Cultura Del Trabajo

Mycological Society of America
Nineteen Years of Culture Work Author(s): J. C. Arthur Reviewed work(s): Source: Mycologia, Vol. 13, No. 1 (Jan., 1921), pp. 12-23 Published by: Mycological Society of America Stable URL: http://www.jstor.org/stable/3753063 . Accessed: 26/11/2012 19:27
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NINETEEN YEARS OF CULTURE WORK1

J. C. ARTHUR
A series of cultureexperiments withthe Uredinaleswas begun by the writerin I899, and continuedunder the auspices of the Indiana AgriculturalExperiment Station without interruption until I917, making nineteen consecutive years in which this methodof researchwas consistently pursued. The resultsof the work were embodiedin fifteen reports, printedin the Botanical Gazette,Journalof Mycology,and Mycologia. It is now proto reviewthe work,in orderto set forthsome posed verybriefly of the objects accomplished, and especiallyto pointout the more of the changingconceptionsof the problemsforming important the groundplan on whichthe work was projected. The cultureswere not undertaken as part of a distinct thesis or circumscribed problem. They were ratherthe aids in a general taxonomic study of American rusts, which was directed toward supplying a technicaldescription as completeas possible for every species of Uredinales in North America recorded in literature or known to the writer. This ambitiousundertaking in the ninetiesat the invitation was definitely of begun sometime the editorsof the NorthAmericanFlora. taxonomicworkon the rustsin I882 it withmyfirst Beginning desirableforthepurposesof a fulltechhad seemedto me highly of species,that everypart of these microscopic nical description should be unicharacters, diagnostic plants,capable of supplying formlyconsidered,quite as much as are the stems,leaves, inflorescence, flowers,and fruitof higherplants, and that every of the means should be taken to arrive at a clear understanding of the various formsand species. No and relationship identity effort should be spared, it was believed,to make the name apand variations plied to any formembracealso thetransformations
1 Contribution fromthe Botanical Departmentof Purdue UniversityAgricultural ExperimentStation.
12
ARTHUR:
NINETEEN
YEARS OF CULTURE WORK
13
which that form undergoes in passing throughits whole life collectedin only one or two cycle. Many rusts are commonly or the of their several stages are taken as development, stages independent objects, and to grow such rusts so as to keep them under directobservationand be able to note the succession of and especiallyso fortheheteroecidesirable, stagesseemedhighly ous species whichpass theirgametophytic and sporophytic stages It unlike and unrelated hosts. was therenatural, upon wholly towardunravelfore,to directchief attention, especiallyat first, ing the tangle of heteroeciousforms. Nobody knew how manyrustswere to be foundon the North Americancontinent and its islands. There were possiblya thousand or more names in existence,but how manyof these names life cycles,and how many to applied to single and independent no one had attempted to say. parts of cycles,or were synonyms, It was, in fact, only with the existingnames that I had to do. It was no part of myproblemto discovernew species,or to give new names,eitherin preparing for the NorthAmerimanuscript can Flora, or in conductingcultures,except in so far as these were required for the systematicdevelopmentof the work. excursionswere made duringtheprogress Many longeror shorter of the cultures, some of thema thousandmilesor more,but they were all forthepurposeof makingfieldobservations upon known species, and in no case for making discoveryof new species. The new species that were found were an incidentalresult. The firstyear of the culture work, that of I899, was very and developed no particulardifficulties encouraging, calling for solution. So far custom was followed in the application of names, and it had not been necessaryto apply any formulato decide what constituted a species. The assumptionthat forms on the same or closelyrelatedhosts,havingno striking morphowere of one species seemed a sufficient logical differences, hyand the followed that cultures would pothesis, corollary necessarily show the range of hosts for each species, as well as serve to demonstrate the stages and spore-forms in the life cycle. Certain featuresin connection withthe commonEuphorbia rust did indicatethat difficulty mightbe found in the applicationof the
14
MYCOLOGIA
corollary,and this indicationbecame more pronouncedduring the year following. In I902 three species of Euphorbia of unlike appearance and rusts, growth habits were found to bear non-interchangeable whichwere tentatively consideredto presentraces of Uromyces because no Euphorbiae C. & P., and with the more confidence be distinctions could detected. Subwell-defined morphological this view of races, and the idea of sequent studies strengthened in mind. races fromthistimeon was constantly keptprominently to evade or simplify taxonomicand culturaldifficulThe attempt ties by treatingsuch races or biological strains as species, as did with these same Euphobia forms Tranzschel2subsequently was not favored. It was also in I902 that the Helianthus rust was grown with in the following indicationof races, developedfurther year,and broughtto a climax in I904, with the conclusionthat a number of moreor less well established races occur in Puccinia Helianthi annuus as a bridgingspecies, followHelianthus Schw., having ing the lead of Marshall Ward3 in his studyof the bromerusts. effort was made to studyraces in autoeciNo further considerable ous species,or to pick out bridging hosts,as it was held that to of species was as great a task as could be ascertainthe identity undertakenin this series of cultures,and that studies leading or other of a speciesintovarieties, to the separation races, forms, subclasses,althoughof much biologicaland often of great ecomustbe left forothertimeand hands. nomicalinterest, The problemsof the Carex rusts came early into view. In I90I and I902 the three remarkableco-species,having telia on various species of Carex and aecia on species of Aster, Solidago were repeatedlygrown from telial and Erigeron respectively P. Caricismaterial,and were called Puccinia Caricis-Asteris, no As P. and single collection Caricis-Erigerontis. Solidaginis, of teliospores was foundthatwould infectmorethan one of the to be species considered generanamed,the formswere tentatively and given distinctivenames, followng the brilliant cultural methods of Klebahn in Germany,Plowright in England, and
2 Ann. 3 Ann. Myc. Myc. 8: I: 1-35.
150. I910.
1903.
ARTHUR:
NINETEEN
15
others,althougha careful comparisonof the three formsmade it seem "not improbablethat the threerepresent more correctly thebiologicalvariationsof one species,"as was statedat thetime. In the further thatthe telial studyof these formsit was thought be foundto be restricted to certainspecies of Carex, stage might or to particularsectionsof the genus,as was believedto be true of the European Carex rusts,which assumptionin the case of the American forms,however,could not be establishedin any definite to be even less reway. The hosts were shown finally stricted than supposed,as the Aster formwas eventually carried over to Euthamia for its aecia and to Dulichiumfor its telia. The necessitysoon became acute to findcriteriaby which to of speciesamongthe rusts,and all themore judge of the standing so because the manuscript was now under preparationfor the North AmericanFlora. It was soon decided that, for the purcharactersmust be the final poses of the Flora, morphological test for species. Yet forpurposesof studyoutsideof taxonomy it mightbe serviceableand desirable to maintainthe so-called biological or physiological species in any rank desired,but they not be to as species properin taxonomicclassiought recognized in 1912 the threeCarex formswere comfication. Consequently bined with certain European forms under the name Puccinia extensicola Plowr., a name which has been supplanted by P. Asterum(Schw.) Kern, sincethe culturalseriesclosed. Furthermore,the culturesof I913 disclosedthat P. vulpinoidiswith its covered telia had its aecia on Solidago, and was a part of this same species heretofore knownonlywithnaked sori,makingthe muchemphasizedcharacter of coveredtelia a secondary one to be with associated principally the host. Thus the idea of species amongthe rustsgrew into a far more than could have been posdefinite, althoughmorecomplex form, sible without the aid of culturalstudies. A liberalview was now also required regardinghosts, and also the stress on certain called for modification, characters but the end was morphological not yet. In I9IO a numberof cultureswith the Carex rust, Uromyces revealeda remarkable betweenthisspecies perigynius, parallelism
16
MYCOLOGIA
and Puccinia extensicola. Aster and Solidago races came to light,not however quite so well stabilizedin some instancesas withthe corresponding Puccinia races,for in one case sowingsof were made to growon teliosporesfromthe same Carex collection bothAsterand Solidago. The two species,one of Uromyces, the otherof Puccinia, were subjectedto an extensivemicroscopical could be found betweentheir study,and no marked differences several corresponding except in the septationof the spore-forms, had already been observed teliospores. This unityof structure culture regardingthe aecia and aeciospores when a preliminary of the Uromyces was made seven years before. From the evidence,unitedwithmuchcollateralevidence,the microscopical statement was made in the discussionof I9IO, which following holds true to the presenttime: "As the aecia and uredinia of the two groups [of host-races],one under the genus Puccinia and the otherunder Uromyces,are indistinguishable, and as the the with of the one-celled Uromycesagree teliospores spores of the Puccinia [mesospores] and also with the two-celledspores in all characters lengthof exceptnumberof cells and consequent the races of latter. doubtlessare morphological spore,the former all of these forms Relationshipcould be shownbetterby putting the several races by under one specificname, and designating varietal names. But in the present state of taxonomyof the rusts it is more convenient to dispose of them under the two Puccinia and Uromyces."4 genera: If any further were needed to show that Puccinia illustration and Uromyceswere not onlyparallelgenerabut actuallyidentical, Durit was suppliedby the culturesof the following year,I9II. ing this season successfulcultureson Atriplex hastata of both each grown from UromycesPeckianus and Puccinia subnitens, fromwidely obtained on the Distichlis spicata, grass, teliospores separatedlocalities,gave rise to aecia that appeared to be indistinguishable. A morphologicalstudy of these two so-called by C. R. Orton in his articleon " Corspecies has been reported in which he findsa relationbetween Puccinia and Uromyces,"5
4 Mycologia 4: 22. 5 Mycologia 4: 199.
1912.
1912.
ARTHUR:
NINETEEN
17
in size of the urdiniospores, and, of course, in slightdifference in numberof cells and consequent the teliosporesa difference are such as size. He pointsout, however,that these differences are to be expected in other similar cases. The comparisonof hosts and these two formsof Distichlisrust,as to morphology, is an interesting distribution, topic, which need not be pursued further here. If the Carex-Aster-Solidago-Erigeron studiessupplemented by studies with the Distichlis rust, opened up new views of the and teliosporicdispecies question in relationto host influence so did the Carex-Ribes studies disclose new views in morphism, another direction. The firstcultures were in Io9I. As the resultsof sowing teliosporeson Ribes gave peculiarlysmall and that an unrecognized pale aecia, it was thought species had been found, which was called Puccinia albiperidia. Whether this form was distinctfrom the common Carex-Ribes rust of the as P. Grossulariae,and whetherAmerican fields,distinguished fromEuropean forms, of whichKlebahn had formswere distinct which received attentionfrom recognizedfive, were questions permitted. In thisstudyDr. Klebahn yearto year as opportunity graciouslyconsentedto lend assistance,and duringthe two seasons of I904 and I906 made cultures at Hamburg, Germany, fromtelial materialsuppliedby the writer. Justas the problemseemed solved,and Dr. Klebahn6and myarrived at the conclusion that in both self had independently and America one heteroeciousspecies occurred, only Europe which possessed a numberof strainsor races, it was discovered by C. R. Orton,7while assistingwith the rust studies,that the originalmaterialof P. albiperidiaonCarex pubescens,as well as that on a numberof American species of Carex similar to C. with only one basal pore, in gracillima, possessed urediniospores at instead the three of usual or four equatorialpores. least, part of Again the Carex-Ribes rusts Amercia seemedto fall into two in the aecia thistime,but on difspecies,not based on differences ferencesin the urediniospores. From I9IO onward the question in this connectionwas whetheror not the same species of rust
6Zeits. Pflanzenkr. 17: 132-134. 7 Mycologia 4: I4, 200. 1912. I907.
18
MYCOLOGIA
could possess urediniospores withone basal pore and partly partly with three or four equatorial pores. The answer involvedthe value and application of pore characters in definingspecies. whichled to both kinds of urediniospores After special search,8 in the same sorus,althoughfor the most being found repeatedly in occurred they part separate sori, it was concludedthat only but with morphoone species of rust was under consideration, races, not well delimited. logical as well as physiological thatthe previously It seemedprobable,furthermore, described, known form Carex as of rust, Uromycesuniporulus one-pored Kern, was a race also belongingto the Carex-Ribes species,but it was not possible to test the matterby cultures. In this conto note, and provocativeof speculation, nectionit is interesting that there is no form yet known with three- and four-pored urediniosporesbelonging under Uromyces in the Carex-Ribes aggregation,to make the parallelism with its Puccinia form complete. In 1917, the last year of the culture series, the principleof with a greateror characters, basing species upon morphological in interpretation, was further illustrated less degree of mobility case Polemonii the of the (Peck) Spartina rust, Uromyces by Barth.,which it was foundcould be segregatedinto four races,9 in morphological separable by small but appreciabledifferences both and charactersof aeciospores teliospores,and by wholly reinforced unrelated aecial hosts,and further by some differences in habitat and geographicaldistribution. The correlatedPucAmericanrust forthiscommon and widelydistributed cinia-form so-calledbecause the typecollecis that of Puccinia Distichlidis, labelled as on Distichlisinstead of on Spartion was incorrectly tina. Its range and aecial hosts,so far as knowncorrespondto only one of the fourUromycesraces. At the time the culture work began the subepidermalrusts on wild grassesin Americawithfew exceptions, passed occurring under the name of Puccinia rubigo-vera, along withpart of the similar leaf rusts of cereals. No criteriahad been found for them,not even those which had received special distinguishing
8 Mycologia 7: 67-69. I915.
Mycologia 9: 309-312. 1917.
ARTHUR:
NINETEEN
19
to make headwas consequently names,and everyeffort put forth The success was in maze of forms. first this into obscure way on with a on and aecia form Elymus uirginicus I902 Impatiens, which became Puccinia Impatientis (Schw.) Arth. The work openedup slowly. In I903 a false move was made in connection with the rust on Bromus, but the year followingthis rust was shown to have aecia on Clematisvirginiana.10 In I907 Puccinia AgropyriE. & E., as it occurredin Coloa was foundto go to Clematisligusticifolia, rado on Agropyron, connection that had been demonstrated by Dietel with European hosts fifteen year Puccinia cinerea years before. The following Arth. on Puccinellia was grown on Ranunculus Cymbalaria,a rustfromKoeleria cristataon Mahonia, fromBromus on Thalictrum, from Agropyronon Aquilegia, the last three being defrom scribed as new species. In 1915 aecia on Hydrophyllum Utah were made to grow on Agropyron and Elymus,givingrise to uredinia and telia similarto those fromthe Ranunculaceous aecia, but believed to constitutea distinct species. In I916 anotherruston Koeleria cristatawas grownon Laciniaria under the name P. Liatridis (Ell. & And.) Bethel. Repeated attempts were made to findthe aecial host of the commonleaf rust of wheat, P. triticina Erikss., but withoutsuccess, althoughthere were manyindications thatpointedto a Ranunculaceoushost,and Anemone. It was thought to Clematis or thata favorespecially able trialon ClematisFlammula would give a measureof success. At any rate it was believedto be one of the numerousraces of the subepidermal leaf-rustof grasses,P. Agropyri, with Ranunits culaceous hosts for aecia.11 The series were discontinued before the study of the subepidermalformswas completed, but ten of themhad been connected with their aecia. The convictionhad been growing for some timethat some of theseten names represented races of Puccinia rather than was stated in disas Agropyri, independent species, cussing the cultures of I912. When the manuscriptwas pre10 For a full account and explanationof the mistake of I903 in supposedly connectingaecia on Dirca with the Bromus rust see Journalof MycologyII:
I905. 62-63. 11 Mycologia 9: 276.
I917.
20
MYCOLOGIA
P. Agropyri, pared for the North AmericanFlora P. tomipara, P. cinerea,P. alternansand P. obliterata, as well as P. triticina, were placed underthe one name of P. Clematidis(DC.) Lagerh. It is considereda great advance to bring fromthe limbo of P. rubigovera,six distinguishable species, some of them having a considerablenumbero.frecognizedraces, and thereby makingit dozen more names that to some or to obscurity possible relegate been encumbering the literature of the rusts. had previously In a somewhatsimilarway the AmericanCarex rustswere in of thecultures. They were quite utterconfusion at thebeginning generallycalled Puccinia Caricis or P. caricina,no cultureswith Americanmaterialhaving been made, and diagnosticcharacters ten species were not having been well worked out. Altogether grownduringthe cultureperiod to show theirfulllife cycle,and .in several of thema numberof races was found,includingthe one-celledUromycesperigynius. Of course,being able to separate theseten species made it possibleto decide upon the identity of otherspecies,whichwere not actuallygrown. A view generallyheld when the culturework began was that the hosts of an autoeciousspecies,or of each of the two parts of a heteroeciousspecies, would be found to be closely related, often,indeed,to be but a single species, or genus, and certainly always within a single family. Consequentlyit was felt that the problem whena grass or sedge rustwas successfully cultured, about hosts for that species was practicallysolved. This complacentopinionwas quite upset in the case of Puccinia subnitens Diet. on Distichlisspicata, which in I902 was firstgrown upon Chenopodiumalbum,. In I904 Rev. J. M. Bates of Nebraska, who had made the field observationsand suggestionsfor this his observations wrotethat he had been continuing combination, of thisspecies and believedthatit had aecia also on hostsbelongwhichseemedto the writerat the time ing to two otherfamilies, on as most incredible. Tests, however,showed it would flourish as well as species of Cleome, Lepidium,Sophia and Erysimum, it that would the admission on Chenopodium, grow compelling "with equal vigor upon species belongingto three families of fact not knownfor any plants,"at the timebeing a " remarkable
ARTHUR:
NINETEEN
21
other species of rust." Additional genera in the same families were added fromtime to time for aecial hosts,until in the culturesof I916 the species was grownon Abronia and Polygonum, thusaddingtwo more families. Mr. E. Bethel,of Denver, Colorado, who made the field observationsand suggestionsfor the later additions,has continuedthe list since the culture series stoppedand broughtthe numberup to 76 species, belongingto I9 families,12 a trulyastonishing and all the more so as showing, no clearly definedraces have so far been detected. The only otherspecies of rust with such a remarkably extendedseries of aecial hosts at all approachingP. subnitensDiet., is that of P. Isiacae (Thiim.) Wint. from the dry trans-Caspianregion of westernAsia, as reportedby Tranzschel.13 This species with telia on Phragmitescommunishas aecia on I9 species of hosts the aecial familiesbeing the same as for to 9 families, belonging P. subnitens. In still another way the conceptionof species was modified when in 1905 teliosporesfromRuellia ciliosa were grownon the same host and also on R. strepens. The latterhost, with loose, tenper cent.largerin every tissues,gave riseto aecia fully watery host with its firm, way than did the former woody tissues,thus showingthatthe formsrecognized by the Sydows underPzccinia lateripesB. & Rav. and P. Ruelliae (B. & Br.) Lagerh.14 represent only a host influence-upon one and the same species, this influence beingtracednot only in the aecia, but also in the other spore-forms. Thus it will be seen that while the main work of the cultures in completing was effective the life cycles for many species,and in some cases extendingand defining the range of hosts, it was at the same timemostprofoundly the current modifying conception of species among the rusts. Instead of a rigid ideal of a few invariablecharactersand a limitedrange of nearly related hosts to be determined a comby cultures,we have substituted of variable somewhat characters the as basis, plex morphological
12 13
Petersb. 3: 40.
Bethel, Phytopathology 9: 193. I919. Beitrage zur Biologie der Uredineen. Trav. Mus. Bot. Acad. Sci. St.
I906; 7: 14. I909.
I902.
14Sydow, Monographia Uredinearum i: 235.
22
MYCOLOGIA
with a more or less extendedrange of hosts in part determined in the fungus. and in partby microscopical similarities by cultures A species at the beginning of the workwas conceivedas a simple and direct succession of individuals of the same appearance, capable of being demonstrated by cultures,but at the close had become a bundle of somewhatmobile characters, oftencomprisand sometimes ing manystrainsvaryingphysiologically morphoby logically,and to a more or less extent not interchangeable cultures. In some other ways than already mentionedthe accepted notionsregarding rustswere modified. It was foundthatteliospores among the grass formswere not all necessarilyresting forms spores,and thatthenon-resting presented special problems, whose solutionwas not far advanced when the work came to a close. Assistancewithfieldobservations and materialpermitted successfulculturesto be made in May, I9II, withthe aeciospores fromArabis sown on Trisetum. The Arabis aecia arise from the stemand leaves of systemic mycelium extending throughout the plant. A monthlater teliosporesresulting fromthis culture, now havingbecomemature, were foundto be capable of germination and were sown on seedlingrosettesof Arabis. The results of thissowingfirst showedwhenthe axis of the Arabis definitely began to elongate as growth started the followingspring. A culturewas similarly carriedout in I903 withPuccinia Eatoniae, the on aecia Ranunculus abortivus,also a formwith difusing fused mycelium, but a reciprocalculturewas not made. These two species of rusts,having a systemicformof aecia, were the only ones of the kind whichwere broughtunder culture. They withsystemic class physiologically, aecia, belongto an interesting which and withteliospores capable of germination upon maturity, the winter,or only through possiblydo not retaintheirviability to an impaireddegree. The culture work began with the too prevalentidea that all in generalto the well known rustscould be expectedto conform that the rusts Puccinia graminis. It closed with the conviction are far too diversified in theirmorphology, theirnumerouscharacters, theirphysiological adaptations,and theirrange of hosts,
ARTHUR: NINETEEN YEARS OF CULTURE WORK
23
to be represented by Puccinia graminisin more than one out of numerousaspects. In this resumeof culturesonlya few of the more prominent that should help to modifythe too developments held have ideas of rust species as commonly rigidand restricted been broughtforward. Yet enough has been said possibly to indicatethe value of what has been accomplishedand the need of more extendedwork along similarlines.
PURDUE UNIVERSITY, INDIANA. LAFAYETTE,

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Mycological Society of America

NINETEEN YEARS OF CULTURE WORK1

YEARS OF CULTURE WORK

YEARS OF CULTURE WORK

YEARS OF CULTURE WORK

YEARS OF CULTURE WORK

YEARS OF CULTURE WORK

14Sydow, Monographia Uredinearum i: 235.

ARTHUR: NINETEEN YEARS OF CULTURE WORK

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