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 THE POLLINATION OF VALLISNERIA SPIRALIS
ROBERT B. WYLIE
(WITH PLATE IX AND SIX FIGURES)

Vallisneria has long been counted one of the classic examples

of cross-pollination. Living vegetativelyas a submersedaquatic,
its dioeciousflowersare broughttogetherat the surfaceof the water
in mostingeniousfashion. These highlyspecializedflowerspresent
the strongestcontrasts, not only in size and structure,but in
behavior as well, and give this plant its rank as one of the climax
types with respect to floral differentiation.Specializations of
such evidentadvantage forcross-pollinationin a formso admirably
situated forvegetative propagation seem to emphasize the impor-
tance of sexuality, or at least of seed production,in the higher
plants.
While the general method of pollination in Vallisneria is well
known,many interestingfacts seem never to have been published,
and the underlyingprinciple has not been emphasized. The
figurescurrentin textbooks are highly generalized, and some of
them are far fromaccurate. The story which they are intended
to illustrateis likewiseincompleteor in some cases highlydistorted.
In any event, neitherfigurenor storyhas done justice to the inti-
mate historyof pollen transferin this remarkableplant.
It will be noted at once that the followingaccount diverges
radically from that suggested by KERNER'S (i) beautiful and
widely copied figure. A comparisonshows that these differences
relatenot only to the size and structureof the flowers,but are even
more fundamental in character. KERNER emphasizes the fact
that pollination is brought about throughthe contact of flowers
floatingon a level water surface; there follows an outline of a
methodof pollen transferthroughthe special agency of the surface
filmof water. The general drawing (pl. IX) is based on photo-
graphs of living flowers,measurements,and camera drawings of
parts.
'351 [BotanicalGazette,vol. 6 3
I36 BOTANICAL GAZETTE [FEBRUARY

The epigynous seed-bearing flowersof Vallisneria are borne

singly,each withinits spathe at the end of a long scape, sometimes
over a meterin length,whichanchorsthefloatingflowerto theshort
uprightstem at the bottom of the pond. Upon reachingthe sur-
face of the water throughthe elongation of this axis, the spathe
opens at its outer end, but remains as a partial investmentof the
ovary until the seeds are nearly mature. The 3 spoonlike sepals
soon separate,disclosingthe 3 bifidstigmaswhich are coiled in the
center of the flower(pl. IX). These fleshystigmas are densely
clothed with the stigmatichairs, and their snowy whitenesscon-
stitutes the most conspicuous part of the flower. Rudimentary
petals and slenderstaminodia are present,but as they seem func-
tionlesstheirdiscussionmay be deferredto a subsequentpaper.
The anchoringscape usually elongatessufficiently to permitthe
opening flowerto assume an inclinedposition in the water as it is
carried to one side by wind or current. The ovary, which is
20-25 mm. long before fertilization,is usually straightuntil the
floweropens and has taken its position at the surface; later it
often curves considerably in response to gravity, thus bringing
the floralparts more nearlyparallel with the surfaceof the water.
This bending of the ovary at this stage is quite marked in plants
growingin aquaria where the flowersare leftundisturbedforsome
time.
The exposed floralparts are waxy and consequently are not
wetted by the water, with the result that the flowercomes to rest
with a portion of its weight restingon the sepals and marginsof
stigmas supported by the surface film. This produces a slight
depression of the water about the flower,perhaps I5 mm. in
diameter,which is abruptly declined at its inner margin next to
the pistillateflower. This sloping surfacefilmplays an important
part in capturing the floating staminate flowers,and later is
intimately bound up with the actual transferof pollen to the
stigmas. Too much emphasis cannot be laid on the complete
dependence of this plant upon the surface film of water for its
pollinationprocesses.
The staminate flowersare borne crowded numerouslywithin
the globose spathe which remains short-stalkedat the bottom of
1917] WYLIE-VALLISNERIA I37

the pond. A count of several of these flowermasses showed an

average of over 2000 flowerspacked withineach spathe, the whole
group the homologueof the singlepistillateflowerwhichis solitary
withinits spathe. The staminate inflorescenceresemblesa large
fernsorus surroundedby an indusium. This similarityis carried
furtherby the strikingresemblanceof the slender-stalkedunopen
staminate flowersto polypod fernsporangia. Massed within the
spathe these flowersare joined to the axis by slender pedicels of
varying length, so as to completely fill the space between the
stem and the spathe.
The pollen-bearingflowersare very tiny, less than i mm. in
diameterbeforeopening,and are simple in structure. The floral
parts consist of 3 sepals, 2 functionalstamens, and rudimentsof
petals. The sepals are of unequal size and are not symmetrically
disposed; 2 are similarand stand nearlyopposite; while the third
and smaller one is placed laterally between them. This reduced
sepal is the firstto open. Numerous tapering and curved hairs
cover the regionabout the base of the stamensand are doubtlessof
some importance, although their functions are not clear. The
2 stamens stand close togetherand have their parallel filaments

unitedup to a point near the anthers (pl. IX).

At maturitythe tip of the spathe opens slightlyand the stami-
nate flowersbegin detachingfromtheirslenderstalks. The upper-
most are the firstto be shed, and 2 or 3 days may be necessaryto
emptya singlespathe. These detached flowersrise slowlythrough
the water to the surface and there very slowly open. In this
respect Vallisneria stands in sharp contrast to the writer's (2)
observations on Elodea canadensis. In that form the staminate
flowersupon release dart to the surface and there fairlyexplode,
scatteringtheir pollen on the surface of the water. In Elodea,
however,it is the freefloatingpollen that functions,whilein Vallis-
neria the pollen retained in the anthers has the better chance of
reachingthe stigmas. SVEDELIUS (3) reportsthat the sepals of the
detached staminate flowersof Enalus acoroides snap back upon
reachingthe surfaceof the water, although the pollen is retained
in the anthers. In Elodea canadensis,and perhaps in Enalus, the
snap of the sepals seems to be due in part at least to the release of
I38 BOTANICAL GAZETTE [FEBRUARY

gases imprisonedbetweenthe floralparts underwater. The writer

(4) has noted elsewherethat in Elodea ioensis, which has a long-
stalked staminate flower,a bubble of gas is generally associated
with the partly opened sepals, giving extra buoyancy to the sub-
mergedflowers,which tug at theiranchorage like captive balloons.
No prolonged observations were made on the possible perio-
dicityin thereleaseof the staminateflowersof Vallisneria,although
doubtless there is a relation between their detachment and the
gases given offby the plant duringtimes of brighterillumination.
On one occasion it was observed that as the sun came up from
behind a building and its direct rays fell on the spathe of the
staminate inflorescencethe rate of detachment was considerably
increased for a time. In Elodea canadensis (2) there is a correla-
tion between the coming of stronglight in the morningand the
rate of detachmentof the staminate flowers. SVEDELIUS reports
that the staminate flowersof Enalus acoroidesare released mainly
(if not exclusively)at periods of low tide. This habit is of peculiar
significancefromthe fact that at high tide the pistillate flowersof
that plant are wholly submergedand pollinationwould be impos-
sible. No explanationof this relationwas suggestedin the paper.
The sepals of the staminate flowerof Vallisneria completely
invest the stamens until some time after the flowerreaches the
surface. They then slowly recurve, the smaller one being first
to open (pl. IX), and as it touches the water it seems to function
in orientingthe flowerso that when the pair of lateral sepals open
there are formed3 boatlike structureswhich engage the surface
film and float the flower. This tiny flower,with its upraised
stamens and pollen mass, is so snuglyfittedto the surfacefilmby
its 3 broad areas of contact that it is kept in equilibriumunder
all ordinarycircumstances. They are rarelyoverturned,even by
rathervigorousagitation of the water, but maintain a strictright
angle to the surfacefilm. So slenderan object as a needle if thrust
into the water among these floatingflowersand slowlywithdrawn
will be covered by the flowersthat have been drawn up with the
film of water about the needle and may be seen standing out
radially fromit on all sides. Once overthrown,however,they are
not again righted,but lie partly under water.
1917] WYLIE-VALLISNERIA I39

This definiteengagement withthesurfacefilmdoes nothinder

the freemovementof the staminateflowerson the water. In
openareastheyare caughtby everypassingbreezeand are hurried
alongthesurfaceof thewater. On windydays theygo scudding
by the observerlike tinyflecksof foam. Wheretheplantsgrow
abundantlytheyoftenmass along the windwardshoresin broad
zonesofsnowywhite(fig.i).
The anthersdehiscebeforetheflowersopen,and thestickypollen
fromthepair of stamensof a givenflowerusuallyformsa single
pollinium(pl. IX). Even if theproductsof the 2 anthersform

_~

FIG. i.-Floatingstaminate
flowers
alongmargin ofEast OkobojiLake in north-
western
Iowa; thedeadfishshownnearcenterofpicturewasabout8 incheslong.

separatepollenmasses,theselie so close togetheras to be prac-

ticallytangential,
and are neverwidelyseparated,as shownby
KERNER (I). Seen under moderate magnificationthese masses
ofpollengleamlikeclustersofpearls. The microspores, whichare
slightlyoval in form,are about65 juin diameterand have a nearly
smoothexine. Theiradhesiveness is ofgreatadvantagein holding
thepollentogether untilit is rubbedoffagainstthe stigmas,and
doubtlessassistsafterpollinationin holdingthesporesto thestig-
matic hairs. The pollenoutputis limited,averagingabout ioo
sporesto theflower, but varyingconsiderably.
The floatingstaminateflowersare carriedalongby the wind,
and comingwithintheradiusofthedeclinedsurfacefilmaboutthe
pistillateflowerslide into the littledepression,wherethey are
I40 BOTANICAL GAZETTE [FEBRUARY

retained. In thismannerpossiblyas manyas 50 staminateflowers

may be caughtin a singledepression,thus forming conspicuous
whitepatcheson the surfaceof the water. It is interesting to
notehowsuccessfully theseareasofassociatedflowersholdtogether
evenwhenthewateris quite rough. COWLES(5) mentionsthese
filmpockets,but givesno detailsotherthanto comparethemwith
thoseof Elodea canadensis. In this
latterform,as previouslydescribed
(2), the floatingpollen grains are
caught in the depressionsthat are
formed about the tiny pistillate
flowers. SVEDELIUS speaksof a sim-
ilar"capturing"ofthepollen-bearing
but does
flowersin Enalus acoroides,
of
not attributethisto the influence
the surfacefilm,althoughobviously
the case closely parallels that of
Vallisneria.
The sepals of the innermostof
the captured staminateflowersin
Vallisneria are of course in contact
on
FIG. 2.*-Flowersfloating with the marginsof the pistillate
surfaceof waterin a smallaqua- flower(fig. 2), but later arrivalsare
rium surroundedby black paper;
pistillateflowerin center. heldback as theyformonlya single
layer in the depression. It should
be notedat thistimethatcontactbetweenflowers on a levelwater
surface, suchas KERNER figures, as
couldnotlead topollentransfer,
thepolliniaareupraisedoverthecenteroftheflower.Butwithany
slightdeclination ofthefilmaboutthepistillateflower, eveninquiet
water, theremightbe contact betweenthe innermostpollinia
and the stigmas (fig. 2). Obviously,however,any movement
resulting in a further depressionofthepistillateflowerwouldcause
the surfacefilmto becomemoreabruptlydeclined,thus tipping
the staminateflowersmoresharplyinward(fig.3) and thereby
* FIGS. 2-6 constitute a series illustratingchanges in relations of associated
flowersfloatingat surfaceof waterwhensubjectedto submergenceby pullingon scape
of pistillateflower.
19 17] WYLIE-VALLISNERIA 14I

makingconditions morefavorable forpollentransfer.The upward

movementof the waterdue to a passingwave mightserve to
temporarily depressthe floatingpistillateflowerweightedby its
longstalk. Shouldthemovementbe sufficient to make taut the
anchoring scape,even for an the
instant, depression wouldbecome
cuplike,withtheinnerstaminateflowers standingat rightanglesto
its nearly vertical walls (fig. 4).
Many of the pollen masses would
thusbe forceddirectlyinto the stig-
mas, althoughthe outerones would
stillbe held back at some distance.
At a certain stage of depression,
however,the lateralpressureof the
waterbreaksthe surfacefilmabove
theflower;thesidesof the cup snap
together, roofingit over; and a con-
siderablenumber ofstaminate flowers,
with the pistillateflower,are thus
shut tightlytogetherin a common
bubble beneath the surfaceof the
water (fig. 5). It should be noted
thatthisprocesshas completely over- FIG. 3.-Positionsassumedwith
turnedthe
thestaminateflowersand on scapeofpistillate
slighttension
turned staminateflowers,andflwr flower.
thattheseare nowinvertedupon the
pistillateflower,with theirpollenmassespressedsharplyinto the
stigmaticsurfaces. The photograph forfig.5 was takenthrough
perhapsa centimeter of water,and showsthe out-turned bases of
thestaminateflowersliningthe bubbleas seen fromabove. To
therightmay be seen a numberof freefloatingstaminateflowers
thatwerereleasedfromthe depressionas the flowersdisappeared
beneaththe surface.
Releaseof thetensionon thescapepermitstheflowers to come
again to the surface; the bubble breaks, and most of the flowers
resumetheiroriginalrelationsat the surfaceof thewater(fig.6).
Examination,however,shows numerouspollen grains or even
entirestaminateflowers scatteredover thesurfaceof thestigmas.
Fig. 6 shows that the groupof freefloatingstaminateflowers,
142 BOTANICAL GAZETTE [FEBRUARY

releasedwhen the groupwas submerged, has again enteredthe

filmpocket. This resultedfrombeingblownintotheradiusofthe
declinedsurface.
Each passingwave thusbringsa shiftin the positionof the
flowersand furthersthe wearingaway of the polliniaupon the
stigmas. Duringthetimethatthepistillatefloweris at thesurface
theeventsoutlinedmay
be repeatedhundredsof
timeswith varyingde-
grees of submergence.
At all times,of course,
theremay be additions
to the group fromthe
free floatingstaminate
flowers. Attention
should be directedto
FIG. 4.-Pistlllate flowerhas been pulled thefactthatanydegree
fartherdownin water;depression is cup-shaped, of depression
is helpful,
and staminate flowersstandat rightanglesto the and that completesub-
verticle
walls.
mergence, althoughthis
probablyoccursfrequently, is notnecessary to adequatepollination.
Whenthepistillateflower is finally
withdrawn intothewaterby
the coilingof the scape, numerouspollen-bearing flowersmay be
trappedin thebubbleof air thatformsabout theretreating floral
parts as theydisappearbeneaththe surface. As the volumeof
imprisonedair is graduallydissolvedby the water,the pollinia
wouldbe pressedmoreand morestrongly intothestigmas. Obser-
vationson isolatedpatchesofpistillateplantsshowthatthescapes
willcoil somewhatwithoutpollination, so theretreatof the seed-
bearingfloweris the finalstepin pollinationif thepollen-bearing
flowers are present. At thefirstfavorableopportunity thewriter
plans to make a studyof the flowering habitsof markedplants
in the fieldto determine the lengthof timetheflowers remainat
thesurface,and theinfluence ofpollinationupon thetimeof their
retreat.
Despite thedioecismand completeseparationof theflowers in
Vallisneria,pollinationseems to take place with remarkable
19I71 WYLIE-VALLISNERIA I43

certainty,providedbothkindsof flowers are near togetherin the

same body of water. No doubt the pollen-bearing flowersoften
ride the surfaceof the waterfor considerabledistances. They
willfloatfordays,and the pollenseemsto withstanddesiccation
fora longtime. In our laboratory aquaria thedeclineof micro-
sporesseems to be due to theattacks from
offungi,and collections
the fieldoftenshowhyphae among
thespores.
From 200 to 450 ovules line the
wallsof theovary,so that the entire
pollen output of several staminate
flowerswould be necessaryforfer-
evenifall thesporesgermi-
tilization,
nated. Fertilizationseems to take
place withcertainty, forfewovules
failto developintoseeds. Scoresof
supernumerary pollen tubes are fre-
quently seen lining the walls or
wandering through theovariancham-
beramongtheovules. Manyofthese
meandering pollentubesformenlarge-
mentsat theirdistalends similarto FIG.5.-With further depression
thosepreviouslyreportedforElodea the water has closed over the
,anadensis flowers, now shut together in a
canadensis(2).
(2). common bubble;out-turnedbases
Turningnow to KERNER'S widely of staminateflowers maybe seen
copiedfigureillustrating his descrip- onall sides,while
pollenmassesare
being
tionof the pollinationof Vallisneria near presseddirectlyintostigmas;
by are somepollen-bearing
spiralis,one is struckby the many flowersthat escapedwhenothers
pointsof contrastwiththe foregoingwerecaughtin thebubble(these
account.tHis illustrationshows a moved duringtime
so areblurred
of exposure
in picture).
~~~~~~~~~and
pistillateflowerwith long slender
ovarywhich,relativeto thespreadof thefloralparts,is onlyabout
one-third of thediameterof thatin ourform. The spatheinvests
onlythebase of theovary,whereasin oursit extendsup almostto
thesepals. The wide-spreading sepalsare shownas straight, while
the broad stigmasare flattened, outstanding, and raisedentirely
above the surfaceof the water. The stigmas,as shownin the
I44 BOTANICAL GAZETTE [FEBRUARY

figure, havemarginsfringed withlonghairsand spreadmuchmore

widelythanin our plant. Finally,the wholepistillateflowerin
KERNER'S figureis placed in such relationto the surfaceof the
waterthatit couldbe sustainedthereonlyon thesupposition that
it is supported by a stiffstem.
Similarly, KERNER has figured thestaminatefloweras markedly
fromours. It is represented
different
as having3 equal and symmetrically
arrangedsepals; the stamens are
figuredwith long filaments at right
angles and protrudingbeyond the
marginsof the sepals. Tiny clusters
of pollen grains crownthese wide-
spreadingfilaments,carriedafterthe
fashionof thespar torpedowellover
the marginsof the flower. Com-
pared with its companionblossom,
the staminatefloweris figuredas
manytimeslargerthanin our plant.
These and many other minor
points of differencesuggest that
FIG. 6.-Release of tensionon KERNER'S accountis highlygeneral-
the scape has permittedsubmerged ized,and perhapsintended to convey
to comeagainto surface;
flowers
onlya generalaccountof thepollia-
pollen and some entire staminate
tion in this plant. It may onlybe
flowersmay be seen on stigmas.
accidentalthat most of the depar-
turesfromtheconditionsfoundin ourplantare necessaryto make
possiblehisproposedplan of pollentransfer.On the otherhand,
it maybe thattheEuropeanplantis essentially fromours
different
ofthesamename; ifso,oursshouldbe describedas anotherspecies.
That theremaybe considerable betweentheforms
difference on the
two continentsis furthersupportedby TURPIN'S (6) figures,which
show slender stamens somewhatdivergent,and stigmas very
fromthose foundin our form. In so far as one can
different
dependupon publishedfigures, it wouldseem that ratherwidely
divergentplants are includedin the species Vallisneriaspiralis.
BOTAtrICAL GAZETTE, LXIII PLATE IX

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WYLIE on VALLISNERIA
I9I7] WYLIE-VALLISNERIA 145

The writerwould welcome photographs,drawings,or specimens

fromdistant regionsforpurposes of comparison.
In conclusion,it seems clear that Vallisneriaoffersa remarkable
seriesof specializationsmainly related to pollinationat the surface
of the water. A few aquatic plants have solved the problemsof
pollen transferunder water, and so may carry out their entire
lifehistoryas submersedplants. A good exampleis seen in Cerato-
phyllumdemersumL., which is pollinated below the surfaceand so
may flourishat considerabledepthsin clear water. Neither Vallis-
neria nor Elodea shows any evidence of transitionto subsurface
modes of pollination,although this would seem to be a desirable
goal for all aquatic floweringplants. On the contrary,they are
perhaps carriedfurtherand furtherfromthispossible habit by their
devices favoringpollinationin air. Their specializationsnot only
bespeak long association with water, but also constitutea remark-
able series of adaptations to pollinationat its upper limit through
the agency of the surfacefilm.
STATE UNIVERSITY OF IOWA
IOWA CITY, IOWA

LITERATURE CITED
i. KERNER, ANTON, Pflanzenleben. Leipzig. 2: I29-I3I. I89I.
2. WYLIE, ROBERTB., The morphologyof Elodea canadensis. BOT. GAZ.
37: I-20. I904.
3. SVEDELIUS, NILS, On the lifehistoryof Enalus acoroides. Ann. Roy. Bot.
Gard. 2:267-297. I904.
4. WYLIE, ROBERT B., A long-stalkedElodea flower. Bull. Lab. Nat. Hist.
State Univ. Iowa 6:43-52. I9I3.
5. COWLES, HENRY C., Textbookof botany. New York. 2:838. i9ii.
6. LOTSY, J. P., Vortrage uber botanische Stammesgeschichte. Jena. 3:642.
fig. 427. I911.

EXPLANATION OF PLATE IX
A general drawingrepresentinga group of associated flowersas they
appear when slightlydepressed; freefloatingstaminateflowersoutside the
depression.