425
Rela ion be ween Supp essi eness o Toma o Fusa ium Wil and Mic obial
Popula ions in Di e en G ow h Media
C. Bo e o, M.J. In an es, E. González
and M. A ilés
Dep . Ciencias Ag o o es ales, E.U.I.T.A.
Uni e sidad de Se illa. C a U e a, km
1, s/n 41013 Se illa
Spain
J.C. Tello
Dep . P oducción Vege al, Uni e sidad
de Alme ía. C a. Sac amen o s/n, 04120
La Cañada de San U bano, Alme ía
Spain
Keywo ds: Compos s, biocon ol, coi ib e, pea , Fusa ium oxyspo um
Abs ac
Toma o Fusa ium wil supp essi eness o h ee g ow h media, wi h espec o
pea and e miculi e, and he ela ionship be ween mic obial popula ions o g ow h
media in he se e i y o his pa hogen we e examined. The g ow h media e alua ed we e
oli e oil husk + co on gin ash compos ed and mixed wi h ice husk (OC+R), spen
mush oom compos ed and mixed wi h pea (SM+P) and coi ib e. To de e mine he
ole o he mic o lo a in he supp ession, he wo compos s we e also hea ed o 60ºC o
6 days. Th ee bioassays we e ca ied ou wi h hese in es ed g ow h media and con ols
no in es ed. Disease se e i y was eco ded du ing 25 days a e oma o seedling
ansplan a ion. In hese se en g ow h media he densi y o cul u able g oups o
mic oo ganisms was de e mined by dilu ion pla ing on semi-selec i e media be o e he
bioassays. The wo compos s showed supp essi eness o Fusa ium wil wi h espec o
e miculi e and pea while coi ib e was conduc i e. Fu he mo e, in OC+R i s
supp essi eness is due mo e o he g ow h media’s non hea -labile p ope ies han in
SM+P. Signi ican nega i e co ela ions we e ound be ween se e i y and Bacillus spp.,
celluloly ic ac inomyce es and bac e ia, oligo ophic ac inomyce es and bac e ia,
copio ophic bac e ia, o al ac inomice es (celluloly ic + oligo ophic + copio ophic)
and o al bac e ia (celluloly ic + oligo ophic + copio ophic) popula ions in hese
g ow h media. Bacillus spp. a e known an agonis s, so hey may accoun o he
supp essi eness in ou compos s. Celluloly ic and oligo ophic ac inomyce e popula ions
we e associa ed wi h Fusa ium wil supp essi eness o ano he g ow h media. High
mic obial ac i i y has been epo ed as a Fusa ium wil supp essi eness ac o .
Copio ophic, celulloli ic and oligo ophic bac e ia con ibu e o his mic obial ac i i y.
INTRODUCTION
Toma o p oduc ion wi h in ensi e sys ems in Spain is inc easingly using po ing
cul u e wi h o ganic subs a es. One o he oma o c op p oblems is Fusa ium wil which
cause impo an yield losses, and o hese p oblems he e a e no e ec i e chemical con ol
measu es. Howe e , biological con ol is becoming an e icien al e na i e o con olling
hese diseases. Some g ow h media o mula ed wi h compos s a e able o supp ess Fusa ium
wil wi h espec o pea (Che e al., 1983; O likowski, 1983; T illas-Gay e al., 1986;
Ga ibaldi, 1988; Pe a and Cal e , 1989; Hoi ink e al., 1991; Se a-Wi ling e al., 1996;
Szczech, 1999; Co xa e a e al., 2002; Reu eni e al., 2002; T illas e al., 2002). The na u e
o soils and compos s wi h supp essi eness o Fusa ium wil is he esul o complex
in e ac ions be ween he abio ic cha ac e is ics o he media and mic obial popula ions
(Hoi ink e al., 1993; Alabou e e e al., 1996; Welle e al., 2002; Bo e o e al., 2004b).
Se e al biocon ol agen s ha e been iden i ied in hese soils and compos s (Hoi ink e al.,
1993; Welle e al., 2002). The mic obiological na u e o Fusa ium wil supp essi eness o
many compos -amended g ow h media has been demons a ed p e iously (Che e al., 1983;
O likowski, 1983; T illas-Gay e al., 1986; Ga ibaldi, 1988; Hoi ink e al., 1991; Se a-
Wi ling e al., 1996; Szczech, 1999; Co xa e a e al., 2002; Reu eni e al., 2002; T illas e
al., 2002). The supp essi eness o soilbo ne plan pa hogenic ungi mechanisms in ol e
gene al supp ession, which is ela ed o high mic obial ac i i y, and speci ic supp ession,
which is ela ed o an inc ease in he popula ion o speci ic mic oo ganisms o g oups o
mic oo ganisms ha ac as an agonis s o he pa hogen (Cook and Bake , 1983). On he o he
P oc. IS on Soilless Cul . and Hyd oponics
Ed: M. U es a azu Ga ilán
Ac a Ho . 697 ISHS 2005
426
hand, gene al disease supp ession can be dependen on communi ies o mic oo ganisms.
These communi ies may be associa ed wi h a subs a e a a pa icula s age o decomposi ion
unde ce ain en i onmen al and managemen condi ions (Boehm e al., 1993, 1997). The
composi ion o unc ional g oups ( a he han indi idual species) may de e mine he cha ac e
o he communi y while indi idual species wi hin a unc ional g oup may be in e changeable
( an B uggen and Semeno , 2000). The aims o his esea ch we e o e alua e h ee g ow h
media Fusa ium wil supp essi eness wi h espec o pea and e miculi e and o s udy he
ela ionship be ween mic obial popula ions o g ow h media on he se e i y o Fusa ium wil
in o de o p edic disease supp essi eness o g ow h media.
MATERIAL AND METHODS
Plan G ow h Media
Two compos ed esidues om ag icul u al and indus ial was es and e ilized coi
ib e (Cocopea , P oja , Valencia, Spain) we e e alua ed o Fusa ium wil supp ession. The
g ow h media made wi h compos s we e oli e oil husk + co on gin ash, 1:1 / , compos ed
and mixed wi h ice husk (1:1 / ) (OC+R), and spen mush oom compos ed (Recomsa,
Quin ana del Rey, Spain) mixed wi h ligh pea (1:1 / ) (SM+P). The OC+R was
compos ed as desc ibed elsewhe e (T illas e al., 2002). Fe ilized ligh pea (Klasmann,
Valinex, Palle e , Spain) and e miculi e (Ve miculi a y de i adas, Gijón, Spain) was used as
a e e ence. Pea was neu alized wi h 4 g L-1 CaCO3. Fe ilized pea and coi ib e had 0.33 g
L-1 o K2O (50% high solubili y g anula ed; Compo Ag icul u a S.L., Ba celona, Spain) and
4.15 g L-1 o P2O5 (18% g anula ed; Fe ibe ia, Mad id, Spain). Plan g ow h media made
wi h compos s we e no e ilized. To s anda dize ini ial condi ions, he g ow h media we e
incuba ed a a wa e ension o 1000 Pa (adjus ed o weigh ) o 14 days a 25ºC. To
de e mine he ole o mic o lo a in supp ession, he wo compos s we e also hea ed o 60ºC
o 6 days (OC+R60 and SM+P60).
Assessmen o Disease Se e i y
Disease supp essi e p ope ies o g ow h media we e measu ed by a Fusa ium wil
bioassay desc ibed elsewhe e (Co xa e a e al., 2002). The bioassays we e de eloped wi h
oma o `Roma´ and a monospo ic isola e o Fusa ium oxyspo um . sp. lycope sici ace 2
(FOL-2). This isola e was ob ained om oma o plan s and s o ed in silica gel. Liquid cul u e,
500 ml 1% mal ex ac (Sigma Chemical Company, S Louis, MO), o FOL-2 was p epa ed
and g own wi h con inuous agi a ion (130 pm) o 10 days a 25ºC. Conidia we e eco e ed
a e il a ion and cen i uga ion a 5000 pm, 15 min, (Eppendo 5810 R, Hambu g,
Ge many) and insed wice wi h s e ile dis illed wa e . The concen a ion o conidia was
de e mined wi h a hemocy ome e . The se en g ow h media we e in es ed wi h FOL-2 (105
mic oconidia ml-1 g ow h media), mixed igo ously and pou ed in o 9 cm diame e plas ic
po s (330 ml olume). Non in es ed g ow h media we e used as con ols. Fou oma o
seedlings (1 o 2 ue lea s age) g own in e miculi e we e ansplan ed in o each po . Plan s
we e i iga ed as needed and e ilized wi h Pe e ’s olia eed 27-15-12 (Sco s Hee len, The
Ne he lands), 0.5 g L-1 en iched wi h 0.6 g L-1 o CaCl2, 0.696 g L-1 MgSO4.7H2O and 0.3 g
L-1 o u ea (46%). They we e g own in a g ow h chambe (27ºC, 280 µE/m2/s PAR in ensi y,
16:8 h ligh :da k pho ope iod and 60% RH). Disease se e i y was moni o ed o 25 days a e
plan ing and was sco ed e e y wo days based on a symp om se e i y scale whe e: 0 =
asymp oma ic plan s; 1 = weakly in ec ed plan s (≤ 50% o lea es chlo o ic o wil ed); 2 =
highly in ec ed plan s (> 50% o lea es wil ed bu plan s no dead) and 3 = dead plan s
(Co xa e a e al., 2002). The a ea-unde - he-disease-p og ess-cu e (AUDPC) was
calcula ed, and AUDPCs was also calcula ed by di iding AUDPC by he o al ime du a ion
(days) o he epidemic o each bioassay o compa ison be ween epidemics in di e en
bioassays (Campbell and Madden, 1990). These bioassays we e epea ed a leas h ee imes
wi h i e po s pe ea men . A each a ing ime, he mean o he disease se e i y and he
pe cen age o symp oma ic plan s pe po we e calcula ed. This mean was conside ed as one
alue. T ea men s we e a anged in a andomized block design.
Mic obial Popula ions
427
The densi y o cul i able g oups o bac e ia and ungi associa ed wi h biocon ol
phenomena was de e mined by dilu ion pla ing on semi-selec i e media acco ding o Tui e
e al. (1998) wi h modi ica ions. Samples we e aken om incuba ed plan g ow h media
be o e he bioassays. Plan g ow h media (5 o10 g) we e suspended in 250 ml o 0.1%
sodium py ophospha e. The suspension was shaken and en old dilu ion se ies we e p epa ed
wi h 0.1% wa e aga . Suspensions we e pipe ed on o h ee pla es pe cul u e medium and
dilu ion. Fou o i e dilu ions pe se ies we e placed on pla es. Fo isola ion o Bacillus spp.,
luo escen Pseudomonas spp., copio ophic, oligo ophic and celluloly ic bac e ia and
ac inomyce es, 100 µg ml-1 o cycloheximide was subs i u ed o 10 µg ml-1 o benomyl
(Ene gía e Indus ias A agonesas, S.A., Mad id) and 0.3 µl ml-1 o P e icu (P opamoca b,
72.2%, Sche ing, Alcáce , Spain). Fungal coun s we e made on po a o dex ose aga amended
wi h 1,000 ppm o Te gi ol-7 (Fluka Chemie AGB, Buchs, Swi ze land) and 50 µg ml-1 o
oxy e acycline hyd ochlo ide (Sigma Chemical Company, S Louis, MO) (Chen e al.,
1988). Analyses we e pe o med h ee imes wi h one sample pe each con ol plan g ow h
medium om each o he h ee bioassays.
S a is ical Analysis
Da a collec ed om all ials we e analyzed wi h S a g aphics Plus, Ve sion 6 (SGS,
1999). The e ec o he g ow h medium on he AUDPCs was analyzed wi h ANOVA.
Signi ican means we e compa ed by Tukey’s whole signi ican di e ence es (P = 0.05).
O e all ela ionships be ween AUDPCs and he mic oo ganism popula ions o he plan
g ow h media we e analyzed wi h eg ession analysis.
RESULTS AND DISCUSSION
The wo compos s wi h and wi hou hea ea men showed a lesse se e i y han
e miculi e, pea and coi ib e (Fig. 1). These esul s indica ed ha he wo compos s had
supp essi e cha ac e is ics o Fusa ium wil wi h espec o he e e ence g ow h media, while
he coi ib e was conduc i e. Fu he mo e, OC+R supp essi eness is due mo e o he g ow h
media’s non hea -labile p ope ies han in SM+P (Fig. 1). In his sense, p elimina y da a we e
exposed by Bo e o e al. (2002 and 2004a). These g ow h media non hea -labile p ope ies
can be physical and chemical ac o s o he mic obial popula ions eco e ed om a hea
ea men a e a ew days. These non hea -labile p ope ies can acili a e a as mic obial
popula ion ecupe a ion, which could aise mic obial ac i i y, associa ed wi h gene al
supp ession (Welle e al., 2002). On he o he hand, an agonis s popula ions can be
des oyed by he hea ea men . This indica es ha OC+R supp essi eness can be due o
gene al supp ession and SM+P supp ession can be due o gene al and speci ic supp ession.
This gene al supp ession can be accompanied by he mophilic an agonis speci ic
supp ession, due o hei esis ance o he hea ea men .
Signi ican nega i e co ela ions we e ound be ween se e i y and Bacillus spp.,
celluloly ic ac inomyce es and bac e ia, oligo ophic ac inomyce es and bac e ia, copio ophic
bac e ia, o al ac inomice es (celluloly ic + oligo ophic + copio ophic) and o al bac e ia
(celluloly ic + oligo ophic + copio ophic) popula ions in hese g ow h media (Table 1). On
he o he hand no signi ica i e co ela ions we e encoun e ed be ween se e i y and
Pseudomonas and ungi densi ies (da a no shown).
The high popula ion le els o Bacillus spp. associa ed wi h he wo compos s, ei he
na u al o hea ed, a e due o he high compos ing empe a u es ha selec he mophilic
mic oo ganisms. They a e known an agonis s o Fusa ium wil (Khan and Khan, 2001), so
hey may accoun o he supp essi eness in ou compos s.
One o he mos impo an g oups con aining an agonis ic mic oo ganisms is he
g oup o luo escen pseudomonads (Ga be a e al., 2004). Al hough, i seems ha hese
bac e ia a e no a main ac o o supp essi eness in ou compos s o , a leas , luo escen
pseudomonads densi y is no a good p edic i e pa ame e . Roo exuda es supply nu ien s ha
p omo e high g ow h a es o pseudomonads (Ga be a e al., 2004), ha can be he cause o
he lack o co ela ion be ween se e i y and pseudomonads popula ions be o e plan ing.
Wi h o he compos s hizosphe e ungi densi y has been nega i ely co ela ed wi h
Fusa ium wil se e i y (Bo e o e al., 2004b). This indica es ha ungi a e no e y impo an in
hese compos s supp essi eness, a leas a he beginning o he c op.
428
Celluloly ic and oligo ophic ac inomyce es izosphe e popula ions we e associa ed o
Fusa ium wil supp essi eness o o he g ow h media (Bo e o e al., 2004b). To al numbe o
ac inomyce es also was posi i ely co ela ed wi h supp ession o co ky oo o oma o (Wo kneh
and an B uggen, 1994). The e o e, ac inomyce e densi ies can be a good p edic i e plan
g ow h media supp essi eness pa ame e . High mic obial ac i i y has been epo ed as a
Fusa ium wil supp essi eness ac o (Bo e o e al., 2004b). The e o e, high ba e ia densi ies
(copio ophic, celulloli ic and oligo ophic) con ibu e o Fusa ium wil supp essi eness. On he
o he hand, oligo ophic o ganisms and a low a io oligo ophic bac e ia / copio ophic bac e ia
ha e been associa ed wi h conduc i eness (Hoi ink e al., 1996; Bo e o e al., 2004b). The plan
g ow h media made wi h compos a e mixed wi h ma e ials wi h low biodeg adable componen s
( ice husk and pea ). These ma e ials suppo ed oligo ophic popula ions. The e o e, he nega i e
co ela ion be ween he se e i y and oligo ophic bac e ia esembles hei mic obial ac i i y.
This indica es he gene al supp ession impo ance in hese plan g ow h media. The e o e,
oligo ophic bac e ia densi ies a e no a eliable supp essi eness pa ame e .
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Tables
Table 1. Co ela ion (simple) among s anda dized a ea unde disease p og ess cu e
(AUDPCs) o Fusa ium wil and di e en mic oo ganisms eco e ed om se en
plan g ow h media (CFU ml-1).
Mic oo ganism g oup1 R2
(%)
P 2 Equa ion
Celulloli ic bac . 87.87 ** AUDPCs = 0.5702-0.03144 · Ln(celulloli ic bac .)
Σ bac . 83.55 ** AUDPCs = 0.665 - 0.03358 · Ln(Σ bac .)
Oligo ophic bac . 82.41 ** AUDPCs = 0.6388 – 0.03392 · Ln(oligo ophic bac .)
Copio ophic bac . 80.50 ** AUDPCs = 0.927 – 0.03302 · Ln(copio ophic bac .)
Celulloli ic ac . 76.33 * AUDPCs = 2.204·celulloli ic ac .-0.3227
Σ ac . 70.69 * AUDPCs = 3.481 · Σac .-0.3318
Oligo ophic ac . 70.23 * AUDPCs = 1.4446 · oligo ophic ac .-0.2994
Bacillus spp. 65.32 * AUDPCs = 0.4074 – 0.02396 · Ln(Bacillus spp.)

430
1 Bac = bac e ia; ac = ac inomyce es; Σ bac = celulloli ic, oligo ophic and copio ophic bac e ia; Σ ac
= celulloli ic, oligo ophic and copio ophic ac inomyce es.
2 Signi icance le els: *, ** and *** indica e P < 0.05, 0.01 and 0.001, espec i ely
Figu es
Plan g ow h medium
OC+R OC+R60 SM+P SM+P60 Coi ib e Pea Ve miculi e
AUDPCs
0.00
0.05
0.10
0.15
0.20
0.25
0.30
a
aa
b
c
bc
c
Fig. 1. S anda dized a ea unde disease p og ess cu e (AUDPCs) o oma o plan s in se en
g ow h media. OC+R: oli e oil husk + co on gin ash compos ed and mixed wi h
ice husk; SM+P: spen mush oom compos ed and mixed wi h pea ; 60: hea ed.
Disease se e i y scale was om 0: asymp oma ic plan s, o 3: dead plan s. Plan
g ow h media we e in es ed wi h Fusa ium oxyspo um . sp. lycope sici (105
mic oconidia ml-1). Da a o AUDPCs we e ans o med o analysis wi h he a csine
√x. Ba s wi h he same le e a e no signi ican ly di e en acco ding o Tukey's es
a P < 0.05. S anda d e o o he mean is indica ed by e ical line.