Enzymes in Farm Animal Nutrition 2010[1]

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    ENZYMES

     

    IN 

    FARM 

    ANIMAL 

    NUTRITION

    ,2ND EDITION

    This  page intentionally left  blank 

    ENZYMES 

    IN FARM ANIMAL

    NUTRITION, 2ND EDITION

    Edited by

    Michael R. Bedford AB Vista Feed Ingredients

    Woodstock Court MarlboroughWiltshire SN8 4ANUK 

    and

    Gar G. !ar"rid#eanisco  Ani!al Nutrition

    "# Bo$  %%%Marlborough

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    Wiltshire SN8 &'NUK 

    $ABI  i% a "radi &a'e of  $AB  I&"er&a"io&al

    CABI Head Office CABI North American OfficeNosworthy Way 875 Massachusetts AvenueWain!ford 7th "oorO#fordshire O$%& 8'( Cam)rid!e* MA &+%,-./ .0A

     1e2 344 &6%4-% 8,+%%% 1e2 3% %7 ,-5 4&5"a#2 344 &6%4-% 8,,5&8 "a#2 3% %7 ,54 875(mai2 ca)i9ca)i:or! (mai2 ca)inao9ca)i:or!We)site2 www:ca)i:or!

    ; CAB Internationa +&%&: A ri!hts reserved: No

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    $o&"e&"%

    $o&"ri(*"or%

    +ii

    !reface

    i

    -. I&"rod*c"io& $*rre&" Mar/e" a&d E0ec"edDe+elo0'e&"%

    -

     A( Barletta

    2. 1la&a%e% a&d $ell*la%e% a% Feed Addi"i+e%

    -2M( "alohei!o) * (  "iironen and  *(Veh!aan+er,

    . Ma&&a&a%e, Al0ha)Galac"o%ida%e a&d !ec"i&a%e

    34M(-( *ackson

    4. 

    S"arch) a&d !ro"ei&)de#radi E&5'e%Bioche'i%"r, E&5'olo# a&d $harac"eri%"ic%Rele+a&" "o A&i'al Feed U%e

    63M(F( Isaksen) A(*( Co.ieson and K(M( Kragh

    3. !h"a%e% Bioche'i%"r, E&5'olo# a&d$harac"eri%"ic% Rele+a&" "o A&i'al Feed U%e

    78/( 0reiner and U( Koniet1n2 

    8. E9ec" of  Di#e%"i+e Trac" $o&di"io&%, Feed!roce%%i a&d Iredie&"% o& Re%0o&%e "oNS! E&5'e%

    -27

    B( S3ihus

    :. !h"a"e a&d !h"a%e

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    -8;"(( Selle) V( /a3indran) A(*( Co.ieson and M(/( Bed5ord

    6. De+elo0'e&"% i& E&5'e U%a#e i& R*'i&a&"%

    2;8K(A( Beauche!in and 6( oltshausen

    v

    vi   Contents

    7. O"her E&5'e A00lica"io&% Rele+a&" "o "he A&i'alFeed I&d*%"r

    2- A( "7ron and 0(0( "artridge

    -;. Ther'o%"a(ili" of  Feed E&5'e% a&d "heir !rac"icalA00lica"io&  i& "he Feed Mill

    247C( 0ilbert and 0( Coone2 

    --. A&al%i% of  E&5'e%, !ri&ci0le% a&d !ro(le'%De+elo0'e&"% i& E&5'e A&al%i%

    28;N( Sheehan

    -2. 

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    A&drea Barle""a* anisco Ani!al Nutrition) "# Bo$ %%%) Mar lborough)

    Wiltshire SN8 &'N) UK(?are&

     

    A. 

    Bea*che'i&* 

    6ethbridge 

    /esearch 

    Centre) 

     Agriculture

     

    and AgriFood Canada) 6ethbridge) Alberta) Canada 9&* 4B&(

    Michael R. Bedford*  AB Vista Feed Ingredients) Woods

    tock  

    Court)Marlborough) Wiltshire SN8 4AN) UK(Gre# $oo&e* anisco  Ani!al Nutrition) :;;8 S( 8th Street ) St  6ouis)

    M#

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    vii

    viii Contributors

    Alea&dre !Cro&* anisco Ani!al Nutrition) "# Bo$ %%%) Marlborough)Wiltshire SN8 &'N) UK(

     @ari !iiro&e&*  AB -n12!es #2) 92kki!,entie &) ;:;; /aa!,ki)

    Finland(el'*r*#* Ra+i&dra&*

     

    Institute 

    o5  

    Food) 

    Nutrition 

    and 

    u!an 

    ealth)Masse2  Uni3ersit2) "ri3ate Bag &&:::) "al!erston Nor th) Ne. ealand(

    Gordo& D. Ro%e&* oloAnal2sis Ser3ices 6td() 

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     1here 

    have 

    )een 

    considera)e 

    deveo

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    ix

    x Preface

    function and di!estive 

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    $rea"i al*e "hro*#h I&&o+a"io&

    "eed en?ymes he< meet consumer demand for safe* hi!h Euaity and aorda)efood: 0ince the ate %-8&s* feed en?ymes have

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    2 A. Barletta

    he< )rea= down antinutritiona factors e:!: fi)re*

    (n?ymes are cate!ori?ed accordin! to the su)strates they act u

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    $ar(ohdra%e%

    Car)ohydrases )rea= down car)ohydrates into sim

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    accordin! to the eves of resistant starch* starch !ranue si?e* starch com

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    + A. Barletta

    it is common 

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    itter was aso a common

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    has increased over time* 

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    Dri+er% for De'a&d

    "eed en?ymes im

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    ietnam:

    Ta(le -.-. orld )eat )ar&ets )illion t$ 20002003 4A5 2001 2003b$.

    20062003"ro7t

    20062003"ro7t

    200020032000

    2008

    esti)ate$ forecast$ 9$ 9$

    Bovine )eat -0.0 -,.1 -+.3 -,.1 0.* 6.,Poultry )eat --.- 30.1 3*.8 3+.8 1.1 +2.2Pi# )eat 31.1 33.6 10*.3 10-.1 2.1 1-.,5vine )eat 11.+ 1+.0 1+.2 1+.2 0., 2+.-!eat 'roduction 2**.+

     

    28+.+ 

    262.1 

    26,.- 

    1.2 

    22.+total$

    Introduction% Current !ar&et and Ex'ected (evelo')ents

    8

    Word feed voumes have risen from  Gust over %& miion t 

    in +&&& toe#ceed 7&& miion t in +&&8: o)a out

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    Because en?ymes im

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    The Re#*la"or E&+iro&'e&" *ali", Ef=cac a&d Safe"

    Most of  the maGor feed mar=ets have a re!uatory a

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    an estimated 7& of the mar=et: 1here are many other

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    (n?ymes can )e added to feed in one of two ways: One o

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    10 A. Barletta

    +&5&: Meat

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    safey and economicay )e added to onfarm fora!e to im

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    I&"rod*c"io&

     1he current !o)a feed en?yme mar=et si?e is estimated 

    to 

    )e 

    a)out.055&J&& miion*

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    en?ymes* thus )ein! a)e to reach the coon* where they wor= asU

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    Ceuose is the most a)undant )io

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    Fi#. 2.-. @ce)atic 'resentation of  cellulose structure courtesy of  te @ (e'art)ent of Ener#y "eno)e Pro#ra)s "eno)e !ana#e)ent Infor)ation @yste) "!I@$ available attt'%#eno)ics.ener#y.#ov$.

    ?ylanases and Cellulases as 4eed Additives

    1,

     Acetobacter $2linu! is a hi!hy crystaine ceuose* whereas mode su)stratesderived from )eached commercia wood

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    used: 1his

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     1he main chain of  #yan is com

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    "i!ure +:+ shows a schematic re

    Ara)ino#yans and mi#edin=ed !ucans are 

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    e#tensive research in many a

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    sou)e in water Chesson* %--56: Aso* heat treatment* such as aminarinase (C 

    ,:+:%:,-6hydroyses aminarin* 

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    of  

    @ 〈68 )arre* whereas cans HB and HC have a Gey ro

     

    structurehtt

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    famiy % entries are isted in the CAy data)ase htt

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    Ce7B and Ce45AL Haa=ana et  al(* +&&46 and 9her!oascus aurantiacusCe7A* Ce5AL Hon! et al(* +&&,a*)6:

    $ell*la%e%

    Fungal cellulases and  βglucanases

    Ony imited hydroysis or reduction of viscosity* rather than com

    (C,:+:%:-%6* endo!ucanases (C ,:+:%:46 and !ucosidases orceo)iases (C,:+:%:+%6L the atter are usuay incuded in the ceuase com

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    ceo)iohydroases fodin! to a sandwich with the e#tended oo

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    ur)ri!!en et al:* %--%L /arsson et al(* +&&+6L 9richoder!a

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    TCC 210-  @CA

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    Le@affre @afi/y) "P "L   "lucanase

    9( longibrachiatu! C

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    However* !enome seEuence annotation su!!ests ei!ht ceuase !enes infamiies H 5*  and 7* and four !enes for e#o%*,!ucanases in famiyH 5 e et al(* +&&76:

    Other fun!a ceuases used in feed incude endo!ucanases derived from

    u!icola insolens* 9alaro!2ces e!ersonii and "enicilliu! 5u

    niculosu!1a)e +:%6: ( insolens 

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    (II6* of which (I and (II were active on aminarin and therefore weredetermined as )eon!in! to cass (C ,:+:%: McCarthy et  al(* +&&,6:  1he4&:7 ='a !ucanase had o

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    on CMC /arsson et al(* +&&+6: A !ene la!&6 for a !ucanase aminarinase6of  the famiy H % (C ,:+:%:6 has )een found in 9( reesei in e#

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    han! and >ynd* +&&46: Most of  the characteri?ed Bacillus !ucanases)eon! to the famiy H %* and have hi!h %*,%*4!ucanase activity (C,:+:%:7,6 e#hi)itin! a strict su)strate s

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    structure of  this )acteria %*,%*4!ucanase and that of  the cosey reatedB( !acerans have )een soved* and they )ear some simiarity with the fun!aCe7@(I endo!ucanases /eite et al(* %--,L Hahn et al(* %--5L /eywe!tet al(* %--76:

    Anaero)ic )acteria ivin! in the di!estive tract of  ruminant

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    severa hemiceuases is needed Cou!han et  al(* %--,6:  1he sidechainceavin! UaccessoryD en?ymes remove the su)stituent !rou

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    acidic as com

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    !eneticay modified strains:  1he we=nown )acteria e#

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    two commercia )a=in! 

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    maGor #yanase activity in at east some of  the first!eneration recom)inantfeed #yanase

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    H %% and one H 56* nine from  A( 5u!igatus four H %&* three H %%and two H 76 and nine from  A( or21ae !enome four H %&* four H %%and one H 76 e et al(* +&&76: ecom)inant A( niger  #yanaseA reAn#A

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    fitration 'Vsterhft et  al(* %--76* su!!estin! that at east two dierent#yanases are aso 

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    shown to efficienty )rea= down #yan su)strates "urniss et  al(* +&&56:However* the s

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    ?ylanases and Cellulases as 4eed Additives

    **

    their activity: $yII in et al(* %---6:

    E&5'e !rod*c"io&

    $ell fac"orie%

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    Introduction

    "eed en?ymes* i=e other industria en?ymes* are currenty

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    1en! et al(* +&&76:roduction hosts can )e divided into two cate!ories )ased mainy on

    re!uatory as

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    e#

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    al(* +&&76:  1herefore* for ma#ima e#

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    detection of the si!na of the !ene of interest in )oth en?yme assays and 0'0A( anaysis:  1he use of  such strain )ac=!round aso acceerates straindeveo

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    isoate Ma* as aready su!!ested )y eary =aryoty

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    s

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    et al(* +&&46:If a

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    One of the maGor tar!ets for the feed a

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    +0 !. Paloei)o et al(

    that their s

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    Fi#. 2.8. !icrosco'ic i)a#e of  endos'er)ic and aleurone 7eat cell 7alls before a$ andafter b$ treat)ent 7it a 9richoder!a xylanase#lucanase 're'aration.

    ?ylanases and Cellulases as 4eed Additives

    +1

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    s

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    en?yme

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    [man* : and raham* H: %-876 Anaysis of  tota and insou)e mi#edin=ed %,6*%46

    '!ucans in )arey and oats:  *ournal o5   Agricultural and Food Che!istr2  ,5*7&4J7&-:

    AM"( +&&-6 >ist of en?ymes Octo)er +&&-L AM"(@&-@&8L htt

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    (#:M:* 

    'avies* 

    :P: 

    et 

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    al( +&&&6  1he $ Uthermosta)ii?in!D domains of  #yanases are car)ohydrate)indin!modues2 structure and )iochemistry of  the Clostridiu! ther!ocellu! $) domain:Bioche!istr2  ,-* 5&%,J5&+%:

    ++ !. Paloei)o et al(

    Cherry*  P:: and "idantsef* A:>: +&&,6 'irected evoution of  industria en?ymes2

     

    an 

    u

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    #yanase en?yme com

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    as 

    feed 

    additive 

    f or 

    chic=ens 

    forfattenin!* chic=ens reared for ayin!* tur=eys for fattenin!* tur=eys reared for)reedin! and

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    >:M: 

    %---6 

    Homoo!ous 

    #yanases 

    from 

    Clostridiu! 

    ther!ocellu!2 

    evidence 

    for)ifunctiona

     

    activity* 

    syner!ism 

    )etween 

    #yanase 

    cataytic 

    modues 

    and 

    the 

    :* 'ean* :* 'rener* 0:* 'unnCoeman*

     

    N:0:*

    +- !. Paloei)o et al(

    oede!e)uur* 

    ":* 

    Houfe=* 

     1:':* 

    (n!and* 

    :P:* 

    /ey* 

    A:0:* 

    Meerman* 

    H:P:* 

    Mitche* 

     1:*Mitchinson* C:* Oivares* H:A:*  1eunissen* :P:M:*  Yao*  P : and Ward* M: +&&,6

     1ranscri

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    +&&86 

    roteomics 

    anaysis 

    of  

    Uova)ioS 

    (#ceD* 

    secreted 

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    Hon!* 

     P:* 

     1ama=i* 

    H:* 

     Yamamoto* 

    /: 

    and 

    /uma!ai* 

    H: 

    +&&,)6 

    Conin! 

    of  

    !ene

     

    encodin!thermosta)e ceo)iohydroase from 9her!oascus aurantiacus and its e#eeuwen* 

    M:P:":* 

    Wood* 

     1:M: 

    and 

    ora!en* 

    A::P:

     

    %--,6urification and characteri?ation of three endo%*46#yanases and one #yosidase from

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     As+ergillus a.a!ori: *ournal o5 Biotechnolog2  +7* ,4-J,5:/uhs*

     

    /: * 

    >iec=fedt* 

    (:* 

    0amues* 

    :P:* 

    /ovacs* 

    W:* 

    Meyer* 

    W:* 

    etrini* 

    O: 

    et  al: %--6Moecuar

     

    evidence 

    that 

    the 

    ase#ua 

    industria 

    fun!us 9richoder!a rees

    ei is a conaderivative

     

    of  

    the 

    ascomycete 2+ocrea  ecorina: "roceedings o5  the Natio

    nal  Acade!2 o5 Sciences USA -,* 7755J77&:

    /u=arni* 

    N:* 

    0hendye* 

    A: 

    and 

    ao* 

    M: 

    %---6 

    Moecuar 

    and 

    )iotechnoo!i

    ca 

    asesa!e* 

    and 

    Bussey* 

    H: 

    +&&6 

    Ce 

    wa 

    assem)y 

    in Saccharo!

     2ces cere3isiae:Microbiolog2 and Molecular Biolog2 /e3ie.s 7&* ,%7J,4,:

    >evasseur* 

    A:* 

    Asther* 

    M: 

    and 

    ecord* 

    (: 

    +&&56 

    Overin* P:* Ndovu* >:M:* 0in!h* 0: and iay* B: %---6 urification and )iochemica characteristics

    of   '#yanase from a thermoinder* M:B:* Mattinen* M:>:* /onttei* M:* >inde)er!* :* 0t]h)er!* P:* 'ra=en)er!* 1: et al:

    %--56 Identification of functionay im

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    )y car)on sources and metas: *ournal o5 Bacteriolog2  7,* +-J+78:MZntyZ* A:>:* aoheimo* M: and 0uominen* : %--86 Industria mutants and recom)inant

    strains of  9richoder!a reesei: In2 Harman* :(: and /u)ice=* C:: eds6 9richoder!a and0liocladiu!E -n12!es) Biological Control and Co!!ercial  A++lications: 

     1ayor and"rancis* >ondon* : 

    and 

    Nevaainen*/:M:H:

     

    %--+6 

    (ectroie)* 

    W: 

    +&&&6 

     1he 

    thermosta)ii?in! 

    domain 

    of  

    themoduar

     

    #yanase 

    $ynA 

    of  9her!otoga !ariti!a re

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    Moers* 

    /: * 

    Ceus* 

    I:* 

    BriGs* 

    /:* 

    Courtin* 

    C:M: 

    and 

    'ecour* 

     P:A: 

    +&&56 

    (ndo#yanase

     

    su)strateseectivity

     

    determines 

    de!radation 

    of  

    wheat 

    watere#tracta)e 

    and 

    waterune#tracta)eara)ino#yan: Carboh2drate /esearch ,4&* %,%-J%,+7:

    Moers* 

    /:* 

    Bour!ois* 

     1:* 

    om)outs* 

    0:* 

    Bein* 

     1:* 

    an 

    Cam: 

    and 

     1aimisto* 

    /: 

    %--46 

    On 

    the 

    safety 

    of  9richoder!a

    reesei: *ournal o5 Biotechnolog2  ,7* %-,J+&&:Niehaus* ":* Bertodo* C:* /Zher* M: and Antrani=ian* : %---6 (#tremo

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    e* H:P:* de Winde*  P:H: * Archer* ':B:* 'yer* :0:* Hofmann* :* 0chaa

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    0chamey* 

    M:* 

    0in!h* 

    A: 

    and 

    Ward* 

    O:: 

    +&&46 

    'eveo

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    ceuosic ethano: Nature /e3ie.s 0enetics -* 4,,J44,:0toner* M::* 'ae* ':A:* uafetti* :P:* Bec=er*  1:* Mannin!* M:C:* Caruo* 

     P: 

    and 

    Wan!* 

     P : 

    +&&76 

    Codon 

    oeuven* Be!ium*

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    eith* 

    B:* 

    Her?)er!* 

    C:* 

    0tec=e* 

    0:* 

    "eesche* 

     P:* 

    Maurer* 

    /:H:* 

    (hrenreich* 

    : et  al( +&&46

     1he com

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    Mannans 

    occur 

    in 

    the 

    forms 

    of  

    !ucomannan* 

    !aactomannan* !uco!aactomannan and !ucuronomannans in nonstarch

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    the coee e#tract dry wei!ht and mannans are a)undant* ma=in! the e#tracthi!hy viscous: Addition of mannanase faciitates

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    mannan content of  a ar!e num)er of  soy)ean mea sam

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    a:siao et al. 200-$.

    Mode of  ac"io&

     1he mode of action of mannanase in mono!astric animas iscom

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    demonstrated to stimuate the innate immune system* resutin! inmacro

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    raded eves of  mannanase were added to mai?eJsoy)ean meaty

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    In the a)sence of !uar !um* mannanase im

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    a!annanase 7as eiter added $ or not added $ to te feed ata rate of  100 )illion units t 1.be!eans in colu)ns 7itout a co))on su'erscri't differsi#nificantly " O0.0,$.

    Ta(le .4. Te effect of  )annanase at #raded levels onbroiler cic& 'erfor)ance to 21 days in diets containin# 19#uar #u) fro) (as&iran et al.  200+$.

    !annanase ! t 1$a B #$ 4CR # # 1$

    0 *+-., 1.**-b

    100 *+-.3 1.*0+c

    200 *+6.1 1.231d

    *00 *+,., 1.26-d

    B body 7ei#tF 4CR feed conversion rate.a! N 10- en/y)e activity units.bd!eans 7itout a co))on su'erscri't differ si#nificantly" O0.0,$.

    the 

    first 

    e#

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    -0 !.E. ;ac&son

    Ta(le .3. Effect of  infectiona )annanase en/y)eb and )edicationc on broiler cic&'erfor)ance fro) 6 to 21 days of  a#e fro) ;ac&son et al. 200*a$.

    Lesion score

    day 1+$d

    Infection En/y)e 

    !edication 

    "ain #$ 4CR # # 1$ 

    !ortality 9$

    ''er 

    Lo7er

          P   ,+0e 1.++-# 0.00f  0.00# 0.00#

         P P   ,+6e 1.+2+# 1.86f  0.00# 0.00#

    P   +23 1.80+e 3.86e 1.*6e 1.,-e

    P P   +30# 1.,*-f  *.8,ef  1.1-f  1.++e

    P       P   ,22f  1.++8# 0.63f  1.0*f  0.66f 

    4CR feed conversion rate.a5rally inoculated 7it a )ixed solution 7it a''roxi)ately 80000 oocysts of  -i!eria acer3ulina and12,0 oocysts of  -i!eria !a$i!a 'er bird on day 8. 5n days 11 12 and 1* birds 7ere #iven brotcultures of  Clostridiu! +er5ringens containin# a''roxi)ately 1., Q 106 cfu 'er bird.b!annanase 7as eiter added $ or not added $ to te feed at a rate of  100 )illion units t 1.c@alino)ycin @ALF -0 # t 1$ 'lus bacitracin )etylene disalicylate B!(F ,0 # t 1$.d''er intestine -( acer3ulinaF lo7er intestine -( !a$i!a.e!eans 7itin colu)ns not sarin# co))on su'erscri'ts are si#nificantly different " O0.0,$.

    -ects on intestinal !or+holog2 and 5unction

     1he eect of  mannanase on intestina function and mor

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    en?yme addition:  1he authors commented that reduced !o)et ce num)ersmay )e e#

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    b!annanase 7as eiter added $ or not added $ to te feed at a rate of  100 )illion units t 1.c@alino)ycin @ALF -0 # t 1$ and bacitracin )etylene disalicylate B!(F ,0 # t 1$.d''er intestine -( acer3ulinaF lo7er intestine -( !a$i!a.ei!eans 7itin colu)ns not sarin# co))on su'erscri'ts are si#nificantly different" O0.0,$.

    authors s

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    In addition* the consistency of carcasses or

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    !annanase 〈"alactosidase and Pectinase

    -*

    )ut at 7& days of a!e a si!nificant en?yme j soy)ean mea source interaction

    was o)served: Mannanase si!nificanty increased ive wei!ht )ya

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    Ta(le 

    .7. 

    Effect of   )annanase on tur&ey to) 'erfor)ancea fro) 5detalla et al .  2002$.

    Body 7ei#t 4CR # # 1@B! 9 'rotein$ En/y)eb  day 12-$ 012- days$

    +6 1+.31 2.80++6   P   1,.0* 2.-**

    ++ 1+.+0 2.83+

    ++   P   1+.88 2.-3,

    Avera#e 1+.-- 2.8+3

    Avera#e   P   1+.30 2.--+

    " value for en/y)e 0.001 0.001" value for @B! Q en/y)e 0.068 0.2+1

    @B! soybean )ealF 4CR feed conversion rate.aT7o ex'eri)ents 'ooled eac ex'eri)ent 7it 18 birds 'er 'en seven 'ens 'er treat)ent.b!annanase 7as eiter added $ or not added $ to te feed at a rate of  100 )illion units t 1.

    -+ !.E. ;ac&son

    with 

    mannanase 

    and 

    su!!est 

    that 

    these 

    eects 

    may 

    )e 

    ar!er 

    when 

    ower

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    +&&,)6 re

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    %&:57 to 7:4& " &:&&%6 at * %+ and %8 wee=s of a!e* resayin! hens are e#

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    :i# 81.+3 68.02 60.2* 8+.8,

    (ifference 9$ 1.*6 0.31 0.32 0.00-n12!eb

      80.8, 6-.21 83.2* 8+.00

    P   80.6- 6-.31 60.*0 8,.,0

    (ifference 9$ 0.11 0.80 1.08 1.,0P statistics

    !E level 0.031 0.001 0.001 0.366

    En/y)e 0.66, 0.008 0.001 0.001!E ⋅ en/y)e 0.+23 0.**1 0.,+0 0.,88

    aLo7 !E level is 100 &cal  1 lo7er tan i# !E level.b!annanase 7as eiter added $ or not added $ to te feed at a rate of  100)illion units t 1.

    " &:&56 with the owest amino acid density tested &:7& ysine6: 1his su!!eststhat mannanase may )e instrumenta in im

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    with resuts

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    &:%6 to ,:& " &:&6* as we as a ar!e 86 im

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    A ar!e num)er of  studies have )een re

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    〈aactosidase is a !ycoside hydroase en?yme that hydroyses the termina〈'!aactose moiety from !aactoside oi!osaccharides*

    !yco

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    ener!y contri)ution is imited: (ary studies to determine the M( of  common

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    @ucrose -6

    @tacyose +,Raf finose 12

    Another 

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    on the a

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    A series of  )roier e#

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    res

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    S*''ar

    A

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    are most

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    anima s

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    acid and crude

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    varyin! in cacium eves* ya et  al( +&&&a6 re

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    with #yanase* !ucanase* ceuase* mannanase and !aactanase en?ymes* tono )enefit at a when

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    !annanase 〈"alactosidase and Pectinase

    83

    Refere&ce%

    Adi)moradi* M: and Mehri* M: +&&76 (ects of mannanase on )roier:* 

    A=avanichan* 

    O: 

    and 

    Chen!* 

     1:/: 

    %--&6 

    (ect 

    of  

    oi!osaccharide

    free 

    0BM 

    on 

    true 

    meta)oi?a)e 

    ener!y 

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    in 

    adut 

    roosters:

     "oultr2 Science -* 787J7-,:

    Couch* P::* Ba=ashi* Y:/:* "er!uson* 1:M:* 0mith (:B: and re!er* C:: %-76 1he eect of 

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    'as=iran* 

    M:* 

     1eeter* 

    ::* 

    "od!e* 

    ': 

    and 

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    H:.: 

    +&&46 

    An 

    evauation

     

    of  

    endo'mannanase Hemice6 eects on )roier 

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     Pac=son* M:(: and Mathis* :": +&&6 (conomics of mannanase HemiceQfeed en?yme6 in

    tur=ey hens under varyin! ener!y eves: "oultr2 Science 85 0u: and Mathis* :": +&&4a6 (ects of 

    :

    +&&4c6 

    (ect 

    of  

    mannanase 

    HemiceQ6 

    in 

    commercia 

    )roiers 

    with 

    two 

    cacium 

    and: 

    and 

    Mathis* 

    :": 

    +&&56

     

    (ect 

    of mannanase HemiceQ6 on

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    !annanase 〈"alactosidase and Pectinase

    61

     Pac=son* M:(:* reenwood* M:W:* 0te: et  al:

    +&&%)6 

    〈aactosidase 

    en?yme 

    su

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    ODuinn* 

    ::* 

    "under)ur=e* 

    ':W:* 

    "under)ur=e* 

    C:>: 

    and 

     Pames* 

    :>: 

    +&&+6 

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    of dietary sui* ':* iao* $:* han!* >: and uo* >: +&&+6 (ect of dietary su:A:* Carter* 0:':* 0enne* B:W: and 0hriver* P:A: +&&+6 (ects of

    mannanase additionto cornJsoy)ean mea diets on !rowth

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    hemiceuase: "oultr2 Science 87* 7%,J7%8: 1or=i* M: and Che!eni* A: +&&76 (vauation of dietary re: 

    and 

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     P: 

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    aactomannans 

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    Wi!!ins* H:0: %-846 Nutritiona vaue of su!ars and reated com

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    of soy)ean mea* soy

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    materias and insects: However* in order to satisfy consumer <reference forUve!etarianD anima

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    0tarch consists of  two 

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    even when the amount of  resistant starch com

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    rotein consists of

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     1he

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    was re

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    there were dierences )etween 

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     1he com

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    that the vast maGority of  endo!enous nitro!en is recovered )y the terminaieum* and even more on a totatract )asis _ 8&6* athou!h the authorsconcede that nitro!en recovered in the ar!e intestine is of imitedimmediatevaue to the anima: Nevertheess* it is 

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    en?ymes2 structura anaysis of the residua amyo

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    (n!yst* H:N:* /in!man* 0:M: and Cummin!s* P:H: %--+6 Cassification and measurement of 

    3+ !.4. Isa&sen et al(

    nutritionay im

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    soy)ean mea on !rowth andmana* 

    W:P:M: 

    +&&-6

     

    'ietary

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    Science and 9echnolog2  %++* ,&,J,+&: 1ester*

     

    :":* 

    /ar=aas* 

     P: 

    and 

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    $: 

    +&&46 

    0tarch 

    structure 

    and 

    di!esti)iity 

    en?ymeJsu)strate

    reationshi

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    reduced to dierent de!rees at the su)strate concentration of the assay:

     1o date* ony one a=aine

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    ne!ativey char!ed ceft into a favoura)e environment for the )indin! of 

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     1his en?yme does not need any cofactor for en?ymatic activity: 1he

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    the ran!e of %5& miion Haefner et al(* +&&56: (ven if

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    di!esti)iity eter and Ba=er* +&&%6* others have found im

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     1he a)iity of a

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    Ac"i+i" 0ro=le a&d %"a(ili" of  0<

    With the e#ce

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    lla and "ibson 1368$F yss

    et al. 1333a$F "reiner etal. 2003$yss et al. 1333a$yss et al. 1333a$F Rodri#ue/ et al. 2000$

    Ber&a et al. 1336$Tsen# et al. 2000$

    Lassen et al. 2001$F lla and

    @etu)adavan 200*$

    uan etal. 2002$Ra#on etal. 2006$

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    :& reiner et  al(* %--86* whereas that from  A( niger  %%15,A- showedma#imum

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    20

    01.0 2.0 3.0 4.0 5.0 6.0 7.0

    8.0

    pH

    Pytases

    10,

    (b)

    140

    120

    100

    80

    60

    40

    20

    01.0 2.0 3.0 4.0 5.0 6.0 7.0

    8.0

    pH

    (c)

    200

    180

    160

    140

    120

    100

    80

    60

    40

    20

    01.0 2.0 3.0 4.0 5.0 6.0 7.0

    8.0

    pH

         A    c    t     i    v     i    t    y    r    e     l    a    t     i    v    e    t    o    p

         H     5 .     5     (     %     )

         A    c    t     i    v     i    t    y    r    e     l    a    t     i    v    e    t    o    p     H     3 .     5     (     %

         )

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    Fi#. 3.-. a$ ': activity 'rofiles of  'ytases fro) rye  $ "reiner et al( 1336$  As+ergillus niger 11T,*A3  $ "reiner et al( 2003$ and a !alaysian 7aste7ater bacteriu)  $ "reiner and4arou& 2008$ usin# sodiu) 'ytate as a substrate at *8SC. Te activity at o'ti)al ': 7as ta&enas 1009. Buffers% ': 1.0*., #lycine:ClF ': *.,-.0 sodiu) acetate

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    activity 

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    et  al(* +&&7L reiner and "arou=* +&&7L Huan! et  al(* +&&86:  1he )acteria

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    hi!her

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    fun!a or

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    terreus %%J+, ML Wyss et al(* %---a6* A( 5u!igatus %& ML asamontes et al(* %--7)L Wyss et  al(* %---aL odri!ue? et  al(* +&&&6* Sch.annio!2cescastellii ,8 ML 0e!ueiha et  al(* +&&+6* Klebsiella aerogenes + ML 1am)e et  al(* %--46* catDs tai 

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    initiatin! 46 assen et al(* +&&%6:

     1o )rin! some carification to )iochemica 

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    to )e 

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    Fi#. 3.2. Classification of  'ytases based on te carbon in te !2oinositol rin# of  'ytate at7ic de'os'orylation is initiated   'os'ate residue$.

    112 R. "reiner and . oniet/ny

    Fi#. 3.. !aDor 'ytate de#radation 'at7ays for te four classes of  'ytase fro) :aya&a7a et al( 1330F "reiner et al( 2000a 2001a 2002 2008ab 2003F 

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    )een found in nature to date: 1hus* screenin! nature for

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    intraceuar 

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    we deveo

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    to im

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    of  -( coli 

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    sta)e in aEueous soutions at 7&`C and in feed at 

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    45 of  its initia activity: >ast )ut not east* the 3 ma# vaues at 

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    116 R. "reiner and . oniet/ny

    of a

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    )ioefficacy of  su

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    Andid* 

     1:A:* 

    eide* 

     P: 

    and 

    0and)er!* 

    A:0: 

    +&&46 

    Meta)oism 

    of  

    e#traceuar

     

    inositohe#aei* 

    $:: 

    and 

    Ba=er* 

    ':H: 

    +&&,6 

    (fficacy 

    of  

    an -( coli

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    from Bacillus s

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    and 

    wheat 

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    oovan* 0::* Meidin!er* ::* AGa=aiye* A:* Cottri* M:* Wieder=ehr* M::* Barney* ':P: et  al:

    +&&%6 i!s e#

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    nove* thermosta)e 

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    tota ee*  P:/:* /im* H:/:*  Yu*  P:H: and Oh*  1:/: %--8)6 Conin! of  the

    thermosta)e 

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    ooch* 

    C:* 

    iarou!e* 

    C:* 

    Wyss* 

    M: 

    and 

    van 

    >oon 

    A:::M:+&&&6 (#chan!in! the active site )etween : et  al: +&&+6

     1he consensus conce

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    >indEvist* 

     Y:* 

    0chneider* 

    and 

    ih=o* 

    %--46 

    Crysta 

    structures 

    of  

    rat 

    acid 

    ei* $::* 0ethumadhavan* /:  et al( +&&+60itedirected

     

    muta!enesis 

    of   As+ergillus niger  N> ,%,5 

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    12,

    Nair* :C:* >aFamme* P: and 'uvnGa=* : %--%6 roduction of oon* A:::M:

    %--7a6 

    Conin! 

    of  

    the 

    oon A:::M: %--7)6 ene conin!*

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    intestina :B: 

    +&&8a6 

    /inetics* 

    su)strate 

    s

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    asmussen* 0:/:* Pohansen* /:0: and 0rensen* M:B: +&&76 oynuceotides encodin!

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    Pytases

    128

     1am)e* 0:M:* /a=iG* :0:* /e=ar* 0:M: and are=h* >:P: %--46 1wo distinct moecuar forms

    of eun!* 

     1:Y: 

    and 

    >im* 

    B:>: 

    +&&+6 

    Moecuar 

    conin! 

    and 

    the 

    )iochemicacharacteri?ation of two nove

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    de

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     Yan=e* 

    >:P:* 

    0ein!er* 

    B:>: 

    and 

    Chen!* 

    /:P: 

    %---6 

    hytase 

    activity 

    in 

    Seleno!onasru!inantiu!2 a : %--56 Com

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    8   E9ec" of  Di#e%"i+e Trac"$o&di"io&%, Feed !roce%%ia&d Iredie&"% o& Re%0o&%e

    "o NS!

     E&5'e%

    B. @=I:@

    I&"rod*c"io&

    Nonstarch 

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    ar!e inFuence on the resuts* )ut which wi not )e deat with here:

     1he to

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    al( +&&86 showed that* )etween

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    tem

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    dierent

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    Fi#. 8.-. Cu)ulative excretion rates for broiler cic&ens fed 7eat diets su''le)ented 7itoat ulls and 7itout su''le)entation. Bars indicate standard deviation n N +$. 4ro):etland and @vius 2001.$

    acce

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    %--+6 showed that )irds ada

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    Fi#. 8.2. 

    (ry )atter 'ercenta#e #rey sMuares$ and content asteris&s$ in cro' of  )ealfed20dayold broiler cic&ens at different ti)es after avin# ad access to feed for 1, )in.

    0vihus et al( +&&+6* where 5& of the feed had

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    ++& min )y 'Znic=e et  al( %---6* whie others found retention time in the GeGunum and ieum to vary )etween %, and +& min 0hires et al(* %-87L vander /is et al(* %--&L utierre? de Aamo et al(* +&&-a*)6: 1his a

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    associated with a considera)e fermentation activity dominated )y actic acid

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    res

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    re

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    1*6 B. @vius

    activity of the !ucanases and #yanases added to the diet*athou!h the

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    where surviva durin!

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    retention time throu!h increased voume of  the !i??ard: However* the moreacidic conditions created in the !i??ard due to increased dietary structure woudreduce the activity of the en?ymes in the !i??ard and woud aso increase theris= of their inactivation* )oth throu!h ow

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    at a constant rate* imitin! access to feed wi therefore increase !astric retentiontime: 1his is in accordance with findin!s of a

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    the stomach: 1o concude* an avera!e

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    a

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    It is worth notin! that Mor!avi et  al( +&&%6 concuded that N0ases wereony modesty susce

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    and thus enhance de!radation activity of added en?ymes: "or

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    for a on! retention time in the cro

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    ma=in! it difficut to ascri)e eects such as an increased starch anea et al(*%---L racia et  al:* +&&,6 and@or 

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    has )een shown that sou)e fi)res may aso have ne!ative eects throu!hstimuation of )acteria

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    b=alues are calculated for te 7eat fraction only.cAvera#e value for all batces tested.

    or e#truded diets ranGes et al(* %--L 0cott et al(* +&&,* res

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    discussed )y Cowieson et  al( +&&6: Aternativey* nutritive vaue may )ecom

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     1he !enera mechanisms have )een e#tensivey reviewed )y Adams %--%6

    and >udi=huy?e et  al( +&&,6* and wi )e ony )rieFy outined here: Watermoecues interact with the en?yme throu!h noncovaent van derWaas )onds*and may even contri)ute to conformationa sta)iity )y formin! a 

    mem)rane

    around 

    the 

    en?yme 

    Adams* 

    %--%6: 

    As 

    tem

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    %--46 and 0iversides and Bedford %---6 o)served reduction in en?ymeactivity when the 

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    fi)res does not increase with 

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    1,0 B. @vius

    at 85`C: As aready discussed* however* the window of  time in the di!estivetract where conditions are suita)e for en?yme activity is short: 1hus* a

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    e#o!enous en?ymes* it is reasona)e to assume that even a modestim

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    Bayer* 

    :C:* 

    Hoover* 

    W:H: 

    and 

    Muir* 

    ":: 

    %-786 

    'ietary 

    fi)er 

    and 

    mea 

    feedin! 

    inFuence 

    on)roier !rowth and cro< fermentation: "oultr2 Science 57* %45J%45-:

    1,2 B. @vius

    Bedford* 

    M:: 

    +&&+6 

     1he 

    roe 

    of  

    car)ohydrases 

    in 

    feedstu  

    di!estion: 

    In2 

    McNa)*

     

     P:M: 

    andBoorman*

     

    /:N: 

    eds6 "oultr2  FeedstusE Su++l2) Co!+osition and Nut 

    riti3e Value:CABI u)ishin!* Wain!ford* ./*

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    of ow a

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    iamide* 

    M:P: 

    +&&-a6 

    Wheat 

    starch 

    di!estion 

    rate 

    in 

    )roier 

    chic=ens 

    is 

    aected 

    )ycutivar )ut not )y wheat cro< nitro!en fertiisation: British "oultr2 Science 5&* ,4%J,4-:

    utierre? 

    de 

    Aamo* 

    A:* 

    erste!en* 

    M:W:A:* 

    den 

    Harto!* 

    >:A:* 

    ere? 

    de 

    Ayaa* 

    andiamide*

     

    M:P: 

    +&&-)6 

    Wheat 

    starch 

    di!estion 

    rate 

    in 

    )roier 

    chic=ens 

    is 

    aected 

    )ycutivar )ut not )y wheat cro< nitro!en fertiisation: "oultr2 Science 88* %

    J%75:Hetand*

     

    H: 

    and 

    0vihus* 

    B: 

    +&&%6 

    (ect 

    of  

    oat 

    hus 

    on 

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    In)orr* 

     P: 

    and 

    Bedford* 

    M:: 

    %--46 

    0ta)iity 

    of  

    feed 

    en?ymes 

    to 

    steam 

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    =inetic information to ar)ier* 

    M: 

    and 

    Nitsan* 

    %--56 

    (ect 

    of  

    a!e 

    and 

    e#o!enous 

    amyaseand

     

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    Mi==esen* >:>:* Nau!hton* :P:* Hedemann* M:0: and Pensen* B:B: +&&46 (ects of :* ode* >:M:* McAister* 1:A:* Iwaasa* A:': et 

    al: +&&%6 esistance of feed en?ymes to

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    ersia* 

    M:(:* 

    'ehority* 

    B:A 

    and 

    >i)urn* 

    M:0: 

    +&&+6 

     1he 

    eects 

    of  

    en?yme 

    su: 

    and 

    Banave* 

    ': 

    %-876 

     1he 

    di!estion

     

    of  

    wheat

    starch in )roier chic=ens: Australian *ournal o5 Agricultural /esearch ,8* ,-J4-:0amarasin!he*

     

    /:* 

    Messi=ommer* 

    and 

    Wen=* 

    C: 

    +&&&6 

    Activity 

    of  

    su

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    hysica and nutritiona eects of

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    ranGes* M::* 0i* ': +&&56 roa?arevic* N:* (dvardsen* ':H: and 0vihus* B: +&&86(ects of fi)re

    content in

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    I&"rod*c"io&

    A century a!o*

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    %--6: 1his was mainy in res

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    )etter scientific understandin! of  the 

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    Fi#. :.-. @ce)atic dia#ra) of  te !#'ytate axis as 'ro'osed by Lott et al. 2000$.

    Pytate and Pytase

    1-*

    from nine surveys are summari?ed in 1a)e 7:%: 1he maGority of in feedstusof

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     1he e#cretion of  e#cess and undi!ested  )y 

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    1.3- 1.6-2.20$ -1.0 *0.0!ai/e 8+, 2.-2 2.*0 

    2.30$1.66 1.802.20$ 81.- 1*.0

    @or#u) --+ *.+2 2.81 *.60$

    2.-- 1.30*.2-$ 88.6 20.0

    eat -38 *.08 2.30 +.03$

    2.13 1.602.63$ 81.- +3.0

    Canola )eal +26 3.82 6.83 11.,0$

    -.+, +.008.86$ --.+ 21.0

    Cottonseed )eal

    *21 10.02 -.+0 11.*-$

    8.82 +.33.11$ 88.1 1.0

    @oybean )eal

    -63 -.+3 ,.80 -.3+$

    *.66 *.,++.,*$ ,3.3 28.0

    Rice bran -*8 18.62 1*.+0 28.13$

    1+.18 8.30 2+.20$

    83., 2,.0

    eat bran -2, 10.3- 6.02 1*.81$

    6.*- 8.003.-0$ 8-.* +1.0

    aei#ted )eans derived fro) 

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    hytate e#traction from 

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    is an interestin! a

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    de

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    Moreover* the 

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     1he main sites of

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    e#am

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    of

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    an avera!e of  +:&& ! non

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    mea* the su)seEuent eEuivaency of && "1. of an As+ergillus 

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    studies: 1he i=eihood is that the Ca eEuivaence of  

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    18*

    Relea%e of  0ho%0hor*% a&d calci*'

    0evera e#

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    standard and eevated incusion eves merit more accurate definitions:

    K!ro"ei& E9ec"H of  !h"a"e a&d !h"a%e

    Officer and Batterham %--+a6 were

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    commercia entities for neary two decades* it is not acce

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    Pytate and Pytase

    18,

    increased avera!e AI' coefficients of %8 amino acids )y 4:7* from &:787 to

    &:8+4* 

    over 

    %+, 

    o)servations 

    1a)e 

    7:,6: 

    Amon! 

    individua 

    amino 

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    ar!ey undi!ested in the sma intestine* any increase in mucin secretion woude#acer)ate Fows of  endo!enous amino acids derived from its 

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    )e reduced si!nificanty throu!h the addition of  increasin! eves of  Na* witheves eEuivaent to those found in a &:+ Na diet )ein! sufficientto )rea= u<more than 5 of these com

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    It is noteworthy that chromic o#ide has )een used as the mar=er in

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    a)e to dissociate !ycininJ

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    Interestin!y* A!)ede et  al: +&&-6 su)seEuenty determined the eects of  "(l2cii 

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    of !i??ard function )y feedin! whoe or coarsey !round !rain in turnstimuatesthe deveo

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    done with care: In a recent study Cowieson* +&%&6* the remova of + soy oifrom a mai?e soy)ased )roier diet resuted in a si!nificantdecrease b,6 iniea amino acid di!esti)iity at day +%: Interestin!y* this eect wasnot o)served)y day 4+ chan!e from C to NC k a

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     *ournal o5 Nutrition 4-* 4&%J4%&(han!*

     

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    (#o!enous en?ymes have the a)iity to increase en?ymic activity within therumina environment Mor!avi et  al:* +&&&)L Coom)atto et  al:* +&&,c6:Increased hydroytic ca

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    Le7is et al. 1333$ )l en/y)e 'roduct    1 fora#e (!$b

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    Fi#. 6.-. Effects of  addin# a fibrolytic en/y)e additive to alfalfa cubes fed to #ro7in# beef cattle Beauce)in et al.  133,$.

     

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    ?e ac"i+i"ie% reJ*ired i& feed e&5'e addi"i+e%

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    on a 'M )asis6* N'" di!esti)iity in the tota tract increased )y +L whenadded to a hi!hfora!e diet 44:5 )arey sia!e* %:& afafa hayand ,-:5concentrate on a 'M )asis6* there was no eect on N'" di!esti)iity: >ac= of eect of  the en?yme in the hi!herfora!e diet may have reFected the hi!her

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    dairy cows in eary actation is the )est way to assess whether an en?yme

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    #yanases: 

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    varia)iity and

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    /incaid* :>:* ari=i: 

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    %---6 (ect of directfed fi)roytic en?ymes on the actationa iEuid* 

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    fue sources* such as )iodiese and ethano* has created mutiand 

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    su)strates for )ioethano 

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    more 

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    com9$FA!E% si#nificant i)'rove)ents " O0.01$at 10 days 89$ and 21 days -9$F )ai/e 

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