Shoot Damage Effects on Starch Reserves of Cedrela Odorata Author(S): H

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Shoot Damage Effects on Starch Reserves of Cedrela odorata Author(s): H. L. Rodgers, M. P. Brakke and J. J. Ewel Reviewed work(s): Source: Biotropica, Vol. 27, No. 1 (Mar., 1995), pp. 71-77 Published by: The Association for Tropical Biology and Conservation Stable URL: http://www.jstor.org/stable/2388904 . Accessed: 31/05/2012 15:20

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http://www.jstor.org BIOTROPICA 27(1): 71-77 1995

Shoot Damage Effectson StarchReserves of Cedrela odorata

H. L. Rodgers, M. P. Brakke, and J. J. Ewel Departmentof Botany, Universityof Florida,Gainesville, Florida 3261 1, U.S.A.

ABSTRACT The effectsof shootdamage on starchconcentrations in largeroots and lowerboles of the tropicaltimber tree, Cedrela odorata,were measured over a 30-day periodfollowing mechanical shoot removal. The degreeof damage was based on amountstypically destroyed by Hypsipylagrandella, a seriouslepidopteran pest of C. odorata.From 14-month- old plantationsat La Selva BiologicalStation, Costa Rica, threetreatments were established based on the lengthof shoot and numberof leaves removed:controls (no tissueremoved), moderate damage, and severedamage. Initial starchconcentrations were comparedwith concentrationsobserved 18 to 30 days followingshoot removalusing enzymaticstarch hydrolysis and colorimetry.Root starchconcentrations increased in undamaged trees,decreased slightlyin the moderatelydamaged trees,and decreasedmarkedly in severelydamaged treesduring the sampling period. Starchconcentrations in lowerboles did not decreasefollowing shoot damage, suggestinga specificrole of root starchreserves in respondingto abovegroundcarbon requirements.The dry weightsof regrownshoots were similarfor the moderatelyand severelydamaged trees,but severelydamaged treesproduced numerous basal shoots while moderatelydamaged treesproduced only one or two terminalshoots near the point of excision.Decreases in root starchconcentrations following shoot damage and rapid shoot regrowthsuggest that starchremobilization in rootsof C. odoratamight provide a survivalmechanism after attack by H. grandella.

Keywords: allocation;Cedrela odorata; Costa Rica; herbivory;Hypsipyla grandella; resprouting; roots; starch.

RESERVE CARBOHYDRATES IN TREES serveas a resource cies can be attributedto attacksby a shoot-boring formany carbon requirements, especially early sea- lepidopteranlarvae, Hypsipyla grandella Zeller, to son shoot growth(Eifert & Eifert1963, Priestley excessivesoil moisture,or to attacksby the fungus 1970), wintersurvival (Bonicel et al. 1987, Sauter Armillariamellea Kummer (Whitmore 1981). 1988, Nguyenet al. 1990), regrowthafter pruning Attackby H. grandellabegins when the mature and defoliation(Parker & Houston 1971, Wargo mothoviposits on thehost, typically on a leafletor et al. 1972, Gregory& Wargo 1986, Mika 1986, youngshoot (Holsten 1976). The emerginglarvae Boryet al. 1991), and resproutingafter fire (Ahrens migrateto the succulenttissues of the shoot apex 1989, Pateet al. 1990). Althoughthe role of reserve and boreinto the parenchymatous pith. During their carbohydratesin plant-pestinteractions has notbeen development,the larvae consumethe innerstem studiedextensively, the resultsof some studiesin- tissue,forming a pupation chamber.Further ex- dicate that the relationshipcan be important.For pansionof the chamber by the larva typically results example,Dunn and Potter(1990) foundthat vul- in damage to vasculartissues which subsequently nerabilityof treesto wood borersincreases with killsthe shoot.The extentof shootdamage varies, decreasedroot starchcontent, and Wargo (1972) yetalmost always results in the loss of some shoot showed thatthe severityof fungalattack increases tissue,often including large compound leaves. Ce- in treeswith high levels of carbohydrates.Starch drelaodorata normally responds to Hypsipylaattack reservesmay also enablesome tree species to respond with rapid regrowthof new shoots,but repeated to herbivory,as we hypothesizedwould be thecase attacksmay resultin death of the tree. Mortality with Cedrelaodorata. ratesdecline if treessurvive more than fiveyears The neotropicalhardwood Cedrela odorata L. withoutsuffering repeated attacks (Whitmore 1981). (Meliaceae) is a relativelyfast-growing species that The improvedsurvivorship of older trees is probably is nativeto moistdeciduous forests of Latin America. due to escape fromherbivory since H. grandella This speciesis valued forits high qualitycabinet reportedlyoviposits only near terminalshoots and wood, but problemswith pests and itsexacting site usuallyflies no higherthan 2 m above theground requirementshave minimizedits use in large-scale, (Grijpma& Gara 1970). plantations.Most failuresof plantations of thisspe- Based on the highproportion of totalbiomass allocatedto large-diameterroots in C. odorata(Hag- gar & Ewel, in press); preliminarytests indicating I Received2 June 1993; revisionaccepted 1 April 1994. high levels of starchin theseroots (J.P. Haggar, 71 72 Rodgers, Brakke, and Ewel pers. comm.); and the prevalenceof starchas the water,patted dry, sealed in plasticbags, placed on primaryreserve carbohydrate in rootsand boles of ice,and returned tothe laboratory within two hours. trees(Keller & Loescher1989), it was hypothesized Eighteendays afteradministering the treat- thatthe starchreserves in lowerboles and rootsof ments,resampling of rootand bole tissues was ini- C. odoratafacilitate rapid regrowthin responseto tiated.In an attemptto determineiftissue samples attackby H. grandella.The objectivesof thisstudy wereobtained during the period of maximum de- were to determinethe effectof shoot damage on pletionof starchreserves, a differentset of three starchconcentrations in largeroots and lowerboles treesfrom each treatment was randomlysampled of C. odorataand to determineif the change in everythree days during the 12-day period from 18 starchconcentrations in rootsand lowerboles was to 30 daysafter shoot damage. This post-excision closelyrelated to shoot regrowthfollowing shoot delayof 18 dayswas incorporated into the sampling damage. schedulebecause starch is notimmediately utilized followingpruning (Eissenstat & Duncan 1992). All new shootgrowth was excised(including basal METHODS shoots)at thetime tissues were sampled and dry The studywas conductedat the Organizationfor weightswere determined after oven-drying (70?C) TropicalStudies' La Selva BiologicalStation in the for48 hours.To determinestem growth during the humidtropical lowlands of northeasternCosta Rica sameperiod, terminal shoots of control plants were (10026'N, 84000'W). The averagetemperature is removedat theshoot height recorded at day0. 260C and the mean annual rainfallis 3841 mm (Clark & Clark 1991). Data weregathered on 14- EXTRACTIONAND ANALYSISOF STARCH.-Starch con- month-oldtrees growing in threeplantations (0.12 centrationswere determined using the enzyme di- ha each) locatedon an 8-ha researchsite. All trees gestionand colorimetricmethod (Allen et al. 1988 showedsigns of havingbeen attackedat leastonce modificationof Carteret al. 1973). On returning by H. grandella. Forty-fivetrees were randomly to thelab, thesamples were immediately dried at selectedfrom the subsetof treesthat met the fol- 70?C for40 hours.The twocores from each part lowing criteria:not currentlyunder attackby H. (rootor bole) of each tree were then ground together grandella; 1.3 to 2.2 m tall; 14 to 31 leaves; and usinga Wileymill with a 40 meshscreen. To isolate 3.9 to 7.9 cm basal diameter. starchfrom sugars and other soluble carbohydrates, The 45 selectedtrees were randomlyassigned a subsampleof the ground tissue (0.05 to 0.12 g) in groupsof 15 to each ofthree treatments: controls was refluxedin 30 ml of90 percentethanol for 1 (no tissueremoved); moderate damage (removalof hour,the supernatants were removed, and theex- 0.2 to 0.3 m of terminalshoot, including 4 to 7 tractionwas repeated. The remainingresidue (con- leaves); and severedamage (removalof 0.5 to 0.6 tainingstarch) was thenair-dried overnight. After m of terminalshoot, including 10 to 14 leaves). adding5 mlof deionized water and autoclaving the These treatmentswere based on observationsof the mixturefor 30 minutes,each subsample received 5 degreeof stem and leaf damage by H. grandella ml ofacetate buffer and 1 ml ofamyloglucosidase on othertrees in the plantations. solutionbefore being incubated for approximately Post-treatmenttree heights and numberof re- 48 hoursat 46?C to 50?C. mainingleaves were recordedand dryweights of Followingincubation, the samples were filtered excisedstems, leaflets, and rachiseswere determined throughglass fiber filters, and aliquots of the filtrate afteroven-drying (700C) for48 hours.To protect weretransferred to test tubes. Water (1 ml) and the experimentalplants fromattack by H. gran- alkalinereagent (1 ml) wereadded to eachsample della,the bases of rachises of all leaveswere encircled as wellas to a tubecontaining a known amount of witha stickyinsect barrier (Tanglefoot?) to inhibit glucose(to permitdetermination ofthe fraction of larvalmovement from the leaflets(the siteof most glucoserecovered in the analysis).The solutions oviposition)to shoots. werethen heated in a waterbath (100?C) for22 On the day of treatmentapplication, the soil minutesand allowedto cool.Arsenomolybdate re- aroundthe base of each plantwas removeduntil a agent(1 ml) andwater (7 ml) wereadded to each large(2-6 cm diameter)secondary root was located. samplesolution and to standards containing known A 7 mm diametercork borer was used to remove concentrationsof glucose. The opticaldensities of two cores fromone root and two cores fromthe thesolutions were read on a spectrophotometerat lowerbole (10 to 15 cm above theground) of each 540 nm and convertedto glucoseconcentrations, tree. The samples were immediatelyrinsed with whichwere then converted to starchconcentrations Shoot Damage and Starch Reserves 73

as percentdry weight of tissue,after correcting for percentrecovery. As an overallcheck on the meth- odology,cores of cassavaroot (starch concentrations of about 30%, Kawano et al. 1987) wereprocessed .t~~~~~~~C -C N \0 c) - and analyzedin the same way as the C. odorata samples.

DATA ANALYSIS.-The relativechange in starchcon- centrationof rootsand boles was calculatedas ((sn - = SinitialOinitial )100, wheresn starchconcentration 4 o os so< - 4C ~ o: \ (% drywt) on day n aftershoot damage and sii,,, = starchconcentration (% drywt) immediatelyprior to shoot damage. Treatmentdifferences in the rel- Cd Ov ativechange in starchconcentration in rootsand the 7E amountof shoot regrowth were analyzed using analysis of covariancewith date of sampling as the covariate.When the interactionbetween the treat- menteffect and the covariateprecluded analysis of

t F ~~~~~~+l+l +l +l +l +l covariance,treatment differences in the linearre- :~~~~~~~~C \>0 \09D\ gressionrelation between relative change in starch concentrationin bolesand samplingdate were tested for heterogeneityof slopes using a generallinear model (SAS 1991). Comparisonsof the slopes of the absolute change in root starchconcentration (Sinid, - sn) over amount of shoot regrowthwere made in the same manner.

RESULTS I-- .ri r--1?t o6 i Mean initial starch concentrations(percent dry- weight)in rootsof C. odoratafor the three damage treatmentsranged from 16.9 percentto 18.4 per- zt cent,which are similarto values reportedfor Acer saccharum,(12 to 14%, Wargoet al. 1972), Prunus avium'Bing', (16 to 27%, Keller& Loescher1989), and Mangiferaindica (13.5% in boles and 25.5% t~~~~~~~+ +1 +1 +1+1 4~~~~~~~~~Cr- CI "' o,' p in roots,Whiley et al. 1989), and lie withinthe t_~~~~~~~~~c \6 \6 rangeof values reported for Populus (5.1 to 31.5%, Nguyenet al. 1990). Althoughthe effectof time aftertreatment on therelative change in root-starch concentrationwas notsignificant (P = 0.48), starch concentrationstended to increasein undamagedtrees, +1 0 vo o

1 5 on the lowerbole. This suggestsa specificrole of rootsas sitesfor reserve carbohydrates that are uti- 4 10 lized in the regrowthof shoots.Although the net 0 change in starchconcentrations in boles did not 0 5 v 0 decrease,suggesting that boles do nothave the same 00 o 0 function,information on starchturnover rates in the 0 0 boles is neededto supportthis conclusion. Further 0 (1)

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