conditions in degraded peat swamp forest

ECTOMYCORRHIZAL FUNGI INCRI,ASED EARLY GROWTH OF Shorea balangeran (Korth,) Burck UNDER NURSERY AND FIELD

CONDITIONS IN DEGRADED PEAT SWAMP FOREST

Ftngi

Ektotnikoliza Menihgkatkan Pet'twkbuhan Awal Shorea

Balargeran (Korth.)

Burck

pada

Kondisi

Persentaia dt

^{Lepangtut di}

Httan

Rawa

Ganbut

Terdegradasi)

by / oleh ^:

\laman Turjamanr,

Hideyuki

Saito'?, Erdy

Snnlosol,Agung

Susantor, Sampdng Gamanr, Suwido

Hesler

LinTin3, Masato Shibuya2, Kunihide Takahashiz, Yutaka Tamai2,

Mjtsum

Osaki2 and Keitaro Ta,,varayal

lForest

_and

Nature

Conser_vation Research

&

Development

Centre, Ministry ofForcstry, Bogor l6610,Indoncsia, E-mail: turjaman@yahoo,com.sg lGraduate

_School

ofAgriculture, Hokkaido

UDiversity, Sapporo 060-8589, Japan

rFacultv ofAgriculture, Unirersity

of Palangka Rayn, Palangka Raya

73l12, Indonesia

{Faculty

_of

Agriculture,

Yamagata

Universit}, Tsuruoka

997-8555,

J.pan

ABSTRACT

Seedlings

of Siorza

balangeran

(Ko*h.) Burck

were inoculated

with

spores

of

four :;romycorrhizal

(ECM)

fitr,gi, Calvatia ^sp-, Boletlts sp., Scleroderma sp.

a

^d Strobilotn),ces sp-, r1d \1ere grown in pots containing sterilizedpeat soil for six months under nursery conditions and

-nsplanted in field

conditions.

Four ECM ftrngi were

collected

from

^peat

soil of

Nyaru

\tenieng,

Palangka

Raya, Certral Kalimantan. Six-months-old inoculated

seedlings were -:"nsplanted into degraded peat-swamp forest. The

ECM

fungal colonization was 59-67% in :roculated seedlings under nursery conditions. Colonization

of

S. balangeran

(Korth.)

Burck

:rcreased shoot height, stem diameter, leafnumbet and shoot fresh and

dryweight

6 months a{ter lansplanting

[nder [ursery

conditions. Shoot height, stem diameter,

leafnunber

and survival

-tes

^{of S.} balangeran

(Korth.)

Burck were higher in the inoculated seedlings 40 months after

-nsplanting

^{under field} conditions than in the control seedlings.

It

is suggested that inoculation

,.i

indigenous

ECM fungi in

native tlee species

is

useful

for

reforcstation

of

degraded peaF

$amp

forcsts.

Kellvords

^:

ectomycorrhizu,st)ore,Shorua bala

^gerar.

(Korth,) Burck,

peat-swamp fbrest, Eursery and

field.

ABSTRAK

Blbit

Shorea balangeron

(Kofth.)

Burck telah

diirokulasi

oleh empat

jenis

_ceudawan

:iiomikoriza ^(ECM)

Srr?bilonl'ces sp., Culvatia sp-,

Bolet

s sp., dan Sclerodeirn a sp. sclarna 3nam bulan di persemaian dan ditanam 40 bulan pada kawasan hutan rawa-gambut terdegradasi

:i

Kalimantan Tengah. Empat

jenis

_cendawan

ektomikoriza

berasal

dari Nyaru

Menteng, lalangka Raya, Kalimantan Tengah.

Betlh

S. balangeran

(Kodh.) Burck

diinokulasi dengan .pora empatjenis ECM dan ditumbuhkan pada po)ybagyang berisi gambut selan'la enam bulan di r.rsemaian.

Bibit

S. balangeran

(Kottlt.)

Burck yang telah berumur enam bulan ditanarn pada ..:a\!,asan

hutan

rawa-gambut

yang

terdegradasi.

Kolonisasi ektomikoriza Bolea,s sp.

Dan St leroderma sp. secara signifikan meningkatkan pertumbuhan tinggi dan diameter S. Dnlnrrger-rur

(orth.)

_Bllrck

40

bulan setelah ditanam di lapangan. Dari hasil penelitian ini bahwa diperlukan ::okulasi ECM lokal terhadap perftrmbuhanjenis-jenis Dipterokarpa ^pada kondisi diperscnraian rnruk mempercepat rehabilitasi kawasan hutan rawa-gambut terdegradasi.

f,ata kunci: ektomikoriza,spota,Shorea

baldfigerun

(Korth,)Burck,

hutan

rawa-gambut,

persemaian

drn

lapangan.

I

JURNAT PENELITIAN DIPTEROKARPA Vo 1 No.1, September 2O0T

I. INTRODUCTION

The biological and economic importance of Dipterocarpaceae lies in the extraordinary dominance

ofits

members ovor vast areas in tropical min forests ofSoutheastAsia (Bawa, 1998).

With approximately 5 t 0 species and 16 genera, the

family

may not be particularly large among

hopical woody goups. Other families such as

Euphorbiaceae,

Myrtaceae,

Rubiaceae, Annonaceae, and Lauraceae have

i\ote

taxa than the Dipterocarpace4e, however, they are parl tropical in

distdbution.

Some important generaof

Diplerccttpaceae

^arc Shorca, Dr)'obalanops.

Hopea,

l/atica,

Parashorca

and

Dipterocatprs

(Soerianegara and Lemmens, 1994). Shorea consist

of about

¹⁹⁴ species and be

the

grealest

divercity

occurs

in

Sumatra

(23

species), Mollucas

(l

species) and Kalimantan (62 species). Unfortunately, theit existence is decreasing because ofover-exploitation, forest firg and illegal

logging.

Shorca balangeran

(Kortl.)

Burck is

distibuted

over scattered locations ofBangka, Belitung and Kalimantan islands and attain a height

of30

m and diameter up to 60

cm.

This species is common ^and occurs often gregarious in tropical peat-swamp forest. ^S. balangeran (Korth.) Burck is economically important, because

it

produces a major source

of

durable wood (heavyweight hardwood) and valuable resin can be obtained from ^the wood.

Tropical

peat-swamp forcsts have beeh decreasing

dre to

forest

fire, illegal

logging, convemion offorests into

indust

al and agricultural lands, rubber tr-ee and palm oil plantation or forest plantation estate for pulp arld paper. Degraded peat-swamp forcsts are recognized ^as

low

values of forest resources

without

s[ccessflll natwal regeneration and dominated by gassland including fern, sedge or scrub ^{(Page et} al., 1999; Kobayashi,

2000).

Indonesian gove.nment has a

frogram ofnational

movement to accelerate rehabilitation

of

degmded peat-srvamp forests.

However, it is not easy to rehabilitate this ecosystem immediately because it is necessary to select

and

^produce

high-quality tree

seedUog species

for rehabilitation. Mycofthizas affEcl

lhe mainteoance

of vegetation in various

ecosystems,

and may play an important role

in rehabilitation of degraded peat-swamp forest (Tawaraya et al., 2003).

Dipterocarpaceae, Fagaceae. Pinaceae, and some genera

ofMltlaceae

are known

lo fonn

ectomycorrhizas

(ECM)

(Alexander _and Lee, 2005).

All

Dipterccorpaceae strveyed

lo

date are associated

with ECM tungi

(Hogberg, 1982;

Smits,

1994;

Lee,

1998), and fungal

families with ^greatest diversity in

Southeast

Asian dipterocarps forests include

^the

Sclercdermataceae,lhe

R ss

ldceae,the Boletacede aDdthe Anenildceae (Smits, 1994;Simset

al.,

1997),

ECM firngi may

have

a

great potential

for the regereration of hopical

forests Dipterocatpaceae

is a

large and diverse

family, with

great ecological amplitude, and ECtr{

dipterccarps are found in a wide range foresttypes, growingunderwide range ofnutrient regimes (Smits, 1994; Alexander and Lee, 2005). The ECM formation has been shown to increase platrt

growth of

S&ored

curtisii

^Dyex

^ex King

and S.

Leprcsula Miq. (Lee

and

Lim,

1989), S,

macroptera Dyer (Tufnet et al., 1993), Hopea odora la Roxb. ard H. he lferi Brandis (Yazid ^{et a1.-} 1994) under nrrsery conditions. However, little is known about the effect

ofECM

inoculation otr the growth ofdipterocarps species ^or Siorea species originated from a pgat-swamp forest under nuisery conditions.

In tropical forest, tbe

[se

of ECM Pisolithus

tincto

us (Pers.) Coker

&

Couch has beed tested

with

Prflus

carriraea

Borelet in

field

conditions after three yea[s

o[

savana itr

Libeda Africa (Marx et al., 1985). Five ECM flrngi

increased

plant growth of hybrid elcallpa

(Eucalyptus

urophylla

S.T. Blake

x

E. birtaniand F.

Muell)

50

morths

after outplanting in a nutrient-poor acidic sandy savanna soil in the Congo,Africa (Garbaye

e/a/.,

1988).

Thiskindoi

fieldexperiment has been already described intempemte ^area with

fourECM

species

inoculaloi

to Pseltclotsuga menziesii

(Mirbel)

Franco (Douglas Fir) in

rorthem

Spain (Pera et al., 1999),

ard

2

,oe,o, {knh.l 3urct UNoER NURStRY

I

ECM Luccariu bicctk ^-

(Meile)

P.D. Orto0 inoculatcd to

P

,?.nzle.rli fN,lirbcl) Franco (Douglas Fir) in ccntral Francc (Selosse ^et al.. 2000). Ho\\'ever, thcre are no reports on lhe

elltct ofECM

inoculation

on thc groulh ol

^.\hoteu

:iP((i(l^ origin tiom a

^pcat

s\verl+

t'orcst

undcr icld

conditions.

Scvcral works have been reported on the lypc

of

ECNI inocnlum uscd in tropical trcc spccies. N,lycelia

ofECNl

have been inoculaled to incrcasc

flent

^{e|owth ofHopcrT spp. (Yazid et} ei., 1994).]'ablel spores

ofECM

hilve

contiincd

^to bc cllcctivc on seve ra I eucal)'pt species (de la

Crrz.

1990), Spore sLrspension

oi ECM flrngi

has heen tested

to tropicdl

lJee EucolJpftls

razpllla

S.T Rlakc (C'hcr ct al.. 2006) and cuttings

ofJrored

/eplo-vrl.r ^r'l./lq. (Onron, 2002), but

littlc

\\,orl( has been undeltahen on the inoculaiion ofdiptcrocarps $ ith lirngal spoLe suspcnsion.

Objectives

oI

this study

ucr. to

invesli!:ate lhe

eflecl ol

tbur ECNI spccics inoculation using spore suspension on plant growlh oI _^1r.,/?.r haLotga au

(Korlh.)

Burck under nur-sery and

field

conditions. These ECN{ llnrgiwcrcchoscn bccauscthc)'are native in peal-swxmp lbrest.

II. ^MATERIALS AND METHODS

Seeds o1 S. b4lrurgc|rl, (Korth.) BLrrck were obtarncd ^fi om Nyam l\Ienteng arborettlm in Central Kalimantan,

I

doncsia (2' ^,13'

Si lt

^1"

i8'E).

Pcat soil used

fbr

the pot experiment was collected lrom a peat s\\,amp forest il1 Kalllrnpangln, Palangka 11a-va. Central Kdlimanlan

(2'

13' S;

II3'56'L).Pcatsoil$'assterilizeclinadrurnbr-hcatingoverarvoocl1irelbr thour.ThepH (HrO)olthepcatsoilu,asS.3.availableP(Bray-1)was5.9lmgkg L ,\

p|elirainary

erperilrerlt

sho[ cd that this stcrilizafion procedrre got rid ofmost

ofthe EC\I

rnd palhogenic lr.ingi. Seeds of5'.

,!r/,rrgcl?,

\\'ere soaked ibr two houl's ^and gcntly uashed with rrLnning \\,ater. These seeds

$,ere sou,n io polyethylenc pots (sizc I 5 crn

; l0

cm) containing 500 ^g steriliTcd pcat soils. Pots containing seeds

wcrc

translel.led

to

a

nulsery

d! the Uni\'clsit-v

of

Palangka Raya. Central

Kalinlantan.

One seedlillg\\as

glor!n

pcrpotrmdcra7591shading irltensily n.'t ¹⁰

conuolso]ar

radiation.

fruit

bodics

of

CdlvLltia sp., IioLctus sp., Salcro.lctnta sp. and

Sr?rilolrx,r'cr

sp. wele obtaincd

tiom

basidiomes ^ir1 thc

iicld

Lucler native lrees

of

S. balangcratl at

Nyaru

Mcntcng

arborellnn in Ccntl'al Kciirn.rnlon. Iruil bodics \\'crc idcntified

pr-evior.Lsly

by DI Tanli (llol(kaido

l-lniversity. Japan)

in 2002.

Each

fluit

bodies 01

LCIM sll.cjcs

rvcrc blcndcd in

disrilled\!ater(1:10,\',r,)t'or'60susingablerdero1llo\!spccd..{dropofTrvccnS0rvasadded to

assisl sr.lspe[siou.

Inocu]ation of

seedlings

\\as carricd out l0

^da,vs

alicr

ecrminalion.

Seedljngs in each trcetmcnt l\.erc

gi\en

10

nl ol spor.

sllspcrlsiorl. ^_l'hc sporc

su\len\ion uir\

adde.i in a 2-3 crr rlc.p hole near the seedling using i11lv. nrl

fipcttc. Allpots\\,erefutonbricks

in nurscrJ bcds to evoid cr-oss conlarninaLi,-rn and lLr cnsurc qood draioage. l'he seedlings were

irrigatedrvithlap\!alere\erydd)aId\rcadswcrc.cmovcd.l'hclollo$inglrealmel1lsNereLLsed:

(l)

untreated

(control). (2) Crrlratrr sp.. (l)

^B.r1c1r\

sp-. (1\ Sclcralrrma sp. and

(5) StrobiLont,,t:es sp. Tllcrc

\ ^is

^{no nllxtLlrc i:CX{}

i[

^{this stur11'} bccfltsa lhc study ^\1,ould liko to kno\\' thc bcst one

oiECI1

lirDgi.

Expcriment site \rs ^locrled in Kale.rp:rIgiln zonc ncar Kapuas border,

Cenlral Kalinantan. Thc pctrt soil ai stud,v sitc has vely

lo\\'pH

(.-.1). thi<

i(

assunled

du.

to a high organic rnattcr \\.hich pro,:lucing soine

olgallic

acid colnpounds

llllough

decomposition proccss undcr

$ aterlogged oorditions ^(TLrah e1aI.,2000).

Ainual

precipitation is2776to 3193 mmlyear;mean annual tempemture is 30'C. NatL[xl regelaratiou \\'as vcrv

poor.

Nativc vcgetations

ofthis

site are

doniraled

by- ContbreLocarput t'oturulalus,

MelilelLca cnjuputi, Pol|slichlon

setifbrun,

Irtperut

^(:lLin(/t

ica,

i\|elL1:tonn sp. and veriolrs species

ol

grasses.

In

November 2002, one

I

IURNAL PENELITIAN OIPTERO(ARPA Vol.1 No.l,september2ooT

hundred ofseedlings per treatme[t were planted in Kalampangan. The seedlings were planted

individually in planting

hole

(15

cm

x

15 cm

x

15

cm)

and

a line

spacing

was I m x I

m.

Mainte[ance of the

seedlings

after

traosplanting

was

each

six

months

in the first

^year

of

plantation. Chemical fertilizerwas not added in thenurseryand the field experiment.

Shoots and roots were harvested

six

months after being inoculated

with ECM fiingi.

Shoot height, stem diameter,

leaf

number, and

fiesh

and

dry weight

were measued.

After

measuring the fresh weight ofshoots, its dry weight was determired after drying at 70'C for ?2

hous.

To calculate the percentage

ofECM

colonization, rcots were cleanedusing

runnirgwater

to separatethem from the soil and then the root systems were spread

o[

trays. The total number

of

roottips and th€

numberofECMshort

^roots were counted under a dissectihgmicroscope.

Verification

of ECM colonizatior

was obtained

by

exarnining the

qoss

_section

of

root tips (cut manually) under a compound microscope

lbr

the presence

of

mantle and Hartig net (Brundrett er.r/., 1996). Plant height, leafnumber and survival rates were measured

2,4,6,

8, 14,

16,19,22,and25

nonths after transp lanting in field conditions. Plantheight, stem diamet€r, and survival mtes were measured 40 months after lransplanting

in field

conditions. Survival rates were calculated as

follows;

Survival rates (yo) = number

ofviable

seedlings / number

ofinitial

seedlings 100x

100.

Data were statistically analyzed ttsing analyses ofvariance (ANOVA)

with

the statistical software Statview 5.0 (Abacus Corcepts). Comparison of means was done using the least significant difference

(LSD)

method at 5%

probability

level where the F-value was significatrt.

III. ^RESULTS

A.

Nurseryconditions

At

the end

of 6

months

in

nursery,

Calvatia

sp.,

Boletis

sp., Sclerodenna sp. ard,

Sttobilofiyces

sp. formed

ECM

in S. balangeralt seedlings.

ECM

colonization was higher

thafl 50% in all

inocu]ated seedlings (Table 1.). There was

no

difference

in

^percentage

colonization between four ECM species. Control seedlings were colonized by indigenous

of ECM fungi. In the

nursery conditions,

ECM

colonization

of

S. balangeran using spores suspension

of Calvatia

sp., Sclet'oderma sp. and. Strobilomyces sp. increased plant height, fresh ard dry weight (Table

1.).

ECM colonization of Boletus sp.lncreased

onlyplantheight.

There was no difference

in

stem diameter and leaf number between

four ECM

and control seedlings.

Table

l.

Shoot height, stem diameter, leafnumber and

ectomyco(hizal colonizatiol of

^S.

balangeran

inoculated with four ECM fungi six months after transplanting under nursery conditions in Palangka Raya, Central Kalimantan

Treatmenl

Plant grorvth

ECM colonization

(%) Shoot

height

(c*)

Stem Diameter

(-,,)

Leaf number (/plant)

Fresh weight (s/plan0

Dry weight G/ptano

Unlreated 23.8a* 2.3a 90a 2.0a 0.68a

l2^

31 2b 2.1n 9.7a 3.1h

l.l2c

67b

:8.7b 2.4a 10.0a 2.2a 0.14a 67b

l.,la 9.0a 2.7b 0.88b 60b

9.3a 2.5b 0.91b 59b

EcroM@RqHlzaLru Grr ciE stDEARTYcnowTtoF 5n&.46,/.,rm, ixonh.l su rk ^{u NeR} r UesteY

& ,ar

saiit

toi6r

M.nadTuiamai, Hid.yurisairq Erdy sanro.o, Aguns su5a.ro, $mp,n3 6anan, swido Hener timin, M8at6 shibuya,

I

B.

Fieldconditions

In

the

field

conditions,

ECM

colonization

did not

increased

plant height

and

leaf

number

two, four

and

six

months

afler

transplanting (Table 2.). There was Do significant difference between four ECM species and control seedlings.

Tabte

2.

Shoot height, leaf number and

suliva]

rates

of

^S- balangeran rnocrlated

with four

ectomycorrhizal

fungi two, four

aIrd

six

months after

transplanting

^under

field

conditions in Kalampangan ^zone, Centlal Kalimantan.

Trealment

I

nmnlhs Ileight

rcm)

Leaf (/plant)

Survival (:%)

Height

(cn, (/pLant)

Sunival

IL)

Hcight Leaf (/planr)

Survival (%)

4l.01a* ^{9.2a 100 44-95a 49.46a '7.6a lr8}

Calwtia sp. 13.19a 9.9a t00 ^46.03a 6.5a 9lt 53.32a It.0a

43.28a 8.6a 100 45.45 9l 55.00a 8.3a IJ6

4t.02a 7.la 100 47.18a 6.6a 95 54.18a 9.0a 87

-19.66a l0(l ^{.12.64a 5.2a} 48.22a

9i

.\'alucs qiih tu smc lencrm nor sisnificunlly difieEnl (F0.05)

ECM colonization of Bolenrs sp. and.

Sclercder,a

sp. increased plant height and

leaf

number 8, 14, 16, 19,22, and25 months

aft

transplanting (Fig. I ). There was no difference

in height

and

leaf

number

betweet Calvatia

sp-, Strobilotiryces sp. inoculated and control seedlings. ECM colon

iz,tion

of Boletus ^sp., Sclerodernta ^{sp. and} StroDl/oayces sp. increased plant height and stem diameter 40 months after transplantiog in the

field

(Table 3.). ECM

colodzation of

Calvatia sp. increased

only pla[t

height. There was

not differcnt in

stem diamgterbetweel Calvad4 ^sp. inoculated alld contl ol

Seedlings.

Survival

^rates

of S.

balangeran seedlings

were i[creased with

ECM

colodzation of

C4lva

tia

sp. ahd Strobilomyces sp. 40 months after transplanting in the

field

(Table3.).

50

-45

3+o

E30

t25 )20

15 100

b90

.9)

70 65

,,

.ab

, .-:..4 .4

_l _.j.1:.D.

t,r;."

-':'';J

510 15 20 25

³⁰

Time (months)

5 '10 15 20 25

³⁰

Time (months)

Fig l.

^Shoot height and leafnumber

ofS. ,alangetun (Kotth.)Butck

inoculated

with four

ECM fungi 8, 14, 16, 19, 22 and 25 months after transplanting under field conditions in Kalampangan zone,

Cental Kalimantal

(O

untreated;l

^Calva tia

sp.;a Boktus sp;D

Scletodemu

sp.;a

Strobilonlyces sp.).

5

I

]URNAT PEN€TITIAN DIPTEROKARPA VD 1No.1,Sepkmber2007

Table3,

Shoot height, stem diameter and survival mtes

of

S.

balafigerafi (Korth.) Bulck

inoculated

with

fo.ui ectomycorrhizal

fungi

40 months after transplanting

ulder

fieldconditions in Kalampangan zone, Central Kalimantan

Treatment

Paranleter Hcight

(c-)

Stem diameter (cm)

Survival mles (%\

Untreated ^{206.63a+ 2.74a 83}

Calrdti.l

^{sp. 222.57c 2.83a 89}

Boletus sD. 240.45c 3.03b

8i

Scle,'odentm sp. 233.53c 3.67c ^'79

Snobilottl\,ces sp. 2t3.55b 3.28c 85

*vllues _with lhe satre letcr are nor sisni fi cantly di ileen I (n<01)5)

IV. DISCUSSION

Even though the ECM ol Dipterocarpaceae in tropical rain lorests of Southeast Asia ^has bccn reported (Alexander et Ir/.,1992; Srnits, 1994; Lee and

Alexandet

1996), the role

ofECM

in dipterocarps species originated from peat-swamp forestneedsto clariry. Our results demonstrate

for

the

first time

that

ECM fungi

have

positive

effects on S. balangeran

(Korth.) Burck

six months under nursery conditions and 40 months after trarsplanting in the

field.

Colonization by four ECM species were identified as the most appropriate fungi for improving carly growth

ofS

balangeran

(Korih.)

Burck six months under nursery conditions. Furthermore, ,Rolerrs sp. and Scleroderma sp. proved to be competitive and effective to iDcrease plant grcwth of S. balangeratl (Korth.) Burck 40 months after transplanting in ^the

field.

Thercforc, Boletus sp. aIrd Sclerodenna and field conditions. Ecological

catego

^es of ECM flrngi may be more useful

for

summarizing fungalperformance in ^the field than in the growing conditions ofnurseries ^(B rundrett e/ a /., 200 5;

Omon(2002).

Plants

with Cdlvatia

^sp., Boletus sp., Sclerodernra sp.

d\d

Strobilomyces sp. werc not dift'erent

in

height, and leaf number than untleated seedli[gs during the

2 -

6 months

in field conditions. At

the time, November 2002-May 2003 was rainy season.

It

was possible due to waterlogged conditions

thatall

roots

ofs.

64langeran

(Korth.)Btrck

were in verysatumtedwith water. Water table in expeiment site was ^{close to} or above the peat surface throughout 6 months fluctuates

with

the intensity and frequency

ofrainfall.

Water has become a factor creating other problems i.e. waterlogged stress. Rhizosphere is the part

ofplant

in stress conditions when the oxygen supply

for root

consumption

is

being

red[ced mainly by

microorganism respiration.

Anaerobic respiration lnay leads to the synthesis arld translocation

of

some toxic components,

and affected various

processing

activities within plants (Naiola and Osaki, 2000).

The

transplanting shock and waterlogged conditions by the ECM plants six months

affer transplanting could be critical to the succoss

ofthe plantatior. Altcr

eight months transplanting in tlre

field

(July 2003), Boletus sp. and Scleroclernta sp. incleased the growth of S. balangeran (Korrh.) Burck in drought periods. BothECM species cons istency increased plant growth 14,16, I9.

ll.

^{25, and} 40 months after transplantin _E ln lhe field.

Boleh$

sp. ard Scletoclenna sp. were

:r.'re

adaptable compared

with Sttobilonyces sp., Cahatfu sp., and control

seedlings in :.

::::logged

and drought periods. Sr?b|lorryces sp. and

Calvdtia sp.lately

increased

of

plant

:::_::-: .ri

5. balangeran

(Korth.)

Burck 40 months after transplanting under

field

conditions.

J.: ::r,::riments

are affected by strong variation irr climatic, such as waterlogged conditions in

6

.croMYcoRRHrzat FUNGI lNcitrstD ant cnowrH or 5h0a b,k a.u, ^{txo^h.) Bod UllDER NUnsEiy}

ANDflE$(orOmONstN D[6MO

MamanTu4.m.i,Hideyukieto,€rdysaftoso,Acuncsunnro,sanpangGam:n,suwidoH.n.rrihin,M*:toshibuv., (unhderak.hshi,yurakrsmai,Miisuru0l.ki:ndKena'oraw ry.

I

the

rainy

season

or

drcught periods

in

the

summet

edaphic a]I.d

fiicrobiological

factors can influence

the

spreading

of ECM

inoculated

to

waterlogged

or

drought and cause

a

Iack

of

inoculation effects (Peft et al ., 1999). Accotding to our knowledge, there ate no .efercrlces in the literature indicatitrg

ECM

effect on s, balangeran

(Konh.) Bulck in

Southeast Asia

region. In

Madagascar, the use PdolithLts

microc.trpls

increased plant height of A. crassicarpa 19 months after transplanting

in the field

experiment (Ducousso er

al,, 2004). In a dry tropical

area, Pisolithus albus and

klerodefiM dyctiosporul,

increased plant

growh

of Acacia holosericea after 24 months karNplanting in the field in Senegal (Duponnois e, dl.,

2005).

In temperate a!ea,

Laccaria bicolor mycelia or spore sNpensions of Melanogaster anbigults,

Rhizopogotl colossus, and R. sltbareolahts increased plant gro\ath

of

Pseudots

ga fienziesii

5 years after

tansplanting

in northem Spai\.

Laccaria

brcolor i[creased plaflt

groMh ofP nefiziesii eight

yeals after transplanting

ill

France (Selosse ef

al.,1999).ln

Spain, Rhizopogon spp. increased platrt growth

ofPirrrJpirea

20 motrths after transplanting in formerlyarable land (Parlad6 era1., 2004).

Suryival rates

of

seedlings are important

for

the

first

establishment plant growth

ofs

balangeran (Korth.) Burck after hansplanting in the field. The role

ofECM

in the

suvival

^rates

of ^S. balangeran (Korth.) Burck requires more careful analyses because survival rates were not different between ECM seedlings and control

seedling.

Surwivalrates

ofECM

plants in average were decreased

four

months

(94.5%) ard six

months

(90%)

after transplanting

in

the

field conditions. Conhol

seedlings were also decrcased

foul

months (96%), and six months (88%) after transplanting. It was possible also that wateilogged conditions can atrect the survival rates

ofECM

plants.

Suvival

rates

ofECM

plants were consistency steady in _{8, 14,}

16,19,22 atd25

months (88%) after transplanting andcontrol seedlingswere in 867o.

Moreovet

suwival rates

of

ECM and cofltrol plaDts were not different ^(83olo) in 40 months after hansplantirg in the

field. It

was possible thatthere was cross contamination lvith inoculated plants because line spacing was

only I

m

x I

m.

Ifcontrol

plants were colonized by ECM fungi

ofhoculated

plants they could increase

plant groMh of conhol

plants.

Howevet we didn't

measure

ECM

colonization 40

motths

after transplanting under

field conditiols.

L1 a

dry tropical

area,

survival

rates

of L

holosericea inoc[latedby Pisolithus albus or Sclercdemu dictyosporznr were higher (98- 100%) than control plants (8loZ)

l2

months affer transplanting in the field in Senegal. In the temperate forest, inoculation

vrtlh

Rhizopogon

roseolls

increased

in

20yo

the stwival of

Pinus pinea seedliogs over cotrtrol seedlings after 43 months outplanted

in

Spain (Parlad6 et a1.,2004).

Effofts to inoculate tree seedlings

with ECM

firngi have gcnerally focused on one

f[ngus

or ^a limited number

offungi,

which may not be

approp

ate ifbenefits from increased fungal diversity are expected,

or

improved functional diversity is required (Brundrett e, a/., 2005).

It

may be possible that in disturbed habitats, there is a succession

ofECM

fungi where a few pioneering

fungi are gradually

replaced

by an

increasing

diversity of ^ECM fllngi

characteristic

of

undisturbed habitats (Gardnerand Malajczuk,

1988;Luetai.,

1999). Because

ofthe difficulties

of

monito

ng the

ECM

fungal species in the

field

samples withoul using rrolecular techniques

(DiBallistaet

a1.,2002), no data or the determination ofeach ECM inoculated inthe root systems ofS. balangeranplants are available. Field growth perfomtance of S. balangeran

$orth.)

^Burck

is still observed for several years.

In this study, spore density

ofeach

ECM species was l1ot measur.ed aod

it

is 1lot

know[

how many spores germinated under nursery conditions and resulted in

ECM

formation

with

S.

balangeran

(Korth.) Burck.

Spore density can influence

ECM fonnation (L\t et al.,

1998).

Furthermore, Chen e, a/. (2006) recommended that spore densities ol

Sclercderna

species lln be used

for

inoculation

ofcontainerized

eucalypts and that spores be stored at 4" C

rntil

use. The

mair

disadvantages

ol

spore

inoctium

are genetic

variability, the lack reliable

laboratory methods to determine spore

viability

al1d the delay in mycoffhization compared

with

vegetative

7

I

JURNAL PENELITIAN DIPTEROKARPA

vol.l No,!septenber2007

mycelium (Brundrctt e t

al.,

1996). The use

olrolelr$

sp. spores in nrrseD, scales is not

widcly

practiced however because amount olBol?rr.r sp. sporcs arc limitcd conrparcd wiih Sclerodernn sp. sporcs.

Bolearr

sp.

may

also

bc

^possiblc

to dcvclop \\"ith legetative

mycelium- These vegetative inocula

of

selected fungal sirains have been recommcndcd as thc prel'ened melhod (BrLrndrett er a1., 1996). However,large quantities

olviable

inoculum ale needed for application on an operatjonal scale and the storage

ol

vegetatiye inoculum usuallv

ad!eNely

influences its effectiveness- Spore based inoculum rvas also easier to use and much lcss exfcnsivc to ploduce than mycelial

slulries.,S./ooderTrd

sp. are also good candidates

for inoculaling

^prutrdms in dcgraded pcat- srvamp ibrest sincoihcse spccies apparcntly adapt to many ecological habitats and they arc easy to usc as sporc inoculllm

(Jcffrics.

I999; Chcn ^{cr a1..}

2006).

Furthcr rcscarch on factors that regulate spore

gern

nation is required to improve both the speed and consistency

of ECM fbrmaliorl. It may

also be possible

to iicrease

the

reliabilily of

iDoculiltion

by

^r.lsiig mixtures of nrngal ^taxa (Brundrett.1al.. 2005).

V. CONCLUSION

In conclusion, coionization of

lbur

nafi\,c ^ECI\,1 spccics can incrcaso plant

gro\rih of,l- halangeran (Korth.) Burck 6 months alter

tr-ansplanting

under

greenhouse conditions- Colonization

Boletus

sp. a]nd

S(leto.lerna

sp. can

corsisiellcy iicrease p]ant

grorvth

of

S.

balangeran

(Korth.) Burck

40

rnontls

aI'ter transplanting

in

the

field.

Sr?1,i/.)rryccr sp. and Or1rd1ld sp. was dclay to incrcase only

ofplant

hcight

ol5'.

balungeran

(Korlh.)

Burck

afier40

months under field conditions. Surl ival ratcs

ofS 6olalgerr:,

^(Korth.) Bu.ck u.erc not

dilfcrcnt

between ECM

aid

untreated

plults.

These resulls \\ ere allecled by climatic inlluenoes, such rls

watcrloggcd

or

drought pcriods can rcduce the

first

cstablishment

of

ECI\{ plants

in

lhe

lield

conditions. Bolerr.i sp. (late stage) and Sc/erc,Ierrrd sp. (carly stagc) can bc uscd to inoculatc

S

baldngeran lKot1l1.)

lfurck

in nurseries becausc thcsc

fllngi

^are \\.cll adaptcd to thc cn\ irontrcrrt conditions encountered in this area rvhich could explain this better eflect on plant

growth. It

is suggesled

that

ino,:rulalion

of

irdigenoLrs ECN'I

fungi to native tree

species

undcr

nurscry

co ⁿ ditions is us cful fbr rchab ilitation dcgradcd peat-s$.ainp fbre sts.

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