Wiley

British Ecological Society

Movements of Helianthus annuus Leaves and Heads

Author(s): A. R. G. Lang and J. E. Begg
Source: Journal of Applied Ecology, Vol. 16, No. 1 (Apr., 1979), pp. 299-305
Published by: British Ecological Society
Stable URL: http://www.jstor.org/stable/2402749
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JournalofAppliedEcology(1979), 16, 299-305

MOVEMENTS OF HELIANTHUS ANNUUS LEAVES

AND HEADS

BY A. R. G. LANG AND J. E. BEGG

CSIRO DivisionofEnvironmental Mechanics,P.O. Box 821,CanberraCity,A.C.T. 2601,

Australiaand DivisionofPlantIndustry,
CanberraCity,A.C.T. 2601,Australia

SUMMARY
(1) Diurnalmovements of heads and leavesof sunflower
(Helianthus annuuscv
Hysun30) weredetermined fromthree-dimensional maps of theorgans.
(2) DiurnalE-W oscillationsof the heads occurredinitiallybut ceased as the
flowersopened and anthesiscommenced,leaving the heads facingeast. The
amplitudeof theleafmovements also diminishedwithtime.
(3) The easterlyaspectof theheads reducedthe maximumtemperature of the
florets;thismayincreaseseed set.

INTRODUCTION
The purposeof thispaper is to characterizethe movementof Helianthusannuusheads
and leaves,at about the timeof anthesis,to measuretheeffect ofthismovementon the
temperature of heads, and to considerpossibleadvantagesthatthismovementconfers
on theplant.
Studiesof themovementof sunflower organspredatetheworkof Darwin & Darwin
(1880). The generalformof thediurnalmovementsare known(Heiser 1976),but while
the movementsof leaves have been studiedin detail(Ross & Ross 1969; Shell & Lang
1976),thereare no corresponding studiesof heads.
The tendencyof leaves to face the sun increasesthe irradianceon the leaves and so
increasesphotosynthesis. Also it will reducethe lightreceivedby competitors(Begg &
Torssell1974; Shell& Lang 1976).The movement maytherefore be regardedas adaptive.
The advantageof movementsof sunflower heads is less clear.The earlymovementof
immatureheads,whichtendto facethesun,appearsto giveno photosynthetic advantage
to theplant,because the head is muchless efficient photosynthetically than the leaves
(English 1976). The laterlockingof opened heads to face east would be expectedto
decreasethenetradiationon theface at noon and, sincetemperatures greaterthan30?
reducetheviabilityof sunflower pollenwhenit is stored(A. Low, personalcommunica-
tion),mayincreasefertilization and seed development.

MATERIALS AND METHODS

Helianthusannuus(cv. Hysun30) was used becauseoftheuniformity ofitsdevelopment.
Plants were sown on 24 November 1977 in the fieldat the CSIRO Field Station at
Ginninderra(14904'E, 35012'S) at a row spacing of 750 mm, with 300 mm between
plants.The plotswereirrigated.
?1979 BlackwellScientific
0021-8901/79/0400-0299$02.00 Publications

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300 Movementof H. annuusleavesand heads
Leaf and headmovements
The positionsof leaves and heads were mapped in threedimensionsby measuring
setsof pointson theplants,usingan instrumented, articulatedarmconnectedto a data
logger(Lang 1973; Shell,Lang & Sale 1974). The mapped co-ordinateswererelatedto
the geographicalframeby reference measurements of the directionof truenorth.
A leafwas characterized by a set of tenpoints,consistingof ninearoundthe margin,
includingthetipand thepointof unionwiththepetiole,and one on themidribnearthe
centreof the lamina. A petiolewas characterizedby a pointat each end. A head was
characterizedby fourpointsabout its calyx,selectedto be on a plane parallelwiththe
faceof thehead. Startingfromthetop of theflowering plant,leavessixto fourteenwere
measured.Measurementswere started64 days aftersowing,which was just before
anthesis,and continuedfor 5 days. Eight plants were studiedwithmeasurements at
2-hourlyintervalsfor1 day and thenat dawn and dusk fora further 4 days.
Aftera head had opened fully,anthesisprogressedinwardsover a periodof 7 days.
Colour photographswere taken of the plants each day so that the relativestage of
developmentcould be estimated.
The Cartesianco-ordinatesof theplantswereanalysedto providerepresentations of
leavesand heads as setsof continuoustriangles.The area of each triangleand direction
of thenormal(perpendicular) to thetrianglewerecalculatedfromtheco-ordinatesofits
apices.
Solar irradianceon thehorizontalwas measuredwithMoll-Gorczynski pyranometers,
one ofwhichwas fittedwithan occultingdisk.Mean solarirradiancenearnoonwas 900
W m-2 whilediffuseirradiancewas 240 W m-2.
Head temperature
Threeheads wereconstrainedwithclampsto facevertically upwardsand threeothers
wereleftfreeto move.The surfacetemperatures of boththefaceand back of each head
weremeasuredat noon,whenthe maximumheat load was expectedon the face of the
constrainedheads. The surfacetemperatures were measuredwith an infra-redther-
mometer(BarnesIT3 with30 fieldofview) whenthe sun was shining.The thermometer
was calibratedby pointingthe instrument at a watersurfaceofknowntemperature in a
vacuumflask.The emissivity ofthehead was notmeasured.Assumingtheemissivities of
heads and leavesweresimilar,thesurfacetemperature would be underestimated by less
than2 0C (Fuchs & Tanner 1966). Temperatures of sunlitleaves,selectedmainlyto be
perpendicularto thesun's rays,weremeasuredforcomparison.
Wind speedwas measured1 m above thetops of theheads,usinga cup anemometer.
Air temperature at thelevelof plantheads was measuredwitha mercuryin glass ther-
mometerin a radiationshield.
RESULTS
Leaf and head movement
The directionof a mean leaf normalwas obtainedby weightingeach normalof a
trianglebythearea ofthetriangle, and takingthevectorsumofall theweightednormals
forall thetrianglesof all theleaves. The movementof thismean leaf normalwithtime
was similarto thatdetailedearlierforthecultivarPeredovic(Shell & Lang 1976). The
crossingof the meridianat nightoccurredbetween03.00 and 06.00h, between4 or 5 h
laterthanobservedpreviously (Shell & Lang 1976); howeverthe same principalfeature,
oftheleavesfacingeast beforethesunhad risen,was stillevident.The movementof the

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 A. R. G. LANG AND J. E. BEGG 301
averagenormalsto head and leaves was mainlyfromeast to west,so it is convenient
to presentthis motionin termsof the angle betweenthe normal and verticalplane
alignedN-S (Fig. 1).
As the plantsdiffered
by up to 3 days in theirstage of development,
averageswere
takenof plantsat thesame stage,judgedfromphotographsand usingthetimeofflower
openingas an index.Averageswerebased on one to eightplants.

40 (a) _

420
-C~~~~~~~~~~~~~~~~~
10 (

20
07
0)
0)
<40

60

- 20
20
20

3 E 1 4 5 6

Time (day)

FIG. 1. The east-westangularmotionofnormalsofsunflower heads(a), and leaves(b),

and theirrelativeleafarea (c), aboutthetimeof anthesis.The barsdenote? standard
errorsofthepoints.Blackzoneson theaboissadenotenight.

For comparison,thedevelopment of leafarea was calculatedfromthemeasurements,

and normalizedbydividingthetotalleafarea ofeach plantbyitstotalleafarea,averaged
fordaysfiveand six afterthe startof observations(Fig. 1).
Leaves on easternand westernsides of plantsweremotivatedmainlyby bendingof
midribsand petioles (Fig. 2). This is shown more clearlyby the curvature(i.e. the
reciprocalof the radius-ofcurvature)of the combinedpetiole and midribwithtime
(Fig. 3).
The ratio of the irradianceon the ventral(upper) sides of the petioleto thaton the
midribwas calculatedvectorialy:ifa, h and S are unitvectors,representing theaverage

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302 Movementof H. annuusleavesand heads

og~~~~~~oo
1100~~ -~ z~ ~ =~ ~7 ~ ~ ~ @3 X 2 1.0~~~~~~~~~~900
30

09.00 up00

11.00 =/74z<0 100

09.00~~~~~~~~~A
S~~~

X ASA

11.00 ~ ~ ~ ~ ~ ~ ~ ~ ~~- 50

17.00 > i X ~ x <05.00

. . .
.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~,,.......

FIG. 2. Typicaldailymovement of easterlyand westerlyfacingleavesviewedfromthe

north.The drawings wereplottedfromsetsofmeasuredcoordinates.S is theprojection
of thesun's direction.

directionsof the petiole,midriband sun's beam respectively, then the ratio of the
is givenby
irradilances
[(a x (a x b).S]/[((a x b) x b).S].
The ratiosare presentedin Fig. 3.

of head
Temperature
Figure4 showstheeffectof windspeedon thetemperature of theface (Fig. 4(a)) and
back (Fig. 4(b)) of constrainedand freeheads. The airtemperatures
at noon rangedfrom
22 to 28 0C.

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 A. R. G. LANG AND J. E. BEGG 303

(a)
12 3

4
W~~~~~~~~~~~~~~~~~~
E 0 0}
I~~~~ CI,
TimeI(hours)

82 18 2 1 2
4 1~~~~~~~~~~~~~~~

0 (V

12 18 0 6 12 18

Time (hours)
FIG. 3. The curvatureof petioles and midribs(solid lines) and the ratio of irradiance on
the ventralside of petiole to thaton midrib(dotted lines) foreasterly-facing
leaves (a) and
westerly-facing leaves (b). Dashed lines show the ratio of irradianceformodel, horizontal
leaves with azimuths eithereast or west and withpetioles at 450 to vertical.

DISCUSSION
Leaf and head movement
The modeof movementofleavesdiffered, dependingupon whethertheleaveswereon
theeast and westsidesof theplantor on thenorthand southsides.In thecase of leaves
on theeast and westsides(Fig. 2), movementwas mainlyby bendingand straightening
ofthepetioleand midrib;forthoseon thenorthand southsides,itwas mainlybytorsion
of thepetiole(estimatesof thirteenleavesshowedanglesof twistchangingby 250 to 900
fromdawn tilldusk).
For the bendingmode,the curvatureof one petioleand midribwas increasingwhile
the otherwas decreasing(Fig. 2), whichraisedtheinterestingquestionas to the cause.
The sunobviouslyprovidedtheprimarystimulusforsynchronizing themotion,butthere
was no obvious difference in irradianceupon the easternand westernlaminae which
wouldaccountfortheiralternatebehaviour.However,therewas a difference in thetime
courseoftheratiosofirradianceon theventralsidesofthe midribto thaton thepetiole
(Fig. 3) whichmaybe significant. This difference
was caused mainlybythemovementof
the sun ratherthan of the leaves, as shownin Fig. 3 forfixedhorizontalleaves with
petiolesat 450 to thevertical.
The periodicmotionof thehead decreasedin amplitude,and thenormalassumedan
easterlyaspect,as thefloweropenedand anthesiscommenced(Fig. 1); thisresultedin the

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304 Movementof H. annuusleavesand heads

12 (a)

4-
0O|*

E _. 0

0 0 -

a).-_
o. 12
E

4 008~ |\ =

0~~~~~~~~~~~~
0 2 4 8

Windspeed(m s-1)
FIG.4. Thesurface
temperatureof the face(a) and back(b) of freeheads(w) and
vertically heads(a) andof leaves(A) at variouswindspeed.All measurements
facing
weremadeat aboutnoon.

familiareasterlydisplayof the floraldisks. Heliotropicmotionof the leaves also de-

creasedin amplitudeabout thesame time.The aerialexpansionof theleavescontinued,
at an undiminishedrateof 4.700 per day,untilthestartof anthesisat day seven;thenit
appeared to stop. Thus the movementsof both heads and leaves occurredduringa
periodof leafexpansion,in accordancewiththe long-heldview thatheliotropicmove-
mentsof sunflowers are drivenby differential
growth,as opposed to photonasticmove-
mentscaused by turgorchangesin fullygrowntissues,such as occursin petiolesor the
pulviniof manyleguminousleaves (Begg & Torssell1974).

Head temperature
The naturaltendencyof matureheads to face east resultedin a substantially lower
irradianceupon theflowersat noon,whentheplantsweresubjectedto an air temperature
near the maximumforthe day. Consequently,the surfacetemperatures of thefaces of
freeheadswere3 to 8 0C cooler,at thistime,thanthoseconstrainedto faceupwards.The
freeheads were0 to 4 0C warmerthanambient,but constrainedheads were3 to 12 'C
warmerthanambient.The maximumambienttemperatures duringanthesiswere22 to
28 0C, and thetemperature in floretsevidently
was not sufficiently
highto preventseed

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 A. R. G. LANG AND J. E. BEGG 305
set in the constrainedheads in the presentexperiments. However,sunflowers
thrive,
both naturallyand commercially, in climateswithmuchhigherambienttemperatures
thanrecordedhere,and avoidanceof highirradianceson thefloretsat anthesiswould
thenbe an advantageforfertilization.
Tanner& Goltz (1972) studiedthereasonsforlocalizedabortionof floretson onion
umbels,and foundthatthetemperature was greatestat theoutersurfaceof theumbels
wherethesolar irradiancewas greatest,but dependedalso upon thewinddirectionand
speed. The increasesin temperature, above ambient,forupwardfacingsunflower heads
in thisstudywerewithin1 'C of thosereportedforumbelsof onion at the same mean
windspeedand comparableisolations(Tanner& Goltz 1972).
The easterlydirectionof heads is an obviousadvantagein reducingtheheatload, but
it also has the advantagethatmoreradiationis intercepted duringearlymorning;this
willspeedthedryingof dew and decreasethelikelihoodof fungalattack.Bowingof the
heads also reducesthe interception morerapid drying
of rain by the heads, permitting
and thusreducingthechanceof fungalattack.

ACKNOWLEDGMENTS
Assistancegivenby R. Endersand A. V. Jacksonin the collectionof the co-ordinates
is gratefully
acknowledged.

REFERENCES
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Wellington, pp. 277-283.
Darwin,C. & Darwin,F. (1880). ThePowerof Movement inPlants.JohnMurray,London.
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(Received6 October1978)

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