Histological Study of The Developing Fruit of The Sour Cherry

HISTOLOGICALSTUDY OF THE DEVELOPIN7G
FRUIT OF THE SOUR CHERRY

H. B. TUKEY
ANDJ. ORAN
YOUNG
(WITH EIGHT FIGURES)
Introduction
Earlierworkersin the morphology of drupefruits,as LAMPE(II),
TSCHIERSKE (I9), FARMER (8), and WINTON(24), dealt primarily
withthe structureof the maturefruit,and recordedbut few signifi-
cant structuralchangesin the developingfruit. For these studies,
variousspeciesof Rubuswerelargelyused,asR. ideaus,R.fructicosus,
R. strigosus, and R. occidentaZis,althoughLAMPEincludeda brief
comparative discussionof the sourcherry,Prungs acida.
Later workershave been more concernedwith the externalor
grosschangesof the developingfruits, and have dealt principally
withspeciesof Prunus, as CONNORS (5), BLAKE(2), LILLELAND (I3),
HARROLD (9), DORSEY andMcMuNN(6), andTUKEY(2I) with the
peach(Prunus persicaStokes);LILLELAND (I2, I4) with the apricot
(P. armeniacaL.) and plum (P. domesticaL.); and TUKEY(20, 22)
withthe sweetcherry(P. aviumL.) andsourcherry(P. cerasusL.).
Thesestudiesshowedthreecharacteristic growthperiodsin the de-
velopingfruit,designated(22) as: stageI, a periodof rapidenlarge-
mentbeginningaboutthe timeof full bloom;stageII, a mid-season
periodof retardeddevelopmentvaryingin durationas correlated
withthe classof fruitandthe seasonof fruitripening;andstageIII,
a secondperiodof rapidenlargementextendingto fruit ripening
(fig. I ) .
Furtherstudy showedgross morphological changesof different
partsof the fruit)such as the stony pericarp,nucellusand integu-
ments, endosperm,and embryo. These were correlatedwith the
three majorgrowthstages of the fruit by TUKEY(20, 2I, 22),
LILLELAND (I4), and HARROLD (9). LOTT(I 5), and ADDOMS,
NIGHTINGALE, and BLAKE(I) showed chemicalchangesin the
I JournalArticleno. 298, New York State AgriculturalExperimentStation.
723] [Botanical Gazette, vol. IOO
Botanical Gazette 1939.100:723-749.

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724 BOTANICALGAZETTE
[JUNE
fleshyandstonypericarp,andLOTT(I5) and TUKEYandLEE (23)
in the seedandembryoas relatedto thesestages.
Thehistologicalandstructuralchangeshavebeenlesswellinvesti-
gated. Thepresentstudyis an attemptto showthesechangesin the
developingfruit of the sourcherry(Prunus cerasusL. var. Mont-
morency),fromthe closedbud (I8 days beforefull bloom)to the
ripe fruit. It includesa treatmentof the principaltissuesof the
pericarpduringeachof the threegrowthstagesand duringthe pre-
bloomstage.
MM
FRtJ/T r
STAGE I , SEAGE ff , STAGE z / R/
s 2 RAP/D DEVELOPMEt/T , RErARDED ' F/WAL SWELL /
t ' DEVELOPMSITi /
' l /
I .
+
I + /
/o ' <
r
/ #UaLus AHD /STtSUMESTS
t / / Et/O
Ag/ /
3 /3 MAY 23 2 /2 JUSf 22 JU 2
FIG. I. Growth stages in development of fruit of sour cherry from pre-bloom to

fruit ripening. After TUKEY(22).
Materialand methods
Montmorency, a mid-seasonvariety,ripens57 to 59 days after
full bloom. In this variety stage I is of 20 to 22 days duration;
stageII, I2 to I6 days;andstageIII, 2I to 23 days (22).
MATERIAL. Freshmaterialfromthe seasonof I938 andmaterial
fixedin threedifferentways and duringfourdifferentseasonswere
used,as follows:
I. Seasonof I930, ripefruitin 95 per cent alcohol.
2. Seasonof I93I, at 2 to 3 dayintervalsfromApril20 to May 26, in
Karpechenko's chromacetic and in formalin-aceticalcohol.

I 93 9] TUKEY & YOUNCHERRY 725
3. Seasonof I932, dailyfromJune4 to June 26, in formalin-acetic

alcohol.
4. Seasonof I938, at 2 to 5 day intervalsfromMay 24 to July I9,
in formalin-aceticalcohol.
Althoughtherewas somevariationin the numberof days from
full bloomto fruitripeningin the differentyears,it was possibleto
correlatethe material.A growthcurvewas madefor each season
basedon grossmeasurements of the fruit and examinationof the
development of the embryoandstonypericarp(fig.I). Thedatesof
full bloomand of fruitripeningbeingknown,it was possibleto lo-
cate definitelythe beginning,middle,and end of eachof the three
stagesof fruit developmentfor the materialfromall four seasons,
and to place one curve upon another. Samplesfrom the various
seasonswereaccordingly selectedforstudyat thesepointsandwere
foundto fit togetheralmostperfectlyto forma fourseasonrecordof
development.
FIXATION, SECTIONING, AND STAINING.Materialused in the
studyof thepre-bloomstageandpartsof stageI waskilledandfixed
in Karpechenko's chromaceticand run up into parafiin;then sec-
tionedwitha rotarymicrotomeandstainedwithsafranin,orangeG,
and crystalviolet.
Largerspecimensused in the study of stages I, II, and III to
fruit ripeningwere eitherexaminedfreshor killed and fixedin a
mixtureof IOO CC. of 50percentalcohol,23 cc. of glacialaceticacid,
and67 cc. of formalin,andheldin thismixtureuntilsectionedeither
freehandor on a slidingmicrotome.All freehandsectionsof this
type werestainedin safraninand light green. Most of them were
mountedin balsam,but somewereexaminedimmediatelyin liquid
mountsand foundsatisfactoryfor outlinedrawings.
Whenthe stony pericarphad begunto harden(stageII), there
was somedifiicultyin getting good sectionswhichcontainedboth
stony and fleshypericarp.Attemptsat freezingand holdingthe
fruitin variouspin devicesso as to exertno pressureon the fleshy
pericarpwerenot superiorto makingnumerousfreehandsections
and selectingthe betterones for study. Sectionsmadefromfresh
materialandfromthat in formalin-acetic alcoholwereequallysatis-
factory.
Whenthe stonypericarphadbecometoo hardto securecomplete

[JUNE
sectionsthroughthe entirepericarp,freehandsectionsfor detailed

study weremade separatelyin three regions:the fleshypericarp,
portionsof the adjacentstony and fleshypericarps,and the stony
pericarp.
For completesectionsof the stony pericarpfromripefruit, the
pit was removedand groundsuccessivelyon coarsesandpaper,on
fine sandpaper,on a mediumcarborundum stone,and on a fineoil
stone. Whenthe sectionwas 2 mm.thick,it wasmountedon a glass
slidewithDucohouseholdcementandgroundfurtherto the desired
thickness. This cementhardenedrapidly,held the mountfirmly,
and permittedstainingwithoutseparationfromthe slide.
Theepidermis wasexaminedin surfaceviewby strippingfromthe
fruit. A5ostainwasneededsincethe cellsweresufficientlypigmented
to showgoodcontrast.
MEASUREMENTS ANDDRAWINGS. All comparativeoutlinedraw-
ings weremadefromcrosssectionsthroughthe centerof the fruit
perpendicular to the polar axis. Comparativemeasurements and
drawingsof numberand sizeof cellsweremadefromsectorsat the
cheekof the fruit, as this has been shown(22) to be the point at
whichdevelopmentis mostregular.The cheekdiametermaybe de-
finedas a line passingthroughthe centerof the fruitperpendicular
to both the longitudinalaxisand the suturediameter.
Outlinedrawingswhereno cell detailswere desiredweremade
withthe useof a Bausch&Lombtriplepurposemicroprojector using
a I6 mm.objectiveand a constantset-up. Detaileddrawingswere
madewith a cameralucida,alwaysto the samescale,andwith the
sameset-upof equipmentto give identicalenlargemente
Countsof numbersof cells and measurements of cell size were
madewiththe use of bothfixedandslidingocularmicrometers, and
checkedagainstcameralucidadrawings.For cell size, an average
wastakenof severalcellswhichseemedtypicalof the tissuesunder
consideration.Mitoticfigureswerenotedand recordedas evidence
of cell division.
Investigation
GENERAL CONSIDERATIONS OFTISSUES INVOLVED
Threeprincipaltissues composethe ovary wall of the cherry:
innerandouterepidermis,stonypericarp,andfleshypericarp.The

I939] TUKEY & YOUNG CHERRY 727
grossdevelopmentof the fruit and of these tissuesis shownin the

accompanying seriesof outlinedrawings(fig. 2), beginningI8 days
beforefullbloomandcontinuingto fruitripening.lthe development
at the beginning,middle,and end of eachstageis shown.
The stony pericarpmay be dividedinto an innerand an outer
layer,andthe fleshypericarpinto an innermostlayerof smallthin-
walledparenchyma,a middleregionof largethin-walledparenchy-
ma, and an outeror hypodermallayerof collenchyma.The hypo-
dermallayerandthe epidermistogetherconstitutethe "skin"of the
fruit. Not all of thesetissuesaresharplydefinedin the earlierstages
of development,and they are outlinedin the accompanying draw-
ingsonly whenthey areto be easilyseenat that stage.
In subsequentdetaileddrawings(figs. 3, 4, 5) of the cellular
changeswhichthesetissuesundergo,the samerelativepositionson
the pageareretainedfor drawingsof similarstagesas thoseof gross
developmentin figure2, to facilitatecomparison.In tablesI and 2
the countsof cellnumberandmeasurements of cellsizein the tissues
involvedare for corresponding stagesof development.In the dis-
cussionwhichfollows,the tissuesaretakenup in the followingorder:
stonypericarp,fleshypericarp,andouterepidermis.
STONYPERICARP
PRE-BLOOM.Eighteen days before full bloom the innermost
portionof the ovarywall,whichbecomesthe stonypericarpat fruit
ripening,is composedof the singlerowof cellsof the innerepidermis
and an additionalthreeor four layersof cells of the pericarp(fig.
3A). The cellsof the inner epidermisare block-shaped while the
othersareelongatein the directionof the centralaxisof the fruit.
The line of demarcationbetweenthat portionof the pericarp
whichis destinedto becomestonyandthatwhichis to remainfleshy
is discernibleeven at the beginningof the period,the cells of the
formerbeingsmaller,in general,than thoseof the latter. It is not
possibleat thisdateto separatea singlebordering cellof theonefrom
the other. The relativelysmallersize of cellsof the stonypericarp
persiststo maturity.
Soonafterthe beginningof this periodthe cellsof the innerepi-
dermiselongatetangentially,as seen in crosssection,and just be-
fore full bloom they dividein a transversedirection(fig. 3B, C).

FIG.2. Developmentof cherryfruit as seen in transversesection, from I8 days
beforefull bloomto fruit ripening.A: I8 days beforefull bloom;B: 6 days beforefull
bloom;C: full bloom;D: II days after full bloom;E: 20 days after full bloom;F:
26 days after full bloom;G: 36 days after full bloom;H: 40 days after full bloom;
1: fruit ripening,57 days after full bloom.
728

LAYEQ
BUNDLE
(FIG. 2.-Continqxed)
729

A B's
WosP
osP
DosP
sP
'
AosP
Ae:
' : a
_
--
(camera lucida drawingsof transverse
3. Developmentof stony pericarp pericarp;isp, inner
inner epidermis;OSp, outer stony
FIG.
sections X I63). p, pericarp;ie, before full bloom; C: full
full bloom; B: 6 days
stony pericarp.A: I8 days before E: 20 days after full bloom; F: 26 days after
bloom;
bloom;D: II days after full I: fruit ripening,
bloom; G: 36 days afterfull bloom;H: 4o days after full bloom;
full
57 days after full bloom.
73o

AW
Bg
cX
FIG.4.-Development of fleshy pbricarp,in regionbetweenhypodermallayer and

ring of vascularbundles(cameralucida drawingsof transversesections XI63). A: I8
days beforefull bloom;B: 6 days beforefull bloom;C: full bloom;D: II days after
full bloom;E: 20 days after full bloom;F: 26 days after full bloom;G: 36 days after
full bloom;H: 40 days after full bloom;I: fruit ripening,57 days after full bloom.
73 I

BW R $ K - ww - -
oooD BUCoCo o C
WC¢^ wn
C
CCoooD o cDoo
Doo ooooooDOso c
b- -X- '-
FIG. 5.-Development of epidermisand hypodermallayer (cameralucidadrawings

of transversesections XI63). A: I8 days before full bloom; B: 6 days before full
bloom; C: full bloom; D: II days after full bloom; E: 20 days after full bloom;
F: 26 days afterfull bloom; G: 36 days after full bloom; H: 40 days after full bloom;
I: fruit ripening,57 days after full bloom.
732

Pre-bloom
Full
Middlestage
bloom
.....bloom
I ...........
...........
.... - 6 . +II
o
4-5 -o
5 I 22
I 6 .......
5I9-2I I7I .......
......
I I5-I 620
... ... ...24-27
7 208 22.......
I2 6IOI- 747 I8-I9
-II4 8
939] TUKEY & YOUNG CHERRY 733
TABLE 1
CHANGES IN NUMBER OF LAYERS OF CELLS IN A RADIAL DIRECTION THROUGH
CHEEK OF CHERRY FRUIT FROM I8 DAYS BEFORE
BLOOM TO FRUIT RIPENING
FLESHY PERICARP STONYPERICARP

DAYS
BEFORE
ST.AGEOF OR
EPIDER- LARGE- SMALL
DEVELOPMENT AFTER
MIS AND CELLED CELLED
FULL TOTAL OUTER LKNER TOTAL
HYPO- PAREN- PAREN-
BLOOM
DERMIS CHYMA CHYMA
Pre-bloom........... -I8 4-5 II ....... ... I5-I6 4-5

* --- 4 5
End of stageI . . . +20 5- 6 I8-I9 3 4 26-29 I7-I8 6- 7 23-25

MiddlestageII. . +26 5-6 I9 3-4 27-29 20 6-7 26-27
End of stageII. . +36 5-6 I9 3-4 27-29 20 6-7 26-27
MiddlestageIII. +40 5-6 I9 3-4 27-29 20 6-7 26-27
End of stage III. +57 5-6 I9 3-4 27-29 20 6-7 26-27
TABLE 2
CHANGES IN SIZE OF CELLS IN A RADIAL DIRECTION THROUGH CHEEK OF
CHERRY FRUIT FROM I8 DAYS BEFORE FULL BLOOM TO FRUIT RIPENING
S TONY
EPIDERMlS FLESHYPERICARP
PERICARP
DAYS
BEFORE
PARENCHYMA
STAGEOF OR
DEVELOPMENT AFTER TANGEN-
RADIAL CROSS
FULL TIAL
DIAMETER TANGEN- SECTION
BLOOM ( ) DIAMETER RADIAL
DIAMETER
(p) (p)
Pre-bloom -I8 I7 IO I8 I3 IO
Pre-bloom - 6 I8 IO I8 I6 IO
Middlestage I +II 28 I3 32 32 I2
End of stage I +20 35 22 83 83 28
Middlestage I I +26 35 28 90 90 30
End of stage I I +36 35 3I 92 92 30
Middlestage I I I +40 35 +39 + I80 + IOO 30
End of stage I I I +57 35 +68 35°-500 IOO-250 30

transverse
direction
round
it .................... therestconsistsof fibres
[JUNE
Togetherwith a few adjacentcellsof the pericarpwhicharesimilar
in shapeandorientation,theyforma bandor "hoop"of transversely
elongatedcellsboundingthe innerovarywall. Sincetheylie at right
anglesto the othercellsof the stonypericarp,theboundarybetween
the two is sharpand makesa cleardivisionbetweenthe sections
whichat maturityare designatedinnerstony pericarpand outer
stonypericarp.
TSCHIERSKE (I9) picturesa similarconditionin the pericarpof
Potentillaanserina;andLAMPE(II), WINTON (24), and FARMER(8)
observedan identicalsituationin speciesof Rubus. FARMER re-
cords"a chainof suchfibresrunningaroundthe fruitcavityresem-
bles a segmentedringor hoop .... and the wholeof the interiorof
the ovary appearsto be boundedby these 'rings'whichrun in a
whoselongestaxis runsin a directionat rightanglesto that of the

cellsjust described.Theresultof this arrangement is that the ovar-
ian cavityis protectedby a bandof tissue,of whichonehalfis fitted
to standstrainsand stressesin one direction,whiletheotherhalfis
equallywellcapableof resistingsimilarstrainsandstressesin a direc-
tion at rightangleswith the first"(8).
By rapiddivisionwithinthe tissuewhichis to becomethe stony
pericarp,the numberof cells is nearlytrebledby the time of full
bloom. The diameterincreaseof cellsis, however,only IO per cent.
By thetimeof fullbloomthe tissueis welldefinedas a regionof small
compactcellsscarcelyhalf the diameterof the adjacentcellsof the
fleshypericarp.Theinnerstonypericarpis approximately fourcells
thickandthe outerstonypericarpten cellsthick.
It may be well to emphasizethat the groupsof cellswhichgive
riseto the stonypericarpapparentlydo not give riseto cellsof the
fleshypericarp,or vice versa. It is not a caseof the entirepericarp
beingderivedfroma commongroupof cells,someof whichtoward
the centerof the fruitbecomesclerenchymatous andsomeof which
towardtheperipheryremainparenchymatous andenlarge.Noris it
a caseof progressive hardeningof cellsof a commontissuefromthe
centerof the fruitoutwardin a radialdirectionandthe enlargement
of the remainingunhardenedcells to comprisethe fleshypericarp.
On the contrary,the two tissuesappearto be derivedeachfromdis-

I939] TUKEY & YOUNG-CHERRY 735
tinct groupsof cellswhichareearlyseparatedby characteristic size,

shape,andfrequencyandperiodicityof division.Thereis no transi-
tion in sizefromthe smallercellsof the tissueof the stonypericarp
to the largercellsof the fleshypericarp.Moreover,mitoticfigures
are relativelymoreabundantin the tissueof the stony pericarpat
the timethat greatestincreasein numberof cellsby actualcountoc-
curs,anda similarconditionobtainsin the tissueof the fleshyperi-
carp.
STAGE I. DuringstageI (fig.3C, D, E) the total thicknessof the
wallof the stonypericarpat the cheekincreasesapproximately five
times. Thisis broughtaboutby a largeincreasein bothnumberand
sizeof cells. Enlargement of cellsfor both the innerand outersec-
tions is marked, an increaseof approximatelysixteen times in
lengthandtwoor threetimesin width. In the innerlayer,elongation
takesplacein a tangentialdirection,as seenin crosssection,but in
the outerlayerelongationis in the directionof the centralaxisof the
fruit. Scatteredgroupsof cellsmay be foundwhichareorientedin
planesat variancewith the generalpattern.
By the end of the periodthe numberof cells alongthe radiusof
the pit at the cheekhas increasedfromfourteento twenty-four,the
greatestincreaseoccurringin the outerstony pericarp.Activecell
divisioncontinueslater than in the Reshypericarp.In the inner
stonypericarpno cell divisionshavebeenobservedafterthe middle
of stageI, but in the outerstonypericarpthe greaterincreaseoccurs
in the latterhalf. Likewisethe increasein the sizeof all cellstakes
placelargelyin the latterhalfof the period(fig.3E). By the middle
of stageI the cellshaveincreasedfromIO to I2 p in transversediam-
eter,whileat the endof stageI they measure28 p.
Cellwallsbecomeprogressively thickerbut do not hardenappre-
ciablyuntilnearthe endof the period. At the endof the stagethe
pit is too hard to sectionwithoutspecialtreatment,althoughthe
wallshavenot yet attainedmorethanone-fifththe thicknessfound
in the ripefruit.
STAGE II. Thickeningand hardeningof the cell walls of the
stonypericarparethe most characteristic developments of stageII
(fig.3E,F, G).Althoughmaturation of thetissueis progressing
rapid-
ly as evidencedby thickeningandhardeningof the cellwalls,yet ac-

tive celldivisionoccurshereas late as if not laterthan,in anyother [JUNE
partof the pericarp.Thismaybe explainedby the fact that harden-

ing doesnot occuruniformlythroughoutthe stonypericarp.It be-
ginsnearthe apicalend,at the hilum,andextendsdownwardalong
the ventralsuturestowardthe chalaza,beforefinallyincludingthe
entirestony pericarp.Theseobservationsagreefairly completely
withthoseof MIKI(I6) andRAGLAND (I7) forthe peach,andBRAD-
BURY(4) for the sourcherry. At the completionof the periodthe
stony pericarpis composedof twenty-sixto twenty-sevenlayersof
very thick-walled, lignifiedcells. This is nearlyhalf the total num-
berof layersin the entirepericarp,but becauseof the muchsmaller
sizeof the cellsof the stonypericarp,they representmuchless than
half the thicknessof the ovarywall.
No cell divisionshave been observedin this regionbeyondthe
middleof stageII. The cells areliving,however.Nucleiare abun-
dant,andby theirbrightreactionto safraninsuggestactivity. Such
suggestedactivity wouldseem in agreementwith the tremendous
thickeningof the wallsandtheirimpregnation with materials.The
lumenis frequentlyscarcelylargerthan the nucleus.
STAGE III. DuringstageIII (fig.3G,H, I) thereis slightincrease
in hardnessand brittlenessof the tissue, associatedwith loss of
moisture(I5). Althoughby the time of fruit ripeningmost of the
cellshavebecomematuredas sclerenchyma, it is stillpossibleto find
occasionallivingcellswith activenuclei.
FLESHYPERICARP
daysbeforefullbloomthe fleshypericarp
PRE-BLOOM.--Eighteen
is composedof nearlyisodiametric,parenchymatous cells (fig.4A,
B). Thereis a suggestionof tangentialelongationnearthe epidermis
anda gradualchangein o.rientation to a slightradialelongationnear
the stonypericarp.Increasein the thicknessof the tissue,whichis a
little morethan doubled,is due mostly to cell division,although
there is some very slight cell enlargement.In a radialdirection
throughthe cheek,fromthe epidermisto the stony pericarp,the
cellsincreaseapproximately 4o percentin numberduringtheperiod.
STAGEI. Duringstage I the numberof cells acrossthe cheek
fromepidermisto stonypericarpis increased20-30per cent, most

I939]
TUKEY & YOUNG CHERRY 737
of the increaseoccurringduringthe first half of the period. NO
mitoticfigurescouldbe foundafterthe tenth dav fromfull bloom.
ADDOMS, NIGHTINGALE, and BLAKE(I) and RAGLAND (I7) have
noteda similarrapidcelldivisionin the peachfollowingfull bloom,
and its cessationearlyin stage I; and TETLEY(I8) has foundthat
growthin the appleafterthe time of fruitset is mostlya matterof
cell enlargement.
Celldiameteris quadrupled, from20 to 83 ,u,the increaseoccur-
ringmostlyin the latterhalfof the period. Intercellular spacesbe-
comenumerousandprominenttowardthe endof the period.
A hypodermal layerfiveor six cellsthickis diSerentiatedjustbe-
neaththe outerepidermisduringthis period,by considerable thick-
eningof the cellwalls(fig.5D, E). The cellsattainbut little greater
size than those of the epidermis,but becomesomewhatelongated
tangentiallyas seen in crosssection. The shapeof the cellsof the
thin-wa]ledparenchymaremainsessentiallythe same throughout
theperiod,exceptin the outerportionwheresometangentialelonga-
tiontakesplace (fig.4C, D, E).
A layer of small isodiametriccells three or four cells wide is
formedadjacentto the stony pericarpduringthe latterpartof the
perio-d.Withthe possibleexceptionof this layer,thereis no further
celldivisionin the fleshypericarpaftercompletionof stageI. The
combinedactivityof cell divisionand enlargementthroughoutthe
fleshypericarpresultsin an increasein thicknessof seven times.
Nearlythe same figureappliesto the stony pericarp;hence the
growthis fairlyuniformforbothtissues. Celldivisionoccurslaterin
thestony than in the fleshypericarp,notwithstanding the earlier
differentiation of the former(fig.4E, F, G).
STAGE II. DuringstageII thereis somesmallenlargement of the
cellsof thefleshypericarp,froma diameterof 83 to 92 ,u. Tangential
elongation of the cells just beneaththe outer epidermiscontinues
slowly.Intercellular spacesremainprominent.Theremaybe some
celldivisionin the thinlayerof smallcellsadjacentto the stonyperi-
carp,but otherwisethereis none. Increasein thicknessof the fleshy
pericarp is not morethan6 per cent.
STAGE III. Duringstage III, the finalswell (fig.4G, H, I), four
divisions of tissuemay be recognized,exclusiveof the outerepider-

[JUNE
mis: (a) thehypodermal layerof collenchyma; (b) a peripherallayer
of thin-walledparenchymaextendingfromthe hypodermal layerto
a line just insidethe ringof vascularbundles;(c) a layerof radially
elongatedcellsextendingfromthis line nearlyto the pit; and (d) a
thin layerof smallcellsadjacentto the pit.
While for conveniencean arbitraryclassificationof tissueshas
beenmade,it mustbe realizedthat thereis a gradualtransitionfrom
a tissuein whichone type of cellspredominates to that in whichan-
otheris dominant.As seenin somesections,thin-walledcellsof the
fleshypericarpmay occasionallycome in directcontactwith the
epidermis;or the layersof tangentiallystretchedandflattenedcol-
lenchymaof the hypodermallayermay be wanting. Likewisethe
collenchyma givesway graduallyto the threeor fourlayersof large,
more nearlyroundishcells of thin-walledparenchyma.These in
turngive way to radiallyelongatedbut notablywidenedor obovate
cells, whichpass to progressivelymorenarrow,radiallyelongated
cellstowardthe pit.
The chieffeatureof this stage,often calledthe finalswell,is the
increasein sizeof the individualcellsof the fleshypericarp,the larg-
est indicatingan increaseof as muchas twenty-fivetimesin diam-
eterfromthe size at full bloom. The cellsof the hypodermallayer
becomeenlargedin a tangentialdirection,similarto the adjacent
cellsof the outerepidermis.Thosein the outerportionof the fleshy
pericarpincreaseseveraltimes in diameterand becomeroundish
oval in shape,with the greatestdiameterparallelto the periphery
of the fruit. Theybecomeprogressively largerfromthe hypodermal
layerinwardto the regionof the vascularbundles,andthe tangential
elongationbecomesprogressively lessmarked.Nearthe ringof vas-
cularbundlesthe cellsare essentiallyround.
In the middleregionof the fleshy pericarptremendousradial
elongationoccurs,an increaseof fourto six times,andsomeenlarge-
menttangentially.The tangentialenlargement is progressivelyless
towardthe stony pericarp.As wouldbe expectedfromthe rapid
stretching,thereis a decreasein the thicknessof cellwalls. Intercel-
lularspaces,whichwereso frequentandconspicuous duringstageII,
are now difficultto find. This may be due in part to the relatively
greatersizeof the individualcellsandthe attendantdecreasedlike-

lihoodfor the inclusionof intercellularspacesin a crosssection. Or

it may be that increasein pressurewithinthe cells accompanying
enlargementcausesthe relativelythin wallsto lie morecloselyto-
gether. ADDOMS, NIGHTINGALE, and BLAKE(I) foundintercellular
spacesmuchless conspicuousin the peachthan in the apple,and
suggestedthat in the caseof the peach,largerareasof the cellwalls
are in contactwith the walls of adjacentcells than in the apple.
TSCHIERSKE (I9) observedthe absenceof intercellular
spacesin the
regionof elongatedcellsin the fleshypericarpof the redraspberry.
At all eventsthenatureof the tissueis oneof closelycompressed and
compactedcells with inconspicuous intercellularspaces,a striking
contrastto the descriptionwhichmay be foundin the literatureof
loose-fittingcellswith largeintercellularspaces.
The bandof threeto five rowsof very smallisodiametricparen-
chymacellswhichlie adjacentto the pit makesno notableincrease
in size duringthis period. The cells lie betweenthe largeradially
elongatedcellson the one sideandthe cellsof the stonypericarpon
the other. Althoughthe line of demarcation betweenthe stonyand
the fleshypericarpis distinct,cellsof the one aremoreor less dove-
tailed betweencells of the other, so that the outer surfaceof the
stony pericarpwhenseparatedfromthe fleshypericarpis slightly
roughand somewhatpitted.
The total numberof rowsor layersof cells in a radialdirection
throughthe cheekof the fleshypericarphasbeencountedas twenty-
sevento twenty-nine(tableI). It is of interestto note that LAMPE
(II) givestwenty-eight as the numberfromobservationsof Prunus
acidamadein I886. Further,the numberof layersof cellsis nearly
equallydividedbetweenthe stony pericarpand fleshy pericarp,
beingtwenty-sixto twenty-sevenfor the former a total of fifty-
threeto fifty-six.
VASCULAR
BUNDLES
The ventraland dorsalcarpillarybundleslie whollywithoutthe
stonypericarp,althoughcloseto it. Twobundleswhichsupplythe
two ovules, calledin the peach "funicularbundles"by RAGLAND
(I7), lie at eithersideof the ventralsutureandadjacent
to the ovar-
ian cavity (fig.2). Thereareno "pitbundles"traversingthe stony
pericarpin the cherry,suchas he describesfor the peach.

74o BOTANICAL GAZETTE
[JUNE
Extendingthroughthe fleshypericarpin a directionparallelwith
the centralaxis,as seenin crosssection,is a ringof vascularbundles,
generallyeighteento twentyin number.They are differentiated as
earlyas I8 days beforefull bloom,but the wallsof the xylemele-
mentsdo not becomesufficientlythickenedto becomeconspicuous
until about the time of full bloom. They correspondto the ring
of "pit bundles"in the stony pericarpof the peach(I8).
As the fruitdevelops,the numberof bundlesin the ringis not in-
creased.Theproportion ofvasculartissueto thetotalareaof theperi-
carpas seenin crosssectionthus appearsmuchgreaterin the early
stagesthanin laterones,as shownin outlinedrawings(fig. 2). The
vascularsystem,moreover,is extendedprogressively throughoutthe
tissue as the fruit develops,largelyby transversedivergenceof
smallerbundlesfromthe mainbundles,similarto the conditionre-
portedin the peachby RAGLAND. At maturitythe vascularbundles
ramifythroughoutthe fleshypericarpto give a skeletonnetwork
of conductivetissue.
Throughout the development of the fruituntiljustbeforeripening
therearecertaincellswhichareconspicuous becauseof the granular
natureof theircontentsandtheirabsorptionof safraninandorange
G stains. They tend to forma loosenetworkthroughoutthe fleshy
pericarp,with greatestconcentrationaroundthe vascularbundles
and in a layerseveralcells thick just beneaththe outerepidermis.
Theyareslightlylargerthanthe averagefor cellsof the fleshyperi-
carp. In the ripe fruit the granularnatureof the contentsdisap-
pearsand the cellsbecomeindistinguishable fromthe adjacentpa-
renchyma.
EPIDERMIS
Theouterepidermisis a singlerowof cellscoveredexternallyby
a cuticlewhichis continuousexceptwhereinterruptedby stomata.
The cells of the epidermisare well diSerentiatedin materialcol-
lected I8 days beforefull bloom (fig. sA). At this time they are
elongatedradially,beingabouttwiceas longas broad.Thedevelop-
ment is similarto that in the peach as describedby DORSEY and
POTTER (7), exceptingthat thereareno hairs,of course.Duringthe
entireperiodtheyincreaseslowlyin size,chieflyin a radialdirection

I939] TUKEY & YOUNG CHERRY 74I
(fig. 5B). Eighteendays beforefull blooman averageof 3I2 cells

was countedin the peripheryof a crosssectionthroughthe cheek.
Two days beforefull bloomthis numberhad increasedto 390 cells.
DuringstageI (fig. 5C, D, E) the epidermalcellsincreasedecid-
edly in both numberand size. At I day afterfull bloom,550 cells
werecountedin the circumference of a crosssection. At the endof
stage I this numberhad reachedslightlymorethan double,II87.
The increasein numberstakes place very rapidlyin the first IO
daysof the stage;no divisionswereobservedin the latterpart. Cell
enlargement in both radialand tangentialdiametersoccursmostly
towardthe endof the period.The cuticleattainsfullthicknessdur-
ingthe latterpartof theperiod,andthe radialwallsareconsiderably
thickened.
DuringstageII (g. 5E, F, G) developmentof the epidermiscon-
sists in a slight tangentialelongationof the cells and additional
thickeningof the walls. No cell divisionsoccur.
DuringstageIII (fig. 5G, H, I) thereis greatenlargement of the
epidermalcellsin tangentialdirections,accompanying the largein-
creasein the surfaceareaof the fruit. Sincecell divisionceasesdur-
ing stageI, anyincreasein the sizeof the fruitmustbe compensated
for by a tangentialstretchingof the epidermalcells. Actuallythe
tangentialdiameteris morethan doubledduringthis period,while
thereis a slightdecreasein the radialdiameteras wellas in the thick-
ness of the cuticle. The drawings(fig. 5I) are similarto those by
DORSEY and POTTER (7) for the peach,and KOSEMANOFF (IO)for
the sweetcherry.
STOMATA.-The stomataare unevenlydistributedover the sur-
faceof the cherry. They arenumerousnearthe apexandfew near
the stemend. By I8 days beforefull bloomthey arefully differen-
tiated (fig.6A).
Sincethe epidermalcells increasemateriallyin numberduring
laterstagesandno morestomataareformed,they appearrelatively
numerousandclosetogetherin this earlystage. The guardcellsin-
creasein size as the fruit develops(fig. 6B, C) but the increaseis
muchless than that of typicalepidermalcells. At I8 days before
fullbloomthe guardcellsmeasureI3 p in length;at fullbloom,23 p;
duringthe middleof stageI, 25 p; at the endof stageI, 3I p; during

GAZETTE
BOTANICAL
742 [JUNE
II, 34 p; at the endof

middle
the of stageII andat the endof stage
III, 47-4g,u.
stage I to fruitripening,the
During aXstagesfromthe middleof stage very muchthinner
cellsare
of the cellsadjacentto the guard the ripefruit thereis no
walls
cells. In
thoseof typicalepidermal
than im-
developmentof chromoplastsin the guardcellsnorin the cells ap-
the cellcontents
adjacent.In freshunstainedmaterial brightredcolorof
mediately
in contrastto the
yellowishor straw-colored,
pear
cellswellsupplied with chromoplasts.
epidermal
S%> jt _s V N\Y
cells in sur-
guard cells, and adjacentepidermal
FIG. 6. Developmentof stomata, bloom;B: 20 days after full
A: I8 days beforefull
view
face (cameralucida X480).
full bloom.
C: fruit ripening,57 days after
bloom;
Thegeneralappearance resultingfromtheseareasof contrasting

seenwith the nakedeye, and
colorsis of dots whichmay be easily
and POTTER (7) describeda similarap-
suggests lenticels.DORSEY the
in the peach. No true lenticelshave beenobservedin
pearance mechani-
material studied.Occasional ruptureof the epidermisfrom
areaswerefound, result
the
calcauseshasbeenseen;anddiscoloredinjury. Even in ripe fruit
of spraymaterialsor somemechanical
on the treesuntil September20,
whichhadbeenallowedto remain werestill
60 daysaftercommercial harvest,stomataandguardcells
werefound.
intact (fig.6C) and no lenticels
Discussion
of
The shapeof the cellsof the fleshypericarpfromthe periphery
as seen in cross section,appears
the fruit to the stony pericarp,

I939] TUKEY & YOUNG-CHERRY 743
muchas it mightif each cell was subjectedto relativelyequalin-

ternalforceof expansionin all directions,suchas osmoticpressure.
The cellsnearestthe pit can expandvery little exceptin a radial
direction,becauseof the close positiontangentiallyof other cells
exertingsimilarpressurestowardthem. The cells surrounding the
firstones,besidesmakinga radialelongation,cannowexpandslight-
ly in a tangentialdirection,sincethe diameterof the fruit has be-
comegreaterandthe distancein a tangentialdirectionbetweenthe
theoreticalcentersof the rowsof cellshasbecomegreater,similarto
the increasein distancebetweenthe spokesof a wheelthe farther
fromthe hub. Progressivelytowardthe peripheryof the fruit the
tangentialpressurefromadjacentcellsbecomesless,so that the cells
assumean obovateshape,with the smallend pointingtowardthe
center.Finally,nearthe peripheryof the fruitthe internalcellpres-
sureis equallybalancedby the inwardpressureof the stretching
epidermisand hypodermallayer,by the outwardpressureof inner
cells,andby the tangentialpressureof adjacentcells,so that they
assumea roundishshape. In someinstancesthesecellsmay evenbe
broadlyoval. The cellsof the hypodermallayerandepidermis,be-
ing subjectedto outwardpressurefromthe expandingthin-walled
parenchyma, becomestretchedtangentiallyandflattened.
In figure7 is showna seriesof cameralucidadrawingsof the cells
in a radialsectorof the ripe fruit fromthe stony pericarpto the
outer epidermis,illustratinghow closelythe cell shapesand sizes
conformto the precedingdescription.ADDOMS, NIGHTINGALE, and
BLAKE(I) have remarkeduponthe radialelongationof cellsof the
fleshypericarpof the peachnearthe pit and the moreovoidshape
furtherout.
The grosssize changesin the fruitduringthe threestagesof de-
velopmentare seenfromthis study to be madeup of cell divisions
and cell enlargements in varyingproportionsin differenttissuesat
differenttimes. Figure8 showsthe amountof increasein a trans-
versedirectionthroughthewallof the fruitat the cheek. It hasbeen
preparedby calculationfromthe actualnumbersandsizesof cellsin
the differenttissuesat thebeginning,middle,andendof eachgrowth
stage (tablesI and 2). Whilethe generalnatureof the curveis sim-
ilarto that for the total diameterof the fruitin figureI, it diSersin

FIG. 7. - Size and shape of cells involved in final swell of fleshy pericarpin radial
directionthroughcheek of sour cherryat fruit ripening(end of stage III), showing
transverseelongationin epidermisand hypodermallayer (A); roundishshape in next
underlyingregionbetweenhypodermallayer and ring of vascularbundles (B); oval
shapeandbeginningof radialelongationin next underlyingregion(C); long oval shape
in next underlyingregion(D); and decidedradialelongationin inner region (E). In-
most layer of smallcells adjacentto stony pericarpnot shown (cameralucida X I40).

PRE-BLOOM / ' STAGE I [ STAGEff , STAGE /
I939] TUKEY & YOUNG CHERRY

745
that it representsonlythe actualincreaseof the tissuesof the ovary
wall,whereasfigureI alsoincludesthe ovariancavity besidesboth
sidesof the ovary wall. Sincethe increasein the diameterof the
ovariancavity duringstageI does not involvea corresponding in-
creasein tissue,the steepnessof the curvein stage I of figure8 is
muchless than that of figureI. The figuredoes showgraphically
the relationbetweencell divisionandcell enlargement andthe part
playedby both in the growthof the cherryfruit.
M/CR0fl/5 , I i R/PE-27
8000 1 : I /
, : , /
, [ I /
, i ,. /
, ; ', /
6000 : t I /
I I ' /
I I I /
1, 1 W /
4000 /
2000 <[[11 10:l 10
v z CELL
ENLARGEMENT
9 0 /1 20 26 30 36 X 50 SZ
FIG.8. Increasein thicknessof ovarywall of sourcherryat cheek,frompre-bloom

to fruit ripening,showingamountdue to increasein numberof cells and amountdue
to enlargementof cells alreadypresent. Verticallines= size changesdue to cell enlarge-
ment. Crosshatching=size changesdue to cell division.
The similarityof developmentof the sourcherryto the develop-

mentof otherdrupefruitshas beennoted. The studiesof TSCHIER-
SKE(I9) withthe redraspberry areparticularlynoteworthy.He ob-
servedtwoperiodsof rapiddevelopmentin the drupelets:a firstone
earlyin the seasonduringwhichcell divisionmainlyoccurs,and a
secondjust beforefruitripening,in whichstretchingof the cellsoc-
curs. Particularlyinterestingis his observationthat the finalswell
beginswhenthe structuresof the seed and surrounding tissuesare
differentiated,
as notedforthe cherryandpeachby TUKEY(2I, 22).

[JUNE
At maturitythe fruitbearsa closeresemblance to the descriptions
given by TSCHIERSKE (I9) for the drupeletsof Rubus ideaus, by
FARMER (8) for R. fructicosus,and by WINTON (24) for R. strigosvs
andR. occidentalis.Further,WINTON mentionsa strikingsimilarity
betweenthe red raspberryand the peach.
Summary
I. Thispaperpicturesanddiscusses the grossdevelopmentof the
fruitof the sourcherry(Prunus cerasusL. var. Montmorency) from
I8 days beforefull bloom to fruit ripening,and the histological
changesduringthe pre-bloomstage,stageI (rapiddevelopmentfor
20 days followingfull bloom),stage II (retarded developmentfor
I6 days), and stage III (rapiddevelopment for 2I days to fruit
* .
rlpenlng .
Three principaltissues composethe ovary wall: inner and
2.
outerepidermis,stonypericarp,andfleshypericarp.Thestonyperi-
carpmaybe dividedinto an innerandan outerlayer,andthe fleshy
pericarpinto an innermostlayerof smallthin-walledparenchyma, a
middleregionof largethin-walledparenchyma, and an outeror hy-
podermallayerof collenchyma.
3. The fleshyand the stony pericarpare derivedeachfromdis-
tinct groupsof cellswhichareearlyseparatedfromone anotherby
characteristicsize,shape,andfrequencyandperiodicityof cell divi-
sion.
4. The innerlayerof the stonypericarpis derivedfromthe inner
epidermis,togetherwith a few adjacentcells of the pericarp,and
formsa bandor "hoop"of transverselyelongatedcellsboundingthe
innerovarywall. The outerlayeris derivedfromthe pericarp,and
the cells of whichit is composedare elongatedat right anglesto
thoseof the innerlayer.
5. The cellsof the stonypericarpincreasein numberduringthe
pre-bloomstageandthe firstfewdaysof stageI, andenlargeduring
the latterpartof stageI. Cellwallsbecomeprogressivelythicker,
andby the endof the periodthe maximumnumberandsizeof cells
is attained. DuringstageII the wallsthickenand hardengreatly.
DuringstageIII thereis slightincreasein hardnessandbrittleness.

I939] TUKEY & YOUNCHERRY
747
6. The cellsof the fleshypericarpincreasein numberduringthe
pre-bloomstage and the first half of stage II. In the last half of
stageII they doublein diameter.DuringstageII thereis slighten-
largement.DuringstageIII the sizeof individualcellsincreasesre-
markably.Those of the hypodermallayer becomeenlargedin a
tangentialdirectionsimilarto the adjacentcells of the outer epi-
dermis. Thosein the outerportionof the fleshypericarpbecome
roundishoval, with the greatestdiameterparallelto the periphery
of the fruit;thosenextinwardbecomeroundish;thosenext,obovate
in a radialdirection;next,radiallyelongate;andinnermost,decided-
ly radiallyelongate. At maturitythe largestcells indicatean in-
creaseof twenty-fivetimesin diameterfromthe size at full bloom.
7. The epidermalcells are elongatedradiallyI8 days beforefull
bloom. Theyincreaserapidlyin numberduringthe pre-bloomstage
andduringthefirsthalfof stageI, increasein sizeandwalltliickness
duringthe latterhalf of stage I, changebut little duringstage II,
andgreatlyenlargetangentiallyin stageIII.
8. The stomataarefullydifferentaated I8 daysbeforefullbloom.
Theguardcellsincreasein sizeas the fruitdevelops,but the increase
is less thanthat of typicalepidermalcells.
9. The similarityof other fruits and the mechanismof enlarge-
ment in stage III arediscussed.
NEW YORKSTATEAGRICULTURAL
EXPERI1WENT
STATION
GENEVA,NEW YORK
LITERATURE CITED
I. ADDOMS, RUTHM., NIGHTINGALE, G. T., and BLAKE, M. A., Development
and ripeningof peachesas correlatedwith physicalcharacters,chemical
composition,and histologicalstructureof the fruit flesh. II. Histologyand
microchemistry.New JerseyAgr. Exp. Sta. Bull. 507. Ig30.
2. BLAKE, M. A., Growthof the fruit of the Elbertapeachfromblossombud
to maturity. Proc.Amer.Soc. Hort. Sci. 22:29-38. Ig2s.
3. BLAKE, M. A., DAVIDSON, 0. W., ADDOMS, RUTHM., and NIGHTINGALE,
G. T., Developmentand ripeningof peachesas correlatedwith physical
characteristics,chemical composition,and histologicalstructureof the
fruit flesh. I. Physicalmeasurementsof growthand flesh texturein rela-
tion to the marketand ediblequalitiesof the fruit. New JerseyAgr.Exp.
Sta. Bull. 5 2 5. I 93I .

.
GAZETTE
BOTANICAL [JUNE
748
developingand aborting
DOROTHY, A comparativestudyof the
BRADBURY,
4. Jour.Bot. I6:525-542. I929.
fruitsof Prunus cerasus. Amer. JerseyAgrAExp. Sta.
C. H., Growth of fruitsof the peach. New
5.CONNORS,
Ann. Rept. 40:82-89. I9I9. L., The developmentof the I926.
peachseed
6.DORSEY, M. J., and McMuNN,R. 23:402-4I4.
Proc. Amer.Soc. Hort. Sci. of the skinand
inrelationto thinning. J. S., A study of the structureAgr. Exp. Sta.
DORSEY, M. J., and POTTER,
7. to bruising.Illinois
pubescence of the peach in relation
Bull.385. I932. physiologyof pulpy
J. B., Contributions to the morphologyand
FARMER,
8. I889.
fruits.Ann. Bot. 3:393-4I4. study of the developingand
abortingfruits
HARROLD,
9. T. J., A comparative
GAZ.96:505-520. I935.
of Prurlus persica. BOT. Eigentumlichkeiten im Bau derHaut ver-
KOSEMANOFF, S., Anatomische Sta. Sektion
IO.
Sorten von Kirschen. Arb.Mleew.Gartenbau-Vers.
schiedener
Obstbau.No. 22. I929. der Entwickelungsaftiger
LAMPE, PAUL, Zur Kenntnissdes Baues und
II. 59:295-323. I886.
Fruchte.Zeitschr.Naturwiss. the apricotfruit. Proc. Amer.Soc. Hort.
LILLELAND,
I2. O., Growthstudy of
BC1.27:237-245. I93I- Amer.Soc. Hort. Sci. 29:
, Growth study of the peachfruit. Proc.
I3.
changesin
8-I2. I933-
of the plum fruit. I. The growth and
I4. , Growth study Amer. Soc. Hort. Sci.
composition of the Climax plum. Proc.
chemical
30:203-208. I934. of the Hiley peach
R. V., The growth rate and chemicalcomposition Soc. Hort. Sci. 29:I-7.
I5.LOTT, maturity. Proc.Amer.
fromstoneformationto flesh
reference
I933-
on the development of peachfruitswith special
I6.MIKI, T., Studies I932.
ChibaColl. Hort. Bull. I:I-II8. reference
to the causesof their split-pit. fruit with special
RAGLAND, C. H., The developmentof the peach 3I:I-2I. I935.
I7.
Proc. Amer.Soc. :Hort.Sci.
to split-pitand gumming. anatomical developmentof the apple
and
URSULA, A study of the walls. Jour.
I8. TETLEY, the cell
"pectic constituents"of
some observationson the I930.
Pom. & Hort. Sci. 8:I53-I72. vergleichendenAnatomie
und Entwick-
PAUL, Beitrage zur 59:580-628.
I9. TSCHIERSKE, Zeitschr.Naturwiss.
lungsgeschichte einigerDryadeenfruchte.
I886. of Prurlusaviuml.
H. B., Embryo abortion in earlyripeningvarieties
20. TUKEY,
I933-
BOT.GAZ.44:433-468. to growthof fruit and
, Growth of the peach embryo in relation I934.
2I.
Amer.Soc. Hort. Sci. 30:209-2I8.
seasonof ripening.Proc.

22. , Growthof the embryo,seedandpericarpof the sourcherry(Pruxus

cerasus)in relationto seasonof fruit ripening.Proc.Amer.Soc Hort. Sci.
3 I : I 25-I 44. I 935.
23. TUKEY,H. B., and LEE, F. A., Embryoabortionin the peach in relation
to chemicalcompositionand seasonof fruit ripening.BOT. GAZ.98:586-
597. I937.
24. WINTON, A. L., The anatomyof edibleberries. ConnecticutAgr.Exp. Sta.
26th Ann. Rept. 283-325. I 90 2.


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HISTOLOGICALSTUDY OF THE DEVELOPIN7G

FRUIT OF THE SOUR CHERRY

Botanical Gazette 1939.100:723-749.

FIG. I. Growth stages in development of fruit of sour cherry from pre-bloom to

Botanical Gazette 1939.100:723-749.

3. Seasonof I932, dailyfromJune4 to June 26, in formalin-acetic

Botanical Gazette 1939.100:723-749.

sectionsthroughthe entirepericarp,freehandsectionsfor detailed

Botanical Gazette 1939.100:723-749.

grossdevelopmentof the fruit and of these tissuesis shownin the

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

FIG.4.-Development of fleshy pbricarp,in regionbetweenhypodermallayer and

Botanical Gazette 1939.100:723-749.

FIG. 5.-Development of epidermisand hypodermallayer (cameralucidadrawings

Botanical Gazette 1939.100:723-749.

939] TUKEY & YOUNG CHERRY 733

FLESHY PERICARP STONYPERICARP

Pre-bloom........... -I8 4-5 II ....... ... I5-I6 4-5

End of stageI . . . +20 5- 6 I8-I9 3 4 26-29 I7-I8 6- 7 23-25

Botanical Gazette 1939.100:723-749.

whoselongestaxis runsin a directionat rightanglesto that of the

Botanical Gazette 1939.100:723-749.

tinct groupsof cellswhichareearlyseparatedby characteristic size,

Botanical Gazette 1939.100:723-749.

tive celldivisionoccurshereas late as if not laterthan,in anyother [JUNE

partof the pericarp.Thismaybe explainedby the fact that harden-

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

lihoodfor the inclusionof intercellularspacesin a crosssection. Or

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

(fig. 5B). Eighteendays beforefull blooman averageof 3I2 cells

Botanical Gazette 1939.100:723-749.

II, 34 p; at the endof

Thegeneralappearance resultingfromtheseareasof contrasting

Botanical Gazette 1939.100:723-749.

muchas it mightif each cell was subjectedto relativelyequalin-

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

I939] TUKEY & YOUNG CHERRY

2000 <[[11 10:l 10

FIG.8. Increasein thicknessof ovarywall of sourcherryat cheek,frompre-bloom

The similarityof developmentof the sourcherryto the develop-

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

Botanical Gazette 1939.100:723-749.

22. , Growthof the embryo,seedandpericarpof the sourcherry(Pruxus

Botanical Gazette 1939.100:723-749.

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