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STAMINATE STROBILUS OF TAXUS CANADENSIS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 255
A. W. DUPLER
Introduction
In a previouspaper(8) thewriter describedthegametophytes of
Taxus canadensisMarsh.,withthestatementthatotherphasesof
themorphology wouldbe treatedinlaterpapers. In thispaperthe
staminatestructures with respectto developmentand vascular
anatomyare described. The lack of detailed information con-
cerning thesestructureshas seemedto thewritersufficient
justifica-
tionfortheinvestigation herereported. In viewof thegenerally
recognizedconservativecharacterof the staminatestructures in
conifers,it seemsthata moreextended ofthem,in the
investigation
groupas a whole,wouldbe worthwhile. The description of the
ovulatestructures willbe givenin anotherpaper.
The generalstatement in thepreviouspaperas to materialand
methodswill also apply here. The writeris underobligationsto
ProfessorW. L. EIKENBERRY,of theUniversityof Kansas, forsome
materialcollectedin northernIllinois a numberof years ago.
Acknowledgmentsare also due ProfessorsJOHNM. COULTERand
C. J. CHAMBERLAIN, under whose direction the study of Taxus
canadensiswas begun.
Historical
Whilethe male gametophyte and its attendantfeatureshave
receivedconsiderableattention,apart fromthe generalmore
obviousfeaturesverylittleis foundin theliteraturedealingwith
Taxusas to themorphology ofthestaminatestrobilus itself. The
earlierworkerswho studiedthe staminatestructures of conifers
wereconcerned largelyinattemptsto interpretthemin termsofthe
angiosperm flower,naturallyleadingto confusion as to the true
natureof the structures.These earlierviews have been sum-
marized by VONMOHL (26) in perhaps one of the most important
345] [BotanicalGazette,vol. 68
346 BOTANICAL GAZETTE [NOVEMBER
layersofirregularcells,andstatesthatdehiscence is accomplishedby
theruptureofcellsat thebase and sidesofthepollensac. CHAM-
BERLAIN(3) describesthemicrosporangium of T. canadensisat the
mothercell stage(Octoberi, i897), at whichtimethenucleiare
stillrathersmallin comparison withthesizeofthecell,thetapetum
beingsharplydifferentiated, and its cells showingno tendencyto
plasmolyzelike the cells of the sporangiumwall. PILGER(i8)
describesthegeneralexternalfeatures, largelyfromthetaxonomic
viewpoint,speakingof the "flower"as consistingonlyof sporo-
phyllssurroundedat the base by a scale envelopewhichcom-
pletelyinclosestheflower in thebud state. He regardsthe"leafy
structure"of theanther,whichis yetto be recognized in Torreya
and Cephalotaxus, as being"entirely lost" in Taxus.
In the relatedformsthestaminatestructures of Torreyahave
been describedin a general way by PILGER,based on T. nucifera;
in more detail by Miss ROBERTSON (20) for T. californica; and
by COULTER and LAND (5) for T. taxifolia. In Cephalotaxus
have beendescribed
someofthefeaturesofthespermatogenesis by
STRASBURGER (23), and by ARNOLDI (I). STRASBURGER (24)
pointedout thatthepollengraindividesin thesporangium
before
shedding; LAWSON(14) also confirmsthis in C. drupacea; and
WORSDELL (27) of the generalfeaturesof the
gives a description
"male flower,"based on C. Fortunei,comparingit withthoseof
otherforms(Phyllocladus especiallyin thesporophyll
and Ginkgo),
features.
Strobilibuds
In theaxilsoftheleavesoftheshootofa givenseason3 types
of structures are produced: (i) the vegetativebuds fromwhich
developthelateralleafyshootsof thenextseason; (2) theyoung
staminatestructures, maturing thenextseason; and (3) theovule-
also
bearingstructures, maturing thenextseason. Duringthefirst
seasonall of thesestructuresare in bud form,the staminatebuds
duringthelatterpartofthesummer andwinterbeingmoreglobular
than the othertwo kinds,whichare so nearlyalike in external
appearanceas to maketheirdistinction uncertainexceptby very
carefulexamination.
348 BOTANICAL GAZETTE [NOVEMBER
Sporophylls
PRIMORDIA
MICROSPORANGIUM
ARCHESPORIAL INITIALS.-HOFMEISTER(II) seems to have been
the first to publish with referenceto the microsporangiumof
conifers,reportingthe spore mothercell stage as being reached in
Pinus maritima in November. GOEBEL (9) traced the arche-
sporiumofPinus to a singlehypodermalcell, and claimed a similar
originforthe archesporiumof Thuja. His mostimportantobserva-
tion on thispointwas that the developmentofthe microsporangium
is like that of the eusporangiateferns. COKER (4) in Taxodium
distichum,and NICHOLS (I7) in Juniperuscommunisvar. depressa,
also found a hypodermal origin of the archesporium,in the
latter case consistingof "a plate of radially elongated cells, 4-6 in
I919] DUPLER-TAXUS 35I
EPIDERMIS
100
15.-
t
- ----- - -- - (
t7~~~~~~~~~~~~~~~~o
20----- ~ ~ ----- O
21.--. e
221-__ __ _
9 2
~~~~~~~~~~~1
FIGS. 9-i2.-Fig. 9, median longitudinalsection of maturestrobilusjust before
pollen shedding,showing"elongating region" of axis and 4. sporophylls; xylemof
bundle black; note that xylembecomescentrallyplaced in upper portionsand does
not extendas farintostalksas phloem; numbersat left(W3-22) indicateapproximately
levelsofcross-sectionsofstrobilusshownin figs.I3-22; fig.IO, mediansectionofopen
sporophyll,showingelongatedstalk,open sporangia,and solitarystoma in centerof
depressionofdisk; fig.II, tangentialsectionof matureepidermis,showingmechanical
thickeningson walls; fig. i2, cross-sectionof matureepidermis,with microspore,at
timeof shedding; figs.9, IO, X36; figs.II, I2, X475.
19191 DUPLER-TAXUS 357
Vascular features
Since the reproductiveorgans, and especially the staminate
structures,are regarded as among the most conservativeof plant
organs,a considerationof the vascular anatomy of the staminate
strobilus is not without interest. While the ovulate strobili of
conifershave been the subject of considerable investigationand
discussion,in theirvascular as well as in otherfeatures,the stami-
nate strobili have not received much attention in their vascular
anatomy, probably not as much as they deserve in view of the
conservativenature generallyassigned to them on other grounds.
The only reference to this feature of Taxus is by STRASBURGER(22),
who gave the arrangementof the scales of T. baccata and states
that each stamen contains a bundle which passes into the stalk.
Like any otherbranch,the strobilusaxis receives2 bundlesfrom
the cylinderof the leafy shoot. These are semicircularin outline,
and by meetingat theiredges soon forma closed cylinder,broken
here and thereby the gaps formedby the weak bundle tracesof the
scales. In the lower portionof the strobilus,where the scales are
small and decussate, the small traces oftenend in the cortex and
do not reach the scale itself. The traces for the upper scales are
betterdeveloped and extendforsome distanceinto themidribof the
scale, especiallyin the 2 or 3 uppermostscales. Althoughthe axis
cylinder,as well as the cortical portion of the scale traces, are
1919] DUPLER-TAXUS 359
Gd Q
14 15 16
0/k
Genj
Oi9 ced (>04
17 18 19
e~~~~~~~~~~~~~~~~~~ln
20 21 22
FIGS. 13-22.-Cross-sectionsofmaturestrobilusat approximately levelsindicated
by numbersto leftof strobilusshownin fig.9; branchesof bundlesto various sporo-
phyllsindicatedby a, b, c, etc., xylemindicatedby black; bundles a and b supply
terminalsporophylls;union of bundles indicatedby combiningletters,as jd in figs.
I7-I9; fig.i9 showscompletecylinderbelow lowermostsporophyll;in fig.i8 1and c
united,in fig.i9 separated,in fig.20 c and m united,and in fig.22 lcn one of the 3
large strandsfromsterileportionof axis; fig. 22 also shows traces to 3 uppermost
scales, Sci, SC2,and sc3; note concentriccharacterof terminalportions,as in c, d, and
e, in fig.I5; X36.
onlyofthephloemportion(fig.39), thexylemusuallyendingwithin
thecortexoftheaxis.
The bundlesof thisregionare collateralendarchin the lower
portions. In theupperportions, however,thebundlesfrequently
showcentripetalxylem(figs.40, 44), givingmesarchbundles,and
19191 DUPLER-TAXUS 36i
Discussion
Perhaps the two most important features of the staminate
strobilusof Taxus are the peltate sporophyllsand the characterof
the vascular bundles of the scale and sporophylls. The peltate
(epaulet) type of stamen occurredamong the Paleozoic Cycado-
filicales,in the Crossothecaforms,but the sporangia were bilocular
and dehisced by a longitudinalslit along the adaxial face, the
bilocular character being differentfromthat of the moderngym-
nosperms. Peltate stamens are not known in Bennettitales,and
none occur in the Cycadales. The peltate stamen has been carried
forwardto modern plants throughthe Cordaitalean line, in all
probability,althoughso faras is known the stamensin the Cordai-
tales bore terminal erect sporangia. As COULTER and CHAM-
BERLAIN state, however,"it cannot be supposed that the stamens
of so great a group were uniformin type," and it is very possible
that peltate stamens occurred there also. The sporophyll of
Ginkgogives a suggestionof thepeltate type of stamen,in occasion-
ally havingmorethan 2 sporangia,in the regularoccurrenceofmore
than 2 sporangiain fossilforms,and in the possibility,pointed out
by Miss STARR,that the mucilage cavity replaces abortive spo-
rangia. Among Coniferalesthere is a suggestionof the peltate
stamenin the Araucarineae,and stamensof truepeltate formoccur
in such formsas Widdringtonia,Torreya,and Taxus. In Torreya
the truepeltate characteris generallyobscuredin the adult sporo-
phyll owingto the developmentof the resincavity from3 of the 7
sporangiumbeginnings. Hence it is seen that peltate stamens,in
one formor another,are scatteredfromCycadofilicalesto modern
conifers,and there is no necessityof regardingsuch a sporophyll
as that of Taxus as being of recentevolution. Assumingpeltate
362 BOTANICAL GAZETTE [NOVEMBER
Summary
i. The staminate strobilioccur in the axils of the leaves. The
buds can firstbe distinguishedfromother types of buds by the
broad apex.
r9g9] DUPLER-TAXUS 363
LITERATURE CITED
i. ARNOLDI,W., Beitrage zur Morphologie der Gymnospermen.III.
Embryogenie Fortunei. Flora 87:46-63. p/s.I-3. 1900.
von Cephalotaxus
2. CELAKOVSKY, L., Die Gymnospermen:einemorphologisch-phylogenetische
Studie. Abhandl. K6nigl. Bohm. Gesell. Wiss. VII. 4:i-48. i890.
3. CHAMBERLAIN, C. J., Wintercharactersof certainsporangia. BOT. GAZ.
25:I25-I28. pl. ii. I898.
4. COKER,W. C., On the gametophytesand embryoof Taxodium. BOT.
GAZ.36:I-27, II4-I40. pls. I-II. I903.
364 BOTANICAL GAZETTE [NOVEMBER
PLATE XXIV
PLATE XXVI
\J~k~0OQ
~ 24 25 26
0 27~~0
0 ~ ~ ~ ~ 0
~~~~ 3
33~~~~43
DUPLER on TAXUS
BOTANICAL GAZETTE, LXVIII PLA TE XXV
370
40
39am4
0~~~~~~~4
41
DUPLER on TAXUS
BOTANICAL GAZETTE, LX VIII PLA TE XXVI
45
48~~~~4
DUPLER on TAXUS