molecular profile of microbiota (7)
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Applied Soil Ecology 92 (2015) 47–53
Contents lists available at Sciencei!ect
Applied Soil Ecology
"o#!nal $o%epage& '''(elsevie!(co%)locate)apsoil
*olec#la! p!o+le o, %ic!obiota o, -innis$ co%%e!cial co%posts#pp!essive against .yt$i#% disease on c#c#%be! plants
/# a A Sinonen ad # i a * #!ola a S #onen b . .a!ia c * 6estbe!g b* o%antsc$# ad8a
9nive!sity o, elsini epa!t%ent o, Envi!on%ental Sciences ie%enat# 73 -:;15140
>$e #se o, co%post o, va!ied o!igins ,o! t$e %anage%ent o, plant disease $as been !epo!ted aove! t$e'o!ld ,o! decades and %ic!oo!ganis%s a!e s#ggested to be one o, t$e d!iving ,o!ces :n$is st#dyco%posts p!od#ced d#!ing t' o s#ccessive yea!s 'e!e %ied 'it$ peat at 1& 4 vv !atios and seened ,o!c#c#%be! .yt$i#% disease s#pp!ession in t' o epe!i%ents in 2009 ine co%posts 'e!e seleed basedon t$ei! s#pp!essive eFects against .yt$i#% on c#c#%be! plants and t$e!ea,te! 'e!e b"ected to%olec#la! analyses >$e ai % o, t$is st#dy 'a s to co%pa!e t$e bacte!ial and ,#ngal co%%#nits p!esentin st!ongly;s#pp!essive and non; o! 'ealy;s#pp!essive co%posts in an eFo!t to identi,y
bacte!ial and,#ngal g!o#ps associated 'it$ .yt$i#% disease s#pp!ession >$e total geno%ic A ,!o% t$e o%posts
'a s et!acted and a%pli+ed #sing p!i%e!s ta!geting 1@S !A and :>S !A genes o, bacte!ia
nd ,#ngi!espectively p!io! to 45 4 seD#encing SeD#encing !es#lts indicated t$e potential !oles , bacte!ialAcidobacte!ia =p14 and ,#ngal Cystobasidio%ycetes in t$e s#pp!ession o, .yt$i#% 'ilt diseae d#e tot$ei! p!esence and absence in co%posts 'it$ st!ong and lo'e!absent disease s#pp!eson ability!espectively >$e ab#ndance o, Actinobacte!ia 'a s associated positively 'it$ s#pessiveness.$osp$o!#s concent!ation 'a s t$e only abiotic ,acto! t$at 'a s associated 'it$ s#pp!esveness& anegative co!!elation 'a s ,o#nd in bot$ epe!i%ents >$es e !es#lts indicate t$at ce!tain b
te!ial and,#ngal g!o#ps can potentially s#pp!ess .yt$i#% 'ilt '$ile t$e abiotic conditions %ig$t be less %po!tant
G 2015 Elsevie! ?6 All !ig$ts !ese!v
d
1997H oble and Covent!y 2005H .#gliese et al 2011H *e$t
et al2014) 6a!io#s %ec$anis%s $ave been s#ggested to epl
n t$eCo%posting
is
an
ae!obic
%ic!obial
d!iven
p!ocess
t$atdeg!ades o!ganic 'aste into a stable sanita!y $#%#s;lie %ate!ial
t$at can be sa,ely !et#!ned into t$e envi!on%ent
t is alsoconside!ed as an alte!native to pesticides in %eans o, cot!olling
http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://www.sciencedirect.com/science/journal/09291393http://www.elsevier.com/locate/apsoilhttp://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://dx.doi.org/10.1016/j.apsoil.2015.03.005http://www.sciencedirect.com/science/journal/09291393http://www.elsevier.com/locate/apsoilhttp://dx.doi.org/10.1016/j.apsoil.2015.03.005
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soil bo!ne plant diseases '$ile gene!ally boosting plant $ealt$ andp!od#ctivity (ada! and *andelba#% 1992H e$e! et al 2013)epo!ts on t$e #se o, co%posts to s#pp!ess soil;bo!ne plantdisease can be t!aced bac al%ost $al, a cent#!y (oitin et al1977) Since t$en t$is concept $as been etensively !evie'ed inn#%e!o#s p#blications (oitin and -a$y 19B@H oitin et al
8 Co!!esponding a#t$o! at& nive!sity o, elsini epa!t%ent o, Envi!on%
ental
Sciences ie%enat# 73 -:;15140 el& I35B 9 191 20334H,a& I35B 9 191 20331
E;%ail add!ess& %a!tin!o%antsc$#J$elsini + (* o%antsc$#)
ttp&ddoio!g10101@"apsoil2015030050929;1393G 2015 Elsevie! ?6 All !ig$ts !ese!ved
disease s#pp!ession by co%post and its %ic!obiota A%ong $esea!e co%petition
,o!
n#t!ients
and space a%ong
%c!obialpop#lations (iKne et al 2005H 6estbe!g et al 2014) in$itionon pat$ogen g!o't$ by antibiotics p!od#ced by %ic!obiota (*$taet al 2014) and t$e $ype!pa!asitis% '$e!e a %ic!ooanis%di!ectly attacs
a
pat$ogenic
%ic!oo!ganis% and ills
it(de ?e!toldi 2010) St#dies $ave s$o'n t$at $eating o! a#toavingeli%inates t$e disease s#pp!ession capacity o, t$e co%post'$ilet$is capacity can be !ecove!ed by %iing in nat#!al co%pos'it$t$e $eated o! ste!ilied co%posts (elson and oitin19B3Hoitin and -a$y 19B@) :n addition to t$e p!es#%ed actity o, antagonistic %ic!oo!ganis%s abiotic ,acto!s s#c$ as nit!ogentotaland available) C& !atio p $eat %oist#!e as 'ell as deg
ee o, co%post %at#!ity $ave been s#ggested to be associated 'itt$es#pp!ession capacity (#te! et al 19BBH ada! and =o!eci1991Hoitin and =!eb#s 1997)
4B /# et al Applied Soil Ecology 92 (2015) 47–53
.yt$i#% is one o, t$e %ost st#died soil bo!ne plant pat$ogens6a!io#s species o, .yt$i#% ca#se da%ping;oF and 'ilting in a ve!yla!ge n#%be! o, plant species ?ot$ nat#!ally occ#!!ing and
a%ended co%posts s#pp!essive against .yt$i#% plant diseases$ave been desc!ibed Co%posts %ade 'it$ peat and ba! $ave alo'
indigeno#s
s#pp!essiveness
against
pat$ogens
s#c$
as.yt$i#%
spp (?onano%i et al
2010)
and $ave
t$e!e,o!eco%%only been #sed as s#bst!ates to p!od#ce co%post %it#!esC$en et al (19BB) and agn et al (200B) !epo!ted t$e signi+canceo, co%post %ic!oo!ganis%s in a%ended co%posts s#pp!essive
against
.yt$i#%
d#%ping;oF
>$e
%ic!obial
co%%#nity
inco%post depends on bot$ co%post ,eedstoc (eg ing!edientsp
n#t!ients
C&
43) and
co%posting
p!ocess
(egte%pe!at#!e
oygen
a%%onia
%oist#!e
%ic!obial
activity)
>$e
%ic!obial co%%#nity and its %etabolic activity $as ani%po!tant !ole in aFecting t$e disease s#pp!ession ability o, co%post (*cella! and elson 2003H agn et al 200BH e$e!et al 2013)
o'eve!
no'ledge
abo#t
$o '
%ic!obes
cont!o
l soilbo!ne pat$ogens is still sca!ce
>$is is a contin#ation st#dy o, 6estbe!g et al (2011 2014) in'$ic$ t'enty;one -innis$ co%%e!cial %at#!e co%posts o, va!iedo!igins (eg se'age sl#dge bio'aste %an#!e) 'e!e %ied it $
ste!ilied peat and sc!eened ,o! t$ei! ability to s#pp!ess c#c#%be!'ilt ca#sed by .yt$i#% in t' o epe!i%ents in 50 0 %l pots 'it
+ve!eplicates >o inoc#late t$e c#c#%be! plants .yt$i#% %ycei#%and t$e g!o't$ %edia (.A – potato det!ose aga!) 'e!e g!o
ndinto p#lp -ive g!a%s o, .yt$i#% inoc#l#% 'a s added to eac$ pot atplanting dept$ >$e c#c#%be! plants 'e!e g!o'n in a g!een$#seat te%pe!at#!es o, 24 (day) and 1B C (nig$t) and a 1@;$o#day
(6estbe!g et al 2011) >$e!ea ,te! nine co%post %it#!et$ats$o'ed s#pp!essive ability o! 'e!e ne#t!alcond#cive to'a!dsc#c#%be! .yt$i#% disease 'e!e divided into t' o g!o#p,o!%ic!obial and statistical analyses in t$e p!esent st#dy >$e a% o, t$is
st#dy
'a s
to
investigate
t$e
connections
bet'eet$e%ic!obial co%%#nities and t$e disease s#pp!ession abily o, co%post against t$e soil;bo!ne pat$ogen >o t$e best oo#!
no'ledge t$is is t$e +!st st#dy t$at at t$e gen#s;level t!gets
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bot$ bacte!ia and ,#ngi s#pp!essive against .yt$i#% da%ping;oFSpeci+cally t$e ob"ectives 'e!e to (a ) c$a!acte!ie t$e %i
c!obialco%%#nity co%position (b) identi,y t$e speci+c bacte!ial and,#ngal g!o#ps (eg at gen#s;level) associated 'it$ t$e s#pp!es;siveness against c#c#%be! .yt$i#% 'ilt and (c ) ea%ine t$econnection bet'een !elative ab#ndance o, bacte!ia and ,#ngi
andt$e disease s#pp!ession ability o, co%posts
2 *ate!ial an d %et$ods
21 S#pp!essiveness o, selected co%post
ine co%posts o, diFe!ent so#!ces ( >able 1) p!od#ced in t$es#ccessive yea!s 200B and 2009 'e!e selected ,o! t$e p!esent%ic!obial dive!sity st#dy based on t$ei! !espective s#pp!essive;ness c$a!acte!istics
d#!ing
t' o la!ge
sc!eening epe!
i%entsagainst c#c#%be! .yt$i#% 'ilt '$ic$ 'e!e ca!!ied o#t in t' ope!iods& ,!o% -eb!#a!y to *a!c$ 2009 (1st epe!i%ent beginning;–end 6estbe!g et al 2011) and ,!o% A#g#st to Septe%be! 2009(2nd epe!i%ent beginning–end 6estbe!g et al 2014) .!io! tobot$ s#pp!essiveness epe!i%ents co%posts 'e!e %ied 'it$ a%it#!e o, 75L stea% ste!ilied nat#!al Sp$agn#% peat and 25Lsand at 1& 4 vv !atios – called Mco%post %it#!esN in t$e p
!esentst#dy >$e stea% ste!ilied and nat#!al non;ste!ilied peat;sand%it#!es 'it$o#t co%posts 'e!e applied as cont!ols in bot$epe!i%ents
S#pp!essiveness capacity 'as calc#lated as t$e L diFe!ence ins$oot d!y 'eig$t (O) bet'een non;inoc#lated and .yt$i#%inoc#lated c#c#%be! plants O 'as %eas#!ed as desc!ibed by
>able 1Co%post %it#!es 'it$ st!ong disease s#pp!ession ability (=!o#p A) and co%st%it#!es 'it$ lo ' o! absent disease s#pp!ession ability (=!o#p ?) t'o
epe!i%ents >'o M=!o#p AN co%post %it#!es 'e!e o%itted ,!o% eac$ epe!i%
ntd#e to t$ei! lo' seD#ence !eads a' %ate!ials o, co%posts& 1 – po#lt!y %an# 3 –bio'aste 5 – $o!se %an#!epape! %ill sl#dge @ – cattle %an#!ega!den 'aste7 –bio'aste 13 – bio'aste 14 – sl#dge 17 – sl#dgebio'aste 20 – bio'aste
=!o#p A co%post %it#!es
=!o#p ? co%post %it#!es
?acte!ia 1st?a 5 14 20 1 3 @ 7
1stEb 5 14 17 1 3 @ 2nd?a 14 17 20 1 3 @ 2ndEb 14 17 20 1 3 @
-#ngi 1st?a 5 14 20 1 3 @
7
1stEb 5 14 20 1 3 @ 2nd?a 14 17 20 1 3 @ 2ndEb 14 17 20 1 3 @
a ?& beginning o, t$e s#pp!essiveness epe!i%entb E& end o, t$e s#pp!essiveness epe!i%ent
Sinonen
et
al
(2013)
S#pp!essiveness
capacity
val#s
o, co%posts t$at signi+cantly (at p P 005) diFe!ed ,!o% t$e co
!olt!eat%ent (stea% ste!ilied peat) 'e!e !ega!ded as s#pp!esive(6estbe!g et al 2014) ?ased on t$e !es#lts o, 6estbe!g et al (011)a%ong nine selected co%posts seven (n#%be!s 3 5 7 13 11720 in >able 1) 'e!e s#pp!essive to diFe!ent etents one (n#%e!@ in >able 1) 'a s ne#t!al and one (n#%be! 1 in >able 1'a s
cond#cive to .yt$i#% 'ilt disease in t$e +!st s#pp!essivnessepe!i%ent #sing co%posts o, 200B Si%ila! obse!vations e!eobtained in t$e second s#pp!essiveness epe!i%ent #sing c%;posts o, 2009 (6estbe!g et al 2014)
:n t$e p!esent st#dy t$!ee co%post %it#!es 'it$ stongs#pp!ession ability 'e!e catego!ied as =!o#p A – t$e Ms#pp!es
veg!o#pN and t$e !e%aining co%post %it#!es 'it$ lo'e! o! absntdisease s#pp!ession ability 'e!e catego!ied as =!o#p ? –
t$eMot$e! g!o#pN ( >able 1) >$e nat#!al peat (n#%be! 23 ) 'a s 'elys#pp!essive against 'ilt disease '$ile t$e stea% ste!iliedpeat(n#%be! 22 ) 'a s ne#t!al
22 A et!action and D#anti+cation
>o st#dy t$e bacte!ial and ,#ngal co%%#nities total geno
ic
A 'a s et!acted and a%pli+ed #sing p!i%e!s ta!geting 1@S A
and :>S !A genes o, bacte!ia and ,#ngi !espectively Sa%png
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'a s pe!,o!%ed as desc!ibed by 6estbe!g et al (2014) >otal A'a s et!acted ,!o% 03 g o, co%post %it#!es and p#!e peat ,!o%eac$ s#pp!essiveness epe!i%ent #sing a -astA1 S.: it ,o!soil (Qbiogene :nc Ca!lsbad SA) acco!ding to t$e %an#,act#
!e!Nsinst!#ctions >$e concent!ation o, A 'a s %eas#!ed R#o!o;
%et!ically #sing .ico=!een
1
dsA Q#antitation eagent and its(*olec#la! .!obes :nc E#gene SA)
23 A%pli+cations o, bacte!ial 1@S and ,#ngal 1BS :>S ta!get g
enes
#sing 454 seD#encing
All p!i%e!s #sed in t$e p!esent st#dy 'e!e p#!c$ased ,!o%
ligo%e! (elsini -inland) .C a%pli+cations 'e!e cond#
edat t' o stages At t$e +!st stage t$e bacte!ial 1@S !A ta!get ene'a s a%pli+ed #sing p!i%e!s 341- (5 0; CC> AC= ==A ==C A=CA=;3 0) (*#ye! et al 1993) and 907 (5 0; CC= >CA A>> C*> >>= A=>
>> ;30) (:s$ii et al 2000) .C %it#!es incl#ded 1 %l o,
geno%ic
AH 1 %l o, eac$ p!i%e! (10 %*) H 5 %l o, 10S yAy%e ?#Fe!1 %l o, d>. (10 %*) 05 %l o, yAy%es :: A poly%ase(2 %l) ,!o% -inny%es >$e !%o Scienti+c -inlandH 05 %l o, bovine se!#% alb#%in (?SA) (2 0 %g%l) ,!o% -e!%entas >$e%oScienti+c -inlandH and 40 %l o, ste!ile 'ate! Cycling cond
ons'e!e 5 %i n at 94 C ,ollo'ed by 20 cycles o, 20 s at 94 C 20s at
/# et al Applied Soil Ecology 92 (2015) 47–53 49
57 C 30 s at 72 C and a +nal elongation o, 5 %i n at 72
C >$e,#ngal :>S ta!get gene 'a s a%pli+ed #sing p!i%e!s :>S1- (5 0 ;C>>==> CA> >>A =A= =AA =>A A;3 0) (=a!des and ?!#ns 1993) and:>S2 (5 0 ;=C> =C= >>C >>C A>C =A> =C;30) (O$ite et al 1990).C %it#!e ing!edients 'e!e si%ila! to t$at o, bacte!ia 'it$%odi+cations in t$e ?SA (1 %l) poly%e!ase (1 %l) and ste!ile'ate! (3 9 %l) Cycling conditions 'e!e B %i n at 95 C ,ollo'ed by20 cycles o, 1 %i n at 95 C 1 %i n at 55 C 1 %i n at 72 C an
d a +nalelongation o, 10 %i n at 72 C >$e +!st stage .C a%pli+cation 'a spe!,o!%ed in .>C;100 >$e!%o Cycle! (* esea!c$ :nc Oalt$%an*A SA) :n t$e +!st !o#nd t$!ee !eplicates pe! sa%pl
e 'e!epooled and p#!i+ed #sing Agenco#!t A*.#!e T. .C p#!i+cationit (?ec%an Co#lte! ?eve!ly *A SA) p!io! to t$e second !o#nd
At t$e second stage t$e a%pli+cations o, bot$ bacte!i
a and,#ngi 'e!e co%pleted 'it$ 10 additional cycles 'it$ s
a%plespeci+c tag seD#ences and line! A (5 0 ;C=> A>C =CC >CC C>C =C=CCA >CA =;3 0) attac$ed to t$e ,o!'a!d p!i%e! (adapte! A) andline! ? (5 0 ;C>A >=C =CC >>= CCA =CC C=C >CA =;3 0) attac$ed tot$e !eve!se p!i%e! (adapte! ?) .C %it#!es incl#ded 1 %l o, .Cp!od#cts ,!o% t$e +!st !o#nd1 %l o, eac$ Adapte! (10 %*)10 %l o, 5S .$#sion =C ?#Fe! 04 %l o, d>. (10 %*) 05 %l o, .$#sionA .oly%e!ase (2 %l) 125 %l o, *S and 34B5 %l o,
ste!ile'ate! Cycling condition ,o! bacte!ial 1@S !A a%pli+cations 'a s
30 s at 9B C ,ollo'ed by 10 cycles o, 10 s at 9B C 30 s at
@5 C 10 sat 72 C and a +nal elongation o, 5 %in at 72 C O
t$ ,#ngaladapte!s t$e annealing te%pe!at#!e 'a s 55 C >$e secnd stage.C a%pli+cation 'a s pe!,o!%ed in .>C;225 >$e!%o Cy
cle! (*esea!c$ :nc Oalt$%an *A SA) >$e .C p!odcts 'e!ep#!i+ed and D#anti+ed as desc!ibed by osinen et al (20
11) p!io!to 454 seD#encing >$e seD#encing 'a s pe!,o!%ed sing t$e454 =S -$e bacte!ial seD#ence data 'e!e analyed #sing *> (v1350 @4;bit ,o! ,ollo'ing t$e sa%e %et$od as desc!ibed by ?!o'n et a(2013)
>$i s gene!ated a %at!i t$at consisted o, 4537 bacte!ial >s and@4 B ,#ngal >s (ecl#ding singletons – >s 'it$ nly oneseD#ence) and t$is %at!i 'a s #sed in dive!sity analyes %ade'it$ statistical so,t'a!e ($ttp&'''!;p!o"ecto!g)
>$e!ea,te!
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bacte!ial
and
,#ngal
>s
'e!e
classi+ed #sing ib
oso%alatabase .!o"ect (.H Oang et al 2007) and :>E :>S database(ol"alg et al 2013) !espectively >o eplo!e o!ganis%al cove!agea%ong pat$ogen s#pp!essive co%posts !a!e,action c#!ves 'e!ecalc#lated #sing Esti%ateS (Col'ell 2011)
25 Statistical analyses
251 ive!sity analysesSpecies dive!sity indices 'e!e calc#lated as desc!ibed b
y # iet al (2012) >otal n#%be! o, >s 'a s #sed as obse!ved species!ic$ness o, bacte!ia and ,#ngi >otal species !ic$ness o, bacte!ia and,#ngi 'a s esti%ated #sing C$ao1 inde (C$ao 19B4H #g$es et al2001) S$annon dive!sity and Si%psonNs indices 'e!e #
sed toesti%ate dive!sity o, bacte!ial and ,#ngal co%%#nities (C$ao andS$en 2003)
252 S#pp!essiveness analysesCo%ponents o, t$e S.SS (:?* S.SS Statistics A!%on
e' /o! S) p!og!a% pacage 'e!e #sed in statistical aalyses.ai!ed sa%ples t;tests 'e!e #sed to analye i, ti%e Fected.yt$i#% disease s#pp!ession L (ie s#pp!essiveness) and %ic!o;bial dive!sity .ea!son co!!elation analysis 'a s #sed to co%pa!es#pp!essiveness in t$e +!st vs t$e second epe!i%ent co%osts*ann–O$itney tests 'e!e #sed to analye t$e assoiationbet'een s#pp!essiveness and t$e !elative ab#ndances be'eenbacte!ia and ,#ngi .ea!son co!!elation analysis 'a s #ed toassociate s#pp!essiveness 'it$ dive!sity indices and bacte!l and,#ngal ab#ndances (bot$ as pe!centages and as !eads)at t$ep$yl#% and class levels $e %ost ab#ndant acte;!ial p$yla (.!oteobacte!ia ?acte!oidetes Actinobacte!iaAcid;obacte!ia -lavobacte!ia and -i!%ic#tes) 'e!e incl#ded n t$e!eg!ession analysis
:n t$e statistical analyses eac$ epe!i%ent and ti%point(eit$e! t$e beginning o! t$e end o, an epe!i%ent) 'a s l'aysanalyed sepa!ately ie t$e val#es 'e!e not co%bined o! t
eatedas (pse#do) !eplicates Co%posts o, lo' seD#ence !ead'e!eecl#ded ,!o% all statistical analyses As bacte!ial tano%icg!o#ps in t$e beginning o, co%posting and ,#ngal taoo%icg!o#ps 'e!e not associated 'it$ s#pp!essiveness in any , o#!atte%pts to pe!,o!% a p!edictive o! desc!iptive statistical anysis'e s$o' only t$e !es#lts o, bacte!ial co!!espondence analyes int$e end o, t$e epe!i%ents
2@ Q#antitative .C a%pli+cation
>o dete!%ine t$e !atio o, bacte!ia (1@S !A) and ,#ng
(:>S
!A) D#antitative .C (D.C) 'a s done #sing a A ngine.>:C 2 (Contin#o#s -l#o!escence etecto! * esea!c$:ncOalt$a% *A SA) in a +nal vol#%e o, 20 %l containing 1–1
ng o, total A A yA%o
>* S S/?1 =!een D.C it (-inn
%es >$e!% o
Scienti
+c -inland)
'a s
#sed
in all .C
!#ns
?acte!
l and
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,#ngal A 'a s a%pli+ed #sing p!i%e!s pai!s pE and p- :>S3 and:>S4 as desc!ibed ea!lie! ( /# et al 2014)
3 es#lts an d disc#ssion
31 Co%post p!ope!ties and disease s#pp!ession
Co%post o, bio'aste and sl#dge so#!ces s#pp!essed diseasesy%pto%s o, c#c#%be! 'ilt ca#sed by .yt$i#% (6estbe!g et
al2014) -i!st epe!i%ent s#pp!essiveness co!!elated 'it$ secondepe!i%ent s#pp!essiveness (! U 0@7 n U 9 p U 0049) and ave!ages#pp!essiveness 'a s t$e sa%e !ega!dless o, t$e p!od#ction yea!(t U V12 d, U B p U 02@) i!ect biva!iate co!!elations be
t'eens#pp!essiveness and abiotic va!iables (desc!ibed in >able 1 inS#pple%enta!y %ate!ial) 'e!e !a!e .$osp$o!#s concent!ation
t$at va!ied bet'een 2@71–4B@97 %g lV1 in diFe!ent co
%posts(>able
1
in
S#pple%enta!y
%ate!ial)
'a s
associate'it$s#pp!essiveness in t$e +!st (!s U V0B2 n U 11 p U 0007
and t$esecond (! s U V072 n U 11 p U 013) epe!i%ents t$e! coelationsbet'een abiotic va!iables and s#pp!essiveness did not eis
32 Species !ic$ness and disease s#pp!ession
one
o,
t$e
!a!e,action
c#!ves
,o!
bacte!ial
and
,#ngalseD#ences !eac$ed t$e asy%ptote (data not s$o'n) indcatingt$at cove!age o, t$e !ic$ness o, bacte!ia and ,#ngi in c%post%it#!es
st#died
'a s
ins#, +cient o'eve! t$e n#%
be!
o, bacte!ial >s inc!eased d#!ing c#c#%be! g!o't$ in t$
+!stepe!i%ent (t U V37 d, U @ p U 0010) and 'a s indicate in t$e
50 /# et al Applied Soil Ecology 92 (2015) 47–53
second epe!i%ent (t U V22 d, U @ p U 007) de%onst!ating apossible bette! cove!age o, bacte!ial co%%#nity !ic$ness in t$eend o, bot$ s#pp!essiveness epe!i%ents o! a %o!e dive!sebacte!ial co%%#nity >$e n#%be! o, >s o! t$e c$ange in t$en#%be! o, >s $o'eve! did not co!!elate 'it$ s#pp!essivenessat any ti%e point (! 04@ n U @ p W 029) C$ao1 inde S$annondive!sity inde and Si%psonNs dive!sity inde did not co!
!elate'it$ s#pp!essiveness at any ti%e point (p W 011)
33 *ic!obial p!o+ling o, co%posts
>$e biotic co%ponent $as been doc#%ented to play a c!
#cial
!ole in disease s#pp!ession b#t ve!y ,e' st#dies $ave desc!ibedbot$ bacte!ial and ,#ngal dive!sities in plant disease s#pp!essiveco%posts (?onano%i et al 2010) :n gene!al t$e %a"o! bacte!ial
and ,#ngal g!o#ps ,o#nd in t$e p!esent st#dy #sing 454 seD#encinga!e si%ila! to p!evio#s st#dies t$at $ave #sed ot$e! seD#encingapp!oac$es (eg :ll#%ina *iSeD seD#encing Sange! seD#encing)to desc!ibe %ic!obial co%%#nities in co%posts o, si%ila! !a '%ate!ials (e$e! et al 2013)
331 ?acte!ial dive!sity ove!vie' and disease s#pp!essionA total o, 237710 D#ality;+lte!ed seD#ences 'e!e
identi+ed asbacte!ia by . classi+e! >$e n#%be! o, 1@S !A seD
#encesobtained pe! sa%ple va!ied ,!o% 22B4 to 11749 (%edian U 5B93)
Sa%ples 'it$ eceptionally lo ' n#%be! o, seD#ences (1@2–1215)'e!e ecl#ded ,!o% statistical analysis -o!ty;si classes belongingto 20 bacte!ial p$yla 'e!e detected and t$e %ost do%inant clases'e!e !aned acco!ding to t$ei! seD#ence ab#ndance o!de!s (- 1)As epected
.!oteobacte!ia
co%p!ised
t$e
%a"o!ity
ot$eseD#ences in all co%posts (anon et al 200B) acco#ntig ,o!49 –@4L and 34–@1L o, t$e total seD#ences in t$e +!st and seconds#pp!essiveness epe!i%ents !espectivelyH '$ile its s#b$yla=a%%ap!oteobacte!ia Alp$ap!oteobacte!ia and ?etap!oteoac;te!ia 'e!e t$e %ost ab#ndant g!o#ps >$e %ost do%inant clas int$e bacte!ial co%%#nity s$i,ted ,!o% =a%%ap!oteobacte!ia 1stepe!i%ent
beginning&
1B–4BL 2nd epe!i%ent
begi
ning&
X(-i
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@–43L) to Alp$ap!oteobacte!ia (1st epe!i%ent end& 23–4@L2nd
epe!i%ent
end&
34 –50L)
d#!ing
bot$
s#pp!essivenessepe!i%ents .!evio#sly Alp$a; and =a%%ap!oteobacte!ia $avebeen doc#%ented as typical co%post bacte!ia (*ic$el et al 2002Hagn et al 200BH .a!tanen et al 2010) =a%%ap!oteobacte!iaespecially .se#do%onas 'e!e desc!ibed as typical bioc
ont!olo!ganis%s t$at p!otects plant ,!o% t$e oo%ycete .yt$i#% #lti%#%(?oe$% et al 1993H eonico et al 2005) and attained t$e$ig$est pe!centage o, positive co!!elation 'it$ disease s#pp!es;siveness (?onano%i et al 2010) o'eve! in t$e c#!!ent st#dy=a%%ap!oteobacte!ia
seD#ence
!eads
d!opped
d#!ing
bot$disease s#pp!ession epe!i%ents A possible eplanation ,o! t$isobse!vation co#ld be t$e t!op$ic inte!action bet'een t$e ,#
ngalpat$ogen (.yt$i#%) and bacte!ia (=a%%ap!oteobacte!ia) int$e!oot envi!on%ent (!$iosp$e!e) (-!ey;lett et al 2011) t$e!e,o!eca#sing t$e !eplace%ent o, =a%%ap!oteobacte!ia 'it$ Alp$ap;!oteobacte!ia
.$yla ?acte!oidetes Acidobacte!ia and -i!%ic#tes 'e!e co%;%only dist!ib#ted in all co%posts !ega!dless o, t$ei! %at#!ity and
c$e%ical and p$ysical p!ope!ties Oit$in t$ose classes Sp$igo;bacte!ia (5 –42L) and -lavobacte!ia (01–19L) 'e!e n#%
o#sActinobacte!ia 'e!e co!!elated positively 'it$ s#pp!essivenesinbot$ epe!i%ents (! W 0B2 n U 7 p 0029) :n bac'a!d eg!es;sion t$e ab#ndances (L at t$e end o, epe!i%ent) o, Aino;bacte!ia aFected s#pp!essiveness in bot$ epe!i%ents (*S U 300- U 92 d, U 1 @ p U 0029 in t$e +!st and *S U 2@10 - U 23
d, U 1 @p U 0005 in t$e second epe!i%ent) >$e eFects 'e!e positiveie$ig$ actinobacte!ial ab#ndance inc!eases .yt$i#% 'ilt disases#pp!ession :t $as been 'ell doc#%ented t$at Actinobacte!ia avea positive i%pact on plant disease s#pp!ession d#e to its stongability to p!od#ce antibiotic co%po#nds (C!oss 19B2) A !eentst#dy
by *cella!
and
elson
(2003)
also
s#ppo!ted
t$isstate%ent
by
de%onst!ating
a
%o!e
do%inant
p!esenco, Actinobacte!ia in disease s#pp!essive conso!tia t$an in disasecond#ctive
o!
ne#t!al
conso!tia
o'eve!
a,te! epo!ing2423 cases de!ived ,!o% 252 st#dies on disease %anage%ent#sing o!ganic a%end%ents ?onano%i et al (2010) concl#ded $at
-ig 1 >$e ab#ndance o, %a"o! bacte!ial classes (log10 scale) in =!o#p A and =!o#p ? co%post %it#!es in t$e +!st epe!i%ent >$e !an o, bacte!ial class in t$e secondepe!i%ent 'as eactly t$e sa%e as in t$e +!st epe!i%ent :n spite o, t$e potential !oles o, Acidobacte!ia =p14 in t$e s#pp!ession o, .yt$i#% 'ilt diseae its seD#enceab#ndance 'as too lo ' to be visible in t$is +g#!e (a) ?eginning o, epe!i%ent (b) end o, epe!i%ent
/# et al Applied Soil Ecology 92 (2015) 47–53 51
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Actinobacte!ia 'e!e only di!ectly co!!elated 'it$ disease s#pp!es;sion in a li%ited n#%be! o, epe!i%ental cases Actinobacte!ia a!eno'n to $ave a %a"o! !ole in deco%position o, o!ganic %ate!ialspa!tic#la!ly ,o! deg!adation o, %ac!o%olec#les s#c$ as cell#lose$e%icell#lose lignin and c$itin (Stege! et al 2003) >$e!e ,o!e t$ediFe!ences in t$ese obse!vations co#ld be eplained by diF
e!entepe!i%ental conditions s#c$ as co%post o!igin co%post c$e%i;cal and
p$ysical p!ope!ties
co%post %at#!ity co%posting%et$od soil;bo!ne pat$ogens and test plants in t$e above;%entioned st#dies Also t$e epe!i%ental conditions ,avo!ingActinobacte!ia %ig$t be ,o!t#ito#sly det!i%ental to soil;bo!nepat$ogens 'it$ no di!ect ca#se;eFect !elations$ip and t$e!ebynot be based on co%petition bet'een si%ila! %ic!obes :n all
t$ep!esence and do%inance o, Actinobacte!ia appea!s to be casespeci+c and p!one to la!ge va!iations
n t$e cond#cive side constant t!ends 'e!e not detected >$e$#ge taono%ic va!iability at na!!o'e! taono%ic levels (eggen#s) %a y be a !eason '$ y st!aig$t,o!'a!d patte!ns co#ld not bedetected
es#lts ,!o% > analysis #sing 454 seD#encing data !evealedt$at Acidobacte!ia a!e p!esent in all co%posts st#died otablyAcidobacte!ia
=p14
'as
detected
only
in
=!o#p
A
co%post%it#!es – ie co%posts 'it$ st!ong disease s#pp!ession a
bility(0011–001BL) '$ic$ indicated its potential !ole in t$e s#pp!es;sion o, .yt$i#% 'ilt disease Acidobacte!ia $ave p!evio#sly beendetected in 'aste 'ate! sl#dge co%post and bio,e!tilie! t$atassociated 'it$ banana -#sa!i#% 'ilt disease s#pp!ession (Z$anget al 2012H -!acc$ia et al 200@H S$en et al 2014) b#t not itss#bg!o#p Acidobacte!ia =p14 >o o#! no'ledge t$is is t$e ve!y+!st st#dy to note t$e potential !elevance o, Acidobacte!ia =p
14 in.yt$i#%
disease
s#pp!ession
>$e !es#lts
obtained $e!e
a!e$o'eve! not s#, +cient to decla!e Acidobacte!ia =p14 as t$e!eliable indicato! o, .yt$i#% disease s#pp!essive capability *o!eco%p!e$ensive !esea!c$ on its association 'it$ disease s#pp!es;sion needs to be cond#cted
X(-igY2)>[:=\
332 -#ngal dive!sity ove!vie' and disease s#pp!ession >$e n#%be! o, :>S !A seD#ences obtained pe! sa%ple v
!ied,!o% B1B to 7171 (%edian U 2143) >$i !te en classes belongig to 3p$yla 'e!e detected (1217B3 D#ality;+lte!ed seD#ences ac!oss allsa%ples) and t$e ab#ndance !an o!de! o, t$e %ost doinantclasses a!e s$o'n in -ig 2 As epected Asco%ycota !ep!esnted
t$e la!gest p$yl#% acco#nting ,o! B5–97L o, t$e total seD#ence!eads ac!oss all co%post %it#!es in t$e +!st s#pp!essvenessepe!i%ent
Asco%ycota classes E#!otio%ycetes ot$ido%y;cetes
So!da!io%ycetes
and Sacc$a!o%ycetes 'e!e
t$%ostp!edo%inant
g!o#ps
>$e
!es#lts
o,
$e +nding s#ppo!tst$e
p!evio#s
state%ent t$at Zygo%ycota
a!e %o!e
closelyassociated 'it$ co%posts o, %an#!e o!igin t$an co%pots o, ot$e!
o!igins
(e$e!
et
al
2013)
:n
cont!ast
in
$e +!st
s#pp!essiveness epe!i%ent Zygo%ycota co%p!ised only 02–1Lo, t$e total seD#ence !eads in Co%posts 1 and 3 >$e total a%o#ntso, n#t!ients in co%post %it#!es #sed in bot$ s#pp!essieness
epe!i%ents did not diFe! in a signi+cant 'a y ,!o% eac$ot$e!o'eve! in t$e second epe!i%ent Co%post 3 'a s ,o#ndto bei%%at#!e acco!ding to its 4;3; !atio and ottg!ad%eas#!e%ent
'$ile
Co%post 1 $ad clea!ly lo'e! ect!icalcond#ctivity (EC) in co%pa!ison 'it$ Co%post 1 in t$
e +!stepe!i%ent (6estbe!g et al 2014) >$e !elations$ip bt'eenco%post %at#!ity and its %ic!obial co%%#nity is yet too co%pleto be identi+ed
>$e!e,o!e
%o!e
st#dies
a!e needed
to con+!
% t$e
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-ig 2 >$e ab#ndance o, %a"o! ,#ngal classes (log10 scale) in =!o#p A and =!o#p ? co%post %it#!es in t$e +!st epe!i%ent >$e !an o, ,#ngal claes in t$e secondepe!i%ent 'as eactly t$e sa%e as in t$e +!st epe!i%ent (a) ?eginning o, 1st epe!i%ent (b) end o, 1st epe!i%ent
52 /# et al Applied Soil Ecology 92 (2015) 47–53
p#tative !elations$ip bet'een co%post %at#!ityEC and !elativeab#ndances o, Zygo%ycota and Asco%ycota
45 4 seD#encing !es#lts o, t$e +!st s#pp!essiveness epe!i
%entindicated t$at ?asidio%ycota class Cystobasidio%ycetes co#ld$ave cont!ib#ted to t$e s#pp!ession o, .yt$i#% 'ilt disease d#e
to its p!esence (005@–03@L) in al l =!o#p A co%post %it#!es andits absence
in =!o#p
?
co%post %it#!es
:n
t$e
seconds#pp!essiveness epe!i%ent no
Cystobasidio%ycetes 'e!edetected in any co%post st#died Cystobasidio%ycetes $ave !a!elybeen %entioned in p!evio#s st#dies -o! ea%ple Cystobasidio;%ycetes $ave not been !epo!ted as potential indicato!s o, anyplant disease s#pp!ession by co%post alt$o#g$ ce!tain species o,
t$e class s#c$ as t$e pa!asitic ,#ng#s $odoto!#la (Sing$ et al2012) $ave been detected in t$e ea!ly p$ase o, %#s$!oo%co%posting
o'eve!
it
is
co%%only
accepted
t$at
so%e%e%be!s o, t$e sa%e $ig$e! taono%ic level %ay $ave si%ila!activity in plant–%ic!obe–soil inte!actions '$ile ot$e!s %ay not$ave >o i%p!ove #nde!standing t$e potential disease s#pp!essionabilities o, Cystobasidio%ycetes and its %e%be!s ,#!t$e! st#dies
a!e
needed
.!evio#sly
so%e
benign
,#ngi
t$at
a!e
closely
!elated to plant pat$ogenic ,#ngi (eg -#sa!i#% spp) $avebeen!epo!ted to s#pp!ess to plant disease appa!ently d#e to $ei!co%petition ,o! space and !eso#!ces in t$e sa%e plant env!on;%ent (-!avel et al 2003H ?onano%i et al 2010) o'eve! no
benign ,#ngi !elated to pat$ogenic .yt$i#% 'e!e ,o#nd int$ec#!!ent st#dy
-#ngal ab#ndances (L) at t$e class level o, any co%post vaiedetensively bet'een epe!i%ents ,o! ea%ple signi+cant d
Fe!;ences 'e!e ,o#nd in ot$ideo%ycetes >!e%ello% ycetesandSacca!o%ycetes (pai!ed sa%ples t;test t W 29 d, U B p 00
9) >$eab#ndance o, t$e above;%entioned g!o#ps see% to depend %o!eon local envi!on%ent t$an t$e co%post pe! se
34 elative ab#ndance o, bacte!ia and ,#ngi #sing D#antitative
.C
es#lts calc#lated ,!o% D.C standa!d c#!ves indicated t$ainspite o, lo ' seD#ence !eads in so%e co%posts t$e copy n#%be!!atios o, bacte!ial 1@S and ,#ngal :>S !A R#ct#ated 'it$in
t$e!anges o, 7–90 t$!o#g$o#t bot$ sc!eening epe!i%ents allco%posts (24 ] 22@ and 3@ ] 277 in =!o#p A and =
o#p ?co%post %it#!es !espectively) >$e!e 'as no clea! assocation
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bet'een .yt$i#% disease s#pp!ession and t$e !elative ab#ndancesbet'een bacte!ia and ,#ngi ( W 3 Z W V15 n U 9 p W 01
2)
4 Concl#sions
Co%posts %ade ,!o% bio'aste and sl#dge s#pp!essed diseaseca#sed by .yt$i#% Actinobacte!ia en$anced disease s#pp!e
ssionin bot$ epe!i%ents .$osp$o!#s concent!ation 'a s associatednegatively 'it$ disease s#pp!ession in bot$ epe!i%ents >$est#died co%post
%it#!es
contained
ab#ndant
and dive!se%ic!obial
co%%#nities
'it$
.!oteobacte!ia
and
Asco%ycotabeing t$e do%inant bacte!ial and ,#ngal !ep!esentatives !espec;tively t$e! %a"o! evenly dist!ib#ted %ic!obial g!o#ps !evealed bypy!oseD#encing 'e!e bacte!ia o, p$yla ?acte!oidetes Acid
obac;te!ia -i!%ic#tes and ,#ngi o, t$e Asco%ycota classes E#!o;tio%ycetes ot$ideo%ycetes and So!da!io%ycetes *e%be!s o, Acidobacte!ia =p14 and Cystobasidio%ycetes 'e!e only p!esent inst!ongly;s#pp!essive co%post %it#!es and co#ld t$e!e,o!e bes#ggested as t$e !espective bacte!ial and ,#ngal indicato!s o, s#pp!essiveness against .yt$i#% disease on c#c#%be! plantso'eve! t$e !ole o, t$e above;%entioned %ic!oo!ganis
%s indisease s#pp!ession !eD#i!es ,#!t$e! st#dy
Acno'ledge%ents
Oe 'is$ to t$an t$e co%posting plants ,o! giving #s co%pots
>$is st#dy 'a s %ainly s#ppo!ted by t$e Acade%y o, -inland (=anto 121574) /#Ns 'o! 'a s pa!tially s#ppo!ted by t$e -i
nis$C#lt#!al -o#ndation (Cent!al -#nd and .^i"^t;^%e egional -#nd)and t$e EnS>e g!ad#ate sc$ool (-innis$ octo!al .!og!a%%
inEnvi!on%ental Science and >ec$nology)
Appendi A S#pple%enta!y data
S#pple%enta!y data associated 'it$ t$is a!ticle can be ,o#n int$e online ve!sion at $ttp&ddoio!g10101@"apsoil201503005
e,e!ences
?oe$% * *adden #ovin en 19BB >$e !ole o
%ic!obial activity in s#pp!ession o, da%ping oF ca#sed by .yt$i#% #lti%#
.$ytopat$ology 7B 314–322
Col'ell 2011 Esti%ates statistical esti%ation o, species !ic$ness and sa!edspecies ,!o% sa%ples 6e!sion 9 se!Ns g#ide and application p#blis$ed a$ttp&p#!loclco!gesti%ates
C!oss > 19B2 Actino%ycetes& a contin#ing so#!ce o, ne ' %etabolites evnd
*ic!obiol 23 1–1Banon * -!ane;O$ittle : :nsa% C$en / ada! / 200B *olec#la
analysis o, bacte!ial co%%#nity s#ccession d#!ing p!olonged co%post c#!g
-E*S *ic!obiol Ecol @5 133–144de ?e!toldi * 2010 .!od#ction and #tiliation o, s#pp!essive co%post&
envi!on%ental ,ood and $ealt$ bene+ts :n & :nsa% -!ane;O$ittle :=obe!na * (Eds) *ic!obes at Oo! – -!o% Oastes to eso#!ces Sp!inge6e!lag ?e!lin eidelbe!g pp 153–170
iKne - Santos * >ello C 2005 S#pp!esion o, soilbo!ne pat$ogens byco%post& s#pp!esive eFects o, g!ape %a!c co%post on p$ytopat$ogenicsoo%ycetes Acta o!tic @97 441–4@0
-!acc$ia ebbe CC 200@ ?acte!ial div
sityin +nis$ed co%post and ve!%ico%post& diFe!ences !evealed by c#ltivatioindependent analyses o, .C;a%pli+ed 1@S !A genes Appl *ic!obiol?iotec$nol 71 942–952
-!avel livain C Alabo#vette C 2003 -#sa!i#% oyspo!#% and its bioco!ol
e' .$ytol 157 493–502-!ey;lett . ?#!linson . evea# A ?a!!et * >a!a * Sa!nig#et A 2011
?acte!ial–,#ngal inte!actions& $yp$ens bet'een ag!ic#lt#!al clinicalenvi!on%ental and ,ood %ic!obiologists *ic!obiol *ol ?iol ev 75 5B3–
@09 =a!des * ?!#ns > 1993 :>S p!i%e!s 'it$ en$anced speci+city ,o!
basidio%ycetes–application to t$e identi+cation o, %yco!!$iae and !#sts ol
Ecol 2 113–11Bada! / =o!eci ? 1991 S#pp!ession o, ge!%ination o, scle!otia o, Scle!ot%
!ol,sii in co%post Soil ?iol ?ioc$e% 23 303–30@
ada! / *andelba#% 1992 S#pp!essive co%post ,o! biocont!ol o, soilboe
plant;pat$ogens .$ytopa!asitica 20 S113–S11@
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