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Th"s "s a( Hpe( /ccess ar-"cle )"s-r"bu-e) u()er -he -erms o -he 'rea-"%e 'ommo(s /--r"bu-"o( C"ce(se

0h--pcrea-"%ecommo(s.orgl"ce(sesby2.& wh"ch perm"-s u(res-r"c-e) use )"s-r"bu-"o( a() repro)uc-"o( "(

a(y me)"um pro%")e) -he or"g"(al wor, "s properly c"-e).

Abstract

Background

Bacillus thuriniensis "s a bac-er"um ,(ow( or pro)uc"(g pro-e"( crys-als w"-h "(sec-"c")al proper-"es. These

-oA"(s are w")ely sough- a-er or co(-roll"(g agr"cul-ural pes-s )ue -o bo-h -he"r spec""c"-y a() -he"r

appl"cab"l"-y "( -ra(sge("c pla(-s. There "s grea- "(-eres- "( "sola-"(g s-ra"(s w"-h "mpro%e) or (o%el -oA"(

charac-er"s-"cs howe%er "sola-"(gB. thuriniensis rom -he e(%"ro(me(- "s -"me co(sum"(g a() y"el)s rela-"%ely

ew "sola-es o "(-eres-. New approaches -oB. thuriniensis "sola-"o( ha%e bee( a() co(-"(ue -o be sough-. I(

-h"s repor- ca()")a-eB. thuriniensis "sola-es were reco%ere) rom e(%"ro(me(-al samples us"(g a comb"(a-"o(

o a (o%el s-a"( h"gh -hroughpu- a() re)uce) selec-"o(. Isola-es were ur-her charac-er"De) by S$SP/!= l"gh-

m"croscopy P'@ probe hybr")"Da-"o( a() w"-h selec-e) "sola-es $N/ se:ue(c"(g b"oassay or =lec-ro(

M"croscopy.

Results

>ase) o( S$SP/!= pa--er(s a() -he prese(ce o cr< ge(es or a crys-al 7# ca()")a-e (o(clo(al "sola-es o B.

thuriniensis were ")e(-""e) rom 4 samples a() o%er 1&&&& colo("es. /l-hough o(ly 167# 02& o -he

"sola-es showe) $N/ homology by Probe ?ybr")"Da-"o( or P'@ -o commo( cr< ge(es "("-"al charac-er"Da-"o(

re%eale) a surpr"s"(gly r"ch l"brary -ha- "(clu)e) a pu-a-"%e (ema-oc")al ge(e a (o%el "lame(-ous s-ruc-ure

assoc"a-e) w"-h a crys-al a spore w"-h sp",es or"g"(a-"(g rom a %ery small parasporal bo)y a() "sola-es w"-h

u(usually small crys-als. Lhe( compare) -o repor-s o o-her scree(s -h"s scree( was also a-yp"cal "( -ha- o(ly

37# "sola-es 03. pro)uce) a b"pyram")al crys-al a() 247# 03& o -he "sola-esO spores possesse) a(

a--ache) )ar,s-a"("(g bo)y.

Conclusion

@esul-s sugges- -ha- -he scree("(g me-ho)ology a)op-e) "( -h"s s-u)y m"gh- )el"%er a %as-ly r"cher a()

po-e(-"ally more useul l"brary oB. thuriniensis "sola-es as compare) -o -ha- ob-a"(e) w"-h commo(ly repor-e)

me-ho)olog"es a() -ha- by eA-e(s"o( me-ho)olog"es u()ame(-ally )"ere(- rom curre(- me-ho)s shoul) also

be eAplore).

Background

Bacillus thuriniensis "s a bac-er"um ,(ow( or pro)uc"(g pro-e"( crys-als w"-h pes-"c")al proper-"es. These

-oA"(s are w")ely sough- a-er or co(-roll"(g agr"cul-ural pes-s )ue -o bo-h -he"r spec""c"-y a() -he"r

appl"cab"l"-y "( -ra(sge("c pla(-s. /l-hough a(y par-"cular -oA"( has a )es"rably res-r"c-e) hos- ra(ge -here "s a
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large (umber o )"ere(- -oA"(s each show"(g -oA"c"-y -o o(e o ma(y )"%erse pes-s. or -hese reaso(s -here "s

curre(-ly grea- "(-eres- "( "sola-"(g (o%el s-ra"(s oB. thuriniensis w"-h e"-her u(":ue hos- spec""c"-y or

ele%a-e) -oA"c"-y. There are (umerous repor-s o( a--emp-s -o "sola-e (o%elB. thuriniensis s-ra"(s rom -he

e(%"ro(me(- 12. I( ma(y o -hese some orm o e(r"chme(- s-ep "s use) "( co(u(c-"o( w"-h Phase 'o(-ras-

M"croscopy as -he bac,bo(e o -he scree("(g s-ra-egy 34. The resul-s o -hese scree(s ha%e a cur"ously s"m"lar

resul- where a large propor-"o( o -he "sola-es 033.5 -o # co(-a"( a cr-132

show"(g spores 0)ouble arrow a() rela-"%ely l"gh-ly s-a"("(g crys-als 0s"(gle arrow.
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)igure 2.0xa!ples of the 1capped1 spore !orphology. $ar, blue QcapsQ a--ache) -o l"gh-er

s-a"("(g spores. / >-22 > >-14 ' >-256 $ >-146 a() = >-214.

'lass"y"(g "sola-es as clo(al 0s"bl"(gs " -hey were rom -he same sample a() ha) -he same crys-al or cap

morphology 7# (o(clo(al "sola-es were collec-e) compr"s"(g mos-ly rou() or amorphous crys-als w"-h 6

rec-a(gular a() 3 b"pyram")al morpholog"es. Some samples were ou() -o ha%e mul-"ple ca()")a-es o B.

thuriniensis e"-her o -he same or )"ere(- parasporal bo)y morphology.

Two "sola-es 0>-1 a() >-256 showe) appe()ages ema(a-"(g rom parasporal bo)"es. >-1 was

charac-er"De) ge(erally as ha%"(g mul-"ple perhaps -hree lo(g sp",es ema(a-"(g rom a small sporeassoc"a-e)

parasporal bo)y 0"gure 3. S$SP/!= a(alys"s o >-1 re%eale) -wo pro(ou(ce) low molecular we"gh-

pro-e"(s o approA"ma-ely 22 a() 23 ,$a 0)a-a (o- show(. Isola-e >-256 was ou() by elec-ro( m"croscopy -o

co(-a"( a mul-""ber "lame(- ema(a-"(g rom a parasporal bo)y loca-e) w"-h"( a sac,l",e s-ruc-ure presume)

-o be -he eAospor"um 0"gure 4. / prel"m"(ary repor- o( -h"s "sola-e has bee( prese(-e) elsewhere .

)igure ./hase contrast !icrograph sho%ing spiked appendages e!anating fro! the

spores of Bt+-33. Co(g s-ra"gh- sp",es 0a ema(a-"(g rom a small sporeassoc"a-e) parasporal bo)y 0b.
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)igure 4.0lectron 5icrograph of Bt2-#. / Tra(sm"ss"o( =lec-ro( M"crograph o (ega-"%ely

s-a"(e) spores rom >-256 co(-a"("(g a "lame(- 0a a() a sacl",e s-ruc-ure co(-a"("(g a spore 0b a()

parasporal bo)y 0c.

Three samples pro)uce) "sola-es w"-h rela-"%ely small parasporal bo)"es >-126 >-14& a() >-1#& 0"gure 5.

>-126 was "sola-e) rom sa() -a,e( a ew "(ches abo%e a h"gh -")e mar, o( a beach >-1#& rom a resh

ma(ure p"le 0cal sheep a() goa- a() >-14& rom compos-"(g ma(ure collec-e) rom a bar( hous"(g sheep

goa-s ca--le horses a() p"gs. S$SP/!= a(alys"s showe) maor pro-e"( ba()s o a rela-"%ely low molecular

we"gh- be-wee( 11F1# ,$a or each o -hese "sola-es 0)a-a (o- show(.

)igure .Relati"ely s!all parasporal 6odies. S-a"(e) >r"gh-"el) m"crographs 01&&G o >-1

22 0a >-14& 0b a() >-1#& 0c.

Two "sola-es 0>-133 a() >-135 showe) se:ue(ce homology -o a (ema-oc")al ge(e 0See Qcr< ge(e pro"leQ

below. >-133 showe) (o crys-als a() amo(g hu()re)s o spores w"-h"( -he m"croscop"c "el) o %"s"o( o(ly a

ew possesse) a blues-a"("(g cap o( o(e e() o -he spore. /-er repea-e) cul-ure o( me)"a -he rela-"%e (umber

o caps appeare) -o "(crease sl"gh-ly 0"gure6/ wh"ch mos- probably woul) (o- ha%e bee( )e-ec-e) by Phase

'o(-ras- M"croscopy 0"gure 6>. 'rys-als were (o- "("-"ally obser%e) w"-h e"-her Phase 'o(-ras- M"croscopy or

s-a"(e) spec"me(s. ?owe%er whe( spec"me(s were hea-e) s-ro(gly o( -he m"croscope sl")e some spec"me(s

prepare) "( -h"s ma((er showe) o%al crys-als ly"(g us- la-eral -o -he -erm"(al e() o -he spore 0"gure 6'.

I(-eres-"(gly bo-h >-133 a() >-135 came rom -he same sample a( area o )ecompos"(g a("mal ma(ure. >-1

35 )emo(s-ra-e) a %ery small cap howe%er maor pro-e"( ba()s were (o- obser%e) o( S$S gel a(alys"s.

)igure #./hase ontrast 5icroscopy of sporulated cells of Bt+-. / s-a"(e) prepara-"o( o

>-133 %"ewe) u()er >r"gh-"el) m"croscopy 1&&A o"l "mmers"o( > >-133 %"ewe) u()er Phase 'o(-ras-
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m"croscopy 1&&A o"l "mmers"o( ' ?ea--rea-e) prepara-"o( o >-133 show"(g phase)ar, crys-als ly"(g

sl"gh-ly la-eral a() -erm"(al -o -he spore Phase 'o(-ras- m"croscopy 1&&A o"l "mmers"o(.

cry-cyt gene prole

Probe a() P'@ a(alys"s "()"ca-e) -ha- o -he 7# (o(clo(al "sola-es o(ly 16 showe) homology -o a(y o(e o -he

maor groups o cr< ge(es -es-e) 0Table 1. cr< a() c-214 wh"ch was pos"-"%e by P'@ or a cr!S'4/3B. thuriniensis sero%ar thuriniensis ?$2 Score 2.&;

>!S'4$1B. thuriniensis sero%ar #ursta#i ?$1 Score 2.& a() >!S'4E3B. thuriniensis sero%ar ai@a"ai?$133 Score 2.7.

iscussion

L"-h ew eAcep-"o(s #1& -he crys-al a() spore oB. thuriniensis are release) separa-ely )ur"(g sporula-"o(.

I- was -hus surpr"s"(g -ha- 247# (o(clo(al ca()")a-eB. thuriniensis "sola-es possesse) spores w"-h a

Qcappe)Q appeara(ce. Some o -hese caps appeare) -o co(-a"( crys-als s"(ce phase )ar, obec-s w"-h a l"gh-
http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.ncbi.nih.gov/entrez/query.fcgi?db=Nucleotide&cmd=search&term=U13955http://www.biomedcentral.com/1471-2180/5/52/figure/F7http://www.biomedcentral.com/1471-2180/5/52/figure/F7http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.biomedcentral.com/1471-2180/5/52/figure/F7http://www.ncbi.nih.gov/entrez/query.fcgi?db=Nucleotide&cmd=search&term=U13955http://www.biomedcentral.com/1471-2180/5/52#B9http://www.biomedcentral.com/1471-2180/5/52#B10http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.ncbi.nih.gov/entrez/query.fcgi?db=Nucleotide&cmd=search&term=U13955http://www.biomedcentral.com/1471-2180/5/52/figure/F7http://www.biomedcentral.com/1471-2180/5/52/table/T1http://www.biomedcentral.com/1471-2180/5/52/figure/F7http://www.ncbi.nih.gov/entrez/query.fcgi?db=Nucleotide&cmd=search&term=U13955http://www.biomedcentral.com/1471-2180/5/52#B9http://www.biomedcentral.com/1471-2180/5/52#B10


10/19

ce(-er coul) be see( "( ma(y o -he caps "(clu)"(g >-256 wh"ch was subse:ue(-ly co("rme) by =lec-ro(

M"croscopy -o possess a parasporal bo)y w"-h"( -he eAospor"um 0"gure 4. The u(c-"o(al s"g(""ca(ce o

sporeassoc"a-e) crys-als "s (o- -o-ally u()ers-oo) howe%er be(e"-s such as pro-ec-"o( aga"(s-


11/19


12/19

was obser%e). These )a-a woul) sugges- -ha- some cappe) spores co(-a"(e) "( eA"s-"(gB. thuriniensis l"brar"es

m"gh- also possess appe()ages a() shoul) be reeAam"(e) spec""cally or -he"r prese(ce.

Perhaps mos- s-r","(g abou- -h"s scree( was -he low perce(-age o "sola-es pro)uc"(g b"pyram")al crys-als. I(

l"-erally all repor-e) scree(s or e(%"ro(me(-al "sola-es oB. thuriniensis -he perce(-age o b"pyram")al

crys-als "s usually abo%e 4& 11 howe%er o(ly 3 "sola-es or 3. o -he (o(clo(al "sola-es where show(

-o ha%e a b"pyram")al morphology "()"ca-"(g a scree( -ha- )e%"a-e) s"g(""ca(-ly rom -he (orm.

!onclusion

@esul-s sugges- -ha- -here are a)%a(-ages -o us"(g a s-a"( o%er -he commo(ly use) Phase 'o(-ras- M"croscopy

par-"cularly "( -he )e-ec-"o( o small crys-als low (umbers o crys-alscappe) spores a() or "-s appl"cab"l"-y or

use "( -he as- h"gh -hroughpu- )e-ec-"o( oB. thuriniensis ca()")a-es rom -he e(%"ro(me(-. There "s a

co(-"(u"(g (ee) -o eAplore -he be(e"-s o (ew me-ho)olog"es or -he )e-ec-"o( oB. thuriniensis rom -he

e(%"ro(me(-.

"ethods

Sample collection and preparation

Samples were ra()omly collec-e) rom e(%"ro(me(-ally )"%erse sources 0"(clu)"(g beach sa() ores- so"l

a:ua-"c a() "(-er-")al se)"me(-s a() so"ls rom urba( rural a() agr"cul-ural areas place) "( a plas-"c bag a()

hel) a- room -empera-ure u(-"l process"(g. Se%e( )"ere(- me-ho)s 0/! were "("-"ally use) or sample

prepara-"o( 0Table 2 where Me-ho) was use) pre)om"(a-ely 0Table 3.

$a6le 2.Summary o prepara-"o( me-ho)s /!

$a6le .Prepara-"o( me-ho)s a() -he"r use.

B thuringiensis isolation

Samples 0&.25 g were prepare) us"(g o(e or more o Me-ho)s /! 0Table 2 -he( 2&& l a() 2& l were

cul-ure) o( Sporula-"o( pla-es 0Nu-r"e(- /gar w"-h ''Y sal-s 1# -o wh"ch a(-"b"o-"cs were a))e) as "()"ca-e)

"( Table2 a- 3&U' or 2 )ays -o allow sporula-"o(.


13/19

us"(g a 1&&G "mmers"o( o"l obec-"%e. 'ell morphology was (o-e) or all ca()")a-es oB. thuriniensis wh"ch

was )e"(e) as a large sporeorm"(g ro) whose ell"pso")al spore )") (o- swell -he mo-her cell a() wh"ch

pro)uce) a parasporal bo)y upo( sporula-"o( 2&21.

S!S protein electrophoresis

Two loopuls o bac-er"al grow-h were remo%e) rom a Sporula-"o( pla-e -o 5& l o "ce col) Pro-e"( Sample

>uer 22 bo"le) or 2 m"( -he( roDe( u(-"l use. $e(a-ur"(g S$S gel elec-rophores"s was perorme) o(

pro-e"( samples accor)"(g -o -he me-ho) o Camell" 23. The rela-"%e mob"l"-y or each ba() was calcula-e) by

apply"(g -he e:ua-"o( @? V /'/> where / was -he pos"-"o( o -he -op o -he gel > was -he pos"-"o( o -he

-op o -he )ye ro(- a() ' was -he pos"-"o( o -he -op o each ba() o "(-eres-. @ela-"%e mob"l"-y vs -he

molecular we"gh-s o ,(ow( pro-e"(s 0116 ,$a #2 ,$a 66 ,$a 45 ,$a 21 ,$a was plo--e) a() l"(ear

regress"o( perorme) -o )e-erm"(e -he e:ua-"o( or -he l"(e o bes- "- wh"ch was use) -o )e-erm"(e -he

Molecular we"gh-s 0B$a o u(,(ow( ba()s.

"CR of to#in genes

$N/ -empla-e was prepare) by lys"s prec"p"-a-"o( where"( approA"ma-ely hal a loopull o cells rom a

sporula-e) pla-e was place) "(-o 2 mls o C >ro-h "(cuba-e) a- 6&U' or 1& m"( -he( a- 3&U' w"-h sha,"(g a-

25& rpm or approA"ma-ely 5 h. ?ar%es-e) cells were washe) "( wa-er resuspe()e) "( 2&& l o resh lys"(g

solu-"o( 0So)"um )uo)ecyl sulpha-e 3.3; Tr"s &.&5 M p? 12.5 "(cuba-e) or 3& m"(s a- 6&U' a() %or-eAe)

or 1& s -o shear -he $N/. $N/ was collec-e) by e-ha(ol prec"p"-a-"o( 24 a() resuspe()e) "( 6& l o &.1 W

T= buer p?.&. H(e m"crol"-er was use) as -empla-e "( P'@ reac-"o(s. 'ry a() 'y- groups -arge-e) pro-ocolsa() co(-rol s-ra"(s u-"l"De) are g"%e( "( Table4. The ab"l"-y o -empla-e prepara-"o(s -o be ampl""e) "( a P'@

reac-"o( was assesse) us"(g >ac"llus 16S r"bosomal @N/ pr"mers 0Table 4 a- #4U' or 15 sec; 55U' or 3& sec

a() 72U' or 2 m"( or 2 cycles w"-h 1.5 mM Mg'l.

$a6le 4.Pr"mers a() probes u-"l"De).

$y%ridi&ation of cry genes

Probe hybr")"Da-"o( was perorme) us"(g -he $I! ?"gh Pr"me $N/ Cabel"(g a() $e-ec-"o( S-ar-er B"- 0@oche

Molecular >"ochem"cals 'a-X 1 745 32. cr< a() c!S' as -empla-es cr!S'4E3; cr!S'4E3; cr!S'4//1; cr!S'481; cr-133; c!S'481; c!S'481; 15 >!S'4E3
http://www.biomedcentral.com/1471-2180/5/52#B20http://www.biomedcentral.com/1471-2180/5/52#B21http://www.biomedcentral.com/1471-2180/5/52#B22http://www.biomedcentral.com/1471-2180/5/52#B23http://www.biomedcentral.com/1471-2180/5/52#B24http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52#B20http://www.biomedcentral.com/1471-2180/5/52#B21http://www.biomedcentral.com/1471-2180/5/52#B22http://www.biomedcentral.com/1471-2180/5/52#B23http://www.biomedcentral.com/1471-2180/5/52#B24http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4http://www.biomedcentral.com/1471-2180/5/52/table/T4


14/19

!' sequencing

/( />I 'ycle Se:ue(c"(g >"g $ye @ea)y @eac-"o( ,"- 0'a-alogue X 43&35&& was use) accor)"(g -o -he

ma(uac-urerOs "(s-ruc-"o(s a() reac-"o(s resol%e) o( a( />I 377 $N/ Se:ue(c"(g Mach"(e. Pr"mers gral

(em0) a() gral(em0r 15 were use) -o ampl"y a() se:ue(ce -he pu-a-"%e (ema-oc")al ge(e "( >-133.

Bioassay (quick screen)

Car%ae o !odo!tera frui!erda were collec-e) rom a cor( "el) a() use) -o ge(era-e up -o 5 ge(era-"o(s o

lar%ae. The w")-h o -he hea) capsule was use) -o selec- 5-h "(s-ar lar%ae or use "( -he b"oassay 25. Car%ae

were s-ar%e) or approA"ma-ely 1 hour "mme)"a-ely pr"or -o be"(g challe(ge) w"-h o(e loopul o a bac-er"al

"sola-e collec-e) rom a sporula-e) colo(y a() smeare) o(-o a 1 cm2 lea p"ece 0orhum hali!ense "( -r"pl"ca-e.

Car%ae were le- or approA"ma-ely 1& h -he( prese(-e) w"-h a resh p"ece o u("(ocula-e) lea -he( le- or a

ur-her 1&F12 h. Scor"(g was (o- ea-e( a score o 3; par-"ally ea-e( a score o 2; comple-ely ea-e( a score o 1.

The -oA"c"-y %alue o aB. thuriniensis ca()")a-e was -he a%erage o -he scores or -he -hree -r"als wh"ch were

arb"-rar"ly class""e) as 1.& (o- -oA"c; 1.& bu- Z 2.& F u(cer-a"( -oA"c"-y; V 2.& -oA"c. /ll "sola-es "( -he

l"brary were -es-e) "rrespec-"%e o whe-her -hey were co(s")ere) clo(al s-ra"(s as well as -he "%e co(-rol

s-ra"(s ob-a"(e) rom -he >ac"llus !e(e-"c S-oc, 'e(-er.

Authors# contributions

E@ was pr"(c"pally respo(s"ble or -he )es"g( a() eAecu-"o( o -he proec- a() )a-a a(alys"s. $/ co(-r"bu-e) -o

proec- )es"g( a() )a-a a(alys"s. >o-h au-hors rea) a() appro%e) -he "(al ma(uscr"p-.

Ackno$ledgements

Le woul) l",e -o -ha(, -he 'ar"bbea( $e%elopme(- >a(, a() -he ac"llus !e(e-"c S-oc, 'e(-er or co(-rol s-ra"(s.

Re%erences

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against Insect /ests.

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3. Mar-"( P/L Tra%ers @S World%ide a6undance and distri6ution ofBacillus thuringiensis isolates.

!!l Environ +icro%iol1## 824372442.

4. Eoh(so( ' >"shop /? ( techni7ue for the effecti"e enrich!ent and isolation of Bacillus

thuringiensis.

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$ype genes ofBacillus thuringiensis isolates fro! $ai%an.

!!l Environ +icro%iol1##4 #82415242&.

6. >e( $o% = [ar"-s,y / $aha( = >ara, [ S"(a" @ Ma(asherob @ Bhamre% / Tro"-s,aya = $ub"-s,y

/ >ereD"(a N Margal"-h Y 0xtended Screening 6y /R for se"en cry group genes fro! field-

collected strains ofBacillus thuringiensis.

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PubMe) 'e(-ral ull TeA-

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Bacillus thuringiensis.

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fila!ent.

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isolated in apan.

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1&. ?as-owo S Cay >L Hhba M *aturally occurringBacillus thuringiensis in Indonesia.

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11. /ro(so( / Sporulation and delta-endotoxin synthesis inBacillus thuringiensis.

:ell +ol )ife ci2&&2 :8417425. PubMe) /bs-rac-\Publ"sher ull TeA-

12. ?o)g,"ss L )ila!entous appendages on the spores and exosporiu! of certainBacillus species. I(

!ore Research. =)"-e) by >ar,er /N !oul) !L Lol E. /ca)em"c Press Co()o( a() New Yor,;

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16/19

13. M"Du," = Hhba M Ich"ma-su T ?wa(g S? ?"guch" B Sa"-oh ? /,ao T 9ni7ue apppendages

associated %ith spores ofBacillus cereus isolates.

Basic +icro%iol1## 38333#.PubMe) /bs-rac-\ Publ"sher ull TeA-

14. Le" E[ ?ale B 'ar-a C Pla-Der = Lo(g ' a(g S' /ro"a( @9 Bacillus thuringiensis crystal

proteins that target ne!atodes.

-roc Natl cad ci *2&&3 +8276&5. PubMe) /bs-rac-\Publ"sher ull TeA-\


15. >ra%o / Sarab"a S CopeD C H(-"%eros ? /barca ' Hr-"D / Hr-"D M C"(a C 9"llalobos E Pe(a !

Nu(eD9al)eD M Sobero( M 8u"(-ero @ haracteri&ation of cry genes in a 5exicanBacillus

thuringiensis strain collection.

!!l Environ +icro%iol1## #484#654#72. PubMe) /bs-rac-\ Publ"sher ull TeA-\


16. Hhba M /"Dawa B Distri6ution ofBacillus thuringiensis in Soils of apan.

nverte%r -athol1#6 4'827722. Publ"sher ull TeA-

17. >el Y !ra(ero /lberola TM Mar-"(eDSebas-"a( ME erre E Distri6ution, fre7uency and di"ersity

ofBacillus thuringiensis in oli"e tree en"iron!ents in Spain.

Eoh(s-o(e B ?agelberg = =llar $E o!!it!ent of 6acterial spores to ger!inate.

Biochem 1#1 +:381&11&6. PubMe) /bs-rac-\ PubMe) 'e(-ral ull TeA-

2&. $eCucca /E S"mo(so( E! Carso( /$Bacillus thuringiensis distri6ution in soils of the 9nited

States.

:an +icro%iol1#1 2'8657&. PubMe) /bs-rac-

21. Sm"-h @/ 'ouche !/ $he phylloplane as a source ofBacillus thuringiensis "ariants.

!!l Environ +icro%iol1##1 '8311315.

22. Bael"( P Morel P !a)a(" Isolation ofBacillus thuringiensis fro! stored to6acco andLasioderma

serricorne ;)


17/19

!!l Environ +icro%iol1##4 #81#25.

23. Camell" rousseau @ EuareDPereD 9 ru-os @ ( =olistic

(pproach for deter!ining the 0nto!opathogenic potential ofBacillus thuringiensis strains.

!!l Environ +icro%iol1## #4847247. PubMe) /bs-rac-\ Publ"sher ull TeA-\


2#. 'ero( E Hr-"D / 8u"(-ero @ !uereca C >ra%o / Specific /R pri!ers directed to identify cry I

and cry III genes %ithin aBacillus thuringiensis strain collection.

!!l Environ +icro%iol1##5 #+8326331. PubMe) /bs-rac-\ Publ"sher ull TeA-\


3&. S"eer- EC Car"osSa(D M Na,amura CB Slepec,y @/ Paul E? Moore =@> oA != Eur-shu, P Er

/hylogeny of 5arineBacillus Isolates fro! the >ulf of 5exico.

:urr +icro%iol2&&& 4+84. PubMe) /bs-rac-\Publ"sher ull TeA-
http://www.biomedcentral.com/pubmed/5432063http://www.biomedcentral.com/pubmed/5432063http://dx.doi.org/10.1038/227680a0http://dx.doi.org/10.1038/227680a0http://www.biomedcentral.com/pubmed/9251185http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=9251185http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9251185http://www.biomedcentral.com/pubmed/9835562http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=9835562http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9835562http://www.biomedcentral.com/pubmed/8526493http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=8526493http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=8526493http://www.biomedcentral.com/pubmed/10856371http://www.biomedcentral.com/pubmed/10856371http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=10856371http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=10856371http://www.biomedcentral.com/pubmed/5432063http://dx.doi.org/10.1038/227680a0http://www.biomedcentral.com/pubmed/9251185http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=9251185http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9251185http://www.biomedcentral.com/pubmed/9835562http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=9835562http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=9835562http://www.biomedcentral.com/pubmed/8526493http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=8526493http://www.pubmedcentral.nih.gov/articlerender.fcgi?tool=pubmed&pubmedid=8526493http://www.biomedcentral.com/pubmed/10856371http://www.ncbi.nlm.nih.gov/entrez/eutils/elink.fcgi?dbfrom=pubmed&cmd=prlinks&retmode=ref&id=10856371


18/19

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Advantages and DisadvantagesPage histor( last edited b( PB$orks 5 (ears ago

(d"antages and Disad"antages

(d"antages

There are se%eral a)%a(-ages -o us"(g >. -hur"(g"e(s"s. >- o(ly ,"lls harmul "(sec-s u(l",e ma(y o-her

"(sec-"c")es.

I- w"ll (o- ,"ll be(e"c"al "(sec-s such as ho(eybees a() o-her poll"(a-ors. / seco() a)%a(-age o >- "s -ha- "-Os

(o- -oA"c -o huma(s w"l)l"e or pe-s. >ecause o -h"s "- "s use) o( ma(y oo) crops a() o-her se(s"-"%e crops

where

pes-"c")es m"gh- harm -he co(sumer or local w"l)l"e.

Disad"antages
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19/19

>. -hur"(g"e(s"s also has )"sa)%a(-ages. Ma(y s-ra"(s are suscep-"ble -o -he su(l"gh- a() -he"r po-e(cy w"ll

)egra)e

w"-h"( a wee,. Some (ewer s-ra"(s use) o( lea bee-les ca( become "(eec-"%e w"-h"( 24 hours. /lso because

>-

"(sec-"c")es are h"ghly spec""c ma(y %ar"e-"es mus- be use) " you wa(- -o ully pro-ec- your crops. They us-

wo(O-

be -ha- eec-"%e o( crops -ha- ha%e mul-"ple pes-s. >ecause >- "(sec-"c")es are o(ly useul whe( -he"r are ea-e(

some-"mes -hey are "(eec-"%e because some "(sec-s such as co)l"(g mo-h )o(O- ge- -he cha(ce -o ea- "-.

/lso ma(y >- "(sec-"c")es ha%e a shor-er shel l"e -ha( o-her sy(-he-"c "(sec-"c")es. !e(erally a-er -wo or

-hree years -he"r

eec-"%e(ess "s re)uce) grea-ly. The op-"mal s-orage co()"-"o(s or >- pro)uc-s are cool )ry a"r a() ou- o )"rec-

su(l"gh-.

/(-ho(y M. Shel-o(>"olog"cal 'o(-rol / !u")e -o Na-ural =(em"es "( Nor-h /mer"ca

!omments *0+

You )o(O- ha%e perm"ss"o( -o comme(- o( -h"s page.

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