerable number of cultural experiments have been tried. The cul- tureH oi asco.sporeH and eonidia liave been subjected to temperatures nnich below tlie freeziiifr i)oint and to various degrees of heat in the thermostat. Sudden changes of tcmjK'rature have l)een tried. Increased and diminished amoimtsof o.xygen, as con- trasted with that of the normal atmosphere, have been tested. Even a chamber tilled witli nearly ])ure oxygen has produced no ai)parent effect. Water from various sources, such as rain water, dew, ice water, distilled water, tap water, et(;., has been testcfl. Solutions of tlie various sugars, malt extract, sterilized beer, plum extract, etc., were tried. Hanging drop cultures of various nutrient media and plate cultures of potato-peptone-sugar gelatin have not shown germination. Drojjs of variou- nutrient solutions placed upon newly forming leaves dissected from unopened peacii buds and thes(! lield in moist chambers have given oidy negative results. The same is true for peach i)itH brought near to germination and the cotyledons treated with a weak solution of diastase, the spores jilaced between theui and held at various tem- I)(Tatur<'s in moist ehamlM'rs. Sections of such cotyledons with spores placed upon them were also prepared in moist ciiambers. A brief treatment <>f tiie spores with ellii-r was tested without bringing about germination. I'rillieiix states that attempts to artilicially infect the leaves or shoots luivc not thus far succerde.i (>hil. .1. I'lantes Agr., Vol. 1, p. :{!«>). RELATIONS OF THE FUNGUS TO THE HOST. 41 As al ready indicated, the writer's work witli sprays seemed to siiow that not more than a small percentage of each year's infections ordinarily arise from a perennial mycelium. In the Lovell orchard, where the personally conducted work was carried out. it would appear that not to exceed 2 to 3 per cent of the infections could have arisen from that cause. On the other hand it would seem that at least 95 per cent of the infections arose from spores, for, as already stated, H5 to !t8 per cent of the spring infections could be prevented ])y a single spraying, and this was actually accomplished where the spraying was done with sufficient thoroughness. It is believed by the writer, however, that these percentages wnll vary within moderate limits iji diti'erent locali- ties, with different varieties, and in different seasons. The following observations will explain these views. The mycelium of diseased leaves is found to l)e comKH-tcKl through the leaf petiole with the nwcelium of the infected limb. From the writings of Sadebeck and many others it might ])e supposed that the leaves were infected from the perennial mycelium in a majority of cases, and that the mycelium met with in the petiole of the leaf origi- nated from the perennial mycelium of the ])ranch. That such spring infection really occurs from the wintering mycelium of the branch should perhaps be admitted, but that such is the common mode of infection of the leaves is certainly doubtful. The writer's studies have shown that the mycelium in the branch close to a cluster of infected leaves diminishes in amount as it passes upward or downward in the branch from such leaves. This fact is as obvious from microscopic studies of the infested tissues as from the external hypertrophies observable to the e^^e. A macroscopic examination of diseased and swollen branches will show that the enlarged parts may extend upward or downward along the branch from the ])ase of the petioles of the leaves, which seem to represent the center of infection. In a majority of cases these swollen ridges terminate before reaching another leaf bud, though in some instances they are seen to extend along the })ranch throughout the entire length of one or more internodes. and in such cases it is fair to suppose that the mycelium may have infected the young leaves of a second or third bud in its course. It should be remem))ered, however, that this mycelium, in a great majority of instances, indicates no connection with a previous year's mycelial growth, but has evidently just entered the branch from one or more infected leaves. The microscopic evidence supports these conclusions, which are, to some extent, in harmony with B.'nton's observations, to be hereinafter considered, but the writer is scarcely prepared to admit the large percentage of spring infections arising from new myc(>lium entering the branch which the observations of that writer schmu to imply. ^ The microscope shows that the hyph.v whicli ])ass away from 'Pacilic- Knral Press, Aug. 2, 1890, p. 88. 42 PEACH LEAF CURL: ITS NATURE AND TREATMENT. the ba.se of tlie leaf petiole gradually deer(>ase in ininibers as they recede from the leaves, and the}' appear to l)e wholly lost at a short distance from the point of entrance into the shoot. As a rule, little or no niyccliuinhas been found extendino- more than 1 or 2 inches })e3'ond a point where external macroscopic evidence of disease exists. The preceding facts lead to the belief that where mycelial infection of foliage takes plac(^ from the branch it is usually done in the spring from hypha^ arising from spore-infected leaves of the same season, and that this occurs only in comparatively few instances or in bad cases of disease. They also indicate that living perennial nn'celium which succeeds in accomplishing spring infection, is comparatively rare. Badly infested and swollen branches are apt to die and dry out, tluis affording no living tissue for the support of the infesting myce- lium. Such branches, even if living until the following spring, are not apt to produce much growth, and frequently produce none what- ever. Furthermore, the badly swollen mycelium-infested branches are comparatively few, and it is believed that the infested winter buds of these l)ranches very rarely exceed 2 to 3 per cent of the total num- ber of ))uds upon the tree. Most branches appear to suffer from the disease only in an indirect manner, that is, by the fall of attected foliage. It seems probable to the writer, therefore, that the swollen })ranches, in which the swelling is apparent to the eye, constitute the trne and almost exclusive home of the perennial mycelium, and there- fore supply the only possible source of spring infection by the win- tering hyphai, and consequently the only source of infection not (•ontrollal)le by sprays. This is in harmony with the results of wide- spread orchard treatment. All ))ut 2 to 3 per cent of infections have been prevented by a single spraying. (See the results of work on half-s])rayed trees.) That such spraying did not prevent the si)read of the mycelium in the inner tissues of the host is shown by the fact that when it is delayed until the leaves have fairly started and hav(> become infested, the treatment is inefl'ective and the disease will con- tiruie to develop and both foliage and crop may be lost. It is not the checking of the spread of the my('(>liuin fioni tiie bi-ancli to the new leaves, therefoi-e, that results from spraying, but the i)r(>v(Mition of the early spore infections from without; and as all but 2 to 3 ])er cent of the year's infections may he thus prevented, all of such infection^ UHist be considei'ed as arising from s])oj'es. Tln" limitation of tlie peremiial myceliiun of Krod.^cu.^ nriaiis to the swollen l)raM(hes or branch parts, as here held, is in harmony with ohser\('ss tiian normally, some of them l)eing heavier and others lighter than in healthy cells. Tratisverse and longitudijial sections of swollen peach twigs show that the pith cells are greatly injured along the course of the infesting mycelium. The location of the mycelium may often l)e detected by treating transvei'se sections with Bismark l)r()wn, tiie infested nu^dul- lary tissue taking less stain than that not harboring the fungus. The walls of the healthy cells of the medulla become reddish brown, while (hose of the infested tissue assume scarcely more than a light yellow or yellowish brown. The cells of the infested tissue are also much more angular and irregular than those in which the mycelium does not exist, while in some instances the cells c()lla]>se. The action of /'/. (leforiiuni^ on the tissues of the peach leaf has been (considered by ditterent writers, as rrillieux,' Knowles," and others. 'IVilli<(KCut< defor- vimis of the peach,'* l)ut made no recommendations for its treatment. In 1885, however, he again spoke of the plum pocket disease and pointed out that it can not be removed except by severe cutting back of the new and old wood of the affected trees.* In 1880-81 Frank, in the second volume of the first edition of his work on plant diseases, rec- ominends the cutting back of tlie twigs as a cure for leaf curl, and the (|ui