Effect of Night Temperature on Flowering and Fruit Size in Pineapple (Ananas comosus [L.

]
Merrill)
Author(s): D. J. C. Friend
Source: Botanical Gazette, Vol. 142, No. 2 (Jun., 1981), pp. 188-190
Published by: The University of Chicago Press
Stable URL: http://www.jstor.org/stable/2474272 .
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BOT. GAZ. 142(2):188-190. 1981.
? 1981 by The Universityof Chicago. 0006-8071/81/4202-0012$02.00

EFFECT OF NIGHT TEMPERATURE ON FLOWERING AND FRUIT SIZE

IN PINEAPPLE (ANANAS COMOSUS [L.] MERRILL)1

D. J. C. FRIEND
Departmentof Botany, Universityof Hawaii, Honolulu,Hawaii 96822

'SmoothCayenne'pineappleplantsgrownin a greenhouseat between25 and30 C weretransferred

for
16h nightly
to darkroomsmaintained was mostrapidat a nighttem-
at 15,20,25,and 30 C. Flowering
peratureof 20 C and was slowestat 15 and 25 C. Floweringdid not occur after3 yr of growthat 30 C.
Flowernumber
andfruit
weight
weregreatest
forplantsgrownat a nighttemperature
of15C.

Introduction houseat a day temperature of 25-30C and a night

Floweringof commercialcrops of pineapple in
Hawaii is normallyinducedby the use of growth
regulatorsat all timesoftheyear.Naturalflowering
is seasonal,however;theinflorescence is initiatedin
December and flowering occurs about 3 mo later
(BARTHOLOMEW and KADZIMIN 1977).This suggests
that flowerinitiationis inducedby a reductionin
temperature, shortdaylengths, orboth.For theRed
Spanish variety, temperaturealone can control
flowering; 1 mo at a minimalnighttemperature of
16 C inducedinflorescence initiationundernatural
summerdaylengths in PuertoRico (VAN OVERBEEK
and CRUZADO 1948).In SmoothCayenne,daylength
may influence flowering,and GOWING (1961) classi-
fiedthisvarietyofpineappleas a quantitativeshort-
day plant. The flower-accelerating effectsof short
daylengths wereapparentevenunderconstanttem-
peratureconditionsof 25 C (FRIEND and LYDON
1979).
This studywas undertaken to determine whether
natural floweringof Smooth Cayenne pineapple
growingundershortdaylengthsof 8 h was also ac-
celeratedbv lowering thenighttemperature overthe
rangefrom25 to 15 C.
Materialand methods
Plants ofpineapple,Ananas comosus(L.) Merrill
'Smooth Cayenne,'Champaka 153, were obtained
bygrowing rootedcrownsin a mixtureofvermiculite
and gravel watered with Hoagland's solution
(FRIEND and LYDON 1979). To checkthat the vol-
ume ofpottingmediumdid not influence flowering,
two sizes of plastic containerwere used for each
treatment:15-cm diameter,holding 2 liters of
medium,or 21-cm diameter,holding 5 liters of
medium.As therewas no significant effectof root-
ing-medium volumeon theflowering response,data
fromthesetwosubtreatments have been combined.
In the firstyear,plants weregrownin a green-
1 JournalSeriesno. 2535 ofthe Hawaii InstituteofTropical
Agricultureand Human Resources.
ManuscriptreceivedAugust 1980; revisedmanuscriptreceived
January1981.
188
temperature of20-25 C. The photoperiod was main-
tainedat 24 h by supplementing 11-13h ofnatural
daylength withlightfrom500-Wincandescent lamps
mounted3 m above the plants. The radiant flux
densitywas 17 k ergcm-2s-' (700 lx) as measured
by a YSI Model 65 radiometerat the top of the
plants. Because of the distancebetweenthe lamps
and the plants, this level of radiantfluxdensity
given at nightdid not measurablychangeduring
thegrowthoftheplants.The spectralqualityofin-
candescentlightis givenin KLEIN and KLEIN (1970,
p. 177). In the secondand thirdyears,plantswere
moved daily from8 h of natural daylightin the
greenhouse to 16 h ofdarknessin constant-tempera-
tureroomsmaintainedat 15, 20, 25, or 30 + 2 C.
Nine plantswereplaced at each nighttemperature,
and the date of the firstfloweropeningon each
plant was recorded.Ripe fruits,as judged by the
developmentof an orange-yellow colorationand
detectablearoma,wereharvested.Fruitand crown
freshweightsand the numberof flowers(eyes) on
the inflorescence wererecorded.The widevariation
in time for floweringboth within and between
different temperature treatments precludeda mean-
ingfulcomparisonof total plant dry weightsas a
measure of total growthat the different night
temperatures.
Resultsand discussion
Nighttemperature affectedthevisualappearance
of the plants approximately 6 mo aftertreatment.
Red pigmentdeveloped in the leaves of plants
grownat 15 C nighttemperature, and plantsat the
30 C nighttemperature werenoticeablysmallerand
had narrower leavesthanplantsat othernighttem-
peratures.Plants at 30 C had well-developed mar-
ginal serrations(fig.1), ratherthan the "smooth"
leaf margincharacteristic of this varietyof pine-
apple.
The mean numberof days to flowering was 470
aftertransfer to alternatingtemperatures (772 total
daysofgrowth)forplantsgrownat a nighttempera-
tureof20 C, 610 daysat 15 C, and 660 daysat 25 C
(fig.2). Plantsgrownat a nighttemperature of30 C
 FRIEND-NIGHT TEMPERATURE AND FLOWERING OF PINEAPPLE 189

were still vegetative690 days aftertreatment,as 1400 i

shownby dissectionand microscopicexamination

of theirapical meristems.
Directassessmentsof the timeoffloralinitiation 1200-
by periodicdissectionsofplantapiceswerenot car-
ried out in these experiments because of lack of
space forthe largenumberof plantsthatwouldbe 1000\
required.However,some information on relative
nighttempera-
ratesof floralinitiationat different
turesis providedby the finalleaf numbers.For a
i
plant with a terminalinflorescence, such as pine- ." 800 150C 200C 250C

apple,floralinitiationpreemptsthefurther produc-
60 FRUIT

I: 600-

U_

400X

CROWN
200-

15* 200 25? 300

FIG. 1.-Marginal serrationson leaf of pineapple grownat NIGHT TEMPERATURE 0C

30 C nighttemperature(upper) comparedwithsmoothmargin
at 20 C (lower). FIG. 3.-Effect of nighttemperatureon freshweightof ripe
fruitand of crown of pineapple. Vertical lines representthe
SE of the mean. There was no significantdifference in crown
weights.
750 I7

tion of foliageleaves and establishesthe finalleaf

700 - - 70 number.An increasein nighttemperature from25
to 30 C increasedthe finalleafnumber(fig.2) and
delayedthe initiationof the inflorescence, as deter-
FLOWERING - 65
650
minedby dissectionsof plants grownwitha 30 C
nighttemperature. It is thereforeprobablethat the
600 -60 increasein the finalleafnumberresultingfromin-
creasingnighttemperature from20 to 25 C was also
a consequenceofdelayedfloralinitiationratherthan
550 -W5 an increasein the intervalbetweenfloralinitiation
LEAF NUMBER z and anthesis.Periodicdissections ofapicalmeristems
would be needed to establishwhetherthe delayed
500 -50
floweringat 15 C was a consequenceof delayed
floralinitiationor an increasedintervalbetween
450 - 45
floralinitiationand anthesis.
The freshweightof ripe fruitwas greatestfor
plantsgrownat a nighttemperature of 15 C (fig.3).
400 _ 40 Crown freshweightswere not different between
night temperatures of 15 or 20 C and were reduced
by 20%go at a nighttemperature of 25 C (fig.3).
The totalnumberof flowers(eyes) on the inflo-
0
150 200 250 300
0
rescencewas inverselyrelatedto temperature, from
NIGHTTEMPERATURE
OC
102 at 15 C to 74 at 25 C (table 1). This reduction
in flowernumberswas causedby a reductionin the
FIG. 2.-Effect of night temperatureon number of days numberof flowersper vertical row on the inflo-
fromtransferto alternatingtemperaturesuntilfloweringand rescence.The numberof rowsaroundthecircumfer-
on final numbers of leaves produced before inflorescence
formation.Plants at 30 C did not flowerduringthe experi- enceof thefruitwas not different.
mentalperiod.Verticallines representthe SE of the mean. Night temperaturewas clearly an important
190 BOTANICAL GAZETTE

TABLE 1
EFFECT OF NIGHT TEMPERATURE ON NUMBER OF FLOWERS ON INFLORESCENCE OF PINEAPPLE

NIGHT TEMPERATURE ( C)

15?SE 20?SE 25?SE 30

Mean no. of flowersper verticalrow.7......... . 7.7 .3 6.1 .0 5.7 .O 0

Mean no. of rowsaroundcircumference...... 1 3 13.0a .0 1301 .0
Total no. of flowersper inflorescence
........ 102 5.7 79b 1 .9 74b 2.3

NOTE.-Itemsmarkedwithsamesubscript
letterare notsignificantly at P > .05 levelin a pairedt-test.
different

determinant of floweringofpineapple,even though pineappleprobablyresultsfromeffects ofbothtem-

plants were maintainedon a shortdaylength.No
flowering occurredat a nighttemperature of 30 C
even aftermore than 690 days of growthat al-
ternatingtemperatures. Pineapple is not a typical
short-dayplant in this respect. In general,the
flower-promoting reactionstakingplace in thedark
periodof short-dayplantsare acceleratedby a rise
in night temperatureover the range of 10-30C
(VINCE-PRUE 1975).
The effectof night temperatureon flowering
paralleledthatondarkfixation ofCO2 (Crassulacean
acid metabolism[CAM]). The dark CO2 fixation
was greatestat a nighttemperature of 15 C and
absent at a night temperatureof 30 C (NEALES
1973). A directdependenceof flowering on CAM
seemsunlikely,however.Floweringwas greatlyre-
tardedby increasingthe daylengthfrom8 to 16 h,
althoughCAM (measuredas malic acid accumula-
tion) was littleaffected(FRIEND and LYDON 1979).
The influenceof climateon naturalflowering in
peratureand daylength.Short daylengthsaccel-
eratedflowering even at a constanttemperature of
25 C (FRIEND and LYDON 1979), but a nighttem-
peratureof20 C further acceleratedflowering.
Inter-
actionsbetweentheeffects oftemperature and day-
lengthon flowering werenot investigatedin these
experimentsbecause of the slow nature of the
flowering response.Furtherexperiments are needed
to determineifplantsgrownat a lownighttempera-
ture retain the facultativeshort-dayresponseor
behaveas day-neutral plants.
Acknowledgments
I thankMr. G. YAMANE, ResearchHead of Del
Monte Co., at Kunia, Oahu, for providingthe
crownsofpineapplesused in theseexperiments,and
JOHN LYDON forhisskillin growing
plantsand main-
tainingthecontrolled
environmental over
conditions
sucha longperiod.

LITERATURE
BARTHOLOMEW,D. P., and S. B. KADZIMIN. 1977. Pineapple.
Pages 113-156 in P. T. ALVIMand T. T. KOZLOWSKI, eds.
Ecophysiologyoftropicalcrops.AcademicPress,New York.
FRIEND, D. J.C., and J.LYDON. 1979. Effectsof daylengthon
flowering,growth,and CAM of pineapple (Ananas comosus
[L.] Merrill).BOT. GAZ. 140:280-283.
GOWING, D. P. 1961. Experimentson the photoperiodicre-
sponsein pineapple.Amer.J.Bot. 48:16-2 1.
KLEIN, R. M., and D. T. KLEIN. 1970. Research methodsin
plant science.Natural HistoryPress, New York. 756 pp.
CITED
NEALES, T. F. 1973. Effectof night temperatureon the as-
similationof carbon dioxide by mature pineapple plants,
Ananas comosus(L.) Merr. AustralianJ. Biol. Sci. 26:539-
546.
VAN OVERBEEK, J.,and H. J.CRUZADO. 1948. Note on flower
formationin the pineapple induced by low nighttempera-
tures.Plant Physiol. 23:282-285.
VINCE-PRUE, D. 1975. Photoperiodismin plants. McGraw-
Hill, New York. 444 pp.