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CURRENT LITERATURE
NOTES FOR STUDENTS
Crowngall.-Recent developments in the studyof crowngall and its rela-
tion to animal cancerhave been presentedby SMITH in severalpapers. The
first'of these,in pointof completionalthoughnot in timeof publication,is a
succinctaccountof remarkablegrowthphenomenaresultingfromthe action
of Bacteriumtumefaciens wheninoculatedintospecial tissuesof plants. Four
cases are distinguished.
i. Whentheinternodalcambiumis inoculated,thistissuelosesitstendency
to formmature structureshaving definiteorientation. Instead, the cells
continueto divide rapidly,forminglarge masses of mostlyembryonicpar-
enchymawithinwhichscatteredand irregularly arrangedxylemand phloem
elementsare differentiated.The process recalls that describedby LAMAR-
LIERE' in the galls of Gymnosporangium and aptly designatedby him as
"parenchymatization. "
2. When the corticalparenchyma is infected,a somewhatsimilardevelop-
menttakes place. The cell divisionssucceed each otherso rapidlythat the
cells in the proliferating
tissue remainsmall in comparisonwith the normal
parenchyma,and appear to remaincontinuallyin an embryonicstate. In
time,however,thereis a tendencyto developvascularelements,and theseare
thenarrangedin a moreor lesswell definedstele. The vascularsystemofsuch
tumorshas no connectionwiththat of the stem,consequentlythe galls soon
die fromimperfect nutritionand lack of water. The galls of thesetwo types
exhibitno externaldifferentiation.They includeall the formsof crowngall
describedin formerpapers.3
3. A more remarkableconditionis broughtabout when the crowngall
organismis inoculatedinto the leaf axils of younggrowingplants (species of
Pelargonium,Nicotiana, Lycopersicum,Citrus,and Ricinus). The tumots

ISMITH, ERWIN F., Crown gall studiesshowingchangesin plant structures due to


a changed stimulus. Jour. Agric. Research 6:I79-i82. pls. i8-23. i9i6; see also
Le cancer est-il une maladie du regne vegetal? PremierCongress Internat. Path.
Comp. Vol. II. I9I2; Cancer in plants. Proc. I7th Internat.Congressof Medicine.
Vol. III. Pathology. London. I9I3; Furtherevidence as to the relationbetween
gallandcancer. Proc.Nat.Acad.Sci. 2:443-448.
crown i9i6; Further
evidence
that
crowngall ofplants is cancer. Science 43:871i-889. i9i6; Chemicallyinduced crown
galls. Proc. Nat. Acad. Sci. 3:3I2-3I4. 19I7.
2
Rev. BOT. GAZ.42: I53. i906.
3 Rev. BOT. GAZ.52:75. I9II; 55:257. I9I3-

337
338 BOTANICAL GAZETTE [OCTOBER

thusproducedare coveredwithabortiveleafyshootsor withflowershootsifa


flowerincept has been disturbed. On tobacco plants these teratoidtumors
may give riseto secondarytumorssimilarin nature. These daughtertumors
are connectedwith the parent growthsby tumorstrandswhich are quite
different in structureand location fromthose occurringin galls of the first
two classes. The tumorstrandsheretofore describedwerefoundin the Paris
daisy. They arise in the regionof the primaryxylemand consistof par-
enchymatissue. The new tumorstrandfoundin the tobacco occursin the
cortex. It consistsof a concentricbundle withthe xylemsurroundedby the
phloem. The daughtertumorsarise at intervalsalong the strandand often
of the parenttumor.
have all the characteristics
4. The last case, even moreremarkable,resultswhenthe youngleaves of
tobacco plants are infectedwiththe crowngall organism. From such infec-
tionson the midriband lateralveinstumorsarisewhichproduceleafyshoots.
These tumorsthe authorregardsas akin to teratoidtumorsin animals. The
factoftheirdevelopmentis anotherproofthatany plantcell notfullymatured
mayretainthe capacityfordevelopingthe wholeorganism.
In anotherpaper,4writtenformedicalreaders,the subjectof crowngall
is discussedin its relationto the problemsof human cancer. The general
resemblancesin mode of growth,cell multiplication,occurrenceof tumor
strands,and productionof secondarytumorsin the two classesof growthsare
pointedout. The materialspresentedin this paper are essentiallythose of
earlierpapers,togetherwiththe new factsof the paper reviewedabove. The
phenomena, however,are describedingreaterdetail,and consideredwithspecial
reference to theirbearingon animal pathology. Here, as in othercases, the
authorreliesmostlyon numerousexcellentphotographsforthe presentation
of his evidence.
In explaininghis standpointwithreference to the bearingof his workon
the problemsof animal cancer,the author makes no claim that the causal
organismofthecrowngall has anyrelationto humancancer. It is pointedout,
however,that thisorganisminducesin plantsa setofphenomenawhichhave a
strikingparallelin the manifestations of animal cancer. Suchphenomenaare
the growthwithoutfunctionshownby gall tissue,the persistently embryonic
characterof the proliferating cells,the lack of orderlydifferentiation
of the
tumortissues,the existenceof tumorstrandsgivingrise to daughtertumors
repeatingthestructure oftheparentgall,and theoccurrence ofgallsresembling
embryonicteratoids. It is further pointedout that in the crowngall thecell,
althoughapparentlypossessinginvasivecapacity,sis notitselftheparasite,as

4 SMITH, ERWIN F., Studies on the crown gall of plants; its relationto human
cancer. Jour.Cancer Research I:23I-258. pis. I-25. ioi6.
5 It appears that the embryonictissue of the gall to a certainextentpushesin
among the cells of the sound tissue, a phenomenonwhichdistinguishesthis growth
fromotherplant galls inducedby fungouso ranimalparasites. The mode of progress
I9I7] CURRENT LITERATURE 339

JENSENthought. On thecontrary, thebehaviorofthecellis due to an invading


specificmicroorganism.These facts,togetherwith the observationthat in
one case at least (Rous' sarcomaoffowls)theabnormalgrowthcan be produced
by some sortof materialseparablefromthe cells and capable of multiplying
wheninjectedinto othertissues,are regardedby the authoras greatlyadvan-
cingthe contentionthat animal canceris due to an intracellular parasite.
Exceedinglyinterestingresults6wereobtainedin a seriesof experiments
designedto determine themoreimmediatecausesoftumorgrowthin crowngall.
In this investigationthe authorwas guided by the hypothesisthat the sub-
stances producedin the metabolismof Bacteriumtumefaciens must be the
directcause of the cell proliferations.To the end of determining the effects
of such substancesvarious plants were injectedfirstwith compoundswhich
chemicalstudieshad shownto be productsof the causal organism,and finally
witha largenumberofothersubstances.
The firstexperimentswere conductedwith ammonia,which in various
concentrations was injectedinto the stemcavityof Ricinusand into the fruit
cavitiesofyounggreentomatoes. The resultoftheseinjectionswas an abun-
dantformation ofcushion-like intumescenceswithinthe cavitiesin bothcases.
Later,proliferations of the same typewereobtainedby theinjectionof a large
numberof other substances,includingammoniumsalts of organic and of
inorganicacids, dilute solutionsof the acids themselves,salts, glucose,and
saccharose,and in some instancesto a slightextentwithdistilledwater. In
many cases whenthe tissuesof the pith cavitiesof Ricinus wereexposedto
weak ammonia vapors from dilute solutions of ammoniumphosphate or
ammoniumcarbonatein tubessealed intothehollowstems,proliferations were
producednotonlyin thecavitiescontainingthereagentsbut also in manyinter-
nodes above and below the openedone. The actionin thesecases, therefore,
took place at a distancethroughthickpartitions. The most strikingresult
was obtainedfromthe injectionof a 5 per cent solutionof ammoniumdihy-
drogenphosphateintoa veryyounginternodeofRicinus. In thisinstancethe
pith cavitybecame completelyfilledby the proliferating pith, and fromthis
tissue a completevascular cylinderwas differentiated.The orientationof
the new innercylinderwas the inverseof that of the normalcylinder,the
phloembeingat the centerand the xylemoccupyingthe outerregion. Such
a completecylinderwas observedonly once, but in manyinstancesisolated

of the tumor strand throughthe tissues is not yet clear. Whetherthis structure
pushes its way throughthe pith or cortexby apical growthafterthe mannerof the
internalroots of lycopods,or whetherprogressthroughthe tissues is accomplished
of the in-
by successive cell-invasionby the bacteria and subsequent differentiation
vaded cells into the characteristictumorstrand,has not yet been determined. From
studyof his stained sectionsSMITH thinksthat both typesof invasionoccur.
6 SMITH, ERWIN F., Mechanism of tumor growthin crown gall. Jour.Agric.
Research8: 65-i86. pIs.4-65.I9 I 7.
340 BOTANICAL GAZETTE [OCTOBER

concentricbundles were produced in the proliferating pith. In these the


phloemwas always at the centerof the bundle. Such bundles,the author
pointsout,occurnormallyin theaxes oftheinflorescence ofRicinus and in the
nodes. Superficialintumescences similarto thosereportedby VON SCHRENK7
wereproducedon cauliflower by exposureof the plantsto vaporsof ammonia
and of acetic acid mixedwithalcohol.
The outgrowths heredescribedall partakeof the natureof intumescences
frequently observedin plants. In some cases, indeed,as in the instancede-
scribedofthecompletefillingofthepithcavityand thesubsequentdifferentia-
tion of a vascular ring,the outgrowthis excessive. This behaviorleads the
authorto thebeliefthatifthestimuluscouldbe continually applied,one would
have a conditionresultingin theproductionoftissuemassesnotunlikethoseof
crowngalls. Since in his experiments the outgrowthsalso resultedfromthe
presenceofmanysubstancesnottheproductofparasites,theauthoris inclined
not to attributethe effectto thespecificchemicalactionofany compound,but
seeksforan explanationin somepropertycommonto all thecompoundsregard-
less of chemicalcomposition. Such a commoncharacteristic he findsin their
osmoticaction,to which,ratherthanto chemicalstimulation, he ascribestheir
effect. In thisconnection it is ofinterestto recallthatintumescences have been
producedby ATKINSON,8 Miss DOUGLAS,9and STEINERIO by subjectingplants
to conditionsincreasingwaterabsorptionand diminishing transpiration;and
by SORAUER, KtSTER, VONSCHRENK, and othersas a resultof applicationof
solutions. In theauthor'sownworktheintumescences weremostlytheresult
ofinjectionof solutions,but in someinstancestheyresultedfromthe injection
ofwater. It is improbablethattheosmoticdisturbances inducedby theappli-
cationor injectionof waterare the same as thoseeffected by the application
or injectionof solutions. The fact that the various disturbancesproduce
responsesdiffering onlyin degreewouldseemto indicatethatthe causes deter-
miningthe formation of intumescences have not yet been fullyanalyzedinto
theirseparatefactors. It is notunlikelythatdifferent plantsreactdifferently
in thisrespect. The experiments ofSTEINER wouldseemto indicatethatsucha
possibilityexists.-H. HASSELBRING.
Taxonomicnotes.-GATEs" has attacked the genus Polygonatum, which
he says "has been in a verychaoticconditionowingto thelumpingof species,
thetransferenceofnames,and theconfusion ofNorthAmericanwithEuropean
7 Rev. BOT. GAZ.40:390. I905.
8 G. F., Oedema of the tomato. CornellUniv. Agric.Exp. Sta. Bull.
ATKINSON,
53:77-io8. i893.
9 DOUGLAS,Miss G. E., The formationof intumescenceson the potato. BOT.
GAZ.43:233-250. 1907.
10Rev. BOT. GAZ.40:39I. I905.

in North America. Bull.


"I GATES,R. R., A revisionof the genus Polygonatum
Torr. Bot. Club 44: II7-I26. pls. 4-6. I9I7.

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