Quality Timber Products of

Teakfrom Sustainable Forest Management

Proœedmss of the Inremational Conkrcrn e:-t Quality T~ ProdIlct5 of TrU from SustaINbk- Fon>st ManagementPeechl.lndia. 2-5 Deœmber 2D.D

Edilors

K.M. Bhat, K. K. N. Nair, K. V. BhatE. M. Muralidharan and J. K. Sharma

Kerala Forest Research Institute, Peechi 680 653, lndiaAn Institution 01 Keraa State CoociI for Scienœ. T«hrdogy and Environrnmt (KSCSTE)

Intemational Tropical Timber Organization UTTO), Yokohama, Japan

.

02005 Keub Forr'lt Rr'lurch IMljtute

The Kerala Forest Research Instltute (KFRI) is one of the SIX institutions underthe KenlOi StateCounol forSoenœ, Technology iU1d Ennronmenl (KSCSTE)ofthe Go.'emment of Kerala, estOlbhshed in 1975. By ronducting time-boundmulbdlSCIpllnary applled research ln lhrust areasof tropIcal forestry, KFRl hascrt'itecl li mchr ..mong the le..dIng fores! research orgamutions in the IropICl.Th", Institut'" unaertakes mulbd1sclplmary research on ..Il asprcts of troptcalfo~try mduding .....ood samœ and technotogy, wtIdlile biology and socio­fCOnOIJUC5 undrr the Progra1ll/Tll!' DiVlSlOOS. KFRI has aSub-Centn .. t Ni1.l.mburand li Field SUbon atVrl~ for calT}'mg out nursery and pbntabon buis,grrmplasm roli«bon, ~c. KFRI tw the 1argest roUl!'ctJon of bamboo and ratblsprrirs in Inch.a fOI" rrsearm and consernbon~ Also il Ni1ambur, lhereis il Tnk Museum. the onIyone cllts kJnd. ~'Oted to a single tnor sprciI!'S in thewOI"Id, It is opm lopubbc, researchers. fores! offioals and ethers mlen'sted in

""Initmitioui Tropinl1im~rOrpniution

The IntermtK'NI Tl'Of'ICI'l TunbeOrpniuIion. (ITIO) is an inll!'lgO\wnmmtalorgmiubon wttid\ bnngs logdhrfoountnes iNt prodl.lCl!' and consumr tropicalbiTIbft- todl5CUS$ and exdunge information and develop poIiors on ail~cl the woOd troptaI bmbrr~y.The rrro is hudquartered m Yokohama.Jilpan and ln luly 2001 had Sl mrmbrrs. induding the European Commuruty,.....him logether rtpresell95 f'l"I" œnt of .....orid trMte in tropial.~ and 75per C'l!'nt oflhe wotld's lropeal forests. The rrro norognizes lhat 01 conllnuingsupPIyoflroplCillllmber on the world markddependson quahty informlllion.about the lTadeand market plaœ, on ef6orn1 bmbrr produclion. and proœs5U'lgmethods and on susbmllbie forest Iniln.J.grmrnl practices

Publishtd by:

Kerala Fo~1 Reseuch Insblute, Peedti 680 653, Ken!a, lrKfu,Tri: 91-487-2699037, 2699061-64; Fax: 91-487-26992..19Email: UnOkfnC'rS Website: hllp://www.kfri.org...Intemabonal TropIcal TImber OrgamUlJon5"' floor, Pacifico-Yokohama, 1-1-1, M.uuto-Mirai, Nishi-kuYokoha~,~I~Japan

Tel: 81-45223-1110, hx~SI· 45 223- 1111Email: ilto@itto.or.)p Website: tottp://www.iltoor.jp

ISBN 81 -85().l1-63-6

KFRI LibraryCataloguing-in-Publication Daia

QUAlITY TIMBER l'RODUCTS OF TEAK FROMSUSTINABLE FOREST MANAGEMNET

I,Teak products 2. Sustainable forest management-teak3. TImber œrbfinlion 4 Tt'("Iorui grandIS

1. Bhal. K. M.II Kerala Fûffst Research in.<titulr1Il.lnlemab(Xl&1 Tropical TImber Organiulion

582.949

Typeset: KFRlPrintt'd &t LUll\l('ff,lhnssur6lltJU20, IndU.

<.

Selection and Propagation of Superior Teak forQuality Improvement in Plantations: case study of

the ICSB/Cirad-forêt joint project

D. K. S. Cohl, D. AlJoysiusl,}. Gidiman ', H. H. Chan l, B. MalleFand O. Monteuuis2

llMoprise'Corporoltion Sdn Bhd., P. O. Box 60793, 91017 Tawau, Sabah, MalaysiaE·mail: domgoh@pc.jaring.my;dil\·K1@rbjtwu@po.my; jgidiman@holmaiLrom; chanhh@pl",ringmr

'CIRAD-Foret, TA IOIC, 8.JilIarguet. J.l398 Montpellier. Cedex 5. FranceE·mail: mallet@cirad-fr; monteulUS@cir.ldfr

ABSTRACf

Innoprise Corporation Sdn Bhd (JCSS). an in\'estment subsidiary of the Sabah Foundation, ha!>embartœd on an e:densh-{' Research and lÀ'velopmenl program for leak (TtdotUJ SfI",dlS), a highlyprized limbe! species. Through ICSB's joinl collaboriltion with CIRAD-Foriot. a French R and Docganization., a plant improvemenl program was initi.lled in the early 1990'$. Special efforts ha...ebeen devoted 10 vegetative propagation strategies b.lsed on the de'l'elopment al efficleJlt lechniqlll.'S.both al the laboralory and al the nwsery Ieo.'els. ~Ialerials from mature selected plus ltees (ortels) fromil broad genelic base, and seeds of preo>uD"lt"d rugh genelic value bul in reslricled number, weremultiplied by using il weU-deveJoped tissue culture techruque al the PIani Btotechnology uboratory.Additionally, seed lois from natural foresl stands, pl.mtations ilnd a mulli-provenance clonai seedorchard (CSO) were obtained. These Weil! germinated and used 10 establish two main prO'.enancelprogeny trials in two dlfft'reflt locations within ICSBconœssion in easl Sabah. Malaysia, in addition toolher numl'rou.~ conservation and demonstralion pIOIS. Data of six-year old trees from thesepron'nance/pl'Ob~ny triab ilIustrated the great potenlial of the CS(} materials esperi.1Uy in terms ofJ;l1Jwlh rate rompared 10 the other origins under the pre\'aiting conditions. Advanœd selection. ofJ;('fM)lypes rombining v.lrious traits of major ('((Inomical value ie. wood charaderistics, is underway.These genolypes and their progenies will he further tested in different sites in the immed~lte future.The ownership of such a rich germplasm coupled 10 the effJCiency of the develol"-'d techniques offersi\ tremendous potential for genetic improvement, selection and mass production for large-scaleplant.ltions of higher yield and superior quality. COll1JTl('m.ll production of sclcct('(i nmleriills fromthis rich genetic base will !hm ensue as tissue culture-issued materials for overseas markets.

Ktywords: clonaI deployment, field trials, genetic resources. germplilsm, provenilnce/progenytrial, superior planling stock production, vegetative propagillion, tissue culture.

INTROOUcnON

Teak was firsl introduœd in Kota Marudu in the stateofSabah, Malaysia, by the Dulch TobaccoCompanyin the 19205', but did not bccome popular until theearly lC)lX)s (Lapongan, 1998). Since then, privateplanlation owners and oil palm companies havegrown teak mainly by intercropping with oil palmorothercashcrops, with parlicular menlion forthemosaic IOxIOm inlerplanted with IOxIOm oil palmdesIgn experimented olt Balung Plantation in Sabah

(Salleh, 1995; White, 2002). Howevcr, the teolkplanting material uscd WolS not spccificallysclc<:tedfor such plantation systems.

o.·er time, lackof informaI ion on lheoptimal densityand spacing in an inlercropping system and thcreby,fcarof potentiallœses 10 theyieldofassociated crops,resulted in the planling of tcak mainly along bordersof many oil palm plantations and to sorne extent,roadsides. Moroo\'cr, the reputation of long rotationspecie; tradilionally associated to teak began 10slacken

390

furthcr in Sabah and Malaysia asa whole. Asa result,investors nowadaysare morecagerto invcst in theoilpalm industry due to the more lucrative retums thaltan beobtained in a timeperiod asshort as threeyears.Planl~ were no!. evCll encouraged by the finding froma cost analysis of teak planling in Malaysia thatestimaled Icakrould be viable as a plantationaop C\'en

wilh an increase of 20% in the rost of planting and adrop in priee on a 15-year rotation (Krishnapillay dal., 1997). Nevertheless, while the interest in teakplanting within Malaysia declines in the face ofcompetition nomothercash cropssuchas cocoa, riœ,rubber and especially oil palm for whkh the countryhasfo,y.....gamoo theworld production leadefship,the scenario is just the opposite in other parts of theworld.

Countries in the tropical and sub-Iropical zones suchas Central and South America, Arrica, India,Australia, and other parts of South-east Asia beginembarking on large-scaled planting of teak,recognizing it as a prime candidate for forestplantationestablishmenl in termsof valueand worldmarket demand. Accordingly, partiaJiarattention isbeingdevoled to means for improvingquantitativelyand qualitalively the yields in shorter time framesthan tradilionally expeded. Since the early 1990'5,Innoprise Corporation. the investment subsidiaryoftheSabah Foundation, astale-run organization. andCIRAD-Forêt, a French Researçh and J:)evelopmentOrganization have been working together, aimingat this ob;ective.

CONSTITITrING GENFrlCAllY-RICH BASEPOPULATIONS

Securing Ihe access to a genetically broad-basedpopulation is an assel of magnitude importance fordeveloping simple or more sophisticated Ireeimprovement strategies. This is particularly true forleak considering Ihe noticeable differenœs amongprovenances, progenies and even half-sib (Bath,2000). Relevantly, one of our priorilies has been togalher as many leak origins as possible and 10establish properly designed base population plotswithin ICSB'sconœssions.

The first batchofseedsoriginating from the Solomonisland seed-sourœ was introduced in 1989. As for

391

many seed sources, information on the accu ratenatural originofthis plant malerial is Iacking, althoughTenasserim (Myanmar, ex Bunna) seems 10 be themosl Iikely provenance (Kevin White, personalcommunication).The resultingseedlings wcre planledby the Planllmprovem<nl andSœd ProdUctiOll (l'lSPforshort) UJÙt asademonstrational plot in the LuasongForeslry Center, located aboul two hour drive fromTawau,oo theeasl coast ofSabah. This planl materialhas been thriving sÎnœ lhen, showing remarkableperfonnanœs under local conditions with averagediameter and height ol2·3 cm and 4m in the firsl2-3yeats resp«tively, followed by• gradu" evening 001inaverageheightof2 to3m insubsequenl years. Thetrees give rise 10 long c1ear boles with delayedllowering, bearing in mind that in teak. the tater thealtainmentof the flowering stage, the longer the boIeand therefore, the higher the value.

The second introduction of seeds was made in 1995,this lime roII<cted frorn the I..ksœd sbnd beIoogingta the Forest Research lnstitute of Malaysia in Perlîs,Iocated in the north of Peninsular Malaysia close 10theborderwith Thailand. Thereisstill a basic need 10docummtlheaccurateprovenanœsoftheseseeds.ThesmalJ amount ofseeds from this balch wasgerminatedunder in vitro conditions at the Plant BiotechnologyLaboratory (PBL) following the proœduredescribedby Monteuuis d al. (1998), and then planted as yetanotherdemonstrational plot in Luasong.

ln 1996, in order 10 wÎden the existing genetic baseofteak in our project, ICSB jointly with Cirad-Foretprocured seeds from two extensive sources - fromnatural forest standsor plantationsand from progeniesproduced from a multi-provenance clonaI seedorçhard in Ivory Coast. Altogelher, there were 77seedlots comprised of India, Thailand, Papua NewGuinea, Tanzania, Ivory Coast, Salomon Island,Indonesia, Segama (Sabah) and Perlis (PeninsularMli<lysœ)(f.bles hnd 2). TheseedsweregenniNledeither at the PISP nursery in luasong or at the PBL,particularly those with presumably low genninationcapacity. Adetailed report on theseedling procedure., the PBl hasbeen published (Monleuu;set<li., 1998).

However, in the same way as for many speciesintroduœd as exotics, tracing the accurate naturalorigins of every seed-sourœ as weil as the number

Table 1. List orthe \'<lriolls ......--edlotsobtained,germinated and pJantcd within our proJcct

Pro\'t'n·lOCt.....; India ChilOdr<lpur MahMaslrollndia Sakrebail Karnoltaka - 2 batchesIndia Vimoli Vir. Kamalolkiln - 2 boltchesIndia Karadibella Kamat<lkaIndia Gilillegundi Kamat<lkilIndia Maukal Kamataka - 7IMtchesThail.lnd Moile Hualloilmpang (nalural sland)Thaîland ~iat' Huai loiImpang (planled stand)

Bulu Kumba, Il1dont'SiaPapw New Guinea ex Brown Ri\·er· presumably {rom Myanmilr (Bumw)(Cameron 1966, White, personal communication);Salomon Isbnd Arara - prt'Sumably introduœd from Myanmar if not fromIndia or ThaiJand, via Papua New Guinea (Whitt'. personal communication);Salomon Isbnd Vîru-presumably introduced from Myarunar if not fromIndia or Thailand, via Papua New Guinea (White, personalcommunication);Perlis. Peninsular Malaysia - from Forest Research Inslitute Malapia.preswnably (rom ThaibndKota Marudu, Sabah - introduced by the Outch Tobacco Company,presumably from India

Tot.l: 22 ~edlots

392

of progeny and provenances induded, constitule areal problem; information, when reliable, isfragmentaryand a lot of unœrlaintiesremain. In thisrespect, wc arc pldcîng strong hopes on the use ofnlolccul.lr nl,ukers for ,1ITessîng the information thatIS 1.1Cklllg.

CERMPlASM CONSERVATION

The teak genetic resources gathered within theprojett can be conservee! either as seeds, plantedoutdoors or in tissue cultureconditions.

As seells

Seee! lots can be stored in a fridge or cold room, at4°C for sorne lime. However, 10ss of germinationcapacity over time must nol be undereslimated.Several months of storage under such environmenlmay result in a dramatic dccrease in germinationrate, already known 10 be 10w and unpred.ictableunder the best conditions for teak (White, 1991).Dcpending on the challenge, resorting to tissueallture for germinating reca1cilrant seeds can becnvisaged, as already successfully undertaken(Monteuuisl't al" 1998).

ln vivo

Wîthin our project, ln VIVO conservation plotsronsistmainly in the resources existing in the nursery or inthe base and broeding populations, in other words,in demonstration plots, provenance-seed source/progeny trials, clonai tests and seed stands.Practically, mainly gcnotypes exhibiting superiorphenotypes will be preservee!. In that sense, ourconcept of germplasm conservation, deliberatelyoperation-oriented, might be too restrictive. Cost,however, remains (orusa majorconœm. Wecannotafford to set up and maintain ex-situ conservationplots for ail the genotypcs (rom as many differentorigins as wc can gcl, owing 10 variotls factors suchasavailability of sites, costs in silvicultural practicesand manpower for maintenance of the plots

br vitro

ln vitro culture conditions can be a good option forgermplasm conservation (Haines, 1994), in a morerestricted environment than ln VlOO plots wherebyproper maintenance remainsa critical issue(ZobeI and.T,Ibert,I984).1kp'thog<!n-free_oftis5uecultureallow the intemational exchangeo( living plant

material without any sanitary or climatic constraints,contrary to ex-vitro plantsor plant portions.

Table 2. List of Families obtained from the IvoryCoast Clonai Seed Orchard and planted wilhin ourproject

Currently, our in vitro gene pool is limited to thegenotypes under micropropagation. The protocolsdeveloped havebeen conœived as very conservativefor maintaining the various genotypes for severalyears under sustainable sub-culture regimes, whilepreventingsomaclonal variation risks (Monteuuisetal., 1998; Goh and Monteuuis, 2001).

Clonai orgenotypic fidelîty, whieh is a requisite forgene bank, remains acrucial conœm for us. Anotheroption under investigation currently is the resort tocryopreservation, Ihe efficieney of which has beenproven for other tree species (Ashmore, 1997).Developing protocols adapted 10 teak will befacilitated by the possibility to regenerate teak plantfrom in vifro meristemculture, contrary to many treespeeies (Monteuuis, unpublished results). Themainadvantage associated with cryopreservation is thepossibility to storegenotypes for unlimiled periodsof times ina very resITicted envirorunent and wîthoutmaintenance requirements, while preserving theirintegrity, as detailed by Ashmore (1997).

Mass selection prevailsat the firsÎ" step of the selectionprocess. Individual selection is mainly phenotypic,with special attention to traits of major economicalimportance for teak such as: growth rate, smallnodes, baie straightness and basal circularity inabsence of butlress or flules. Wood qua lityassessment including bath aesthetic (wood pattern,lexture) and technologiea! characteristic:s has also 10he taken intoconsideralion for phenotypic selection.

SEED STAND OPTION

STRATEGIES fOR SEITINC-UP IMPROVEDYIELD AND QUALITY PLANTATIONS.

From seeds

Thedifferent origins listed (fables 1and 2) have beenset up on easily accessible ICSB stands according 10planting designs adapted to the conversion of thedemonstration plots or provenances-seed sourcelprogeny trial intoseed stands. The Iwo provenancelprogeny trials were set up in a partially cquilibratedincomplete block design and were comprised of 41and 42seedlots respedively, with 26seedlots rommontobothtrials(WiIliamsandMatheson, 1996). Thetwosites were very dissimilar in terrain in lhat one ison a'hilly area (Luasong ForestryCenter) whereas the otheeis located on lowland (faliwas, Lahad Datu). Datafrom these trials will thereby also allow us to assessthegeneticoriginxenvirorunent interaction. Further,as much as possible,close victnityofgenetically relatedindividuals within eath trial has been avoided. Eachprogeny elementary plot did not indude more than 5sib1ings inorcier to minirnize the lasses when selectingthe best in the plot to he kept as seed producers, whilefelling the others inorder to prevent risks of înbreedingdepression.

Thetraditional meansof propagaling leak is throughseeds, as has been practieed for centuries, with lhepossibility of storing the seedlings in lhe form of"stumps" when necessary (Kaosa-ard, 1986;Tin Tun,2000). Various seed-based strategies have beensuggesled by specialists for improving the genetiequality of teak seeds (Wellendorf and Kaosa-ard,1988; Kaosa-ard, 1998, 1999; Kjaer et Q/., 2000). Theseed stand and the seed orchard options are worthconsidering in ourcontext.

1310323212133332221

Number ofseedlots

Talai: 56 seedlots17 origins

India NellicuthaIndia NilamburIndia Vemolirgetndia VaIndia PurunakoteIvory Coast BamoroIvory Coast KokondekroLaos PaklaySenegal DjibelorTanzania KihuhwiTanzania MtTanzania BigwaThailand Huoi-Nam-QonThailand Maasale ValleyThailand Pong SaleeThailand Ban Pha LayThailand Ban Cham Pui

Total:

Grigin of seed lots

393

ln this respect. wc arc placing a lot of emphasis onthe uhlizalion of non-<:!estruclivc methods for woodqualily analysis (Baillères and Durand, 2000).

Furtner 10 thiSsclectionof plus tœes fordonaltesting.inferior trees and half-sib in close vicinity are culledinorder to lavor the intennatingofsuperior indi\idualnot gmehcally relaled. In the absence ofsufficientlydetailed records, rcsorl 10 molecular biologytechniques such as AFLP dc\'cloped in Cirad-ForêtIaboralory can help in thedetenninatÎon of thegeneticrelaledness among îndividuals withîn the sameooghborllood.The lurtt.... testing of thesclectedsœdproduœr.;01 the sœdstmd00sed on the perlonnanœsof thcir progeny will indicale lheirrombiningabilitywhich can be used for rcfined or more advanœdsclectooandcullingactivilies(lqaerand Faster, 1~;Mandai and Chawhaan. 1999). A final densityof 120- 180 seed produœrs per heclare of seed stand îsexpected at lhe end of lhese selection and roguingactîvilîes(Figure 1).

SEED ORCHARD OPTION

The besl combining genolypes or "combiners" canhe asexually propagatcd or duplicated 10 be mixed

according to a well-suitcd plantmg design within a\-egelalive seed orchard consisting clones fromdifferent familiesand provenances, as illustrated inFigure 1.

Thesc clones will be produœd on their own rootsystem as either rootcd cuttings. microcullings oreven layerîng (Lahirî. 1985; Monlcuuis,1 al , 1995;Monteuuis ri a/., 1998) in order to prevcnt graftingmcompalibility problems and Ihe consequenlialproduction of "ilIegitimates". These are likely 10dcpreciate the genctîcquality of the seeds produœd.firstly, whencollected directly from suffi unexpeeled'mothers' sinœ iIIegitimatesare most of the lime hard10 dîslinguîsh from the grafts as Ihey look similar.and secondly. as these will pellute the genetîcqualityof theseeds produœd by lhe "Iegitimates" around.

However, the numerous question marks andunœrtaintiesassocialed with lhe real benefits thal canbeexpected from such orchardshave 10 beserîouslypondered (Kjaer and Foster. 19%; Kasoa-ard ri al..1998; Kjaer ri al.• 2(00; While and Gavinlcrtvatana.1999).The implcmentationof trus strategy is thereforeonly 10 beconsidered for the long lerm.

IMPROVED QUALITY SEED PRODUCTION STRATEGY

U~

"_"l'I••h

OCA ....1.......

,_.,._n_...... ..·'.. 00__..'010_ ...."-....

n~,,-".. , .

GCft. ...i ' ••

$_ podue!l_

--. B ...

CtonalseedEliteTree --Sad St.nd (ta1 u ....em...n)

r ....., .....

Q~ ••.....n....... 0.__ 0'bo........... •... E....

=> •=>

•po_...... 00"'''''''' Qo"hoI' • ..._Clpt...........

Plus T.eeSad Stand

S9...c1.,.0duc4 .. _

....... " "'...

..~." .(Nol..... ) ...do.. ...., ...

"*"""'9.' ,.....­........,.."',....

_....'_Iv· ....,..........,...-

Hoo.............._ .._<l

......".,.-.."".

" ... '''"",Iio"Cil •• T'.....

N'.""'ph_l_

BasePopula'.on(P'o-.enanœ

'po-ogenylnab)

394

Figull' ): Illustration of the project striltegy aiming al producing improved qualityseeds in the shorlest delay slartîngfrom genetic.lll)'-rich base populations. Ad\'anœd generations of clonai seed orchArds can he envîsaged for theI\ln~ lerm

395

FROM CUlTINGS OR MICROCUlTlNGS

Production of teak planting stock by seeds, althoughprimarily cmployed. remains severely handicappedby the followings factors:

1. Quantilalively limited seed production(Wellendorf and Kaosa-ard. 1988, White,1991 ).

IL laie flowering. Il is noteworthy that in teak,straight bole length, which affedsdirectly ilsmarket value. is strictly dependcnt on thecapadty of the terminal meristem 10 remainvegetative as long as possible (White. 1991).Ils conversion iota the f10wering stageinduœs a lork formation, as the result of atrue dichotomy proœss.

iü. Lowgermination rates (Kaosa-ard ttal., 1998;White, 1991). In Thailand, for e.xample, onIy5 plantable seedlings on average can bep<actically "'J"'CIed (rom lOOseeds ID' Luge­scale nursery (Kjaerand Fosier, 1996; Kaosa­ard,I998).

iv. Substantial variability among individuals,even among half-sibs, within progenie<>, withregard 10 traits of major economicalimportance such as growth, form,technological and aesthetic characteristics(wood pallem) (Dupuy and Verl>aegen. 1993;Kaosa-ardet al., 1998; Bath, 2<XXI);

v. Limited accurate genetic knowledge aboutthe inheritance of such economicallysignificant traits, and consequently, sorneuncertainty for the ultimate gain,notwithstanding the time conslraintsassociated with sound breeding programs(Wellendorfand Kaosa-ard, 1988; Kjaer andFoster, 1996; Kaosa-ard, 1998).

The hindranœsand uncertaintiesassociated with longtenn breedingstrategies for teak greatly penalized bylow seedling pnxiuctivity havebeen deve10ped (Kjaeret al., 20lXl; White and Gavinlertvatana, 1999).According to lheseauthors, lhe magnitudeof the realgenetic gain associated with the seedling route hasyet to be c1early defined, and the basic question toknow whether ail the efforts invested during the pastdeeades are worthwhile remains. This isundoubtedly

amajorconœm for potential investors, forwhich rapidpay-off is a decisive argument. From practical andtheorelical infonnation, il can beob;ectively assumedthat for leak, greatergeneticgaincan beexpectcd fromthe cuHing foreslry option, especially whcn clonai,!han !ha, from theseedling 10"",')' (ZOOeI .ndT.1bert,1984; Ahuja and Libby, 19933 and b; MonteuuisandGoh,I999).

Two different strategies can he considered forestablishing wood production populations fromvegetatively mass-produced teak planting stock,either in bulk fonn or by clonai propagation.

TIlE BULK OrnON

Bulk propagation consists in vegelalivelypropagaling a group of mixed genolypes withoulmainlainingany individual identification. Thiscanbe useful for increasing the number of a limiledquantity of juvenile genotypes of presumably highand. sirnilargenetic value, derived for example, fromcontroUed pollination.

The main advantage of the bulk propagationoptionlies in the absence of a need to strictly and c1earlyidenlify by proper Jabeling one genolype fromanother. This option will also maintain a certaindegree of genelic variability depending on thenumberofgenotypes involved at the beginning, andwhich in tum may inducean overall heterogeneityin the resulling wood production populations. Thisseems espe<:ially true for teak considering thevariability among genotypes. However, successivegenerations of seriai propagation may eventuallyresult ina significant reduction of the original geneticbase due to genolypic differences in themultiplication rates, i.e. in the number of shootsproduœd to beusedascuHings withsuffidently highability for adventilious rooting.

Bulk propagation has been reslricted in our projectto superior quality, quantitatively limited or rareseed lots propagated under in vitro conditions. Atlhis stage, the genotypes are too young 10 expressany individual differenœ wanantingaselection. Theoriginal geneticbase in optimized micropropagationconditions, contrary to nurseryenvironment, is thllSmaintained at 1east during the first several subcuJture

cycles. Howevcr, over lime, with littleor no conlrolon lhe gcnolypc-dependenl capacity for axillaryshoot production, the risk of gradually losing cloneswith rcduœd capacity hasto beacknowledged.

THE CLONAL OPTION

Talbcrt. 1984; Ahuja and Libby. 1993a and b). Thepossibilily 10 reproduce. lheoretically in unlimitednumhers. the best Irces. from quality and yieldstandpoints, for large-scale plantation uses. offerstremcndous prospects.

ln clonai propagation. contrary to the bulkpropagation strategy. the genotypic identity isrigorously and individually preserved throughsuccessive propagation c)'des. which may last5e\'eral œnturies III œrtaincases. Eachcloneronsistsof a.sexually-derived offspring with virtually thesame genetic make-up. rcgardless of the numberofils represcntatives.

Oonal propagation im'olvingscrupulousgenotypicidentification ensures a better control of the plantmaterial propagated by cuUings than by the bulkoption. in addition toa numberof olher advantagesassociated with doning of foresttrees (Zobel and

The basic requisite for sucœssfultcak propagationby rooted cullings is the exi:.tcnœof a good capacityfor adventilious rooting (Monlelluis ri 0/.• 1995).Fromour first field observations. it appears lhat oncerooted. teak cullings de\'elop Htrue-to--I)'J)(''', and agood wilhin-done uniformily can he ultimatelyexpeeted. Increased yield. higher uniformity foreconomically important traits such as growth rate.trunk form, straight bole length. woodcharacteristics. and shorler rotations constitutestrong inœntivcs todevelop teak donal plantations(Wellendorf and Kaosa-ard, 1988; Bath. 200). 50chunifonnity can not objectively bc expected fromplantations set up from seedlings, or even fromcuUings issucd from bulk propagation by virtueof

CLONAL STRATEGY

3%

Figure 2; Illustration of the pro;ect strategy for Iarge-scale cml! plantations olll.'.ll MolecuL.u markets can hdp with

l''-'dJgree information, certîfK'atÎQn as weU as for genetk reLlledness between the dones used for c\on.ll deploymenl

Advanced soedioll

Clonai perfonnances IcharacleristlCSacCOl'ding 10 site conditions

Additional 'NOod (desruclt-/e) analyses

Genetic relatedness (rmlecUlar markers)

~D-_~Mlmo_

Mass selection of .... Trec:s ..

based on phenotypic criteria

Famly. 'tNithin family selection

Non-deshuctlve wood analysismethods

Base Population(Provenance/progeny lrials)

the arguments devcloped previously. Properlyselected and wisely dcployed clones will thusmaximize the shorHerm returns From suitableplanting sites, which are dramatically reduang insurface area (Bali dol" 2(0)).

Utilizalion of clones can he profitably adapled 10inlercroppingafter properselection particularly,oncrown form. Planling densily and silviculturepractiœs can he adapted ta inlensive managemenlsystems. with the p<xssibility to harvest severallimesfrom the same slump, laking advantage of theexcellent coppicingability (Martin tt al., 2(0)). Sochpraclices look very attractive for enhancingplantation yield while significanUy reducing theplanling and lhe (micro)cutting rosts. With regardto these financial aspects, it has ta he emphasizedthatclonal plantationsgener.illy require le;,; plantingslock than plantations eslablished from seedlings,whichcompensate, laacertainextmt. the highercœtof (micro)cuttings. Figure2summarizesthedonalstralegy adopled for our project:.

Elficitnt techlliquu10rmassfJt!gt!latroe production01 silpaiorquality planting stock

Mass production of rootnJ cuttings in nursny

Mass propagation of leak of any age by rooledcultings in nursel)' conditions has hem developedand become fully operational withinour project sinœ1992. The requisites, as weil as the advantages andIimils of Ihis propagation option have beenexlensively prescnted and confirmed by a decade ofexperience (Monteuuis, 1995; Monteuuisd 01., 1995;Monleuuis, 2(0)). Average rooting rates of8O%areroutinely oblained From malure genotypesintensively managed as container-grownstock plantsonce the mobilizalion phase has been sucœssfullycarried out. In such conditions, annual productionof600 rooted cuttings per square meterof stock plantarea (15 stock plants persquare meler) q1n he easilyoblained (Montcuuis el al., 1995).

ln vitro mass propagation

Theavailabilityofa well-equipped Iaboralol)'withinour joint pro;eet prompted us to explore the prospectsof propagaling teak in lissue rulture conditions via

microcuttings (Bon and Monteuuis, 1996). Thissuceessful application has already becn reported(Monteuuisetal., 1998;Goh and Monteuuis, 2IXlI). Theconœived tissue culture protocols wcremadeas simpleas possible in order 10 beeasilyapplicableand tocopewith the constraintsof large-scaleapplication in tennsof cosl efficiency and high produclivity. Massmicropropagation of any genotype, either in bulk orcIonaUy, throughaxill<uy-produœd rrticrœhoots withan exponential multiplication rate of three to four atlheendof f!Verysix week-durationsub<ultures is nowpossible. The rooting-acclimatization phase isadvanlageouslyachieved in nurseryconditionsunderamisI-system with more thanm SlJ<.'ŒS501l average.After weaning and raisins onder the same intensivenursery conditions as for the cutting.œnved plants,the tissue-cu1ture plants develop iolo vigorous andtrue-to-type vegetalive plants. More than 400.00>miaocu~have been produœd up tonow forkx:alplanlationsasweil as for oversea markets byapplying!lUs tedm;que.The pœsibilily ID send the in "",,,plantsoff ta differenl destinations, regardless ofdistance, inthe absence of phyto-sanitation restrictions,considerably expands lhe market prospects andconstitutesanoutstandingasset.

The in vioo rooted cutting and in vitro microcuttingoptions forvegetatively mass propagating superiorteak genotypes with the respective pros and conshave been extensively reviewed, leading to theconclusion that the best oplion, in many respects,consîsts in the combination of the Iwo techniques(Monteuuis,2OO)).

CONCLUSION AND PROSPECTS

The Research and Development collaborativeprojectbelween less and Cirad-Forêt was iniliallyimplemented towork primarilyon rattanspecies. Theshift and the emphasisgiven to teakstarted in the carly1990's as a result of the unexpected sucœssobtainedfrom the vegetative propagation lechniquesdevelopedat the nursery and tissue culture levels. The possibility10 mass produœ clones by rooted cuttingsor in vitromicrocuttings in cost-effective conditions from anyselected mature superior teak tree, as weil as fromgood-performing teak Irees introduced locally,prompted us to inves! more on these adivities. Therationale and the prospects of using clonal materia1s

397

for leak have been indœd advOGlled for a long lime.Beingawareofour vegetative propagation assets, thenexl step !lasbeen 10gatherourownbasepopulations....Tilh agcnetic background as rich as possibleThisaim!las been attained to a large exlenl by our possesslonof a highly diverse teak genetic resourœ thal coversIhe range of adaplabilily of Ihe spedes. Thisnoleworthy genetic richness combined 10 efficienlpropagation and diffusion lechniques are Iikely 10satisfy any plant malerial arder, in the form ofseeds,or clones in tissue culture conditions for over5eamarkets, with Ihe possibilily 10 œrtify Ihe plantmaterial by DNA fingerprinting.

Keeping up wilh the Ialest advances in lechnologywhich can further bcnefit lhe projet! rcmainsa majorconccm. In this respect, we are placing emphasisonthe application of motecular markers such asmicrosatellites and AFLP lechniques currentlycmployed for leak by Cirad-Forêt, as weil as non­deslructive woocl analysis mclhods for moving oneslep forward in the selection of superior pIanimalerial and clonai dcployment. Proper site xgenotype matching indeed deserves particularattention with respect to pest and disease(defoliatorsand borers) aspects, as weil as to heartwoodformation, controlled by genetîc and environmentalparameters, and which delennînes log qualily(Bailleresand Durand, 2OOl}

The need to intensify the production of premiumquaüty lùgh v.>Iue tin>œs..eakbeing the most prizedone, with a worldwide demand far grea!er lhan thesupply available, isa slrong inœntivealong this Hne.

REFERENCES

Ahuja, M.R. and Libby, W.J.I99,Ja. C/01l~i Forrslry l, Grnrtics~ndBiotrrhnoiogy. Springer.Yerlag, Berlin, Heidelberg, NewYork, l.ondon, Paris, Tokyo, Hong Kong, Barcelona,Budapest, 277p.

Ahuja. M.R. and Libby, W.J.I993b.C/otW/ fomtryll.CortsrmltlO1land AWirlltlOll. Springer-VerLtg.. Berlin, Heidelberg. New·York, London. Puis, Tokyo, Hong Kong, Barcelona,Budapest,24Op.

Ashmore, S.E. 1991. StatuJ report on th~ devdopmenl andappliation of ill tll'Iro t«hniques for the conservatiŒl and~ofrlantgmftic~ IPGRI.~,67p.

&t1,J-B.,P~, o. and Hini.5. 2OXl.GIobaI O\·~ofl~akplantations. In: S"~, Trdl/sology and ProJlldWlty rf Tt'lItPWrIllIIllll$. TEAKNET PubliaIbon No. 3, 11.J3.

a.iUms,H. and Durand,. P.Y. 2OXl.N~v~ kduUquesfor wood qu.ality assessrnerol ofpLtntabon-grown luk. Bols

398

rf foritsdrs TropiqutS 263,17·"19.Bhat, K M 200(1, Tlmber quahly of teak from maniged

ptantatKJns of the tropiC'i. Bols rf fo"lsdrs Trtlp'lqllN 263, G­16

Bon. t-t. ..c. and MŒlteuws. O. 1996 Btot~ knst~

au SAbah; pmruer bilan. &llSrf forfudrs TnlJ"'l1ll'S 248,31­4L

Cameron. A.L 1966. GmftJc lOlprovtmml of It'ak m NtwGUU'M'a Allst. fortslry 30 CI): 76-Ç1,

Duruy, B.andv~ 0_1993, lzltddt pLtntabon TrctonoIgnrndas ftl COted1~Vlre.&IlS ri rom, lits Tn"fIIII'lIrS 235: 9-24

Goh, 0 and Monleuuis, O. 2001. Produdlon of Iwut rultum1teak: the pLtnt btol~ogy laboritory upmmœ. In:Potrnhall7ltlOppommÎtItS m Mlrrb'frllg,1Id T,,,,,, rfPlalllat/OflTt'lIk:Chilllrngrp IlrNnvMi/lrnlllm. Proct«hng of the 1lurdRtgionalSmUnaron Tealt, Yog)'OIlarta, Indooe51a,31 July.4 August 2000, 237-247.

l-JaultS, R. 1994. Biotechnology in forest trtt Improvemml. FAOForest')' piper 118, Rome, 230p.

Kaosa-ard, A. 1986. Tuk, Ttcton~ grlndu l.mn. f.: nurserytechniqu~, with ~al l"('ferenœ to Thailand, OANIOAr>QrestSeedCenter, Seed leaOet No. 4, Novembcr 1986, 42p.

KaO$Ol-ard, A.I998. Teak breedingand improvement strategies.ln: Tratp Iht flllll". Pme. oftheSecond Regional Seminaron Tt'ak, Yangon.lndonf5ia,"19 May - 3 June 1995,41-81.

Kaosa-ard, A.l999. Teak (Ttd01lllgnlluirs L.mn.f.): Oomesticahonand bre«ling. Teaknet publication No. 5/1999, 86p.

~-arti, A., Suangtho. V. and Kjaer, E.O. 1998. ExptnenœlTom Irt>e unpruvtrnent ai !l'ak (Ttd(lllllpndu) Ul ThidOind.DANIDA Ttdmial/ N,,"No. SO, 14p.

KJBtr, E.O. and Fosttr, GS. 1996. The KOnomia or Irft'~aiTsk(Ttdor\IIgrand"L~DANIDATtdm..Ndt ND. Q, 23p.

Kjaer,E.D~ I<MJsa-ard,.A. andSüangtho, v. 2OXI.~tIonai tuk through Irft impruvtmml Optlons, possiblt gaUlSand ailical fillClOB. ln: Sdr, Trdlndogy.Jul Prodlldwrty rfTrlÙc PinlllltlllflS. TEAKNET Publiabon No. 3, 161-189.

ICnshnaptllay, 8., Norini, H.. Mohd Noor, M., ZuhadJ, A.Y.•Gharu. A !lA. and MMunud, A.W 1991. P\ilntUlg !l'ak Ul

~uLtysu for rornmerci..llISt -1$ Il vublt? Paper prtsmltdal the AsIa P;aafic f'lanbhon Fure;try Confer~, KWlI;aLumpur, Mala}'Sia, 26- 27 May, 1991, 13 p.

iAhiri, A.K.. 19&5. A note on possiblhties of mound LtytllJ\g 01teak. TIlt flldlllll fortslrr 111(10): 870-871.

Lapongan, J. 1998. Siaius 01 teak plantallon managemml Hl

Sabah. In: Hrgh Valllr Trlllbu SpiCIt, for PlalllatrOIlEsla"h,hlllrlll- TtIl1: IInd Mllhogillllrs .ProcCf'dmgs of tht'Seminar, 1-2 December 1998, Tawau. Sabah. JIRCASWorking Report No. 16, pp. 19-25.

Martill, B., Kadio, A. and Offi, K. 2000. Towards Teak cultureintt'llsification in Côte d'Ivoire. In: SIIr, Trchllology IIndProdru1ivityofTtrl1: Pla71tatioll5. TEAKNET Publication No.3, 151-160.

Mandai, A.K.. and Oul'o"hann, P.H. 1999. Progrny Iesttd phutret'5 in Ttik (Ttdorur gralllllSl. Tl'Ilknrl NtwSlrtttr 15: 1-3­

Monleuuis, 0.1995. Reœnl OIdvUK1'S in c10ml propagation 01teak.1n: Pnx. of the Inltmational Work!;hop 01 BIO-REroR.Kangar, MIlaysia, Nov. 28 - OK. 1, 1994, 117-121

MontnulS, O. 2000. Propagatmg Itak by cutlLngs andrnKTOCUttmgs.. In: sm, TrduloIogy and ProdNdrrlrty rfTNkPlantmlllflS. TEAXNET Publicabon No. 3, 209-222.

Monleuuu, O.• VaiLtun, 0 .. Poupud,C Huard, L, YU5Of, Y,Wahap, l.A., GUCiOl, C. and Chauvlèrt, M 1995.

Propag~tion clona le de tecks matures par bouturagehorticole. Boist! FQrtls dts TropiqutS, 2n: 25-39.

Monteuuis, O., Bon, M.-C. and Goh, O.K.5. 1998. Teakpropagatioo by ln vitro rulture. Bois ri FQms dis Tropiquts256,43-53.

Monteuuis, O. and Goh, O.K.5. 1999. About the use of clones inteak. Bois tl Fortts dts TroplqUtS 261: 2&-38.

SaIJeh, H. 1995. TtQbll Silbah-Q suslalnabiragrojiJl'tstry: /lIt' HamsSaI/th tXPfritllcr. Published by Sejati Sdn Bhd, Sabah,Malaysia, np.

Tin Tun. 2000. StOled teak stump, simple tedmology but veryreliable results can be amieved. Tmnt/ Nt'WSltlltr 19: 5-6.

Wel1endorf, H. and Kaosa·ard, A. 1988. Teak improvementstrategy in Thailand. foust Tm /mprlJV('mrllt 21, Op.

White, K.J.I991, Teak: some aspcctsof rcsearchand develomenl.

FAO Regional Offiœ for Asia and the Pacifie (RAPAj,publication 1991/17,53p.

While, K. 2002. General plantation management. TtaknrtNrU's/rl/rr 26: 4..

White, K.J. and Gavinlertvatana, P. 1999. Vegetativereproduction ofTeak: the future to increased productivily.Papel presented at the Regional Seminar on "Site,technology and productivity ofTeak plantations~,ChiangMai, Thailaru:l, 26-29.1.1999.

Williams and Matheson. 1995. Experimental design and analysisfor use in tree improvement. C5IRO Information Service,314. Albert Street, East Melbourne, Victoria 3002, Australia.

Zobel, B. and Talbert, J. 1984.. Applit'd fortSI Tru /mprovrnrtlli.John Wiley and Sons, New York, Chichester, Brisbane,Toronto, Singapore, 505p.

399