Nutritional Value and Physiological Effects of D-Xylose and L-Arabinose in Poultry and Pigs

NUTRITIONAL VALUE AND PHYSIOLOGICAL
EFFECTS OF D-XYLOSE AND L-ARABINOSE

IN POULTRY AND PIGS
; LANDBOUWCATALOGUS
0000 0456 7893

Promotoren : dr.ir. M.W.A. Verstegen,
buitengewoon hoogleraar ophet vakgebied van de veevoeding
in het bijzonder de voeding van de eenmagigen.
dr.ir. S. Tamminga,
buitengewoon hoogleraar ophet vakgebied van de veevoeding
in het bijzonder de voeding van herkauwers.
Co-promotor: dr.ir. E.J. van Weerden,

voormalighoofd van het TNO-instituut voor diervoeding en
fysiologie (ILOB)te Wageningen.
B1BL10THEEK:
fAMDBOUT!7UN"'ERSITEH
pôliP^1^
J.B. Schutte
NUTRITIONAL VALUE AND PHYSIOLOGICAL

EFFECTS OF D-XYLOSE AND L-ARABINOSE
IN POULTRY AND PIGS
PROEFSCHRIFT
ter verkrijging van de graad van
doctor in de landbouw- en milieuwetenschappen,
op gezag van de rector magnificus,
dr. H.C. van der Plas,
in het openbaar te verdedigen
opmaandag 16december 1991
des namiddags te vier uur in de aula
van de Landbouwuniversiteit te Wageningen.
t5rt ,55/e^
Omslag: Henk Stappers
Schutte, J.B., 1991.Nutritional value and physiological effects ofD-xylose and

L-arabinose in poultry and pigs.(Nutritionele waarde en fysiologische effecten
van D-xylose en L-arabinose bij pluimvee en varkens). The main sugars other
thanglucose,whichwillbereleasedfromahydrolysisofnonstarchpolysaccharides,
are D-xyloseand L-arabinose. The results ofstudies on the fate ofboth pentose
sugars after oraladministration inpoultry and pigsarepresented. Digestibility
ofL-arabinoseattheendoftheileumwasfoundtobelowerthanthatof D-xylose.
The presence of D-xylose or L-arabinose in the diet increased the ileal flow of
volatilefatty acidsinpigs,suggestingmicrobialdegradationofbothsugarsinthe
small intestine. Both pentose sugars were partly excreted in the urine. The
extent of this urinary excretion, as a percentage of intake, increased as the
dietary inclusion of either of the two sugars was increased. At equal dietary
levels,morexylosethanarabinosewasexcretedintheurine.FeedingofD-xylose
andL-arabinosetopigsandpoultrycausedsomephysiologicalchanges.Inchicks
these changes resulted in wet droppings. The net energy value ofboth pentose
sugars was calculated to be approximately 25 to 35% of that of D-glucose.
Doctoral Thesis,WageningenAgricultural University. TNO-Institute ofAnimal
Nutrition and Physiology (ILOB). P.O. Box 15, 6700 AA Wageningen, The
Netherlands.
The studies described in this thesis were carried out at the TNO Institute for
AnimalNutritionandPhysiology(ILOB),Wageningen,andfinancially supported
byTNO Nutrition and Food Research, Zeist, The Netherlands.
STELLINGEN
1. Het feit dat varkens en pluimvee niet over een enzymsysteem beschikken
voorhet afbreken van niet-zetmeelkoolhydraten, berust niet geheel op
toeval.
(Dit proefschrift)
De ileale verteerbaarheid en de benutbaarheid van suikers kunnen sterk

uiteenlopen.
(Dit proefschrift)
De waarde van de xylose absorptietest als indicator voor een verstoorde

darmfunktie dient ter discussie te worden gesteld.
(Dit proefschrift)
In het onderzoek naar de toepassingsmogelijkheden van enzymen in de

voeding van pluimvee dient de nadruk te liggen ophet elimineren van de
anti-nutritionele effekten van niet-zetmeelkoolhydraat frakties.
(Dit proefschrift)
5. Volwassen hanen voorzien van een ileostomie canule, kunnen als model
dienen voor het schatten van de ileale verteerbaarheid van eiwit en
aminozuren van grondstoffen voor het varken.
Ibepassingvanvrije aminozuren invoedersvoorvarkens enpluimvee kan

een wezenlijke bijdrage leveren tot het reduceren van de N emissie.
7. Debewering van Pinchasov et al., dat bij kuikens minder goede produktie
resultaten worden bereikt met vrije aminozuren dan met eiwit-gebonden
aminozuren isonterecht, daar dezegebaseerd isopeenonvolledige toevoe-
gingvan vrije aminozuren aan het rantsoen.
(Pinchasov et al., 1990.Poultry Sience 69: 1950-1955)
De stelling in de Consumentengids van September 1991, dat veel veevoer

bestaat uit afvalprodukten van devoedings-engenotmiddelenindustrie en
zodoendeleidttotgoedkoopvoer,isonvolledigendientuitgebreidte worden
met "en tot goedkopere levensmiddelen voor de mens".
9. Veelvan de huidige problemen in de dierlijke produktiesektor kunnen
worden opgelost door devarkensstapel te vervangen door pluimvee. Het
enige bezwaar van deze vervanging is dat er een tekort aan poten kan
ontstaan.
10. Desuggestiedateenlinolzuur-rijk eigezonderisdaneennormaal ei,berust

op een misverstand.
11. Mettuinieren alshobbylooptmenzeldenpersoonlijk gezichtsverlies op;de

schuldvaneenslechteoogstligtimmersaltijd bijhetuitgangsmateriaalen/
ofhet weer.
12. Debevolkinginhet westenvan onsland dient zichmeerbewust te worden

van het feit, dat zijhaar welvaart voor een belangrijk deelte danken heeft
aan de in Drente en Groningen aanwezige bodemschatten.
Proefschrift J.B. Schutte.

Nutritional value and physiological effects ofD-xylose and L-arabinose in
poultry and pigs.
Wageningen, 16december, 1991.
AanAnny
AanIrma,Hans enAstrid
Aanmijnmoeder
Iternagedachtenis aanmijn vader
VOORWOORD
Gaarne maak ikvan dezegelegenheid gebruik omalientebedanken die direkt

of indirekt aan het tot stand komen van dit proefschrift hebben bijgedragen.
Allereerst ben ik veel dank verschvildigd aan mijn leermeesters, Dr.Ir.E.J. van
Weerden enwijlen Dr.Ir.P.van derWal.Mijnwetenschappelijke vormingheb ik
vooreenbelangrijk deelaan hen te danken. Het wasvoormij dan ook bijzonder
plezierigdatDr.Ir.E.J.vanWeerdenalsco-promotorvanmijn promotieonderzoek
wildeoptreden.Uwkritischeinstellingenervaringhebbenbijdebeoordelingen
correctie van de manuscripten er medetoebijgedragen dat deze als onderdelen
van dit proefschrift gepubliceerd konden worden. Bijzonder hartelijk bedankt
vooruwgoedeenplezierigebegeleiding,temeeromdatUhiereenbelangrijk deel
van uw VUT tijd aan opofferde. Op het gebied van de begeleiding van het
proefschrift benikvoortsveeldankverschuldigdaanmijnpromotoren Prof.Dr.Ir.
M.W.A. Verstegen en Prof.Dr.Ir. S. Tamminga. De energieke en niet aflatende
steun die ik van Ubeiden heb mogen ondervinden, heb ik ophoge prijs gesteld.
Het onderzoek was zeer gevarieerd. Bij de aanvang van het onderzoek (eind
1987) had ik nog geen vermoeden van de problemen die zich zouden kunnen
voordoen. Een van deze problemen lag op het vlak van het analyseren van de
suikers xyloseen arabinose in darminhoud, urine en faeces. Dankzij de inspan-
ningen van de voormalige IGMB-TNO afdeling Chemie en Granen werd dit
probleem tot eenoplossinggebracht. BesteRobHamer,Maarten van Oort,Wim
Lichtendonk en Henk van Lonkhuijsen, ik heb het jullie niet altijd even
gemakkelijk gemaakt met mijn twijfels aangaande de nauwkeurigheid van de
analyses. Bedankt voorjulliebijdragen aan het onderzoek. Deoverige analyses
en het onderzoek met varkens en pluimvee werden uitgevoerd ophet ILOB.
Velemedewerkers van het ILOBhebben openigerleiwijzemet het onderzoek
temakengehad.GijsvanKempenwilikbedankenvoordevrijheiddieikvanhem
kreeg om het geheel vrij snel af te ronden. Een bijzonder woord van dank aan
Johan deJong.BesteJohan,inveelgevallenwasjijmijn steunentoeverlaat. Het
organiserenvandedierproevenenhetverzamelenvandegegevensheeftjouheel
wat inspanningen gekost. Nogmaals bijzonder hartelijk bedankt voor de goede
enplezierigewijzewaaropjijdezetakenvervulde.Tevensgaathierbijmijn dank
uit naar Gerard Beelen, die in voorkomende gevallen deze taken van jou
overnam. Piet Roeleveld en Karel Siebers zegik dank voordezorgvuldige wijze
waarop zij de proefvoeders hebben bereid, en Piet van Leeuwen en Martje
Fentener van Vlissingen voor de chirurgische ondersteuning. De technische
uitvoering van de onderzoekingen met varkens en pluimvee was in de ver-
trouwdehandenvanKasperDeuring,JohandeZeeuw,JanvandeBroek,Rikvan
de Heuvel, Dick van Kleef, Anne Hoek en Jan van Harn. Bedankt voor jullie
bijdrage en develeopgeofferde nachtelijke uren. Martin van Baak, Rolf Coolen,
ChrisRietveld,RonaldKramerenAnnemarievandenDriesschewilikbedanken
voordeveleanalyses diezijinhetkadervan ditonderzoekhebben uitgevoerd,
enJanWiebenga,GerritHaakenGiljamDerksen(TNO-ITI)voordesteunbdjhet
statistisch verwerken van deuitkomsten.
Sake Bakker ben ik erkentelijk voor zijn steun bij het bewaken van het
projectbudget. Een woord van dank ben ik zeker verschuldigd aan Renee
Polziehn (Canada) voor de geboden hulp bij het uitvoeren van de proeven, en
HuugBoervandeVakgroepVeevoedingvoorzijnadviezenopanalytischgebied.
Verder dank ik dedames van de administratie, met name VickyKokvoorhet
typenvandemanuscripten.BesteVicky,zonderjouwsteunzouhetboekjezeker
enige weken vertraging hebben opgelopen. En last but not least wil ik in het
bijzonder mijnvrouwAnnybedankenvoordemorelesteunenhet overbrengen
vanmijn handgeschreven teksten indecomputer.
Hetonderzoekwasniettot stand gekomenzonderfinancieleondersteuning.
Verschillende fondsen hebben een bijdrage geleverd. Deze bijdragen waren
afkomstig van TNO-Voeding, het Produktschap voor Veevoeder, het Fonds
Mestonderzoek envan deProgrammacommissie Landbouw Biotechnologie.
Ben Schutte
CONTENTS
General introduction 1
PartA.Studies withpoultry 21
Chapter 1. Nutritional implications and metabolizable energy 23

value ofD-xyloseand L-arabinose in chicks
Chapter 2. Ileal digestibility and urinary excretion ofD-xylose 39

and L-arabinose inileostomized adult roosters
Chapter 3. Nutritional value ofD-xyloseand L-arabinose for 57

broiler chicks
Part B.Studies with pigs 77
Chapter 4. Nutritional implications ofD-xyloseinpigs 79
Chapter 5. Urinary excretion ofD-xyloseinpigsasaffected 99

byage,frequency offeeding and dietary level
Chapter 6. Nutritional implications ofL-arabinose inpigs 117
General discussion 141
Summary 161
Samenvatting 167
Curriculum Vitae 173

GENERAL INTRODUCTION
GENERAL INTRODUCTION
Traditionally pig and poultry diets are mainly based on cereals and
soyabean meal. However, for economical or political reasons, the use of
alternative products such aspeas,beans,sunflower meal,rapeseed meal,
lupins, cereal by-products and sugarbeet pulp has received increasing
interest inrecentyears.Inprinciple,theseproducts havethe potential to
replace wholy or partly cereals and/or soyabean meal in pig and poultry
diets. Several factors such as the presence ofharmful compounds orlow
digestibility, however, limit the level of their inclusion in the diet. The
harmful compounds are mainly anti-nutritional factors (ANF) such as
trypsin inhibitors, lectins, tannins and antigenic proteins. These ANF,
which are mainly found in legumes, can depress protein digestion and
performance of pigs and poultry (Huisman, 1990). In products such as
sunflower meal, rapeseed meal,lupins,cereal by-products and sugarbeet
pulp,thelowdigestibilityoftenlimitstheirinclusioninpigandpoultrydiets
in appreciable quantities. This low digestibility is associated with the
composition ofthe carbohydrate fraction inthese products,which mainly
consistsofnonstarchpolysaccharides(NSP).Asthetermimplies,theseNSP
exist in a form other than starch. It is well known that starch can be
hydrolysed bypancreatic oc-amylaseand maytherefore bedigested in the
smallintestineofpigsandpoultry,andabsorbedasglucose.However,NSP
arenotsusceptibletopancreaticenzymes(Trowelletal.,1976)andcanonly
beutilizedafter fermentation bygutbacteria.Thisfermentation processis
notonlycoupled with considerable lossesin energy,but dietary NSPmay
alsoinduceadepressionofthedigestibilityofotherdietarycomponents.In
principle both problems, the lowdigestibility of NSP,and their negative
effect onthedigestibilityofotherdietarycomponentscanbeovercomebya
treatmentwithenzymeswhichcanhydrolysetheNSP.Aswillbediscussed
inmoredetailinthischapter,thishydrolysiswillnotonlyreleaseD-glucose,
butalsoothersugars(e.g.D-xyloseandL-arabinose)ofwhichthenutritional
value forpigsand poultry isnotfully understood.
NATURE OFNONSTARCH POLYSACCHARIDES
The nonstarch polysaccharides (NSP) are complicated compounds, both

from thepointofviewofphysicalstructureandchemicalcomposition, and
include cellulose, hemicelluloses, pectins, oligosaccharides and lignin
General introduction
(Figure 1).Infact,ligninisnotapolysaccharidebutapolymerofoxygenated
phenylpropane unitsincludingconiferyl, sinapyl,and p-coumarylalcohols.
Because of the low levels of lignin in common feed ingredients, its
encrustration with structural carbohydrates and for the sake of simplicity,
this fraction is also included in the group of NSP. The basic structural
material ofplant cellwalls iscellulose.It is composed oflinear chains ofJJ-
(1-4) glycosidically bound glucose, arranged in fibrils in a crystalline or
amorphousregion.Beforetheycanbedegradedenzymatically,the crystalline
regionshavetobeconvertedintoamorphousones.Hemicellulosesaremade
up ofa complex ofpentoses and hexoses and frequently show branched as
well aslinear chains.The chain structure varies in the different feedstuffs
ofplant origin.Moreoverhemicelluloses areoften encrustrated with lignin.
Thisencrustrationinparticular seemstopreventattackbyenzymesaswell
asbymicrobes.Dependingontheirstructure,hemicellulosescanbe present
in soluble or insoluble forms. In cereals,the hemicelluloses mainly consist
ofarabinoxylans,often referred toaspentosans.Pectinsin plants form the
socalled middle lammela between cells. The main component of pectine
is a-(l-4)-glycosidically linked galacturonic acid. Oligosaccharides of
the raffinose family (e.g. raffinose.stachyose, verbascose) can form an
Carbohydrates 1
Crude fibre I Nitrogen free extract |
Cellulose Hemi- Pectin Oligo- Starch+

Cellulose Sacch. Sugars
Nonstarch Polysaccharides
FIGURE 1. Classification of carbohydrates in vegetable feeds

important constituent ofthe NSP in legume seeds.The raffinose family is

constituted ofsugars related to raffinose by the fact that they have one or
more a-D-galactopyranosylgroups in their structure. Thebasicunit of the
raffinose family is sucrose, and a-galactose units are bound to glucose.
The contents ofNSP in feed ingredients ofvegetable origin vary widely
(Table 1). From the data presented in this Table, it is obvious that the
carbohydrate fraction ofmaize and wheat is predominantly starch-based.
This isin contrast tothe carbohydrate fraction ofthe protein-rich products
sunflower meal, soyabeanmeal,groundnut meal and rapeseed meal which
existsmainlyasNSP.Thesameistrueforthecerealby-products,lupins and
sugarbeet pulp. Compared with these products, the contents of NSP in
beans and peas are relatively low.
Regardingthe composition ofNSPinthevariousfeed ingredients,there is
still a lack of precise analytical data on the quantitative amounts. Infor-
mation about the contents ofcellulose, hemicellulose and lignin from
plant residues canbederived from the acidand neutral detergent methods
developed and modified by Van Soest and co-workers (Robertson & Van
Soest, 1981).The acid-detergent fibre (ADF)residue provides an estimate
ofthe cellulose and lignin content. Estimates ofhemicellulose content are
obtained by difference using the amounts ofADF and neutral-detergent
fibre (NDF)present. Ligninsmustbedetermined inan additional step.Van
Soest'smethod doesnotmeasure solublesubstances,suchaspectins,which
are extensively removed by NDF extraction. Broadly, it can be said that
celluloseandhemicellulose arethemostimportant constituents ofthe NSP
fraction in cereal by-products (Theander et al., 1989).In products such as
sunflower meal, soyabean meal,beans and peas, the NSP fraction mainly
consists of pectins, oligosaccharides and cellulose, and in sunflower meal
alsooflignin (Taufel et al., 1960;Rackis, 1974;Cerning et al., 1975;Voseet
al., 1976;Reichert, 1981;Brillouet &CarrS, 1983;Carre"&Brillouet, 1986).
The NSP fraction of lupins and sugarbeet pulp is characterized by high
contents ofpectins (Carre"&Leclercq, 1985;Beldman,1986).
TABLE1.Contents ofcarbohydrates, starch (+sugars) andNSP in

somefeedstuffs (% of product)x)
Ingredient Ibtal Starch NSP

carbohydrates (+ sugars)
Maize 72 64 8
Wheat 71 63 8
Maizegluten feed 62 20 42
Wheat bran 62 18 44
Beans (Phas.vul.) 60 43 17
Peas 58 40 18
Lupins 44 7 37
Soyabean meal 36 9* 27
Sunflower meal (dehulled) 44 + 44
Groundnut meal 45 15* 30
Rapeseed meal 45 13* 32
Sugarbeet pulp 70 15 55
1)Valuesderived from figures givenbyAnonymous(1988).

*)Vervaekeet al.,(1989).
DEGRADATIONOFNSP
TheabsenceofindigenousNSPdegradingenzymesandthelowdensityof
micro-organisms inthe smallintestine ofpigsand poultry,mean that the
NSPlargelypasstothehindgut.HeretheNSParedegraded toa greater
orlesserextentbythemicrobesforwhichtheyarethemajor carbonsource.
Theendproducts ofthis microbial degradation (lacticacid,volatile fatty
acids) are readily absorbed and can be used as an energy source by the
animal(Imoto&Namioka, 1978;Kassetal., 1980;VanEs, 1987;Vervaeke
etal.,1989).Thisfermentationprocess,however,iscoupledwithconsiderable
losses in energy, assumed to vary in pigs between 33% (Agricultural
Research Council, 1981) and 50%(Just et al., 1983;Van Es, 1987). No
literature data for poultry are available, but energy losses of a similar
magnitude are assumed.
Theextentoffermentation ofNSPinthehindgutisquitevariable between
various constituents of NSP. Lignin and cellulose are poorly degraded,
whereas pectin, oligosaccharides and to some extent, hemicelluloses are
reported to be easily fermented (Keys et al., 1969,1970; Cummings et al.,
1978;Nyman&Asp,1982;Stanogias&Pearce,1985;Longland&Low,1988;
Trevino et al., 1990).It also appears that microbial degradation ofNSP in
poultry is less important than in pigs (Longstaff & McNab, 1989). As
demonstrated byCarre'et al.(1990),birdscanonlydigest thewater soluble
fraction oftheNSPin significant amountsthrough bacterial fermentation,
but inpigsalsoa significant part ofthewater insolubleNSPfraction canbe
digested (Dierick et al., 1989).
Apartfrom thelowdigestibilityandutilization of NSPbypigsand poultry,
theNSP-inducedmicrobialactivitymaydecreaseilealandfaecal digestibility
(Saueretal., 1980;Diericketal., 1983;Justetal., 1983;Grahametal., 1986;
Siriwan et al., 1990).In addition NSP may inhibit lipid absorption (Just et
al., 1980; Kay, 1982; Kies, 1985) and decrease digestibility of minerals
(Partridge, 1978; Graham et al., 1986; Ward & Reichert, 1986). Viscous
polysaccharides, such aspectin substances and B-glucanshavebeen shown
todepress chickperformance and tocausesticky,wetdroppings (Wagner&
Thomas, 1977;Gohl et al., 1978;Day &Thomas, 1980;White et al.,1981;
Annison, 1990).
HYDROLYSIS PRODUCTS OF NSP
From aliteraturereview,Rexen(1981)and Chesson (1987)concluded that

the digestibility ofNSP canbeimproved bytreatment with enzymes which
can hydrolyse the NSP to monosaccharides. This has been confirmed in a
recent study at our institute (Schutte et al., 1990).Our study showed that
in addition to an increase ofthe digestibility ofcell wall components, the
digestionofprotein andfat wasalsoimprovedinpigswhenwheatbran was
treated with a cellulolyticenzymepreparation. Thebenefits ofa hydrolysis
ofNSP,however,arenotdeterminedonlybyanimprovementindigestibility,
but alsobythe potential ofthe animal toutilize the hydrolysis products. In
addition toglucose,other sugars and uronicacidswillalsobereleased by a
completehydrolysisofNSP(Table2).Fromtheproductsofhydrolysis listed
inTable 2,the sugars glucose and fructose arewellutilizedin monogastric
animals (Demetrakopoulos &Amos, 1978;Miles et al., 1987).The same is
true for galactose,but in chicksonlyat lowdietary inclusion levels (Rutter
etal.,1953;Longstaffet al.,1988);athighdietaryconcentrationsgalactose
caused kidney damage, convulsions and death (Sondergaard et al., 1957;
Rigdonetal.,1963).Thenutritionalvalueofuronicacidshasbeenreported
tobeverylowinchicks(Longstaff etal.,1988).Thiswassupportedbyour
owndata,(J.B.Schutte,unpublisheddata).Wefoundalsothatadministering
galacturonicacidtochicksresultedinincreasesinwaterintakeandcaeca
weights. Noliterature data on the utilization ofuronic acids in pigs are
available.
TABLE2.Themostimportantsugarswhichwillbereleasedfroma
complete hydrolysis ofthe NSP fractions.
NSP fraction Hexoses Pentoses Other

(C6 H12 0 6 ) H
<C, i o 0 5 )
Cellulose D-glucose
Hemicellulose D-glucose D-xylose

L-arabinose
Pectins D-galactose L-arabinose Uronic

acids*
Oligosaccharides D-glucose
D-fructose
D-galactose
*Mainlyasgalacturonic acid(=derived form ofD-galactose ofwhich the

CH2OH-groupisoxidizedto-COOH).
In addition to glucose, the pentose sugars xylose and arabinose are in

quantitative termsthemostimportant oneswhichwillbereleasedfrom a
complete hydrolysis ofthe NSP.Carre'&Brillouet (1986)determined the
composition of NSP of various feedstuffs commonly used in diets for
monogastricfarm animals(Table3).Fromtheir data itcanbe calculated
that inpractical dietsforgrowingpigsa completehydrolysis ofNSPwill
result in the release of each of both pentose sugars (D-xylose and L-
arabinose) in quantities equal toabout 4%ofthe diet. In Dutch practical
poultry diets this amounts toabout 2to 3%for each ofthese twopentose
sugars.
TABLE3.Composition of NSP isolated from various feed in-

gredients (%)".
Glucose Xylose Arabi- Galac- Mannose Uronic Lignin

nose tose acids
Maize 24.2 24.0 19.1 4.1 0.8 6.0 5.4

Wheat 26.0 28.9 17.4 0.9 0.7 3.4 7.2
Barley 30.4 31.9 14.6 0.9 1.0 2.1 6.6
Oats 32.5 29.0 5.0 0.9 0.3 3.3 7.8
Wheatbran 24.4 18.3 18.3 1.1 0.2 4.2 7.2
Soyabean meal 18.9 5.4 13.6 24.9 5.4 15.2 1.9
Sunflower meal 27.9 13.0 7.7 2.2 3.4 11.5 21.1
Rapeseed meal 17.3 4.8 12.8 4.2 0.3 17.3 25.2
Peas 41.6 7.9 19.7 1.7 + 15.5 2.4
Field beans 37.4 5.2 8.9 1.6 + 15.0 8.2
White lupins 32.4 11.4 9.7 21.9 0.5 10.8 2.3
Lucerne meal 42.2 11.7 3.2 1.7 1.2 13.0 17.2
DDatafromCarre'&Brillouet(1986).Inthedicotyledonousplantmaterials,
a small amount (0.1 -1.7%)ofrhamnose andfucose wasalsofound. Data
regardingthecontentsofstarch,proteinandashintheNSPfractions are
omitted.
+Traces.
ABSORPTION AND UTILIZATION OF D-XYLOSE AND L-

ARABINOSE
KnowledgeaboutthenutritionalvalueofD-xyloseandL-arabinoseinpigs
andpoultryisincomplete.Thisisnotsurprising,sinceintheirfreeformboth
sugars, having five carbon atoms (Figure 2),are normally present in the
smallintestine inonlyverylowconcentrations.Itiswellknownthat both
pentose sugars are absorbed from the intestinal tract of monogastric
animals (Cori, 1925;Miller& Lewis, 1932;Bogner, 1961;Wagh&Waibel,
1967a;Arnal-Peyrot&Adrian,1974).Itappears,however,thatbothpentose
sugars,in spiteoftheir identical molecular size,have a different modeof
transport inthe smallintestine.Itisgenerally acceptedthat L-arabinose
is passively absorbed from the small intestine of animals (Crane, 1960;
Herman, 1974). However, the mode of transport of D-xylose from the
intestinal tract is still controversial. The idea that D-xylose crosses the
10 General introduction
intestinal mucosa by simple diffusion was initially reported (Cori, 1925;

Wilson & Vincent, 1955; Crane, 1960). The bulk of evidence, however,
suggests that this sugar shares the same transport mechanism with D-
glucose (Salomon et al., 1961;Csaky & Lassen, 1964;Alvarado, 1966;
Caspary, 1972;Ohkohchi &Himukai, 1984).This willmean that glucose,
galactose and xylosewillcompetefor the same transport system, so that
absorption ofxylose,which has the least affinity for this system, will be
inhibited(Kidder etal.,1968).
Research ontherate and extent ofxyloseand arabinose absorption from
the intestinal tract ofmonogastric animals is limited. Bogner (1961) and
Wagh &Waibel (1967a) studied the absorption rate of D-xylose and L-
arabinose in youngchicks by analysing the gastro-intestinal contents 30
minutesafter oraladministrationbycroptube.Theirdata showedthatL-
arabinosewasabsorbedatalowerratethanglucoseandxylose,whereasthe
rate ofabsorption ofxylose was only slightly lower than that ofglucose.
Similar results were found in rats by Cori (1925), reporting rates of
absorptionofvarioussugarsinthesequencegalactose>glucose>fructose
>mannose>xylose>arabinose.Thissequencewouldbethesameforman,
rabbit and the dog(Herman, 1974).
FIGURE2.Structural formulae of D-glucose, D-xylose and L-

arabinose
H H H
I i
I
i
i
C= 0 C= 0 C= 0
I i
i
i
i
H - C- OH H - C- OH H- C- OH
I i
i
Ii
HO- C- H HO- C- H HO- C- H
I i
i 1
H - C- OH H - C- OH HO- C- H
I i
i
i
i
H - C- OH H - C- OH H - C- OH
I i
i
i
i
H - C- OH H H
I
H
D-glucose D-xyl()se L-arabinose

Themetabolicpathways ofbothpentose sugarsisstill under discussion.

Itiswellrecognizedthatbothpentosesugarsarepartlyexcretedintheurine
ofman and monogastric animals (Wise et al., 1954;Segal &Foley, 1959;
Finlayetal.,1964;Arnal-Peyrot&Adrian, 1974;Haeneyetal.,1978;Craig
&Atkinson, 1988).Considering the literature, there are indications that
renal excretion of D-xylose is affected by several factors. These factors
includeintestinalbacterialgrowth,dietarylevelandage.Becketal.(1962)
reported low xylose excretion in patients with intestinal diverticulli,
suggesting that part ofthe ingested xyloseisconsumed bythe intestinal
microbes. This hypothesis is supported by data of SchifFer et al. (1962),
Cookeet al.(1963)and Goldstein et al.(1970),whoreported an increased
urinaryxyloseexcretionafter antibioticsinpatientswith small intestinal
diverticulosis. Wagh and Waibel (1966) reported that in chicks, the
metabolizableenergy(ME)valueofxylosedecreasedwhenthedietarylevel
increased.Theirfindingmayprovidesomeevidenceofanincreasedurinary
excretion ofxylose in percentage ofintake when the dietary level of this
sugar is increased, since LongstafF et al. (1988) reported that apparent
digestibilityofD-xyloseinchickswasalmost100%.Ithasbeenreportedthat
renalxyloseoutputinmandeclineswithage(Fowler&Cooke,1960;Finlay
etal.,1964).Thereasonforthisisunknown,butithasbeenpostulated that
renalfunction, andconsequentlyxyloseexcretion,isaffected bytheageing
process (Kendall, 1970). On the other hand altered intestinal mucosal
function associated with ageing, and/ or a decrease in arterial bloodflow
throughthesmallbowelduetoatherosclerosishasbeensuggested(Sappet
al.,1964).Noliteraturedataareavailableonfactorswhichmayaffect renal
excretion ofarabinose.
Thexyloseabsorptiontestiswidelyusedinthediagnosisofmalabsorption
in man and animals. This test, in which the renal excretion ofxylose is
measuredafter anoraldoseofthesugar,isbasedontheassumptionthatD-
xyloseisnotmetabolizedtoanysignificant extentinthebody.Inman, the
xyloseabsorptiontestisoften standardizedtoanoraladministration of25
g D-xylosefollowed by a 5hour urine collection.Alow renal excretion of
xylose has been connected with mucosal diseases. The validity ofthe D-
xyloseabsorptiontesthasbeenquestioned(Sladen&Kumar,1973;Kwawitt
&Beeken, 1975;Hindmarsh, 1976).AccordingtoCraig&Atkinson (1988)
thisisnotsurprisingsincethexylosetesthasbeenderivedempiricallyand
only a few formal kinetic studies have been made to provide a rational
physiologicalbasisfortheir interpretation. Theassumption thatD-xylose
isnotmetabolized inthe bodydoesnotholdwhenconsidering the dataof
Wyngaarden et al. (1957). These investigators observed a rise in blood
glucose level after infusion ofD-xylosein man. The same was true after
infusion of L-arabinose in man. Hiatt (1957) observed in mice a slight

conversion of D-xylose-l-C14 to glycogen via the pentose phosphate
scheme, but virtually no conversion of L-arabinose.
The known biochemical pathways of metabolizing D-xylose, suggest
conversion to D-xylulose followed by the phosphorylation to xylulose-5-
phosphate,withsubsequentconversiontohexoseviathepentose phosphate
pathway. Enzyme systems responsible for such metabolic pathways have
been demonstrated to be present in bacteria (Hochster, 1955; Stumpf &
Horecker, 1956;McCormick &Touster, 1957),but not in pigs and poultry.
Consideringthe data of Wijngaarden et al.(1957),metabolism ofD-xylose
inhigherorganismsseemstobepossible.Theseinvestigators reported that
from an intravenous doseofC14labeled D-xylose given toman, about 14%
couldberecovered ascarbon dioxideintheexpired air.Similarresults were
reportedbySegal&Foley(1959)whofound that 16%ofanintravenous dose
of C14 D-xylose given to man was metabolized to carbon dioxide. Radio-
isotope studies with D-xylose in monogastric animals have onlybeen done
withchicks.Wagh&Waibel(1967b)reportedthatfromasubcutaneous dose
ofC14 labeled D-xylose, 4%was recovered in expired carbon dioxide.
According to Segal & Foley (1959), metabolism to carbon dioxide for
arabinose is of lower magnitude than for xylose. When an intravenously
infused dose of C14 L-arabinose was given to man, only 0.8% could be
recovered as carbon dioxide in the expired air. In bacteria, the pathway L-
arabinose -> L-arabinolactone -> L-arabonic acid -> a ketoglutarate has
been described (Weimberg &Doudoroff, 1955).Since a ketoglutarate gives
rise to carbon dioxide, if this pathway were operative in monogastric
animals the end product would be prior to the keto acid.Apparantly, the
sequence L-arabinose ->L-ribulose -> D-xylulose-5-phosphate reported in
bacteria(Heathetal.,1958),seemstobeabsentinmonogastricanimals,for
this would give rise to C14 carbon dioxide via the pentose phosphate
pathway.AnotheralternativeforthefateofL-arabinosemaybeitsconversion
toL-arabitol,a substance which hasbeen isolated from pentosuricurine of
man (Touster &Harewell, 1958).The L-xyluloseformed from the oxidation
ofL-arabitol maybereducedtoD-xylitolandthenoxidizedtoD-xylulose, as
found in animal cells (Hollman &Touster, 1957).
Studiesperformed byDarby&Day(1939)andBoothetal.(1953)withrats,
and Wise et al.(1954)with pigs,indicated that xylose can cause cataracts,
diarrheaandsevereanorexiawhenfedathighdietaryinclusionlevels.Wise
et al. (1954) also observed that retention ofN was significantly decreased
when pigswere fed a diet containing 560gD-xylose/kg.Theybelieved that
thiswasaresultofanenergydeficiencyontheD-xylosediet,and consequently
greater N catabolism. This is supported by data ofWagh &Waibel (1966)
whofound that plasma uric acid was significantly increased when chicks
werefedondietscontaining200and400gD-xylose/kg.Similarresultswere
foundwhenfeedingdietscontaining200and400gL-arabinose/kgtochicks.
They reportedalsothatfeedingdietscontainingthesehighlevelsofxylose
orarabinose tochicksresulted indecreased liverweights.
AIM AND OUTLINE OF THE STUDY
Theaimofthestudiesdescribedinthisthesiswastoinvestigatethe fate
ofD-xyloseand L-arabinose after oraladministration inpoultry andpigs.
Themainemphasishasbeenonthedigestionofthesesugarsinthe different
parts ofthe gastro-intestinal tract and on the renal excretion. From the
resultsofthestudies,themetabolizableenergyvalueofthesugarscouldbe
estimated, and the potential ofthe animals toutilize the energy of these
sugars is discussed. In addition some other nutritional implications of
xyloseand arabinoseindietsforpoultry and pigshavebeen examined.
Atotal ofsix studies were performed, three with poultry and three with
pigs.Thebasaldietinmostofthesestudieswascomposedoffeedingredients
ofsemi-purified origin.Thereason for this wastoavoid interference with
xylose and arabinose from dietary NSP material. Todetermine the ileal
digestibility of xylose and arabinose in pigs, a post valvular T-caecum
cannula (PVTC) was used (Van Leeuwen et al., 1988). This cannulation
technique was developed at ILOB as an alternative for the re-entrant
ileocaecalcannula.Beforestartingthepresentpigstudies,bothcannulation
techniques were compared in which D-xylosewas used as test sugar.No
significantdifferencesinapparentilealdigestibilityofD-xylose,drymatter,
Nandenergybetweeneithercannulationtechniquewereobserved.Based
ontheseresults,itwas decidedthat the PVTCcannula shouldbeused in
the present pigstudies.
InpartA(Chapters 1,2and 3),theresultsofthe studieswithpoultry are
presented.Inthefirst study,theeffect ofgradeddietarylevels(25to150g/
kg) of either D-xylose or L-arabinose on chick performance, bloodsugar
concentrations,caecalengthandweight,andliverweightwasinvestigated.
In addition, the metabolizable energy value of both pentose sugars was
determined.Thesecondstudymainlyfocussedonthedeterminationofthe
ilealapparentdigestibilityandurinaryexcretionofbothpentosesugarsby
usingileostomizedadultroosters.Thethirdstudydealswithtwoexperiments
inwhichthenutritionalvalueofbothpentosesugarsandtheireffectonbody
composition wasinvestigated byusingtwodifferent typesofbasal diet.
TheresultsoftheexperimentswithpigsarepresentedinpartB(Chapters
4, 5and 6).The main objective ofthe first study was todetermine the ileal
and faecal digestibility of D-xylose. In addition, urinary excretion of D-
xylose was examined, and the effect ofthis pentose sugar on ileal flow of
volatile fatty acids, and ileal and faecal digestibility of other dietary
components was investigated. The second study was carried out with L-
arabinose ofwhich the design was similar to that done with D-xylose. In
the third study, the effects ofsomefactors which may affect urinary excre-
tion ofxylose were investigated. The factors tested included frequency of
feeding, age ofthe pigs and dietary inclusion level of D-xylose.
Inthegeneraldiscussion,variousaspectsoftheresultsrelatingto animal
species differences, and the practical consequences of the application of
enzymes in pig and poultry diets are discussed.
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White,W.B.,Bird,H.R., Sunde,M.L.,Prentice,N.,Burger,W.C.&Marlett,
J.A. (1981).The viscosity interaction ofbarley beta-glucan with
Trichoderma viride cellulase in the chick intestine. Poultry Science 60:
1043-1048.
Wilson, T.H. &Vincent, T.N. (1955).Absorption ofsugars in vitro by the
intestineofthegoldenhamster.JournalofBiologicalChemistry 216:851-
866.
Wise, M.B.,Barrick, E.R., Wise, G.H. &Osborne, J.C. (1954).Effects of
substituting xylose for glucose in a purified diet for pigs.Journal of
Animal Science 13:365-374.
Wyngaarden, J.B.,Segal,S. &Foley,J.B. (1957).Physiological disposition
and metabolic fate ofinfused pentoses in man. Journal of Clinical
Investigation 36: 1395-1407.
Yang,M.G.,Manoharan,K.andYoung,A.K(1969).Influenceand degradation
ofdietary cellulose in cecum ofrats. Journal ofNutrition 97: 260-264.
21
PART A
STUDIES WITH POULTRY

23
CHAPTER 1
NUTRITIONAL IMPLICATIONS AND

METABOLIZABLE ENERGY VALUE OF
D-XYLOSE AND L-ARABINOSE IN CHICKS
J.B. Schutte
TNO-InstituteofAnimalNutrition and Physiology(ILOB),

P.O.Box 15,Wageningen,TheNetherlands.
Published inPoultry Science69: 1724-1730(1990)

Reproducedbypermission ofThePoultry ScienceAssociationInc.
25
NUTRITIONAL IMPLICATIONS AND
METABOLIZABLE ENERGY VALUE OF
D-XYLOSE AND L-ARABINOSE IN CHICKS
J.B. Schutte
TNO-Institute ofAnimal Nutrition and Physiology (ILOB),

P.O. Box 15,6700AA,Wageningen, The Netherlands.
An experiment was conducted to examine the effects of graded dietary

levels (2.5,5.0, 7.5, 10.0 and 15.0%) ofdietary D-xylose or L-arabinose on
chickperformance.Asreference D-glucosewasincluded inthe experiment.
A second experiment was performed to determine theAME n of D-xylose
and L-arabinose. Results of Experiment 1 showed a significant linear
decrease(P<0.05)inweightgainandefficiency offeedutilizationwhen the
dietarylevelofeitherD-xyloseor L-arabinosewasincreased.Thesamewas
true for daily feed intake of the D-xylose treatments. Water intake was
linearly(P<0.05)increasedasdietarylevelofbothpentosesugars increased
and, as a result, drymatter content ofthe droppings decreased. Results of
Experiment 2 showed that the AMEn value ofeither pentose sugar was
doserelated.The AMEn valuesfor D-xylose at 5and 10%dietary inclusion
were 2,660 and 2,020 kcal/kg,respectively. Those for L-arabinose at these
inclusion levels were 2,300 and 1,360 kcal/kg, respectively. Feeding equal
dietary levels ofeither pentose sugar resulted in higher concentrations of
xylosethan ofarabinose inbloodplasma. Concentration ofglucosein blood
was not affected by feeding either D-xylose or L-arabinose. Cecal length
and weight were markedly increased by feeding L-arabinose and
intermediately by D-xylose. (Key words: pentose sugars, D-glucose, D-
xylose, L-arabinose, chicks)
INTRODUCTION
Theprotein-richingredientsusedinpoultrydietsaremainlyofvegetable
origin. The carbohydrate fraction of these ingredients consists mainly of
nonstarchpolysaccharides(cellulose,hemicellulose,pectins,etc.)which are
practically indigestible by poultry because birds do not possess the
appropriate gastrointestinal tract enzymes. The microflora in the large
intestine of birds seem to play only a minor role, thus digestibility of
26 Chapter 1
nonstarch polysaccharides (NSP) by microbial fermentation is also low.

Apart from the low digestibility, higher dietary inclusion of some NSP
fractionslikecellulose,betaglucansandpectinsmayalsoreducedigestibility
ofotherdietarycomponents andperformance ofchicks(Saitoet al.,1959;
Wagner&Thomas, 1977;Gohletal., 1978;Day&Thomas,1980;Nahm&
Carlson, 1985).
Improvementofthedigestibility andutilizationofNSPcouldbe attained
by including enzymes in the diets that can hydrolyze the NSP to
monosaccharides. However, a completehydrolysis oftheNSPwill release
not only glucose, but also other sugars such as xylose, arabinose and
galacturonicacid.Thenutritivevalueofthesesugarsinpoultryarereported
tobelow(Waghand Waibel, 1966, 1967a,b,Baker, 1977;Longstaff et al.,
1988) at dietary inclusion ranging from 10 to 60% of the diet. Little
information isavailable from research donewith lowerinclusionlevels.
Thetwoexperimentsreportedhereinwerecarried outtostudythe effect
of D-xylose and L-arabinose in comparison with D-glucose at dietary
inclusion rangingfrom 2.5to 15%onchickperformance and to determine
themetabolizable energyvalue ofbothpentose sugars.
MATERIALSAND METHODS
Experiment 1:Growth.
Thisexperiment wasdesignedtocomparethe effects ofgraded (2.5,5.0,

7.5, 10.0 and 15.0%) dietary D-glucose, D-xylose or L-arabinose on body
weightgain,feedconsumption,feedutilization,waterconsumption,anddry
mattercontentofexcreta.Day-oldfemalebroilerchicks("Hybro",Euribrid
BV,Boxmeer, Holland) were used. The birds were housed in electrically
heatedbattery cagesof 975cm2 offloorspacewithwirefloors.Thecages
weresituatedinaninsulatedroomwithfacilitiesforcontroloftemperature
and humidity. Chicks were subjected to continuous artificial fluorescent
illumination.Astandard diet was fed for the first three days,followed by
anotherthreedaysofamixtureofthestandarddietandthebasaldietused
inthisexperiment.Atsixdaysofage,fourteen birdswere allotted toeach
of60cagessuchthat averagebodyweight(156g)andweightrange(135to
180g)weresimilar.Treatment groupsconsisted offour cagesarranged in
a randomizedblock design.
Thenutritionallycompletebasaldietusedwasbasedoncorn,cornstarch
and isolated soyaprotein (Table 1).Thethree sugars (D-glucose,D-xylose
Pentose sugars in chicks 27
TABLE 1. C o m p o s i t i o n of t h e b a s a l diet.
Ingredient %
Corn 28.65
Corn starch 28.70
Soy oil 4.00
Animal fat 4.00
Isolated soya protein (88%CP) 22.33
Cellulose ("Akufloc") 6.00
Ground limestone 1.25
Monocalcium phosphate 2.10
Salt, iodized 0.30
Potasium bicarbonate 1.50
Vitamin-mineral premix 1 ' 1.00
DL-methionine 0.17
Calculated contents:
ME, kcal/kg 3460
Crude protein, % 22.5
Calcium, % 0.90
Phosphorus, % 0.70
Lysine, % 1.33
Methionine plus cystine, % 0.90
1) Supplied perkilogram ofdiet:vitaminA, 10,000IU;vitamin D,

2,000 ICU;dl-a-tocopherylacetate, 30mg;menadione,5mg;thiamin,
2.5mg;riboflavin, 5.5mg;d-pantothenic acid, 15mg;niacinamide,
50mg;cobalamin, 15ug;cholinechloride, 1,850 mg;pyridoxine,
3mg;biotin,0.15mg;folic acid,0.75mg;ascorbicacid, 50mg;
inositol, 100mg;para-amino-benzoic acid, 2.5mg;ethoxyquin,
100mg;avoparcin, 15mg;Mg,420mg;Mn,95mg;Zn,50mg;Fe,
40mg;Cu,40mg;Co,2mg;Se,0.1mg.
andL-arabinose),suppliedasanhydrousmonosaccharidesweresubstituted
by weight for corn starch. The rations were pelleted utilizing a labor-
monoroll press (Simon Heesen, Boxtel, Holland) at an approximate
temperature of50°C.Both feed and water were available for ad libitum
consumption duringthe days6to25posthatching growth period.
At the end ofthe trial, chicks were weighed individually and feed con-
28 Chapter 1
sumption for each cage was recorded. During the last two days of the
experimental period water consumption was measured, and excreta were
collected,bothforeachcageatintervalsof12h.Waterintakewas measured
as the difference between a predetermined volume of water in the water
pans andthat remaininginthepans.Excretawerecollected quantitatively
intocontainers and stored at 4C.Immediately after the two-day collection
period the excreta were pooled per treatment group and analyzed for dry
matter content.
Experiment 2: Balance
This experiment was conducted toestimate theAME ofD-xyloseand L-

arabinose at 5and 10%dietary inclusion levels.In addition, blood samples
were collected from the birds for glucose, xylose and arabinose analyses;
gross morphological observations on the gastrointestinal tract, liver and
kidneys were performed.
Thebalancetrialwascarried outwithbirdsfrom Experiment 1consuming
rations containing 10% D-glucose, 5 or 10% D-xylose and 5 or 10% L-
arabinose, respectively. During the balance trial, rations with the same
type and levelofsugar werefed (Table 2).Thirty-twobirds (25days ofage)
TABLE 2. D e s i g n of e x p e r i m e n t 2 (balance trial)
Treatment Diets fed in the Diets fed in the

preceding period balance period
A 10% D-glucose Basal *

B 5% D-xylose 95%basal + 5% D-xylose
C 10% D-xylose 90%basal + 10% D-xylose
D 5% L-arabinose 95%basal + 5% L-arabinose
E 10% L-arabinose 90%basal + 10% L-arabinose
* Contained 10%D-glucose
fromeachofthefiveprecedingtreatmentgroupswereselectedsuchthat the
differences in body weight between these groups at the end ofthe growth
trial were maintained. In the balance trial each treatment group included
four cages, each with 8 birds.
The birds were housed in conditions similar to those described in the
growth trial.The experimental dietswere again freshly prepared. The 10%
D-glucose diet was used as a control (basal) diet. In the other diets a
proportionalpartofthebasaldietwassubsitutedbyD-xyloseorL-arabinose
(Table 2).
The balance trial included a pretest period of 10days and a test period
of4 days. During the last 3 days ofthe pretest period and the 4 days test
period, equal amounts offeed were fed toall cages (800gfeed per cage per
day). Feed was supplied as pellets four times daily. These portions were
quantitativelyconsumedwithinhalfanhour.Duringthe4x24htestperiod,
the excretawere collected quantitatively from glasstrays at intervals of12
h. Contaminants, such as downand scales,werecarefully removed and the
excreta were then stored in closed containers at -25C.In addition to feed,
water intake was alsorecorded during the test period.Thelatter was done
according to the same procedure as followed in Experiment 1.
The five experimental diets and the excreta were analyzed for dry mat-
ter, nitrogen and gross energy (GE).TheAME ofeach diet was calculated
from thefigures for GEofthefeed and excreta;eachreplicateof8birds was
assessed separately. The AME values were corrected to zero nitrogen
balance (AME n ).Thecorrection factor of8.22 kcal/gofretained nitrogen as
proposed by Hill andAnderson (1958) was used.
When the balance trial was finished, blood samples were taken from 8
randomly selectedbirdspertreatment group.Thiswasdonetwice,the first
time without fasting and the second time after a 12-h feed deprivation
(waterremained available duringthefeed deprivation).Theblood samples
wereobtained from theulnarveinusingheparinized syringes.Plasma was
analyzed for glucose, xylose and arabinose.
Furthermore 16randomlyselectedbirdspertreatmentgroupwerekilled
after afeeddeprivationperiodof12h.Waterremainedavailableduringthis
period. The birds were killed by injection of T61 (Embutramide-
Mebezoniumiodide-Tetracainhydrochloridemix,Hoechst,F.R.G.).Thesmall
intestine and the ceca were removed immediately and their length was
measured. In addition, the cecal weight (including contents) was deter-
mined.Grosspathological examination oftheliverandkidneyswas carried
out, and weight ofthe liver was recorded.
Chemical analysis
An IKA-C4000 adiabatic bomb colorimeter was used to determine the

gross energy energy content ofthe diets and excreta. The nitrogen content
ofdietsandexcretaweredeterminedbytheKjeldahlprocedure (Association
ofOfficialAnalytical Chemists,1975)usinganautomaticanalyzer(Technicon
Instruments systems, Tarrytown, NY).
30 Chapter 1
Plasma sugar concentrations were determined as silyl derivativesof

monosaccharidesbygas-liquid chromatography (Sweeleyet al., 1963).One
ml of plasma was diluted with distilled water (1:10), deproteinized with
potassium ferrocyanate and zinc-acetate and desalted bypassing through
a mixture of anion (Biorad AG 3x4) and cation (Biorad AG 50 W x 4)
exchanger. Next, the samples were derivatized in pyridine with hexame-
thyldisilazane andtrimethylchlorosilane and analyzedbyusinga Hewlett-
Packard (Model HP 5890) gas-liquidchromatograph with a Chrompack
capillaryWCOTfused silicacolumncoatedwith CPsil5CBof50m length.
Statistical analysis
A two-way ANOVA was used for Experiment 1, which had a factorial

arrangement oftreatments.Aone-wayANOVAwasused for Experiment 2.
Forexperiment 1,the sum ofsquares forlevelswaspartitioned intoa setof
orthogonal linear and quadratic polynomial regression components. The
interaction sum ofsquares was partitioned bythe same set ofcomponents.
The computer program GENSTAT 5 (Reference Manual 1987, Oxford
University Press, New York) was used to calculate the ANOVA. The
significance ofdifferences betweentreatmentmeanswastestedbyusingthe
LeastSignificanceDifference test(Snedecor&Cochran,1980).All statements
ofsignificance are based on a probability ofless than 0.05.
RESULTS
Experiment 1: Growth.
The results ofthis experiment are summarized inTable 3.At all dietary
levels ofD-glucose,almost identical growth rate and feed conversion were
achieved.Water intake and drymatter content ofthe excreta alsowere not
affected by the added level ofD-glucose in the diet.
Weight gain and efficiency offeed utilization decreased linearly (P<0.05)
as the level ofD-xylose or L-arabinose increased (Table 3).
Pentosesugarsinchicks 31
TABLE3.The effect ofdietary inclusion levels ofD-glucose,

D-xyloseandL-arabinoseonchickperformance from6to25
days of age with ANOVAsummary (Experiment 1).
Sugar Dietary Weight Feed Feed/ Water Dry matter

level gain intake gain intake content 2
(%) (g) (g/bird (g:g) (g/bird excreta(%)
per day) per day)1
D-glucose 2.5 860 65.8 1.46 154 26.6
5.0 852 66.3 1.48 144 26.0
7.5 860 67.4 1.49 151 26.3
10.0 870 67.6 1.48 148 26.4
15.0 858 66.6 1.48 146 25.2
D-xylose 2.5 817 63.6 1.48 186 19.1
5.0 802 62.6 1.49 203 16.4
7.5 770 61.4 1.51 246 13.3
10.0 726 61.1 1.60 252 13.0
15.0 671 58.1 1.64 291 9.7
L-arabinose 2.5 826 65.0 1.50 209 17.1
5.0 802 63.8 1.51 222 16.6
7.5 781 63.9 1.56 281 12.9
10.0 761 63.6 1.59 303 12.1
15.0 739 65.4 1.68 377 10.0
ANOVA summaries.
Source of variation Probability
Df
Sugar(S) 2 ** ** ** **
Level(L) 4 ** ** ** **
Linear 1 ** ** ** **
Quadratic 1 NS NS NS NS
SxL 8 ** ** ** **
Linear 2 ** ** ** **
Quadratic 2 NS * NS NS
Residual MS 45 369 1.76 0.0006 637
1Meansoffour pensper treatment ofDays24plus25.

2Pooledsamplespertreatment ofDays24plus25;
figures not analysed statistically.
* P<0.05.
** P <0.01.
32 Chapter 1
ThesamewastrueforthedailyfeedintakebybirdsfeddietscontainingD-
xylose,butnotforintakeofchicksfedL-arabinose.Thedifferencesinweight
gainbetweenbothpentosesugarsweresmalluptoadietarylevelof7.5%,
but at 10and 15%,weight gainofthebirdsfed the L-arabinose diets was
significantly (P<0.05)greaterthanthosefedtheD-xylosediets.However,
feedconversionefficiency,wassomewhatbetterwithD-xylosethanwithL-
arabinose.Onacompositebasisthedifferences infeedconversion efficiency
betweenthe twopentose sugars were significant (P<0.05).
Asignificant linear increase (P<0.05)indailywater intake occurred
as the dietary level ofeither D-xyloseor L-arabinose increased (Table3).
However,ateachdietarylevel,waterintakeofthebirdsfedtheL-arabinose
diets wasmuchhigher than that ofthe birdsfed the D-xylosediets.On a
composite basis, the difference in water intake between the two pentose
sugarswassignificant (P<0.01).Asaresultoftheincreaseinwaterintake
fromincreasingdietaryD-xyloseorL-arabinose,drymatter contentofthe
excreta decreased steadily.
Mortality rate wasverylow;only 1%ofthebirdsdied.No appreciable
differences inmortality amongthe treatments were observed.
Experiment 2:Balance.
The values determined for the AMEn of the experimental diets are
presentedinTable4.FromthesevaluestheAMEn contentof D-xyloseand
TABLE 4. Determined AMEn values of the experimental diets

(Experiment 2).
Diets AMEn (kcal/kg)
Basal 3505+ 2a
95%basal + 5%D-xylose 3463±12b
90%basal + 10%D-xylose 3357±14c
95%basal + 5%L-arabinose 3445±10b
90%basal + 10%L-arabinose 3291±25d
ad
Means(±SD)withnocommonsuperscript differ significantly (P<0.05).
L-arabinose was derived by comparing theAMEn ofxylose and arabinose

containing diets with that ofthe basal diet (Table 5). The AME n value of
bothpentose sugarswasdose-related.Atanyspecific dietarylevel,a higher
AMEn value was observed for D-xylosethan for L-arabinose, but only the
difference at the 10%dietary inclusion was significant.
TABLE 5.D e r i v e d AME n v a l u e s for t h e s u g a r s ( E x p e r i m e n t 2).
Sugar Dietary (%) AMEn (kcal/kg)

inclusion
D-xylose 5 2660 ±243 a

10 2020 ±141 b
L-arabinose 5 2300 ±204 a
10 1360 ±251°
ac
Means (± SD)with nocommon superscript differ significantly (P<0.05).
Data regardingbodyliveweight, dailywater intake,drymatter content

ofexcreta, length ofsmall intestine and ceca, weight ofliver and ceca and
sugarconcentrationsinbloodplasmaareshowninTable6.Resultsfor daily
water intake and dry matter content of excreta agree well with those
obtained in the growth trial. The length of the small intestine was not
affected byfeedingdietscontainingeitherD-xyloseorL-arabinose.Thececa
from chicks fed diets containing D-xylose or L-arabinose were not only
longerbutalsoheavierthan thosefrom chicksfedthebasaldiet.The length
andweightofthececaofthebirdsfedtheL-arabinosedietswas dose-related;
cecal length and weight of the 10%L-arabinose group were significantly
different from thoseofthe5%L-arabinosegroup.Also,therewasa tendency
forcecaweightofthebirdsfedtheD-xylosedietstoincreaseasdietary level
increased from 5to 10%.
Relative liver weights of the birds fed either D-xylose or L-arabinose
were not significantly different from that ofchicks fed the basal diet
(Table 6).
34 Chapter 1
TABLE 6. Effect of dietary inclusion of D-xylose and L-

arabinose on water intake, dry matter content of excreta,
and physical and biochemical characteristics of chicks
(Experiment 2).
D-xylose L-arabinose
Parameter Basal 5 10 5 10
Live weight attheendof 1750* 1661 b : L593c 1663b 1526d

the experiment, g
Daily water intake, g/bird 179* 226b 292c 274c 365d
Dry matter content excreta, % 26.6* 19.0b 15.4C 16.l c 11.6d
Small intestine length,cm 124* 129* 129* 126a 135*

cecal length,cm 29 a 31 b 31 b 30*b 35c
cecal weight (incl. contents),
g 7.4" 9.2ab 10.6b 10.3b 14.5C
%ofLW 0.42a 0.55*b 0.67b 0.62b 0.95
Liver weight,
g 45.0" 40.4ab 42.2ab 47.5* 35.l b
%ofLW 2.6ab 2.4b 2.6*b 2.9a 2.3b
Blood plasma values, mmol/1

Without feed deprivation
Glucose 12.9* 12.6a 13.5* 13.5* 13.4*
Xylose 0 4.6* 8.2b 0 0
Arabinose 0 0 0 0.9* 1.5b
After 12h feed deprivation
Glucose 10.6* 11.2* 11.1" 10.9* 11.2*
Xylose 0 0 0 0 0
Arabinose 0 0 0 0 0
a-d
Meanswith nocommonsuperscripts within a row differ
significantly (P<0.05).
Pentose sugars inchicks 35
Gross pathological examination of the liver and kidneys did not show
abnormalitiesinanyofthetreatments.Noindicationswereobserved that
the D-glucose level of the blood plasma was influenced by feeding diets
containingD-xyloseorL-arabinose.However,whenequallevelsofD-xylose
or L-arabinose were fed, concentrations of xylose in the plasma were
significantlyhigherthanthoseforarabinose.Xyloseplasmaconcentrations
were almost doubled byincreasingthe levelofdietary inclusion from 5to
10%.Thesamewastrueforarabinose.After12hfeeddeprivation,thexylose
and arabinoselevelswerereducedtozero.
DISCUSSION
The results of Experiment 1 show that even at the very low dietary
inclusion of 2.5%, both D-xylose and L-arabinose affected performance
adversely.ThiscouldbeexpectedsincetheAMEn valueforD-xyloseandL-
arabinosearemuchlowerthantheAMEn valueofD-glucose(3640kcal/kg,
Andersonetal.,1958).Furthermore,theAMEnvalueofbothpentosesugars
decreasedwhenthedietarylevelwasincreased.Thisfindingisinagreement
withthoseofWagh&Waibel(1966),showingthat theAMEn valuesforD-
xyloseandL-arabinosewerereducedtozerokcal/kgatdietaryinclusionof
40%.Thedecreaseinenergyvalueofbothpentosesugarsbyincreasingthe
dietarylevelsmayhaveresultedfrom decreasedabsorptioncapacity,oran
increased urinary excretion, orboth ofthese.Adecrease inthe utilization
of the other energy bearing components in the diet may be a further
possibility,resultinginalowerderivedAMEnvalueforbothpentosesugars.
Results ofa recently performed study (Schutte et al., unpublished data)
indicatethat allthreefactors aremoreorless responsibleforthe decrease
in energy as dietary of D-xylose or dietary L-arabinose were increased.
However,inthat studythe increaseinsugar excretionwith theurine was
the most important factor. This may also be the reason for the excessive
waterconsumption ofbirdsfedthehigherdietarylevelsofeitherD-xylose
orL-arabinose observed inthe present studies.
TheAMEn value ofD-xylosewashigher than for L-arabinose.The feed
conversionefficiencydataofExperiment1wereconsistentwiththisfinding,
showingabetter efficiency offeed utilization ofthe birdsfed theD-xylose
diets than those fed the L-arabinose diets. The higherAMEnvalue ofD-
xylosewasnotreflected intheresults forweight gain duetothe depres-
sioninfeed intakecausedbydietary D-xylose.Depressed feed intake as a
resultoffeedingD-xylosedietstochicksalsowasobservedbyBaker(1977).
36 Chapter 1
ResultsweresimilarwhenD-xylosedietswerefedtopigs(Wiseetal.,1954).
The appearance of higher concentrations of xylose than arabinose in
plasmaobservedinthepresentstudyagreeswiththedataofWagh&Waibel
(1967a),suggesting a faster absorption ofthe former. This is supported by
the gastrointestinal absorption data of Wagh & Waibel (1967b), showing
that D-xylose was more efficiently absorbed from the small intestine than
L-arabinose. The more efficient absorption ofD-xylose over L-arabinose is
supported further bythe heavier cecalweights ofbirds fed the L-arabinose
diets than those fed the d-xylose diets,indicatingthat a greater part ofthe
former is fermented by the cecal microflora.
The observation that blood glucose was not affected in chicks when D-
xylose or L-arabinose were included in the diet, agrees with the finding of
Wagh & Waibel (1967a). These investigators also studied the effect of
feeding D-xylose and L-arabinose diets on liver weight and liver glycogen.
Atdietaryinclusionof10to40%,liverweight and liverglycogen decreased,
indicatingadepletionofliverglycogenasareflection ofenergy deprivation.
The results of the present study (Experiment 2) do not support this
conclusion.InExperiment 2,relativeliverweightwasonlyslightly reduced
when 10%L-arabinose was included in the diet.
In conclusion, the nutritional values ofD-xylose and L-arabinose are less
than that of D-glucose and are dose-dependent. In addition both pentose
sugars may induce unwanted nutritive problems and wet droppings.
ACKNOWLEDGMENTS
The author wishes to thank J.M. de Zeeuw and J. de Jong for technical
assistance. The assistance of G.B. Derksen for the statistical analyses is
gratefully acknowledged.
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39
CHAPTER2
ILEAL DIGESTIBILITY AND URINARY

EXCRETION OF D-XYLOSE AND L-ARABINOSE
IN ILEOSTOMIZED ADULT ROOSTERS
J.B.Schutte,P. vanLeeuwen,andW.J.Lichtendonk
TNO-InstituteofAnimalNutritionandPhysiology(ILOB),
P.O.Box 15,6700AAWageningen,TheNetherlands.
PublishedinPoultry Science70:884-891(1991)
Reproducedbypermission ofThePoultryScienceAssociationInc.
41
ILEAL DIGESTIBILITY AND URINARY

EXCRETION OF D-XYLOSE AND L-ARA-
BINOSE IN ILEOSTOMIZED ADULT
ROOSTERS
J.B. Schutte, P.van Leeuwen, and W.J. Lichtendonk

P.O.Box 15,6700AAWageningen, The Netherlands.
Anexperimentwithileostomizedadultroosterswasconductedtodetermine
the ileal digestibility and urinary excretion of D-xylose and L-arabinose.
As a reference D-glucosewas included in the experiment. The sugars were
tested at graded dietary levels of 2.5, 5.0, 7.5 and 10.0%. Mean ileal
digestibility ofD-glucose and D-xylose was nearly 100%.Ileal digestibility
ofL-arabinose decreased linearly(P<0.05)withincreasingdoselevel. The
corresponding ileal digestibilities for L-arabinose at dietary levels of2.5,
5.0,7.5and 10.0%were95.5,93.6,80.3and74.6%.Bothpentosesugars were
partly excreted in the urine. The extent of this urinary excretion in
percentage of intake increased linearly (P < 0.05) as the dietary level
increased. In roosters fed the 2.5% D-xylose diet, 7.2% of the D-xylose
consumed appeared in the urine. This level increased to 20.2% when
roosters were fed a diet containing 10.0%D-xylose. Corresponding values
for L-arabinose at these dietary inclusion levels were 8.7 and 16.6%.
(Key words: pentose sugars, D-xylose, L-arabinose, ileal digestibility,
urinary excretion, adult roosters).
INTRODUCTION
The feed ingredients used in poultry diets are mainly from vegetable
origin.Thecarbohydratefraction ofmanyofthesefeedingredients,including
soybean meal, sunflower meal, rapeseed meal, lupins and wheatbran,
consists mainly of non-starch polysaccharides (Carre & Brillouet, 1986;
Brillouet et al., 1988; Schutte et al., 1990), which are resistant to the
digestive enzymes. Moreover, degradation of non-starch polysaccharides
(NSP) in the hindgut ofbirds by microbes is also low (Vogt &Stute, 1971;
42 Chapter 2
Carre"&Leclercq,1985;Longstaff&McNab,1986,1989).Fromaliterature
review,Chesson(1987)concludedthatthedigestibilityoffeed ingredients
containinghighlevelsofNSPcanbeimprovedbytreatmentwithenzymes
whichcanhydrolyzetheNSPtomonosaccharides.However,hydrolysisof
NSP will release not only glucose, but also sugars such as xylose and
arabinosewhicharenormallynotencountered inthe small intestine.
Itiswellrecognized that both pentose sugarsare absorbed from the
intestinal tract inrats (Cori, 1925; Miller &Lewis, 1932;Arnal-Peyrot&
Adrian, 1974).The study reported byArnal-Peyrot &Adrian (1974) also
showed that part ofthe ingested xylose and arabinose is excreted in the
urine.StudiesperformedbyDarby&Day(1939)andBoothetal.(1953)with
rats, and Wise et al. (1954) with pigs, indicated that xylose can cause
cataracts,diarrheaandsevereanorexiawhenfedathighdietaryinclusion
levels.Inthe studiesreported ontheutilization ofbothpentose sugars in
chicks, mainly feed intake, weight gain, and metabolizable energy were
measured (Wagh &Waibel, 1966; 1967a; Longstaff et al., 1988;Schutte,
1990).Fromthesestudies,itappearsthatbothpentosesugarsarelesswell
utilized than glucose by poultry and may induce unwanted nutritive
problemstogetherwithwetdroppings.Researchontheabsorptionofxylose
and arabinosefrom theintestinal tract ofchicksislimited. Bogner(1961)
andWagh&Waibel(1967b)studied absorption ofxyloseand arabinose in
youngchicksbyanalyzingthegastrointestinalcontentsat30minutes after
oral administration by crop-tube.Their data showed that arabinose was
absorbed at a lower rate than glucose and xylose, whereas absorption
velocityofxylosewasonlyslightlylowerthan that ofglucose.
The present trial was designed to get more information on the quan-
titative aspectsofdigestionandutilizationofD-xyloseandL-arabinose at
dietaryinclusionlevelsof2.5,5.0, 7.5and 10.0%infowl.Forthis purpose
ileostomized adultroosterswere usedinordertomeasurethe disappear-
ancerateof D-xylose and L-arabinose atthe endoftheterminal ileum,
and to determine the urinary excretion of these sugars. D-glucose was
includedinthetrial asa reference.
Birdsand Housing
Adult ileostomized roosters ("Lohmann Meat", Lohmann GmbH,

Cuxhaven,Germany)of3.5to4.0kgbodyweightwereused.Thebirdswere
housedindividually inwire-meshcages(40x75x60cm,widthxheightx
depth), which were provided with a feed and water bowl.The cages were
Pentose sugars inadult roosters 43
situatedinaninsulatedroomwithfacilities forcontroloftemperature and

light.Birdsweremaintainedundera16hlight:8htwilightcyclethroughout.
Thetemperature intheroomwaskept at approximately 20°C.
Surgical procedures
Birds were ileostomized by a modification of the simple T-cannula

procedure of Raharjo &Farrell (1984). Prior to the surgery, birds were
starvedfor12handpremedicatedforanesthesiabyintramuscular injection
of0.8mL/birdofROMPUN (BayerAG.Leverkusen,Germany).Anesthesia
was induced and maintained byinhalation ofhalothane evaporated in02
andN20.Allfeathers wereremovedfrom theabdominalwallcaudaltothe
sternum,andtheareawasdesinfectedwithaBetadinesolution.Laparotomy
wasperformed bya3cmincisionventraltothetipoftherightpubicbone.
Thedistalileumwasexteriorizedandtransectedabout5cmanteriortothe
ileocaecaljunction.Next,thestumpswereclosedusing"Vicrylsurgical gut
(size 4).Alongitudinal incision of minimum length, to receive the ileal
flange,wasmadeintheantimesentericborderoftheileumnearthesealed
stump.Nextapurse stringsuture ofsize00surgicalgutwas incorporated
into the sub-mucosa of the ileum proximal to the incision. The silicone
rubber cannula, with an inside diameter of approximately 8 mm, was
insertedintheintestinallumenthroughtheincision,andthepurse string
suturewasdrawnsnuglyaroundthebarrelofthecannula.Asecondpurse
stringplacedabout 1mmfrom thefirst one,completedtheinversionofthe
secretorymucosa.Theintestinewasthenreplacedinthebodycavityandthe
muscletissuewassuturedusingVicrylsurgicalgut(size4).Finallytheskin
was closed with non-absorbable sutures (size 4).Arubber "O" ring was
placedatthebaseofthecannulatoprevent movementofthecannula. For
collectionofdigesta,ascrew-capvialwasattachedtotheendofthecannula
barrel. For collection ofurine, a funnel was fitted into a flask and placed
directlybeneath eachcage.
Postoperative care included keeping the birds warm and withholding
feed for 24h.Birdswereallocated tothetest 4weeksafter surgery.
Diets
Thebasal dietusedwasbasedoncorn,wheat starch andisolated soya

protein(Table1)andcalculatedtobeadequateinallnutrientsfollowingthe
recommendations ofAgricultural Research Council(1975)and National
Research Council (1984). The three sugars (D-glucose, D-xylose and L-
arabinose)were tested at dietary inclusion levels of2.5,5.0, 7.5and10%.
44 Chapter 2
Thesugars,supplied asanhydrousmonosaccharides,weresubstituted by
weightforwheatstarch.Thefeedwassuppliedasdrymashandfedatadaily
rate of 140g(= 125.6gofdrymatter) toeachbird throughout. The daily
amount of feed wasoffered as two equal meals at 08.00hand 20.00h.
Waterwasavailable adlibitum access.
TABLE 1.Composition of the basal diet.
Ingredient Percentage
Corn 39.00
Wheat starch 38.75
Soy oil 2.00
Animal fat 2.00
Isolated soya protein (88%CP) 12.00
Ground limestone 1.25
Monocalcium phosphate 2.10
Salt, iodised 0.30
Potassium bicarbonate 1.50
Vitamin-mineral premix 1 1.00
DL-methionine 0.10
Calculated contents:
ME, kcal/kg 3390
Crude protein, % 14.40
Calcium, % 0.90
Phosphorus, % 0.70
Lysine, % 0.78
Methioninepluscystine,% 0.59
Supplied perkilogram ofdiet:vitaminA,10,000IU;vitamin D 3 ,

2,000ICU;dl-a-tocopherylacetate,30mg;menadione,5mg;thiamin,
2.5mg;riboflavin, 5.5mg;d-pantothenicacid, 15mg;niacinamide,
50mg;cobalamin, 15ug;cholinechloride, 1,850 mg;pyridoxine,3mg;
biotin,0.15mg;folic acid,0.75mg;ascorbicacid, 50mg;inositol,
100mg;para-amino-benzoic acid,2.5mg;ethoxyquin, 100mg;avoparcin,
15mg;Mg,420mg;Mn,95mg;Zn,50mg;Fe,40mg;Cu,40mg;Co,2mg;
Se,0.1mg.
Pentose sugars inadult roosters 45
Experimental design
Asisillustrated inTable2,duringtheexperimental periodthesametype

ofsugarwasfedtoeachbirdinthesequenceof2.5(phase 1),5.0(phase2),
7.5 (phase 3) and 10.0%(phase 4) of the diet. Each of the four phases
consistedofapre-testperiodof7daysandatestperiodof4days.Duringthe
first 3daysofthepre-testperiod,birdsweregraduallychangedtothenext
dietcontainingahigherinclusionlevelofthesamesugar.Duringthe4x24
h test period digesta and urinewere collectedquantitatively for eachbird
separatelyatintervalsof2h.Thedigestaandurinewereweigheddailyand
stored at -25C.
TABLE 2.Experimental design
Treat- Sugar Number Dietary sugar level(%)

ment of phase 1 phase 2 phase 3 phase 4
birds (l-lldays) (12-22days) (23-33days) (34-44days)
A D-glucose 3 2.5 5.0 7.5 10.0

B D-xylose 3 2.5 5.0 7.5 10.0
C L-arabinose 3 2.5 5.0 7.5 10.0
ChemicalAnalysis
Eachdietwasanalyzedfordrymatterandnitrogen(N);thedietscontaining
5 and 10%of the test sugars also were analyzed for gross energy (GE).
Digesta and urine wereanalyzed for the content ofN,glucose,xylose and
arabinose.The digesta were analysed for drymatter. In addition, digesta
and urine ofbirds fed diets containing 5and 10%ofthe test sugars were
analyzedforGE.Theanalysesofdigestaandurineweredoneseparatelyfor
eachbird.
DrymatterandNanalyseswereperformed accordingstandardmethods
(AssociationofOfficialAnalyticalChemists, 1984).GEwasdeterminedby
using an IKA-C4000 adiabatic bomb calorimeter (IKA-Analysentechnik
GmbH,8044Unterschleissheim, Germany).
Sugarconcentrationsindigestaandurineweredeterminedassilylderi-
vatives ofmonosaccharides bygas liquid chromatography (Sweeley et al,
46 Chapter2
1963).Aknownamountofwetdigesta(lg)orurine(1mL)wasdilutedwith
distilled water(1:10). The diluted samplewasthendeproteinized with
potassium ferrocyanate and zinc-acetate and desaltedbypassing through
amixture(1:1w/w)ofanion(BioradAG3x4)andcation(BioradAG50W
x4exchanger.After centrifugation, 200uLofthe supernatant was freeze
dried.Tothefreeze driedsamplephenylglucopyranoside (0.4mgina 1mL
pyridinesolution)wasaddedasaninternalstandard.Thesamplewasthen
derivatized by the addition of 0.6 mL hexamethyldisilazane and 0.3 mL
trimethylchlorosilane. Next the contents were stirred ona Vortex stirrer.
Afteranincubationperiodof30minatroomtemperature,thereagentswere
removedbyevaporationwithnitrogenat40°C.Theresiduewasredissolved
in0.5mLethylacetate.Fromthissample,2uLwasanalyzedbyusingagas
liquid chromatograph (Model HP 5890, Hewlett-Packard Co., PaloAlto,
CA.), equipped with a flame ionization detector and integrator (Model
3396A,Hewlett-Packard Co.,PaloAlto,CA.).Thecarbohydrate derivatives
were separated with a Chrompack capillary WCOT fused silica column
(Chrompack,Middelburg,TheNetherlands.) coatedwithCPsil5CBof50
mlength.Hydrogenwasusedascarriergas.Theoventemperaturewasheld
for3minat190°C,thenraisedattherateof5°C/mintoafinal temperature
of265° C,which washeld for 5min.Thetemperature ofthe injector and
detector was 240°and 300°C,respectively.
Calculations
Ileal digestibilities ofsugars, drymatter, nitrogen and energywere

expressed asdigestibility coefficients (DC)usingthe equation:
j-jp _ Nutrient consumed -nutrient excreted in digesta .,«~

Nutrient consumed
Rates of urinary excretion of sugars and energy were expressed as

percentage ofthe intake ofsugars and energy,respectively.
Alldata wereanalyzedbyanalysis ofvariance usinga split-plot design

model,inwhichtheanimalsarethewholeplots(Cochran&Cox,1957).The
computerprogramGENSTAT5(ReferenceManual1987,OxfordUniversity
Press, New York) was used to calculate the analysis of variances. The
treatment factors were typeofsugar and the dietary levelofsugar.If the
Pentose sugars in adult roosters 47
number ofdietary sugarlevelswasgreaterthantwo,the sum ofsquares for

levelswaspartitionedintoasetoforthogonallinearandquadraticpolynomial
regression components. The dietary level was confounded with time and
age. It was assumed however, that differences between treatment groups
wereduetotheincreaseindietarysugar.Statementsofstatistical significance
were based on P <0.05.
RESULTS
Themeanvaluesfordrymatter andwaterintake,outputofwetdigesta and

urine,and drymatter contentofdigestainbirdsfedtheD-glucose,D-xylose
and L-arabinose diets are given in Table 3.The birds consumed their
TABLE 3.Intake of dry matter and water, output and dry matter
content of digesta, and output of urine in adult roosters fed
on D-glucose, D-xylose, or L-arabinose diets.
Sugar Dietary Intake Intakeof Output of Dry matter Outputof

level ofdry water wet digesta content urine
matter digesta
% (g/bird/day) (%) (g/bird/day)
D-glucose 2.5 115.8 ± 7.0 888 ± 2 1 9 110.4 ± 26.4 12.0 ± 3.3 622 ± 2 1 4
5.0 123.6 ± 3.0 862 ± 48 122.0 ± 32.8 10.8 ± 2.3 663 ± 1 1 4
7.5 124.9 ± 2.2 898 ± 2 1 9 124.3 ± 16.2 10.9 ± 1.8 661 ± 1 4 8
10.0 122.0 ± 4.5 859 ± 2 2 0 125.8 ± 25.1 10.7 ± 1.0 603 ± 1 1 9
X 121.6 a 877" 120.6 a 11.1" 637 a
D-xylose 2.5 113.8 ± 2.1 933 ± 1 9 0 113.2 ± 30.0 11.8 ± 3.8 616 ± 1 7 0
5.0 116.4 ± 8.2 1054 ± 1 5 2 125.9 ± 29.9 10.2 ± 1.8 733 ± 94
7.5 119.2 ± 5.6 1144 ± 1 5 0 136.2 ± 25.5 9.9 ± 1.5 767 ± 1 3 0
10.0 122.3 ± 6.0 1375 ± 41 160.8 ± 15.2 8.6 ± 1.2 915 ± 93
X 117.9 a 1127 a 134.0 a 10.l a 758 a
L-arabi- 2.5 113.1 ± 2 . 8 744 ± 147 136.8 ± 15.7 10.4 ± 3.1 448 ± 168
nose 5.0 121.0 ± 2.1 939 ± 100 182.8 ± 21.0 8.7 ± 1.9 555 ± 169
7.5 120.0 ± 2.4 1020 ±211 223.0 ± 35.4 8.1 ± 1.8 542 ± 1 5 1
10.0 118.2 ± 4.3 1086 ± 172 277.9 ± 23.8 6.7 ± 1.0 565 ± 100
X 118.0 s 947 a 205.l b 8.5 a 528 a
ab
Means (± SD) within a column with no common supercript differ
significantly (P 0<.05).
48 Chapter 2
dailyfeedallowanceof125.6gdrymatterperbirdwell.Waterintake, output
ofwetdigestaandurine,anddrymatter contentofdigestawerenot affected
significantly by the dose level ofD-glucose. When roosters were fed diets
containing D-xylose,the water intake and output ofwet digesta and urine
werelinearly(p<0.05)increased,whereasdrymattercontentofdigestawas
linearly(p<0.05)decreasedasthelevelofthispentosesugarincreased. The
same pattern of response were observed for water intake, output of wet
digesta and dry matter content of digesta when roosters were fed diets
containing L-arabinose. The output of urine was not affected by the dose
level of L-arabinose.
Ileal digestibilities of D-glucose, D-xylose and L-arabinose, and the
urinary excretion ofthese sugars are shown in Table 4.Digestibility ofD-
glucose and D-xylose was nearly 100%. However, ileal digestibility of L-
arabinose decreased linearly (P <0.05) as the dietary level ofthis pentose
sugar increased. The mean difference in digestibility between L-arabinose
and the other two sugars was significant (P < 0.05). Data for urinary
excretion ofsugars are alsoshown in Table 4.Small traces ofglucose were
found in the urine of all experimental treatments, suggesting that these
originatedfrom thebasaldiet.Bothpentosesugarswerepartlyexcretedvia
the urine.The extent ofthe urinary excretion ofD-xylose and L-arabinose,
in percentage ofintake, was linearly (P<0.05) related todose.In birds fed
the 2.5%D-xylosediet, 7.2%ofthexyloseconsumed appeared inthe urine.
This level increased to 20.2% when the 10% D-xylose diet was fed. The
urinary excretion ofL-arabinose, as a percentage ofintake, increased from
8.7to16.6%whenthedietarylevelofthispentosesugarwasincreased from
2.5 to 10%.
Ileal digestibilities ofdry matter and N are shown in Table 5. Ileal
digestibility of dry matter in birds fed diets containing L-arabinose was
consistently lower as compared to birds fed D-glucose or D-xylose diets.
Digestibilities ofthe latter twowere not different. Dry matter digestibility
ofthebirdsfed L-arabinose decreasedlinearly(P<0.05)asthedietary level
was increased. The mean difference in dry matter digestibility between L-
arabinoseandtheothertwosugarswassignificant (P<0.05).The differences
in N digestibility among the sugar treatments were not significant.
Results for ileal digestibility and urinary excretion ofenergy are shown
in Table 6. Ileal digestibility of energy of the diets containing either D-
glucose or D-xylose was veryhigh with no appreciable differences between
TABLE4.Ilealdigestibilityandurinaryexcretionofglucose,xylose
and arabinose in adult roostersfed the experimental diets.
Sugar Dietary Digestibility Urinary excretion

level sugars Glucose Xylose Arabinose
(%) (%) (%of intake)
D-glucose 2.5 98.0 ± 0.3 +1)

5.0 99.3 ± 0.0 + - -
7.5 99.5 ± 0.2 + - -
10.0 99.8 ± 0 . 1 + - -
X 99.2*
D-xylose 2.5 99.7 ± 0.4 + 7.2 ±2.2 .

5.0 100.0 ± 0.0 + 13.1 ±1.9 -
7.5 99.7 ± 0.2 + 16.9 ±2.7 -
10.0 99.6 ± 0.5 + 20.2 ±2.0 -
X 99.8 a 14.4
L-arabinose 2.5 95.5 ± 4.0 + _ 8.7 ±3.9

5.0 93.6 ± 5.4 + - 11.2 ±3.6
7.5 80.3 ± 7.9 + - 13.9 ±7.1
10.0 74.6 ± 7.0 + - 16.6 ±5.2
X 86.0 b 12.6
1) Smalltraces(0.09-0.13%)ofglucosewerefound intheurine ofall

experimental treatments.
a,b Means (±SD)within acolumnwith nocommonsuperscript differ
50 Chapter 2
TABLE5.Ileal digestibility ofdietary drymatter andnitrogen in

adult roostersfed the experimental diets
Sugar Dietary Digestibility

level Dry matter Nitrogen
(%) (%)
D-glucose 2.5 89.0 ± 1.2 89.6 ± 1.5

5.0 89.7 ± 1.4 90.0 ± 0 . 8
7.5 89.1 ± 2.3 90.0 ± 1.4
10.0 89.1 ± 1.0 90.1 ± 1.1
X 89.2 a 89.9 a
D-xylose 2.5 88.9 ± 0.7 89.2 ±0.7

5.0 89.2 ± 0.9 90.3 ±0.2
7.5 88.7 ± 0.8 89.8 ±0.5
10.0 88.7 ± 1.9 90.3 ±2.4
X 88.9 a 89.9 a
L-arabinose 2.5 87.7 ± 2.7 88.0 ±4.1

5.0 87.1 ± 2.2 87.1 ±3.6
7.5 85.2 + 1.7 87.1 ±3.7
10.0 84.5 + 1.2 88.1 ±2.8
X 86.l b 87.6 a
1,b
Means (±SD)within a columnwith nocommonsuperscript are
significantly different (P<0.05).
Pentose sugarsin adult roosters 51
TABLE6.Ilealdigestibilityandurinaryexcretionofdietary energy
in adult roosters fed the experimental diets
Sugar Dietary Ileal digestibility Urinary excretion

level(%) energy ofenergy
(%) (%) (%of intake)
D-glucose 5 91.7 ± 0.2 2.8 ± 0.8

10 91.6 ± 0.9 2.4 ± 0.9
X 91.6 a 2.6a
D-xylose 5 91.7 ±0.3 4.3 ± 0.9

10 91.4 ±1.6 5.2 ± 0.4
X 91.6 a 4.7 b
L-arabinose i 5 88.9 ±2.9 3.7 ±1.2

10 87.9 ±1.6 4.3 ±1.8
X 88.4b 4.0 ab
a,b
Means(±SD)within a columnwith nocommonsuperscript are
theD-glucoseandD-xylosediets.Comparedtothesediets,significantly (P
<0.05)lowerdigestibilityvaluesfordietaryenergywerefound when diets
containingL-arabinosewerefed.Theurinaryexcretionofenergyofbirdsfed
dietswitheitherD-xyloseorL-arabinosewasgreaterthanthatofbirdsfed
dietssupplemented with D-glucose.However,onlythe difference between
the D-xyloseand D-glucose treatments wassignificant (P<0.05).
DISCUSSION
Thechoiceofthe experimental designneedstobeconsideredfirst. Latin

squares are often used asan experimental designinbalance studies with
animals,inwhichthe dietsarefed insequence,with the diet sequence
beingdifferent foreachanimal.TheadvantagesofusingLatinsquares are
thatvariationbetweenanimalsandthosearisingfromacommontimetrend
canbeequilibrated.However,thisisonlytruewhentherearenocarry-over
52 Chapter 2
effects. Theresults ofaprevious study (Schutte, 1990)showedthat water

intake and ceca weight of birds fed on D-xylose or L-arabinose diets
increasedmarkedly.Thus,carry-overeffects ofbothpentosesugarscannot
becompletelyexcluded.Therefore,inthepresenttrial,eachbirdwasfedthe
same type of sugar in increasing dietary levels during the experimental
period. Moreover, in previous studies performed at the authors' institute
(Schutte, J.B. &P.van Leeuwen, unpublished data),noindications were
found that digestibility of nutrients in ileostomized adult roosters was
changed with increasing age.The results ofileal Ndigestibility (Table5)
showingthatthecoefficients ineachsugarblockwerealmostconstant,also
indicatethatthepresentdatafordigestibilitywerenotconfoundedbytime.
Themainobjectives ofthepresent trial weretodetermine the ileal
digestibilityandurinaryexcretionofD-xyloseandL-arabinoseinbirds.The
dietaryinclusionlevelschosen(2.5to10.0%)wereapproximatelyonetofour
timeshigherthanthelevelswhichwouldbereleasedbycompletehydrolysis
oftheNSPinpoultrydietsbasedoncereals,soybeanoilmealandcerealby-
products.Theilealdigestiondataofthetestsugars(Table4)indicated that
D-xylosewasaswellabsorbedfromthesmallintestineofbirdsasD-glucose.
Theilealdigestibilityvaluesfordrymatterandenergy(Tables5and6)were
consistentwiththisfinding,becausetherewerenodifferences betweenthe
D-glucoseandD-xylosetreatmentgroups.Theilealdigestibilitydataofthe
present study also demonstrated that L-arabinose was not absorbed
completelyfrom thesmallintestine.Inaddition,absorptionofL-arabinose
wasdoserelated. Theileal digestibility valuesfor drymatter and energy
(Tables5and6)wereinagreementwiththisfinding, showingadecreasein
drymatter and energy digestibility when the dietary level ofL-arabinose
was increased. The decrease in ileal digestibility of L-arabinose, as the
dietary level increased, was associated with an increase in output ofwet
digesta.Similarresultswerefoundinastudyperformed recentlywithpigs
(Schutte, J.B. et al., unpublished data). This phenomenon of increased
outputofwetdigestacausedbyincreasingthedietarylevelofL-arabinose
may be connected with the osmotic properties of unabsorbed arabinose
resulting in an inflow of water into the intestinal lumen. However, the
presence of unabsorbed arabinose in the intestinal tract may stimulate
microbial activity, which also may result in an increased output of wet
digesta.Theincreaseinwetdigestaoutputbyincreasingthe dietarylevel
ofD-xylose(Table3)mayprovidesomeevidenceofanincreased microbial
activity whenincludingpentose sugars inthe diet. Longstaff et al.(1988)
and Schutte (1990)reported that ceca ofbirdsfed L-arabinose diets were
heavier than those ofbirds fed diets containing D-glucose.The same was
truewhenbirdswerefeddietscontainingD-xylose,buttheeffects were less

pronounced than on diets containing L-arabinose. Thus, these results
suggest that inclusion of either L-arabinose or D-xylose in the diet may
result in an increased microbial activity in the intestinal tract ofbirds.
The digestibility data collected in the present study agree with the
absorption data reported by Bogner (1961) and Wagh & Waibel (1967b)
despite the differences in experimental technique.However,their data are
not strictly comparable with the results ofthe present study,because their
observations are based on an absorption period of only 30 min. Studies
performed with rats (Cori, 1925: Miller & Lewis, 1932) suggested that
absorption ofsugars will not be completed fully during this period.
Theresults discussed, suggest that D-xylose and L-arabinose,inspiteof
theiridenticalmolecularsize,haveadifferent modeoftransportinthe small
intestineofbirds.Itisgenerallyaccepted(Crane,1960;Herman, 1974) that
L-arabinoseispassivelyabsorbedinthesmallintestineofanimals.However,
the mode of transport of D-xylose from the animal intestine is still
controversial. The idea that D-xylose crosses the intestinal mucosa by
simple diffusion (Wilson &Vincent, 1955;Crane, 1960;Finlay et al., 1964)
wasinitiallyreported;however,thebulkofevidencesuggeststhatthis sugar
sharesthesametransportmechanismwithD-glucose(Salomonetal.,1961;
Csaky &Lassen, 1964;Alvarado, 1966;Bihler et al., 1969).
It isgenerally accepted that D-glucose canbeutilized almost completely
inhuman and animals (Demetrakopoulos &Amos,1978),soonly negligible
amountsofglucosewillbefoundintheurine.Thelatterisinagreement with
the finding in the present study.It iswellrecognized that inman, rats and
pigs, part of the ingested D-xylose and L-arabinose appears in the urine
(Loos, 1954;Wiseet al., 1954;Arnal-Peyrot &Adrian, 1974;Haeney et al.,
1978).However, little information is available onthe relationship between
the dietary concentration of D-xylose or L-arabinose and the urinary
excretion ofthese pentose sugars. In the present study, this relationship
was clearly demonstrated and supported by the results of the urinary
excretion ofenergy (Table 6).The results ofan energy balance study with
chicksofWagh&Waibel(1966)areconsistent withthis,showinga decrease
in metabolizable energy ofthe two pentose sugars with increasing dietary
inclusionlevel.Thelatterwasconfirmed inarecentstudyofSchutte (1990).
Only a few reports are available concerning the metabolism of pentose
sugarsinbirds.Longstaff etal.(1988)reportedthatchickswereabletogrow
well on diets containing D-xylose or L-arabinose at dietary concentrations
of 5%. Radioisotope studies of Wagh and Waibel (1967a) showed that L-
arabinose was better metabolized than D-xylose by chicks, but neither
pentose sugar was metabolized to C0 2 as rapidly as D-glucose.
54 Chapter2
ACKNOWLEDGMENTS
The authors wish to thank J.M. Fentener van Vlissingen for surgical
assistance. The assistance ofG.B. Derksen for the statistical analyses is
gratefully acknowledged.
REFERENCES
AgriculturalResearchCouncil(1975).TheNutrientRequirementsoffarm
livestock.No.1:Poultry.HerMajesty's Stationary Office, London.
Alvarado,F.(1966).D-xyloseactivetransport inthehamster small
intestine.Biochimica etBiophysicaActa 112: 292-306.
Arnal-Peyrot,F.&Adrian,J.(1974).Metabolismedespentosanesdecergale
chezlerat. (metabolism ofcereal pentosans inrat). International
Journal forVitamin and Nutrition Research 44:543-552.
Association ofOfficial Analytical Chemists (1984).Official Methodsof
Analysis, 14thed.Washington,DC:Association ofOfficial Analytical
Chemists.
Bihler,I.,Kim,N.D.&Sawh,RC.(1969).ActivetransportofL-glucoseand
D-xyloseinhamster intestine invitro.Canadian Journal ofPhysiology
and Pharmacology 47:525-531.
Bogner,P.H.(1961).Alimentaryabsorption ofreducingsugarsbyembryos
andyoungchicks.Proceedings ofthe SocietyofExperimental Biology
and Medicine 107: 263-267.
Booth,AN.,Wilson,R.H.& Deeds,F.(1953).Effects ofprolonged
ingestion ofxyloseonrats.Journal ofNutrition 49:347-355.
Brillout,J.M.,Rouau,X.,Hoebler,C, Barry,J.L., Carre,B.& Lorta, E.
(1988).Anewmethod for determination ofinsoluble cellwalls and
solublenon-starchy polysaccharides from plant material.Journalof
Agricultural andFoodChemistry 36:969-979.
Carr6,B.&Leclercq,B.(1985).Digestionofpolysaccharides, protein
and lipidsbyadult cockerelsfed ondietscontainingapecticcell-
wallmaterial from whitelupin (Lupinus albusL.)cotyledons. British
Journal ofNutrition 54:669-680.
Carre\ B.&Brillouet,J.M. (1986).Yieldandcomposition ofcellwall
residues isolated from variousfeedstuff's used fornon-ruminant farm
animals.Journal ofthe ScienceofFoodandAgriculture 37:341-351.
Chesson,A.(1987).Supplementary enzymestoimprovetheutilizationof
pigand poultry diets.Pages 71-90in:RecentAdvancesinAnimal
Nutrition. (W. Haresign & D.J.A.Cole,editors).London, Butterworths.
Cochran, W.G. &Cox, G.M. (1957).Experimental Designs, 2nd ed. John

Wiley and Sons,Inc.NewYork.
Cori, F.(1925).The fate ofsugar in the animal body. 1.The rate of
absorption ofhexoses and pentoses from the intestinal tract. Journal
ofBiological Chemistry 66:691-715.
Crane,R.K.(1960).Intestinal absorption ofsugars. Physiological Reviews
40: 789-825.
Csaky,T.Z. &Lassen, U.V.(1964).Active intestinal transport ofD-xylose.
Darby, W.J. &Day, P.L. (1939).Xylose as a cataractogenic agent.
Proceedings ofthe Society ofExperimental Biology and Medicine41:
507-508.
Demetrakopoulos, G.E. &Amos,H.(1978).Xyloseand xylitol, metabolism,
physiology and nutritional value.WorldReview Nutrition and Dietetics
32:96-122.
Finlay,J.M.,Hogarth,J. &Whightman, K.J.R. (1964).Aclinical evaluation
ofthe D-xylose tolerance test.Annals ofInternal Medicine 61: 411-422.
Haeney, M.R., Culank, L.S.,Montgomery, R.D.&Sammons, H.D. (1978).
Evaluation ofxylose absorption as measured in blood and urine:Aone-
hour blood xylose test in malabsorption. Gastroenterology 75:393-400.
Herman, R.H. (1974).Hydrolysis and absorption ofcarbohydrates, and
adaptive responses ofthejejunum. In: Sugars in Nutrition, pp. 145-186
(H.L. Sipple and K.W. MacNutt, editors). NewYork,Academic Press.
Longstaff, M.&McNab,J.M.(1986).Influence ofsiteandvarietyon starch,
hemicellulose and cellulose composition ofwheats and their
digestibilities by adult cockerels. British Poultry Science, 27:435-449.
Longstaff, M., Knose,A. & McNab,J.M. (1988).Digestibility of
pentose sugars and uronic acids and their effect on chick weight gain
and caecal size. British Poultry Science 29:379-393.
Longstaff, M. &McNab,J.M. (1989).Digestion offibre polysaccharides of
pea (Pisum sativum) hulls, carrot and cabbage by adult cockerels.
British Journal ofNutrition 62:563-577.
Loos, M. (1954). Studies in the utilization ofpentoses in diabetes.Acta
Medica Scandinavica 148:425-431.
Miller, M.M. &Lewis, H.B. (1932).Pentose metabolism. 1.The rate of
absorption ofD-xylose and the formation ofglycogen in the organism
ofthe white rat after oral administration ofD-xylose.Journal of
Biological Chemistry 98:133-140.
NationalResearchCouncil(1984).Nutrientrequirementsofpoultry.National
Academy Press, Washington, DC.
56 Chapter 2
Raharjo, Y.&Farrell, D.J. (1984).Anew biological method for

determining amino acid digestibility in poultry feedstuff's by using a
simple cannula, and the influence ofdietary fibre on endogenous amino
acid output.Animal Feed Science Technology 12:29-45.
Salomon,L.L.,Allums,J.A.&Smith,D.E.(1961).Possiblecarriermechanism
for the intestinal transport ofD-xylose. Biochemical and Biophysical
Research Communications 4:123-126.
Schutte, J.B., Kempen van, G.J.M. &Hamer, R.J. (1990).Possibilities to
polysaccharides for poultry. In: Proc. VIII. European Poultry Confe-
rence, p.p. 128-135.Feria de BarcelonaAvda.Reina M .Cristina, s/n.
Schutte, J.B. (1990).Nutritional implications and metabolizable energy
value ofD-xylose and L-arabinose in chicks. Poultry Science 69:1724-
1730.
Sweeley, C.C., Bentley, R., Makita, M. &Wells,W.W.(1963). Gas-liquid
substances. Journal oftheAmerican Chemical Society 85: 2497-2507.
Vogt,H &Stute, K.(1971).Uber die Verdaulichkeit einiger
Kohlenhydratfraktionen (Zucker, Starke, Pentosane, Rohcellulose,
Lignin) im Huhnerfutter. Archiv fur Geflugelkunde 35:29-35.
Wagh,P.V.&Waibel,P.E.(1966).Metabolizability and nutritional implica-
tions ofL-arabinose and D-xylose for chicks.Journal ofNutrition 90:
207-211.
Wagh,P.V.&Waibel,P.E.(1967a).Metabolism ofL-arabinose and D-xylose
Wagh,P.V.&Waibel,P.E.(1967b).AlimentaryabsorptionofL-arabinose and
D-xylose in chicks. Proceedings ofthe Society ofExperimental Biology
and Medicine 124:421-424.
Wilson, T.H. &Vincent, T.N. (1955).Absorption ofsugars in vitro by the
intestine ofthe golden hamster. Journal ofBiological Chemistry 216:
851-866.
Wise,M.B.,Barrick, E.R., Wise, G.H. &Osborne, J.C. (1954).Effects of
57
CHAPTER3
NUTRITIONAL VALUE OF D-XYLOSE AND

L-ARABINOSE FOR BROILER CHICKENS
J.B.Schutte,J.deJong,E.J.vanWeerdenandM.J.vanBaak
TNO-InstituteofAnimalNutritionandPhysiology(ILOB),
P.O.Box 15,6700AAWageningen,TheNetherlands
Acceptedforpublicationin:BritishPoultry Science
Reproducedbypermission ofTheBritishPoultryScienceLtd.
59
NUTRITIONAL VALUE OF D-XYLOSE AND

L-ARABINOSE FOR BROILER CHICKS
J.B. Schutte, J. de Jong, E.J. van Weerden and M.J. van Baak

P.O.Box 15,6700AA Wageningen,The Netherlands
Twoexperiments wereconducted toexamine the effect offeeding D-xylose

and L-arabinose on broiler performance, body composition, caecal length
and weight, and liver weight. Graded amounts (25, 50 and 75 g/kg) of D-
xyloseorL-arabinosewereaddedtoeitherapracticaltype(ExptA)orasemi-
purified (Expt B)basaldiet.Asreference, a diet containing 75gD-glucose/
kg was included in the experiments. Both experiments were conducted in
batterybrooders,andbirdsreceivedtheisocaloric(onMEbasis)dietsas dry
mash ad libitum from 6to 27days ofage.Anegative dose-dependent effect
ofbothpentosesugarsonweightgainandfeedutilizationwasobserved.The
samewastruefordailyfeedintakeoftheD-xylosetreatments.Water intake
increased linearly (P<0.05) asthe dietarylevelofbothpentose sugars was
increased.Consequently,drymattercontentofthedroppingsdecreased. Fat
content of the chick body tended to decrease when either D-xylose or L-
arabinosewasincludedinthediets.Caecalweightwasincreased markedly
byfeeding L-arabinose. Liver weight was not affected byfeeding either D-
xylose or L-arabinose to birds. From data for ME intake and gain in body
energy it was estimated that utilization ofthe ME ofboth pentose sugars
was inferior tothat ofD-glucose. (Keywords:pentose sugars, D-xylose, L-
arabinose, nutritional value, chicks)
INTRODUCTION
Inmonogastricanimals,themajor partofdigestiontakesplaceinthe small

intestine by the digestive enzymes of the host. These enzymes hydrolyze
most of the alimentary components with the exception of the nonstarch
polysaccharides (cellulose, hemicellulose, pectins, oligosaccharides, etc.).
The absence ofnonstarch polysaccharide (NSP) degrading enzymes in the
hostandthelowdensityofmicroorganismsinthesmallintestinemean that
the NSP will largely pass to the hindgut. In the hindgut the NSP are
degraded to a greater or lesser extent by the microbes for which they are
the major carbon source. The end products of this microbial degradation
(lacticacid,volatilefatty acids)are readily absorbed and canbea potential
60 Chapter 3
energy source for the animal. Contrary to pigs, in poultry microbial

degradationofNSPinthecaecaandcolonappearstobelow(VogtandStute,
1971;Carre"and Leclercq, 1985;Longstaff and McNab, 1989;Carr<5 et al.,
1990).
Fromaliterature review,Chesson(1987)concludedthat the digestibility
of feed ingredients containing high levels of NSP can be improved by
treatmentwithenzymeswhichcanhydrolyzetheNSPtomonosaccharides.
However, hydrolysis ofNSP will release not only glucose, but also other
sugarssuchasD-xyloseandL-arabinose,whicharenormallynotencountered
in the small intestine. Knowledge about the utilization of these pentose
sugars in poultry is incomplete. It is well recognized that both pentose
sugarsareabsorbedfrom theintestinaltractinbirds(Bogner, 1961;Wagh
andWaibel,1967;Schutteetal.,1991a).Ourdata(Schutteetal.,1991a)also
showed that part ofthe ingested xylose and arabinose is excreted in the
urine.Theextent ofthisurinary excretionofbothpentose sugars,in%of
intake, was increased linearly as the dietary level of these sugars was
increased (Schutte et al., 1991a). This may explain the decrease in
metabolisable energy(ME)valueofD-xyloseandL-arabinose inchicksby
increasing the dietary inclusion level, as observed by Wagh and Waibel
(1966)and Schutte (1990).This dosage-related MEvalue ofboth pentose
sugars in chicks was reflected in the results for weight gain and feed
conversion efficiency (Schutte, 1990).
Themain objective ofthe present study wasto investigate the effect of
dietaryinclusionlevelsof25,50and75g/kgofbothpentosesugarsonchick
performance, wherebythedosage-related MEcontent ofthese sugars was
taken intoaccount. Performance was studiedbyusingtwodifferent types
ofbasaldiet;apracticaltypeandasemi-purifiedbasaldiet.Inaddition,the
effect ofdietaryD-xyloseandL-arabinoseoncaecallengthandweight,and
liverweightwereexamined.Furtheronbodycompositionofbirdsfedonthe
practical dietswas determined.
Experimental
The composition of the two basal diets is presented in Table 1 and

calculatedtobeadequateinallnutrientsfollowingtherecommendationsof
theNationalResearchCouncil(1984).Withbothbasaldietsseventreatment
groupswereformedcontaining75gD-glucose/kg(referencediet)and25,50
and 75gofeither D-xyloseorL-arabinose/kg,respectively.
Thesugars,suppliedasanhydrousmonosaccharides,wereonaMEbasis,
exchangedfortapiocaandcelluloseinbasaldietAandforcornstarch and
celluloseinbasal diet B.TheMEvalue ofthe pentose sugarswas derived
from those determined in a previous study with chicks(Schutte, 1990) at
dietaryinclusionlevelsof50and 100g/kg.Basedonthesevalues,theME
contentofD-xyloseatdietarylevelsof25,50and75g/kgwasestimated to
be 12.6,11.1and 9.8MJ/kg,respectively.Thecorrespondingvalues for L-
arabinoseatthesedietarylevelswereestimatedtobe11.7,9.6,and7.7MJ
ME/kg.TheMEvalueofD-glucosewasassumedtobe15.1MJ/kg(Anderson
et al, 1958). The diets were fed ad libitum as dry mash. Water was also
availablefor adlibitum consumption.
Thetwotrials (codedasExptAandExpt B)were run parallel. Day-old
female broiler chicks ("Hybro") were used. The birds were housed in
electricallyheatedbatterycagesof0.975squaremetresoffloor spaceeach
withwirefloors.Thecagesweresituatedinaninsulatedroomwithfacilities
tocontroloftemperatureandhumidity.Chicksweresubjectedtocontinuous
artificial fluorescent illumination.Astandard dietofpracticalcomposition
wasfedforthefirst threedays,followedbyanotherthreedaysofamixture
ofthestandard dietandthebasaldietfedduringtheexperimental period.
At6daysofage,14birdswereallottedtoeachof56cagessuchthataverage
bodyweight(130g)andweightrange(120to150g)weresimilar.Treatment
groupsconsistedoffourcages,eachwith14birds,arrangedinarandomized
blockdesign.Theexperimental dietswerefedforaperiodof21days (from
6to27daysofage).
At the end ofthe trials, chicks were weighed individually, and feed con-
sumptionofeachcagewasrecorded.Duringthelast2daysoftheexperimental
period,waterconsumptionwasmeasured,andexcretawerecollected,both
foreachcageseparatelyatintervalsof12h.Waterintakewasmeasured as
thedifference betweenapredeterminedvolumeofwaterinthewater pans
andthatremaininginthepans.Excretawerecollectedquantitatively from
glasstraysintocontainersand storedat 4°C.Immediately after the2day
collectionperiodtheexcretawerepooledpertreatmentgroupandanalyzed
for drymatter content.
After termination of the trials, 100chicks ofExpt A and 40chicksof
Expt Bwere killed by injection ofT 61(Hoechst, Germany) after a feed
deprivation of 12 h. Water remained available during this period. The
sacrificed chicksoriginated from the treatments fed dietscontaining 75g/
kgofD-glucose,and25gand75g/kgofeitherD-xyloseorL-arabinose.The
birdswereselectedinsuchawaythattheyrepresentedthepopulationofthe
treatment groups. From 40 chicks of each experiment (= 8 chicks per
treatment group)thecaecawereexcisedandtheirlength andweightwere
62 Chapter 3
measured individually .Gross pathological examination ofthe liver and

kidneyswascarried outandweight oftheliverwasrecorded individually.
Theremaining60sacrificedchicks(=12chickspertreatmentgroup)ofExpt
Awere used for determination ofthe dry matter, nitrogen and crude fat
contents of the entire carcasses with feathers, blood and residual gut
contents.Bodyanalyseswereperformed inpooledsamplesof3chickseach.
Inordertoprovideinitialdataforbodycomposition,at6daysofage20birds
were sacrificed according to the same procedure as described previously.
Bodyanalysesinthesechickswerecarriedoutinpooledsamplesof5chicks
each.Thechickbodieswere stored at -20°C.
Chickbodysampling and analysis
The frozen chick bodies were minced in a MoermanAlexander mincer

(model,SSK45,TheNetherlands).Next30%(w/w)ofdiatomaceous earth
(celite)wasaddedtothemixturetoaidinbindinganddispersingthebody
fat.Thenthemixturewaspassedthroughamechanicalgrinderwitha2mm
dietoachieveadequatemixinganddesiredfineness.A500grepresentative
samplewastakenfromeachmixingbatchandthenfreeze-dried. Whendry,
the mass was equilibrated with air humidity, weighed and the weight
correctedforcelite.
TABLE 1.Composition ofthe basal diets (g/kg)
Ingredient Basal Basal

ExptA ExptB
Maize 158.6 287.3

Wheat 155 -
Tapioca 160 -
Wheat starch - 287
Soybeanoil 50 40
Animal fat 20 40
Soybean oilmeal (500g CP/kg) 240 -
Isolated soyprotein (880 gCP/kg) - 223
Sunflower meal (320g CP/kg) 20 -
Peas (220g CP/kg) 75 -
Herring meal (720g CP/kg) 20 -
Meat meal tankage (580g CP/kg) 30 -
Cellulose ("ArbocelB 800) 30 60
Molasses 10 -
Dicalcium phosphate 10 -
Monocalcium phosphate - 21
Limestone 6 12
Iodized salt 3 3
Potassium bicarbonate - 15
Vitamin-mineral mix 1 10 10
DL-methionine 2.4 1.7
Contents:
Crude protein (analysed, g/kg) 211 225
Metabolisable energy (calculated, MJ/kg) 12.4 14.1
Ca (analysed,g/kg) 9.3 8.9
P (analysed,g/kg) 6.2 5.9
Lysine (calculated,g/kg) 12.1 13.3
Methionine +cystine (calculated,g/kg) 8.8 9.0
11
Supplied per kilogram ofdiet:retinol 3mg,cholecalciferol 0.05mg,
dl-a-tocopherylacetate30mg,menadione5mg,thiamin2.5mg, riboflavin
5.5mg,d-pantothenic acid 15mg,niacinamide 50mg,cobalamin 0.015
mg,cholinechloride 1,850 mg,pyridoxine3mg,biotin0.15mg,folicacid
0.75mg, ascorbicacid50mg,inositol 100mg, para-aminobenzoicacid2.5
mg, ethoxyquin 100mg,avoparcin 15mg,Mg420mg,Mn95mg,Zn50
mg,Fe40mg,Cu40mg,Co2mg,Se,0.1mg.
64 Chapter 3
Finally the dry mass was further homogenized with a Brabander blender
(model, 880802, Germany), with a 1 mm die and then analyzed for dry
matter, fat and N.
Drymatter was determined bydryingthe samples toa constant weight at
101°C. Fat (ether extraction) and protein (N x 6.25) were determined by
standard methods (A.O.A.C., 1984). Standard methods were also used for
determining the percentage ofN, Ca and P in the basal diets.All analyses
were carried out in duplicate.
Calculations
In ExptA,gain in energy ofchickswas calculated from the differences in

protein andfat composition ofthechickbodiesbetweentheinitial and final
samples. Energy gains were computed from protein gain at 23.7 k J per
gram,andfat gainat39.0kJpergramasproposedbyAndersonetal.(1958).
Efficiency of ME utilization for gain in the dietary treatments was
calculated by using the following equation:
,,_. , . , . „ . - . Gain in energy

ME utilization for gain = w | _ wg
intake maintenance
Inthis equation the chick dailymaintenance need for energy was assumed
at 600 kJ ME per kg metabolic weight (Scheele et al., 1987).
The results of both trials were combined in the analysis of variance

(Cochran and Cox, 1957).Thiscanbejustified becausebothtrialswere run
parallel in the same experimental room.The computer program SPSS/PC
+V2.0 (Norusis, 1988)was used to calculate the analysis ofvariance. The
treatment factors were type ofdiet and sugar, and dietary level ofsugar. If
thenumber ofdietary sugarlevelswashigher than two,the sum of squares
for levels was partitioned into a set of orthogonal linear and quadratic
polynomialregressioncomponents.Allstatements ofsignificance are based
on a probability ofP <0.05.
RESULTS
ExptA
Table 2 summarises performance data collected. Weight gain and feed

utilizationwerereducedwheneitherD-xyloseorL-arabinosewasincluded
inthe diet.Thisreduction wassmall at a dietaryinclusionof25g/kg,but
increased linearly (P < 0.05) as the level ofD-xylose or L-arabinose was
increased. The same was true for daily feed intake of chicks fed diets
containingD-xylose,butnotforintakeofchicksfedL-arabinosediets.Water
intakeofchicksfedondietscontainingthepentosesugarswasatalldietary
inclusion levels significantly higher than those fed onthe reference diet.
Moreover,waterintakewasincreasedlinearly(P<0.05)asthedietarylevel
ofeither D-xyloseorL-arabinose wasincreased.However,theincrease in
water intake ofchicksfed onthe L-arabinose diets wasmore pronounced
thanofchicksfedontheD-xylosediets.Asaresultoftheincreaseinwater
intakefromincreasingdietaryD-xyloseorL-arabinose,drymatter content
ofthe excreta decreased steadily.
Mortalityrate wasverylow,only 1.5%ofthebirdsdied.Noappreciable
differences inmortality amongthetreatments were observed.
Drymatter, fat and protein contents ofthe chickbodiesarepresented in
Table3.Therewasatendencytoadecreaseddrymatter andfat contentof
thebodywhenchickswerefed ondietscontainingthepentose sugars.No
systematic effect ofD-xyloseorL-arabinose onthe protein content ofthe
bodywas observed. Utilization ofMEfor gain was decreased when birds
werefedondietscontainingeitherD-xyloseorL-arabinose.Thisdecreased
MEutilizationwasmorepronouncedinchicksfedontheL-arabinose diets
than in those fed diets supplemented with D-xylose. Utilization of crude
proteintendedtodecreasewheneitherD-xyloseorL-arabinosewasincluded
inthe diet.Becausefeed intakeofbirdswasnotrecordedindividually, the
dataforMEandproteinutilization couldnotbeanalyzed statistically.
66 Chapter 3
TABLE2.The effect of dietary inclusion levels ofD-xylose and

L-arabinose on chick performance from 6to27days of age by
usinga practical typebasal diet (ExptA).
Sugar Dietary Weight Feed Feed: Water Drymatter

level gain intake gain intake content
(g/kg) (g) (g/bird (g:g) (g/bird excreta
per day) per day) 1(g/kg)2
D-glucose 75 809 a 61.8 ab 1.605a 166a 279
D-xylose 25 776abc eo.o1* 1.6248 192b 226

50 751 c 58.6cd 1.638ab 206bc 203
75 716d 57.0d l^l1* 216cd 159
L-arabinose 25 806 ab 62.2a 1.620a 207bc 213

50 775bc 60.2abc 1.632ab 229d 191
75 750cd 60.3 abc 1.686c 257e 142
SEM(df42) 3 11.8 0.8 0.014 5.5
1
Means offour pensper treatment ofdays 26plus27.
2
Pooled samples per treatment ofdays 26plus 27;figuresnot analysed
statistically.
3
Standard errorofthe mean.
a,b,cMeanvalueswithnocommonsuperscripts within a column differ
TABLE 3.Effect ofdietary inclusion levels of D-xylose andL-

arabinoseon bodycompositionandutilizationofdietaryMEand
protein in chicks by using apractical typebasal diet (ExptA).
Sugar D-gluc. D-xylose L-arabinose Pooled

Dietary level (g/kg) 75 25 75 25 75 SEM
(df30)
Body composition (final

Dry matter (g/kg) 318" 308" 307" 304" 316" 5.3
Fat (g/kg) 107" 106" 102" 106" 102" 4.0
Protein (g/kg) 169"b 166"b 169"b 163" 172b 2.3
Utilization ofME for gain

Intake ofME (MJ/bird) 16.1 15.6 14.8 16.2 15.7
Maintenance (MJ/bird) 7.4 7.2 6.9 7.4 7.1
1
Gain in energy (MJ/bird) 6.8 6.5 5.9 6.6 6.2
Utilization (%)2 78.2 77.4 74.7 75.0 72.1
Utilization ofprotein
Intake ofprotein(g/bird) 274 266 252 276 267
Gaininprotein (g/bird)1 140 131 124 134 132
Utilization (%)3 51.1 49.2 49.2 48.6 49.4
11
Birds sacrificed at 6days toprovide initial data had following
composition;averagebodyweight 129g,fat 78.7g/kg,protein (Nx6.25)
149g/kg.
2)
Calculated accordingtoformula as described inthe sub-heading
"Calculations".
3)
Calculated as: protein gain
protein intake
ab
Meanvalueswithnocommonsuperscriptswithinarowdiffer significantly
(P<0.05).
68 Chapter 3
Data regarding caecal length and weight, and liver weight are shown in
Table4.Therewasatendencythatcaecalweightwasincreasedwhenchicks
werefeddietscontainingD-xylose.Thecaecafromchicksfedon L-arabinose
diets were not only longer but also heavier than those from chicks fed the
reference diet. The differences in caecal weights between birds fed on the
reference dietand thosefedontheL-arabinose dietsweresignificant. Liver
weight in % of live weight was not affected significantly by the dietary
treatments.Grosspathologicalexaminationoftheliverandkidneysdidnot
show abnormalities in birds fed on the different diets.
TABLE 4.The effect of dietary inclusion levels of D-xylose and L-

arabinoseoncaecallength andweight, andliverweight of chicks
by using a practical type basal diet (ExptA).
Sugar D-gluc. D-xylose L-arabinose SEM

Dietary level (g/kg) 75 25 75 25 75 (df70)
Live weight (LW) at 896 874 842 901 855 20.2

autopsy, g
Caecal length, cm 26 a 26 a 26 a 28 a 28 0.81
Caecal weight (incl. contents)

g 5.5a 5.6" 5.8a 7.2b 7.6b 0.47
%ofLW 0.62a 0.64ab 0.69ab 0.80bc 0.89c 0.06
Liver weight
g 19.9a 21.2a 20.2 a 22.3 a 21.2 a 0.93
%ofLW 2.2a 2.4a 2.4a 2.5 a 2.5 a 0.10
a,b,c
Mean values with no common superscipts within a row differ
significantly (P < 0.05).
ExptB
The performance data are summarised in Table 5.Almost similar results

for weight gain, daily feed intake, feed conversion efficiency, water intake
and drymatter contentofexcretawereobtained asinExptA.Again weight
gain and feed utilization were reduced when chicks were fed on diets
containing eitherD-xyloseorL-arabinose.Weightgainand feed utilization
was decreased linearly (P < 0.05) as the dietary level of D-xylose was
increased.DailyfeedintakeofbirdsontheD-xylosedietswasnotonlylower
as compared to birds fed on the reference diet, but also dose-dependent.
Water intake ofchicksfed either D-xylose orL-arabinose diets showed the
same pattern ofrespons as in ExptA.The increases in water intake when
chickswerefed thepentose sugarswerereflected inthe drymatter content
ofthe excreta.
Mortality rate was again low.As a mean 0.6%ofthe chicks died with no
appreciable differences among the treatments.
Results for caecal length and weight, and liver weight are shown in Table
6.Caecal length and weight were not clearlyaffected when chickswere fed
ondietscontainingD-xylose.However,whenfeddietscontainingL-arabinose,
caecalweightincreasedascomparedtothecaecalweightofchicksfedonthe
reference diet. The difference in caecal weight between the reference
treatment and the 75gL-arabinosse/kgdiet treatment was significant. In
addition,caecaofchicksfedonthe75gL-arabinose/kgdietwere significantly
longer than those ofchicks fed on the reference diet. Liver weight was not
affected significantly by the inclusion ofeither D-xylose or L-arabinose in
the diet. Gross pathological examination of the liver and kidneys did not
show abnormalities in any ofthe treatments.
70 Chapter 3
TABLE5.The effect of dietary inclusion levels ofD-xylose and

L-arabinose onchick performance from 6to27days ofage by
usinga semipurified basal diet (ExptB).
Sugar Dietary Weight Feed Feed: Water Dry matter

level gain intake gain intake content
(g/kg) (g) (g/bird (g:g) (g/bird excreta
per day) per day) 1 (g/kg)2
D-glucose 75 770a 56.l a 1.531a 143 a 267
D-xylose 25 737 ab 54.0ab 1.538ab 169b 208

50 719 b
52.8bc 1.544ab 174b 153
75 682c 51.4C 1.584c 193c 132
L-arabinose 25 761 a 55.7a 1.537ab 1 7 9 b c 174

50 748 ab 55.7a 1.565abc 191 c 156
75 738 ab 55.4a 1.576bc 233 d 123
SEM(df42) 11.8 0.8 0.014 5.5
x)
Meansoffour penspertreatment ofdays26plus27.
2)
Pooledsamplespertreatment ofdays 26plus 27;figures not analysed
statistically.
a,b,c
Meanvalues with nocommonsuperscripts within a column, differ
TABLE 6.The effect of dietary inclusion levels of D-xylose and

L-arabinose on caecal length and weight, and liver weight of
chicks by using a semipurified diet (Expt B).
Sugar D-gluc. D-xylose L-arabinose SEM

Dietary level (g/kg) 75 25 75 25 75 (df70)
Liveweight (LW)at 866 835 818 848 832 20.2

autopsy, g
Caecal length, cm 23 a 23 a 22 a 23 a 26 b 0.81
Caecal weight (incl. contents)

g 3.9a 4.1 a 4.0a 4.7 ab 5.8b 0.47
%ofLW 0.45a 0.50a 0.49a 0.56ab 0.70b 0.06
Liver weight
g 21.6a 21.0a 20.3 a 20.0a 20.8 a 0.93
%ofLW 2.5 a 2.5 a 2.5 a 2.4a 2.5 a 0.10
a b
- Meanvalueswithnocommonsuperscriptswithinarowdiffer significantly
(P < 0.05).
DISCUSSION
Although the experimental diets were formulated to be balanced in ME

content,weightgainandfeedutilizationinbirdsfedthepentosesugarswere
inferior to those fed the reference (75gD-glucose/kg)diet (Tables 2and 5).
The extent ofthis reduction in weight gain and feed utilization was small
when birds were fed the pentose sugars at dietary levels of 25 g/kg, but
became larger when fed dietary inclusion levels of 50 and 75 g/kg. This
negativedosage-dependent effect ofbothpentosesugarsonweightgain and
feed conversion efficiency may have resulted from several factors.
In the case of D-xylose, one of these factors relates to feed intake.which
decreased linearly(P<0.05)asthedietarylevelofthis sugarwas increased
(Tables2and5).Consequently,weightgainofbirdsfedonD-xylosedietswas
moredepressedthanofthosefedondietscontainingL-arabinose. Depressed
feed intake as a result offeeding D-xyloseto chicks was also observed in a
72 Chapter 3
previous study (Schutte, 1990). The birds' dislike for D-xylose had been
demonstrated by Kare and Medway (1959), who showed an almost total
rejection ofwater solutions containing this pentose sugar. In similar tests
performed by these investigators, other sugars such as lactose, galactose,
rafiinose and arabinose had only minor effects on water consumption.
Second,the pentose sugarsmayhave had anindirect negative effect on the
utilization ofthe ME ofother dietary energybearing components, and as a
result depressed performance. Some evidence of such an effect may be
providedbythe data ofa recent study (Schutte et al., 1991b),indicating an
increased microbial fermentation ofNin pigsfed ondiets containing 200 g
D-xylose/kg.Athirdfactorwhichneedsconsiderationrelatestotheutilization
oftheMEofbothpentose sugars ascomparedtoD-glucose. Without taking
into account a possible indirect negative effect ofboth pentose sugars, the
utilization ofthe MEofD-xyloseand L-arabinose canroughlybe estimated
from the differences in ME utilization among the 75 g sugar/kg diet
treatments (Table3).Thiscalculation pointedoutthat theutilization ofthe
MEofD-xyloseforgainwasonlyapproximately 20%.Inthiscalculation the
difference in ME utilization between the reference diet and the 75 g D-
xylose/kgdietwasfully attributed toD-xylose.Calculationssimilartothose
for D-xylose were made for the 75 g L-arabinose/kg diet treatment. This
calculation resulted in a zeroutilization ofthe ME ofL-arabinose for gain.
The estimated lesswellutilization ofthe ME ofD-xylose and L-arabinose
probably relates to the metabolic pathways of these sugars. It is well
recognized that the ME ofD-glucose ishighly utilized in chicks (Anderson
et al., 1958.).Knowledge about the utilization ofthe ME ofD-xylose and L-
arabinose in chicks and also in other monogastric animals is limited. In a
previous study (Schutte et al., 1991a)with adult roosters it wasfound that
about 15%of the ingested D-xylose or L-arabinose was excreted into the
urine.Theremainingpartnotaccountedformayhaveeitherbeenmetabolised
to carbon dioxide or fermented in the intestinal tract, or by both of these.
AccordingtoSegalandFoley(1959)metabolismofthesetwopentose sugars
tocarbon dioxide appears tobeonlyofsignificant importance for D-xylose.
Whengivenanintravenouslyinfused doseofeitherC14-labeledD-xyloseor
L-arabinose toman, 16and0.8%,respectively,couldberecovered as carbon
dioxide.Iftheir data are transferable tochicks,this would mean that both
pentose sugars mainly mightbefermented in the intestinal tract when fed
orally.This hypothesis is supported bythe results ofprevious studies with
pigs (Schutte et al., 1991b,c)showing substantial increases in ileal flow of
volatile fatty acids when they were fed on diets containing either D-xylose
orL-arabinose.Fromtheseresultsitwasconcludedthat atleastpart ofthe
ingested pentose sugars was degraded microbially. It seems likely that
microbial degradation ofD-xylosein chicks occurs mainly in the crop and

small intestine, since ileal digestibility ofthis pentose sugar was found to
benearly 100%inadultroosters(Schutteetal.,1991a).Inthatstudyanileal
digestibilityvalue ofabout85%wasfound forL-arabinose,indicating that
thispentosesugarwillbeatleastpartlyalsofermented inthehindgut.This
was supported by the absence of arabinose in the faeces of colostomizt;]
cockerels fed on L-arabinose diets (unpublished data, Schutte et al.).
MicrobialdegradationofL-arabinoseinthehindgutisalsosupportedbythe
observed increaseincaecalweight inbirdsfedonL-arabinose diets (Tables
4and 6).Increased caecalweightas a result offeeding L-arabinose diets to
chickshas beenpreviously reported (Longstaff et al., 1988;Schutte, 1990).
It is well known that microbial degradation ofdietary energy bearing
substances is coupled with considerable losses ofenergy,not accounted for
intheMEdetermination. Inpigstheselossesinenergyareassumedtovary
varybetween 33%(Agricultural Research Council, 1981)and 50%(Just et
al., 1983;Van Es, 1987).No published data for poultry are available, but
energy losses of a similar magnitude may be assumed. In addition to the
energylossesasaresultofthefermentation process,metabolismofD-xylose
andL-arabinose per semayalsobeassociated withlossesnotaccounted for
in the ME determination.
The higher intakes ofwater inbirdsfed oneither D-xyloseor L-arabinose
diets (Tables 2 and 5) are in agreement with the observation in a previous
studywithchicks(Schutte, 1990).Thisphenomenonmaybeconnected with
theosmoticproperties ofunabsorbed pentose sugars andincreased volatile
fatty acids concentrations in the intestinal tract resulting in an inflow of
water into the intestinal lumen (VanWeerden, 1959;Hof, 1980).
Wagh and Waibel (1966)reported that plasma uric acid was significantly
increased when chickswerefed ondiets containing 200and 400gof either
D-xyloseorL-arabinose/kg.TheirresultssuggestanincreasedN catabolism
whenbirdsarefedonD-xyloseorL-arabinose diets.Theyreported also that
feeding these diets to birds resulted in decreased liver weights. In the
present study some indications were achieved that even at low dietary
inclusion levelsboth pentose sugars mayinfluence Nutilization adversely.
However,noeffect onliverweightwasobservedattheapplieddietary levels
ofeither D-xylose or L-arabinose in the present study (Tables 4 and 6).
In conclusion it canbe stated that D-xyloseand L-arabinose may provide
only some energy tobirds.Besides,these sugars induce increases in water
intake and asresult wetdroppings.Consideringthese aspects,the benefits
ofhydrolysingofNSPfractions whichwillreleasemainlythese sugars(e.g.
hemicellulose) are very doubtful.
74 Chapter 3
ACKNOWLEDGEMENTS
TheauthorswishtothankJ.M.deZeeuwfortechnicalassistance,andG.B.
Derksen andJ. Wiebenga for statistical analysis ofthe data.
REFERENCES
Agricultural Research Council(1981)TheNutrient Requirements ofpigs.

(Slough,CommonwealthAgricultural Bureaux).
Anderson,D.L.,Hill,F.W.&Renner,R.(1958)Studiesonthemetabolizable
and productive energyofglucoseforthe growingchick.Journalof
AOAC(1984).Official MethodsofAnalysis, 14thEdn (Washington.DC,
Association ofOfficial Analytical Chemists).
Bogner,P.H.(1961)Alimentary absorption ofreducing sugarsbyembryos
andMedicine 107: 263-267.
Carr6,B.&Leclercq,B.(1985)Digestionofpolysaccharides, protein and
lipidsbyadult cockerelsfed ondietscontaininga pecticcell-wall
materialfrom whitelupin(LupinusalbusL.)cotyledon.British Journal
ofNutrition 54:669-680.
Carr6,B.,Derouet, L.&Leclercq,B.(1990)Thedigestibility ofcell-wall
polysaccharides from wheat (bran orwholegrain),soybeanmeal, and
whitelupinmealincockerels,muscovyducksandrats.Poultry Science
69: 623-633.
Chesson,A.(1987)Supplementaryenzymestoimprovetheutilizationofpig
and poultrydiets,in:W. Haresign & D.J.A.Cole(Eds)RecentAdvances
inAnimalNutrition, pp.71-90(London, Butterworths).
Cochran,W.G.& Cox,G.M.(1957)Experimental Designs,2ndedn(New
YorkWileyand Sons,Inc).
Hof, G.(1980)Aninvestigation intothe extenttowhichvarious dietary
components,particularly lactose,are related totheincidenceof
diarrhoea inmilk-fed calves.Ph.D.thesis,Agricultural Universityof
Wageningen,The Netherlands.
Just,A.,Fernandez,J.A. &Jorgensen, H.(1983)Thenet energyvalueof
dietsforgrowth inpigsinrelationtothefermentative processesin the
digestive tract andthe siteofabsorption ofthe nutrients. Livestock
Production Science 10:171-186.
Kare, M.R.&Medway,W.(1959)Discriminationbetweencarbohydratesby
thefowl.Poultry Science38:1119-1127.
Longstaff, M.A.,Knox,A.& McNab,J.M.(1988)Digestibility ofpentose
sugarsanduremicacidsandtheireffect onchickweightgainandcaecal
size.British Poultry Science29: 379-393.
LongstafF, M.&McNab,J.M.(1989)Digestionoffibre polysaccharidesof
pea(Pisum sativum)hulls,carrot andcabbagebyadultcockerels.
BritishJournal ofNutrition 62:563-577.
National Research Council (1984)Nutrient Requirements ofPoultry, 8th
revised edn(Washington, DC,NationalAcademyofSciences).
Norusis,M.J.(1988).BaseManualfortheIBMPC/XT/ATandPS/2(SPSS
inc..Chicago,Illinois)
Scheele,C.W.,VanderHel,W.,Verstegen,M.W.A.&Henken,A.M.(1987)
Climaticenvironment and energymetabolism inbroilers,in:M.W.A.
Verstegen &A.M.Henken (Eds)Energy Metabolism inFarmAnimals,
pp.217-260(Dordrecht,TheNetherlands,Martinus Nyhoff publisher).
Schutte,J.B.(1990)Nutritional implications and metabolizable energy
value ofD-xyloseand L-arabinose inchicks.Poultry Science 69,1724-
1730.
Schutte,J.B.,VanLeeuwen, P.&Lichtendonk,W.J.(1991a)Ileal
digestibility and urinary excretion ofD-xyloseand L-arabinose in
ileostomized adult roosters.Poultry Science 70,884-891.
Schutte,J.B.,DeJong,J., Polziehn,R.&Verstegen, M.WA.(1991b)
NutritionalimplicationsofD-xyloseinpigs.BritishJournalofNutrition
66,83-93.
Schutte,J.B.,DeJong,J., VanWeerden,E.J. &Tamminga,S.(1991c)
Nutritional implications ofL-arabinose inpigs.BritishJournalof
Nutrition (inpress).
Segal,S.&Foley,J.B.(1959)Themetabolicfate ofC14labeled pentoses
inman.Journal ofClinical Investigations 38: 407-413.
VanEs.A.J.H.(1987)Energyutilizationoflowdigestiblity carbohydrates,
in:D.C.Leegwater,V.J.Feron&R.J.J.Hermus(Eds)LowDigestibility
Carbohydrates,pp. 121-127(Wageningen, Pudocpress).
VanWeerden,E.J. (1959)Theosmoticpressure and the concentrationof
somesolublecomponents ofthe intestinal contents and thefaecesof
the cow,inrelation tothe absorption ofminerals.Ph.D.Thesis,
Agricultural University ofWageningen,The Netherlands.
Vogt,H.&Stute,K.(1971)Uber dieVerdaulichkeit einiger
Kohlenhydratfraktionen (Zucker,Starke,Pentosane,Rohcellulose,
Lignin)imHuhnerfutter. Archivfur Gefliigelkunde 35:29-35.
Wagh,P.V.&Waibel,RE.(1966)Metabolizability and nutritional
implicationsofL-arabinoseandD-xyloseforclucks.JournalofNutrition
90:207-211.
Wagh,P.V.&Waibel,RE.(1967)AlimentaryabsorptionofL-arabinose and
D-xylosein chicks.Proceedings ofthe SocietyofExperimental Biology
and Medicine 124: 421-424.
77
PART B
STUDIES WITH PIGS

79
CHAPTER 4
NUTRITIONAL IMPLICATIONS OF
D-XYLOSE IN PIGS
J.B. Schutte,*J.de Jong,*R.Polziehn* andM.W.A.Verstegen**
*TNO-InstituteofAnimal Nutrition and Physiology (ILOB),P.O.Box15,

6700AA Wageningen, The Netherlands
**Department ofAnimal Nutrition,WageningenAgricultural University,
P.O.Box338,6700AHWageningen,The Netherlands
Published in:BritishJournal ofNutrition 66:83-93(1991)

Reproduced bypermission ofTheNutrition Society
81
NUTRITIONAL IMPLICATIONS OFD-XYLOSE

IN PIGS
J.B. SCHUTTE,*J. DE JONG,* R. POLZIEHN*
AND M.W.A.VERSTEGEN**
*TNO-Institute ofAnimal Nutrition and Physiology (ILOB), P.O.Box 15,

••Department ofAnimal Nutrition, WageningenAgricultural University,
P.O. Box 338,6700AH Wageningen, The Netherlands
Hemicellulose consists primarily of pentose sugars,joined together in a

polysaccharidechainwithD-xyloseasthemostabundant component. Ileal
digestibilityand urinary excretion ofD-xyloseand associated effects ofthis
pentose sugar on ileal and faecal digestibility ofdry matter (DM), organic
matter (OM), gross energy (GE) and nitrogen (N) were studied in pigs.
Castrated pigs were prepared with a post-valvular T-caecum cannula to
measure ileal digestibility. Faecal digestibility was measured in non-can-
nulated pigs.D-xylosewas given at dietary inclusion levels of100and 200
g/kg, and the control sugar, D-glucose, at a rate of 200 g/kg diet. Ileal
digestibility ofD-xyloseaswellasthat ofD-glucosewasfound tobecloseto
100 %.The presence ofD-xylose in the diet decreased ileal digesta pH and
increased ileal flow of volatile fatty acids, suggesting the occurrence of
microbial degradation ofD-xylose in the pig small intestine. In pigsfed on
the 100 g D-xylose/kgdiet, 44.5 %ofthe D-xylose intake appeared in the
urine.Thispercentage increased significantly to52.6%when pigswere fed
on the 200gD-xylose/kgdiet. Ileal and faecal digestibility ofDM, OM, GE
and N,as well as Nretention decreased significantly in pigsfed onthe 200
g D-xylose/kg diet. (Key words: D-xylose, digestion, excretion, pig).
INTRODUCTION
Cellulose and hemicellulose form the bulk ofthe cell wall constituents of
feedingredientsofvegetableorigin.Bothcarbohydratefractions areresistant
to the digestive enzymes ofpigs,and pass to the hind gut where microbial
degradation takes place. The microbial degradation of cellulose and
hemicellulose in the hind gut ofpigs leads to the production of absorbable
volatile fatty acids which provideenergy tothe animal (Imoto&Namioka,
82 Chapter 4
1978;AgriculturalResearchCouncil,1981;VanEs,1987).Thisfermentation
process,however,iscoupledwithconsiderablelossesinenergy,assumedto
varybetween33%(AgriculturalResearchCouncil,1981)and50%(Justet
al. 1983;VanEs,1987).
Improvingthe utilization ofcelluloseandhemicellulose maybe attained
byanenzymetreatmentwhichcouldhydrolysethesecarbohydrate fractions
tomonosaccharides.Thereislittledoubtthatthemonosaccharideunitsin
cellulose,i.e.glucose,areanexcellentsourceofenergyforpigs.Hemicellulose
primarily consists ofpentose sugars,joined together in a polysaccharide
chain withD-xyloseasthemost abundant component.
ThestudiesreportedontheabsorptionofD-xyloserelatetoother animal
species than pigs. These studies have shown that D-xylose is readily
absorbedfromtheintestinaltractbyrats(Cori,1925;Miller&Lewis,1932;
Fowler &Cooke, 1960;Arnal-Peyrot &Adrian, 1974)and chicks (Wagh&
Waibel,1967a).ThesestudiesalsoshowedthatpartoftheingestedD-xylose
isexcreted intheurine.Findings ontheutilization ofD-xylosemainlyre-
latetochicks.Longstaff etal.(1988)reportedthatchickswereabletogrow
well on diets containing D-xylose at a dietary concentration of 50 g/kg.
Radioisotope studiesbyWagh&Waibel(1967b)inchicks,showedthatD-
xylosewasmetabolizedtocarbondioxide,butlessrapidlythan D-glucose.
The present studies were designed to obtain information on ileal di-
gestibility (absorbability) and urinary excretion of D-xylose at dietary
inclusionlevelsof100and 200g/kginpigs.Theeffects ofdietaryD-xylose
ontheilealandfaecaldigestibilityofdrymatter(DM),organicmatter(OM),
gross energy (GE) and nitrogen (N) were also examined. D-glucose was
included inthe trials asa reference.
Animals and diets
Two separate trials were conducted with growing castrated male pigs
(DutchLandracexDutchYorkshire):onetrial withcannulated pigs(Expt
A)andonewithnon-cannulatedpigs(ExptB).Inbothtrialsthepigswere
individuallyhousedinmetabolismcagesundera 12hlighten12htwilight
cyclethroughout.Thenutritionallycompletebasal dietusedwasbasedon
maize,wheatstarchandisolatedsoyaprotein.Thecompositionofthebasal
dietanditschemicalcharacteristicsareshowninTables1and2,respectively.
D-xyloseinpigs 83
TABLE 1.Composition ofthebasal diet (g/kg)
Maize meal 287

Wheat starch 287
Soya-bean oil 40
Animal fat 40
Isolated soya protein (880 g protein/kg) 223.3
Cellulose* 60
Monocalcium phosphate 24
Limestone 10
Potassium bicarbonate 15
Iodized salt 3
Mineral mix* 5
Vitamin mix++ 5
DL-methionine 0.7
* ArbocelB800(Rettenmaier,FRG)
+ Provided(mg/kgdiet):magnesium400,zinc110,copper25,manganese
45,iron 80,cobalt0.5,selenium 0.1.
++ Provided (mg/kgdiet):thiamin 2,riboflavin 5,nicotinamide30,
pantothenic acid 12,pyridoxine 3,cyanocobalamin 0.04,biotin0.1,
folic acid 1,menadione 3,ascorbicacid 50,retinol3.1,
cholecalciferol 0.045,vitamin E40,cholinechloride 1000.
TABLE2.Chemical composition of thebasal diet (analysed, g/kg

unless otherwise stated)
Constituent
Dry matter 909

Ash 47
Crude protein (nitrogen x 6.25) 218
Crude fibre 56
Crude fat 86
Gross energy (MJ/kg) 17.9
Calcium 9.2
Phosphorus 8.1
84 Chapter 4
The test sugars (D-glucose and D-xylose), supplied as anhydrous

monosaccharides, were substituted by weight for wheat starch.
In both trials the experimental diets were fed at a daily rate of0.9 MJ
metabolizable energy (ME)/kg metabolic body weight, assuming that D-
xylosehasthe sameMEcontent asD-glucose.Thedailyamount offeed was
offered at two equal meals at 08.00 and 20.00 hours, and adjusted weekly
according to body-weight. The feed was mixed with water (1part feed + 1
part water). In addition, water was freely available.
Experimental protocol
ExptA.The ileal digestibility ofD-xylose, and the effect ofthis pentose

sugaronilealdigestibilityof DM,OM,GEandNweremeasured. Moreover,
digesta pH, concentrations ofvolatile fatty acids (VFA)in the digesta, and
ileal flow ofVFAwere investigated.
Four pigs, 9 weeks old at the start ofthe trial, were involved. The pigs
were surgically fitted with a post-valvular T-caecum cannula (PVTC)
according to the procedure described by van Leeuwen et al. (1988). Post-
operativecareincludedkeepingthepigswarm(25°)andwitholdingfeed for
24h.Duringthe3-weekpost-operativeperiod,thepigswerefedonthe basal
diet (Table 1).The experimental period lasted 24 d and consisted of three
phases,duringwhichtimeeachpigwasfedconsecutivelyonadietcontaining
200gD-glucose/kg(Gluediet),100gD-xylose/kg(LL-Xyldiet)and 200gD-
xylose/kg(HL-Xyldiet),witha4dadaptation and a4dcollectionperiod for
each diet.
Atthe start ofthe experimental period, the pigsweighed on average 24.3
(SD 2.6)kg and at the end ofthis period 31.8 (SD 3.4) kg.
Expt B.The objectives of this trial were to determine the urinary ex-
cretionof D-xylose,andtostudytheeffect ofD-xyloseonfaecal digestibility
of DM, OM, GE and N, and N retention. This trial, involving four 9-week
old pigs,was run parallel with ExptA.The pigs were accustomed to cages
and the basal diet (Table 1)for 3 weeks before starting the experimental
period. The experimental design of Expt B was similar to that of Expt A.
Duringthe24dexperimentalperiodthesamethreedietsandbatchesoffeed
were used, and fed in the same order as described for ExptA.
At the start ofthe experimental period, the pigs weighed on average 25.0
(SD 0.7) kg and at the end ofthis period 34.4 (SD 0.8) kg.
D-xylose in pigs 85
Digesta collection
Ilealdigestaduringeach4dcollectionperiodwerecollected quantitatively
from individual animals over a 12h period per day (08.00-20.00 hours). In
this procedureitwasassumed that ilealdigestibilitywascompleted within
12h.Thisassumption wasbased onpreviousstudies (E.J. VanWeerden, J.
Huisman&P.vanLeeuwen;unpublishedresults)indicatingthatthere were
no significant differences in ileal digestibility when digesta were collected
over a 12h or over a 24h period per day.
The digesta were collected continuously in dry ice, weighed daily and
stored at -20°.Atthe end ofthe experiment, thefour 12h collected portions
werepooledforeachpigseparately,homogenized and sampled.ThepH and
VFA determinations were performed in the digesta as such, the other
measurements in freeze-dried samples. Until analysis all samples were
kept at -20°.
Faeces and urine collection
Faeces were collected directly into a bag fitted around the anus, and the
urine was collected using a funnel fitted under the cage.Total collection of
faeces andurinewascarriedoutduringthefour 24hcollectionperiods from
the individual animals. The faeces were collected at intervals of 12h, and
storedat -20°.Allfaeces producedduringeachcollectionperiodwerepooled
for each pigseparately, homogenized and sampled. Then the samples were
freeze-dried.
Urine was collected in containers provided with merthiolate sodium
(Thiomersal, BDH chemicals Ltd, Poole, England) at intervals of
approximately 4 h. The portion from each interval was pooled daily from
individualanimals.Arepresentative sampleof10%ofthepooledurine was
taken and frozen at -20°.The 4dsub-samples ofurine werepooledfor each
animal separately, homogenized and sampled. Faeces and urine were kept
at -20°between sampling and before analysis.
Analytical methods
Samples offeed and freeze-dried digesta and faeces were milled to pass
through a 1.0 mm screen (Retsch mill ZM1,Retsch B.V., Ochten) before
analysis.Allanalyseswerecarriedoutin duplicate.DMwasdetermined by
drying the samples to a constant weight at 101°. Inorganic matter and N
were determined by standard methods (Association of Official Analytical
86 Chapter 4
Chemists, 1975),GEwas determined usingan IKA-C4000adiabaticbomb

calorimeter.
Concentrations ofVFAin wet digesta were determined by a modification
of the gas-liquid chromatographic method of Imoto &Namioka (1978).A
known portion (about 20 g) of the digesta was centrifuged. Immediately
afterwards the supernatant fraction (5 ml) was acidified with 500 ul
phosphoric acid (850 ml/1, reagent grade), 3 ml of an aqueous solution of
isocapronic acid (4.0193 g/1)was added as an internal standard. Distilled
water was then added tothe mixture to obtain a final volume of 10ml. A 1
ulsampleofthefinal solutionwasinjected intothe columnofthe gas-liquid
chromatograph. The gas-liquid chromatograph was fitted with a flame
ionization detector (Packard 419,USA).Aglass column (1850 mm x 2 mm
i.d.)packedwithChromosorb 101of80/100meshwasused.Thecarrier gas
(N 2 )was saturated with formic acid,and had a flow rate of25ml/min. The
oven temperature was set at 190°,and the inlet and detector temperature
at 225°.Standard solutions containing acetic acid, propionic acid, butyric
acid,isobutyric acid,valericacid and isovalericacidwereprepared for gas-
liquidchromatographyinthesamewayasdescribedpreviously. Calibration
curvesforthese acidswerethen madebyobtainingthe peak-heights ofthe
acids:thatofisocapronicacid.Recoveriesvaluesbetween95and 100%were
found for the individual VFA and the internal standard. Total VFA was
represented as the sum ofall six acids.
Concentrationsofglucoseandxyloseindigestaandurinewere determined
as silyl derivatives of monosaccharides by gas-liquid chromatography
(Sweeley et al. 1963). A known amount ofwet digesta (1g)or urine (1ml)
was diluted with distilled water (1:10 v/v). The diluted sample was then
deproteinized with potassium ferrocyanate and zinc-acetate and desalted
bypassingthrough amixture(1:1w/w)ofanion(BioradAG3x4)and cation
(Biorad AG 50 W x 4) exchanger. After centrifugation, 200 ul of the
supernatant fraction was freeze-dried. To the freeze-dried sample
phenylglucopyranoside (0.4mgina 1mlpyridine solution)wasaddedas an
internal standard. The sample was then derivatized by the addition of 0.6
ml hexamethyldisilazane and 0.3 ml trimethyl-chlorosilane. Then the
contentsweremixed usingaVortexstirrer.After an incubation periodof30
min at roomtemperature, the reagents were removed by evaporation with
N 2 at 40°.The residue was then redissolved in 0.5 ml ethyl acetate. From
this sample,2ulwasanalysed usingaHewlettPackard HP5890,equipped
with a flame ionization detector and a Hewlett Packard 3396A integrator.
The carbohydrate derivatives were separated with a chrompack capillary
WCOTfused silica column coated with CP sil5CBof50m length. H 2 was
D-xyloseinpigs 87
usedascarriergas.Theoventemperature washeldfor3minat 190°,then

raised at therate of57mintoafinal temperature of265°,whichwasheld
for 5min.Thetemperature ofthe injector and detectorwas240and300°,
respectively.
Allvalues wereanalysed bymeans ofanalysisofvariance.Arandomized

blockdesignwasused,inwhichtheanimalsweretheblocks(Cochran&Cox,
1957).Althoughthetreatments areconfounded bytimeitisassumed that
differences are due to the test sugar. The Genstat 5 package (Oxford
UniversityPress,1987)wasusedtocalculatetheanalysisofvariance.The
treatmentfactorswerethecombinationofthetypeofsugarandthedietary
level.ThenthetreatmentmeanswerecomparedusingtheLeastSignificance
Differences test.Allstatementsofsignificance arebasedonaprobabilityof
P <0.05.
RESULTS
Thepigswerehealthyandconsumedtheirdailyfeedallowancecompletely
for allexperimental treatments.
ExptA
Intake ofDMand water, output ofdigesta, and DMcontent of digesta

measured in cannulated pigs onD-glucose or D-xylosediets are given in
Table3.Sincetheoutputofdigestawasmeasuredover12h/d,intakeofDM
and water is also presented over a 12 h period. There were significant
differences in DMintake among the treatments. These differences were
caused bythe feeding system applied, sincethis system was coupled with
liveweight ofthepigs. Waterintake ofpigsfed onthe Gluediet (200gD-
glucose/kg)wassignificantly lowerascomparedwiththeLL-Xyl(100gD-
xylose/kg)and HL-Xyl(200gD-xylose/kg)diets.Pigsfed onthe Gluediet
produced onaverage 255gwetdigesta/12h,whichvaluewasincreased to
326 and 547g/12h when pigs were fed onthe LL-Xyl and HL-Xyl diets,
respectively.TheamountofdigestaproducedinpigsontheHL-Xyldietwas
significantly different from that ofpigsonthe Glueand LL-Xyldiets.The
increase in digesta output in pigs on the LL-Xyl and HL-Xyl diets was
associatedwithadecreaseinDMcontentofthedigesta.However,thelatter
wasmorepronounced onthe HL-Xyldietthan onthe LL-Xyldiet.
ApparentdigestibilityvaluesforDM,OM,GE,N,D-glucoseandD-xylose
88 Chapter 4
are shown in Table 4. In pigs fed on the Glue and LL-Xyl diets, similar
digestibilitycoefficients for DM,OM,GEandNwereobserved. However, in
pigs fed on the HL-Xyl diet digestibility ofDM, OM, GE and N decreased
significantly. Theapparent ilealdigestibilityofD-glucoseandD-xylosewas
found tobe close to 100%.
Digesta pH,VFAconcentrations in the digesta, and ileal flow ofVFAare
given in Table 5.ThepH decreased significantly from 6.5in digesta ofpigs
on the Glue diet to 6.2 and 6.0 when they were fed on the LL-Xyl and HL-
Xyl diets, respectively; the latter two values being also significantly diffe-
rent from each other. The decrease in pH on the LL-Xyl and HL-Xyl diets
concurred with the appearance ofgreater amounts ofVFAin the digesta.
The increase in total VFAconcentrations in pigs on the LL-Xyl diet was
about 50 %, but not significant. The latter due to the large differences
between animals within the treatments.Whenpigswerefed onthe HL-Xyl
diet,totalVFAconcentrations indigesta increased significantly with about
110%when compared with the Gluetreatment. The increase in total VFA
concentrations on the HL-Xyl diet was reflected in all individual VFA
fractions. In terms of ileal flow of VFA the differences between the
treatments are much greater, since pigs on the LL-Xyl and HL-Xyl diets
produced greater quantities ofdigesta than when fed on the Glue diet.
TABLES. ExptA. Intake (g/12h) of dry matter (DM) and water,

output (g/12 h) of ileal digesta, and DM content (g/kg) of ileal
digesta, measured in cannulated pigs fed on D-glucose and
D-xylose (100 (LL-Xyl) or 200 (HL-Xyl) g/kg) diets
(Mean values offour pigs per treatment)
SEM
Diet Glue LL-Xyl HL-Xyl (df6)
Intake ofDM 316a 333 b 349c 0.3

Intake of water 817 a 1029b 1156b 53.4
Output ofwet ileal digesta 255 a 326a 547b 45.9
DM content ileal digesta 172a 15l a 118b 9.0
a,b,c
Within a row, mean values with different superscript letters were
significantly different (P <0.05).
D-xyloseinpigs 89
TABLE4.ExptA. Apparent ileal digestibility coefficients of dry

matter (DM),organic matter (OM),gross energy (GE),nitrogen,
D-glucoseandD-xylose,measured incannulated pigs fed on
D-glucoseandD-xylose (100(LL-Xyl)or200 (HL-Xyl)g/kg) diets
(Meanvalues offour pigsper treatment)
Diet Glue LL-Xyl HL-Xyl SEM df
DM 86.2a 85.7a 81.9b 1.00 6

OM 87.6 a 87.2 a 84.7b 0.71 6
GE 87.6 a 87.5 a 84.5 b 0.62 6
N 90.3 a 89.l a 87.2b 0.47 6
D-glucose 99.3 . _ . _
a a
D-xylose - 98.7 98.6 0.23 3
,,b
Within a row,meanvalues with different superscript letters were
90 Chapter 4
TABLE5.ExptA.DigestapH,concentrations ofvolatilefatty adds

(VFA)in digesta and ileal flow ofVFA,measured in cannulated
pigs fedonD-glucoseandD-xylose (100(LL-Xyl)or200(HL-Xyl)
g/kg) diets
(Meanvaluesoffour pigsper treatment)
SEM
pH, and concentrations of

VFAin digesta (mg/100 g)
pH 6.5a 6.2b 6.0C 0.06
TotalVFA 437 a 644 ab 930b 86.6
Individual VFA
Acetic acid 270 a 478 ab 654 b 61.2
Propionic acid 77 a 79 a 121 a 17.9
Butyric acid 51 a 49 a 75 a 11.4
Isobutyric acid 12a lla 22 b 2.0
Valeric acid 13 a 14 a 27 b 2.4
Isovaleric acid 14a 13 a 31 b 2.8
Ileal flow ofVFA(mg/12 h)

Total VFA 1106a 2062b 4888 c 123.7
Individual VFA
Acetic acid 684a 1508b 3447c 99.2
Propionic acid 196a 253 a 630b 42.0
Butyric acid 125a 171a 386 b 37.6
Isobutyric acid 31 a 34 a 115b 6.4
Valeric acid 33 a 48 a 146b 10.3
Isovaleric acid 37 a 48 a 164b 7.7
a,b,c
Within arow,meanvalues with different superscript letters were
ExptB
The mean values for DMand water intake, output offresh faeces, DM
contentoffaecesandoutputofurineinpigsfedontheGlue,LL-XylandHL-
Xyldiets over a 12h period, are given in Table 6.There were significant
D-xyloseinpigs 91
differences inDMintakeamongthetreatmentgroups.However,asalready
stated in Expt A, these differences were caused by the feeding system
applied. Water intake andurine output tended toincreaseand theDM
contentoffaeces tended todecreasewhenthepigswerefed ontheLL-Xyl
diet.Whenfed ontheHL-Xyldiet,water intake aswellasoutput ofurine
andfreshfaecesincreasedsignificantlycomparedwiththeGlueandLL-Xyl
diets.Inaddition,the DMcontentof faeces inpigsfed ontheHL-Xyldiet
was significantly lowerwhencomparedwiththe Gluediet.
Apparentfaecaldigestibilitycoefficients forDM,OM,GEandN,retention
of N,andtheurinaryexcretionofglucose,xylose,energyandNaregiven
inTable7.Similarresultsforapparent faecal digestibilityofDM,OM, GE
andNwereachievedontheGlueandLL-Xyldiets.WhenfedontheHL-Xyl
diet,digestibilitiesofallfoursubstancesdecreasedsignificantly.Nretention
wascalculated from theintakeofNandthelossesofNintothefaeces and
urine.WhenfedontheHL-Xyldiet,significantly lessNwasretained than
when feeding the Glue and LL-Xyl diets. This is due to both a lower N
digestibilityandahigheramountof NexcretedintheurineontheHL-Xyl
diet.Thelossesofxyloseintotheurinewereconsiderable.Whenpigswere
fed onthe LL-Xyldiet, 44.5%ofthe D-xyloseintake was excreted in the
urine.Thispercentageincreasedsignificantly to52.6%whenpigswerefed
onthe HL-Xyldiet.Asa result ofthexyloselossesintotheurine, urinary
excretion ofenergy alsoincreased significantly in pigsonthe LL-Xyl and
HL-Xyldiets.
TABLE6.Expt B.Intake (g/12h) ofdrymatter (DM)and water,

output (g/12h) offaeces andurine, and DMcontent (g/kg) of
faeces, measured in non-cannulated pigsfed on D-glucose
and D-xylose (100 (LL-Xyl)or200 (HL-Xyl)g/kg) diets.
SEM
Intake ofDM 322a 339 b 355 c 0.1

Intake of water 655 a 736a 963 b 36.6
Output offresh faeces 35 a 34a 68 b 3.4
DM content faeces 482 a 448 ab 419 b 14
Output of urine 326a 407 a 676 b 28.8
i,b,cWithinarow,mean valueswith different superscript letters were

92 Chapter 4
TABLE 7. Expt B.Apparent faecal digestibility coefficients of dry

matter(DM),organicmatter(OM),grossenergy (GE)andnitrogen,
retention of N (% of intake), and urinary excretion (% of intake)
of glucose, xylose, energy and N, measured in non-cannulated
pigs fed on D-glucose and D-xylose (100 (LL-Xyl) or 200 (HL-Xyl)
g/kg) diets
Diet Glue LL-Xyl HL-Xyl SEM df
Digestibilities
DM 94.9 a 95.5 a 92.2 b 0.37 6
OM 95.8 a 96.5 a 93.5 b 0.31 6
GE 95.2 a 96.0a 92.7b 0.34 6
N 96.1" 96.l a 93.5 b 0.44 6
Urinary excretion
Glucose + + +
Xylose - 44.5 a 52.6b 1.44 3
Energy 2.2a 6.1 b 9.9C 0.83 6
N 34.9 a 35.l a 38.7b 0.71 6
Retention of N 61.2 a 60.9a 54.8b 1.00 6
+ Small traces (0.2 -1.2 g/1)ofglucose were found in the urine of all
a,b,cWithin a row,mean values with different superscript letters were
significantly different (P <0.05).
DISCUSSION
The choice of the experimental design needs to be considered. Latin
squares are often used as an experimental design in balance studies with
pigs, especially in trials in which the diets are fed in sequence, the diet
sequence being different for each pig (Goedhart, 1990).The advantages of
using Latin squares are that variation between animals and those arising
from a common time trend between periods can be equilibrated. However,
this is only true when there are no carry-over effects. For D-xylose, the
results ofa previous tentative study showed that carry-over effects of this
pentose sugar cannot be excluded. Therefore, for the sake ofsafety in the
present trials each pig was fed on the sugars in the sequence of D-glucose
(Glue),low-level D-xylose (LL-Xyl; 100g/kg) and high-level D-xylose (HL-
D-xylose in pigs 93
Xyl;200g/kg).Onedisadvantage offeeding experimental dietsin sequence

is that faecal digestibilities may be affected by a time period x treatment
interaction,becausethedigestivecapacityofthepiglargeintestine increases
with increasing age. Considering the results reported by McConnell et al.
(1971,1972),Hennigetal.(1979)andGoedhart(1990),thechangesin faecal
DM,OM,GE and N digestibilities in Expt Bas affected byage would have
beenlessthan 1%.Furthermore,it shouldbenotedthat the feeding system
applied in our design may have induced changes in intake of water, and
output ofwet digesta, fresh faeces and urine. However, when corrected for
thedifferences inDMintakeamongthetreatments,itcanbecalculated that
there are stilllargedifferences inthese characteristics between the D-Gluc
and HL-Xyl diets.
The results obtained in the present study indicate that D-xylose was
digested almost completely at the terminal ileum; this would suggest an
almost completeabsorption ofthispentose sugarperse.Ontheother hand,
administration ofD-xylose to pigswas associated with an increase in ileal
flowofVFAanda decrease inpH.Both symptoms pointtoamore extensive
microbial activity in the small intestine ofpigs on the D-xylose diets. This
mayhaveresultedfrom thedifferences inratesofabsorptionfrom the small
intestine between D-glucose and D-xylose as reported by Miller & Lewis
(1932) in rats, and Bogner (1961) and Wagh & Waibel (1967a) in chicks.
These authors showed that absorption velocity ofD-xylose was lower than
that ofD-glucose.The presence ofunabsorbed xyloseinthe small intestine
may stimulate microbialactivity.Thus,the observedhighileal digestibility
of D-xylose in the present study could partly be due to a microbial
degradation ofthis sugar. The extent ofmicrobial degradation of D-xylose
in the pig small intestine cannot be derived simply from the differences in
ileal flow ofVFAbetween the D-glucose and D-xylose treatments, because
someoftheVFAwillbeabsorbed alreadyinthe smallintestine.In addition
to D-xylose, other readily fermentable components in the diet may also be
attacked by an increased intestinal bacterial activity. It is likely that the
depression in apparent ileal digestibility ofNinpigsonthe 200gD-xylose/
kgdietis,atleastpartly,aresultoftheincreasedmicrobialactivitywith this
diet. As protein is part of DM, OM and GE, this will also affect the
digestibilityofthesesubstances.However,thereductioninileal digestibility
ofDM, OM and GE on the HL-Xyl (200 gD-xylose/kg)diet can only partly
beexplainedbythe depression inN digestibility. Additionally,the increase
inilealdigestaflowmayalsoberesponsibleforthedepressionindigestibility
ofDM,OMand GEonthe HL-Xyldiet.Thishigher ilealdigestaflowcanbe
explainedbythepresenceofunabsorbed xyloseinthesmallintestine which
94 Chapter 4
will lead to an inflow of water into the intestinal lumen in order to keep
osmolality constant (VanWeerden, 1959;Hof, 1980).
The magnitude ofthe difference in ileal DM, OM, GE and N digestibility
between the treatments was maintained at a similar level in the faecal
digestibility values (Table 4 v. Table 7). These results may suggest that
microbialactivityinthepiglargeintestinewasnotmarkedlychanged when
theD-xylosedietswerefed.Theobserved depressed Nretention ontheHL-
Xyl diet is a result ofthe depressed N digestibility on the one hand and of
a higher urinary excretion on the other. Since the experimental diets were
fed in sequence,the higher urinary excretion ofN onthe HL-Xyldiet could
be partly due to an age effect (Carr et al. 1977).
It is well recognized that a portion ofthe ingested D-xylose appears in
the urine ofman (Loos,1954;Folwer &Cooke, 1960),rats (Arnal-Peyxot&
Adrian, 1974)and pigs (Wise et al. 1954).This observation is confirmed in
the present work. However, there is a scarcity of information about the
relationship between the dietary inclusion level of D-xylose and the
urinary excretion of this sugar. Wagh & Waibel (1966) reported, that in
chickstheMEvalueofD-xylosewasdecreasedwhenthedietarylevelofthis
sugar was increased. Their finding may provide some evidence of an
increased urinary excretion of D-xylose in percentage ofintake when the
dietarylevelofthissugarisincreased,sinceLongstaff etal.(1988)reported
that apparent digestibility ofD-xylose in chicks was nearly 100 %.In the
presentstudy,urinaryexcretionofxyloseasapercentageofintake increased
whenthedietarylevelofD-xylosewasincreasedfrom 100to200g/kg.When
fedonthe LL-Xyl(100gD-xylose/kg)diet,44.5%oftheD-xyloseintake was
excreted via the urine pathway. This percentage increased to 52.6 %when
fed onthe HL-Xyl (200gD-xylose/kg) diet.This dosage dependent urinary
excretion ofD-xyloseinpercentage ofintakemaybe connectedwiththelow
renal threshold for this sugar as suggested by Loos(1954).The differences
in urinary excretion ofxylosebetween the twoD-xylosetreatments are not
reflected intheurinaryexcretionofenergy.Calculationshaveindicated that
when the increases in urinary excretion of energy over the D-glucose
treatment werecontributed toD-xylose,thiswouldrepresent about 45%of
the D-xylose intake at both dietary levels.
Inconclusion,itcanbestated that utilization ofD-xyloseinpigsat dietary
inclusion levels of 100 gand 200 g/kgis low.Apart from the great losses of
D-xylose into the urine, at least part ofthis pentose sugar is fermented in
theintestinaltractofpigs,whichprocessiscoupledwithconsiderable losses
inenergy.Inaddition, at highdietarylevelsthis pentose sugar may induce
unwanted nutritive problems together with ahigher excretion ofurine and
faeces. Considering these aspects, the benefits of a hydrolysis of the
D-xylosein pigs 95
hemicellulose fraction in pig diets seem tobe doubtful as compared with a

fermentation ofthis fraction in the hind-gut ofpigs.There are no findings
available onthe extentofrelease ofD-xyloseinthegastrointestinal tractof
pigs as a result of enzyme inclusion in pig diets. Carre' &Brillouet (1986)
determinedthecontentandcompositionofvariousfeedstuffs usedbysingle-
stomach farm animals. From their findings it can be calculated that by a
complete hydrolysis of non-starch polysaccharides in pig diets based on
cereals,soyabeanoilmealandcereal-byproducts,about4%D-xylosewillbe
released. Considering the results ofWagh &Waibel (1966) with chicks, it
might be expected that also in pigs utilization of D-xylose will be much
better at low than at high dietary levels.This was confirmed in a recently
performed study (J.B. Schutte, G. Beelen, G.B. Derksen & J. Wiebenga;
unpublished results) with pigs, in which D-xylose was tested at graded
dietary levels of 25 - 100 g/kg. The results ofthat study showed that ME
contentofD-xylosewassignificantly decreasedwhenthedietarylevelofthis
pentose sugarwasincreased. Further studieswillberequired toclarify the
utilization and metabolism of D-xylose in pigs in relation to the dietary
inclusion level.
ACKNOWLEDGMENTS
TheauthorswouldliketothankMr.P.vanLeeuwenforsurgical assistance,
Messrs. G.B. Derksen and J. Wiebenga for statistical analysis ofthe data,
andDr.E.J.vanWeerdenandProfessorS.Tammingaforvaluablediscussions.
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Carr,J.R., Boorman, K.N. &Cole,D.JA. (1977).Nitrogen retention in the
96 Chapter 4
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Carr6, B. &Brillouet, J.M. (1986).Yield and composition ofcell wall
residues isolated from various feedstuffs used for non-ruminants farm
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Cochran,W.G.&Cox,G.M.(1957).Experimentaldesigns,2nded.NewYork:
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Cori,F.(1925).The fate ofsugar in the animal body. 1.The rate of
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Fowler, D. &Cooke,W.T.(1960).Diagnostic significance ofD-xylose
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Goedhart, P.W.(1990).Experimental design for comparative digestibility
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Hennig,U.,Bock,H.D.,Wunsche,J. &Kreienbring, F.(1979).Einfluss des
Geschlechtes und der Lebendmasse auf die wahre Verdaulichkeit des
proteins und derAminosauren verschiedener Futtermittel bei
Schweinen. (The influence ofthe sex and live mass ofpigs on the actual
digestibility ofprotein and amino acids ofvarious feeds.)Archiv fur
Tierernahrung 29,1-8.
Hof, G. (1980).An investigation into the extent to which various dietary
components, particularly lactose, are related to the incidence of
diarrhoea in milk-fed calves. PhD Thesis,Agricultural University of
Wageningen, The Netherlands.
Imoto, S. &Namioka, S. (1978).VFAproduction in the piglarge intestine.
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Just,A., Fernandez, J.A. &Jorgensen, H. (1983).The net energy value of
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Longstaff, M.A., Knox,A. &McNab,J.M. (1988).Digestibility of pentose
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Loos, M. (1954). Studies in the utilization ofpentoses in diabetes. Acta
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McConnell, J.C. Barth, KM. &Griffin, S.A. (1971).Nutrient digestibility
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McConnell,J.C.Barth,K M . &Griffin, S.A.(1972).Nitrogenmetabolism at

three stages ofdevelopment and its relationship to measurements of
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Miller, M.M. &Lewis,H.B. (1932).Pentose metabolism.1.The rate of
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Van Kleef, D.J., Van Weerden, E.J. &Den Hartog, L.A. (1988). Anew
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Wagh,P.V.&Waibel,P.E.(1967a).AlimentaryabsorptionofL-arabinose and
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99
CHAPTER 5
URINARY EXCRETION OF D-XYLOSE IN

PIGS AS AFFECTED BY AGE, FREQUENCY
OF FEEDING AND DIETARY LEVEL
J.B. Schutte, G.M. Beelen, G.B. Derksen and J. Wiebenga

P.O. Box 15,6700AA Wageningen, The Netherlands
Publishedin:ProceedingsoftheVthInternationalSymposiumonDigestive
PhysiologyinPigs(M.WA'\ferstegen,J.HuismanandLAdenHartog,
editors).EAAPPublication54:411-421(1991).PudocPress,Wageningen.
ReproducedbypermissionofPudoc,Wageningen
101
URINARY EXCRETION OF D-XYLOSE IN

PIGS AS AFFECTED BY AGE, FREQUENCY
OF FEEDING AND DIETARY LEVEL
J.B. SCHUTTE, G.M.BEELEN, G.B.DERKSENANDJ. WBEBENGA
TNO-Institute ofAnnimal Nutrition and Physiology (ILOB),

P.O. Box 15,6700AA Wageningen, The Netherlands
Thepentose sugarD-xyloseisoneofthemostabundant components which

will result from a complete chemical or enzymatic hydrolysis of nonstarch
polysaccharides of feed ingredients of vegetable origin. Because of the
uncertainties about the nutritional value ofD-xylose, two trials with pigs
were conducted toinvestigate the urinary excretion ofxylosein relation to
the ageofpigs,frequency offeeding and dietary inclusion levelofD-xylose.
Moreover the effect of inclusion of D-xylose in pig diets on N and energy
utilization was examined.Urinary excretionofxylosewasnot significantly
affected by age and frequency offeeding. The extent ofurinary excretion of
xylosein%ofintake increased linearly(P<0.05)asthe dietarylevelofthis
sugarwasincreased.Inpigsfedonadietcontaining25gD-xylose/kg,about
20%ofthe D-xylose consumed appeared in the urine. This level increased
to about 43%when pigs were fed on a diet containing 100 g D-xylose/kg.
Retention ofNwasslightlydecreased whenpigswerefed 100gD-xylose/kg
diet.Urinary excretion ofenergybearing components tended toincrease in
pigs fed on D-xylose diets. Liver and kidney weight, pH ofurine and blood
composition were not significantly affected by inclusion of D-xylose in the
diets. (Key words: D-xylose, excretion, pigs)
INTRODUCTION
Nonstarch polysaccharides (NSP) can form a major fraction of the

carbohydrate content of practical diets for pigs. These NSP include a
mixture of substances such as cellulose, hemicellulose, pectin and
oligosaccharideswhichcontainhexoseandpentosesugarsanduronicacids.
It is wellknown that NSP are resistant tothe digestive enzymes of saliva,
stomach and small intestine ofpigs.As a result they pass to the hind gut
where microbial degradation takes place.The end products ofa microbial
degradation ofNSP (lactic acid, volatile fatty acids) are readily absorbed
andcanbeutilizedbythepigasanenergysource,butwithalower efficiency
102 Chapter 5
thane.g.glucose(AgriculturalResearchCouncil,1981;Justetal.,1983;Van
Es, 1987).
In a literature review, Chesson (1987) concluded that the digestibility of
feed ingredients containing high levels of NSP can be improved by
treatment withenzymeswhichcanhydrolysetheNSPto monosaccharides.
Thiswasconfirmed ina recently performed study at ourinstitute (Schutte
et al., 1990).Our study showed that in addition to an improvement of the
digestibility ofcell wall components, digestion ofprotein and fat was also
improved in pigs when wheat bran was treated with a cellulolytic enzyme
preparation. However, it remains an open question as to what extent
pentosesugarsanduronicacidscanbeutilizedinpigs.NexttoD-glucosethe
pentose sugar D-xylose is one of the most important components to be
released in an enzymatic hydrolysis of NSP (Carr£ & Brillouet, 1986;
Brillouetetal.,1988).Itiswellrecognizedthat D-xyloseisreadily absorbed
fromtheintestinaltract ofmonogastricanimals(Cori,1925;Miller&Lewis,
1932;Arnal-Peyrot &Adrian, 1974; Schutte et al., 1991a, 1991b). These
studies also showed that part of the ingested D-xylose is excreted in the
urine.Theextentofurinaryxyloseoutputmaybeaffected byseveral factors
like intestinal bacterial growth, state of health, age and dietary level
(Hindmarsh, 1976).
In the two trials reported herein the influence of frequency of feeding,
ageanddietaryD-xyloselevelonurinaryexcretionofxylosewas investigated
in pigs. In addition, in these trials the effect of D-xylose on nitrogen and
energy utilization was examined.
MATERIALS A N D METHODS
Animals and diets
(DutchLandracexDutchYorkshire).Inbothtrialsthepigswere individually
housed in metabolism cages under a 12 h light - 12 h twilight cycle
throughout.Thenutritionallycompletebasaldietusedwasbasedonmaize,
wheatstarchandisolatedsoyaprotein.Thecompositionofthebasaldietand
its chemical characteristics are shown in Table 1.
The test sugars (D-glucose and D-xylose), supplied as anhydrous mono-
saccharides,weresubstituted byweightforwheat starch.Inbothtrials the
experimental diets were fed at a daily rate of approximately 0.9 MJ
metabolizable energy (ME)/kg metabolic body weight, assuming that D-
xylosehasthe sameMEcontent asD-glucose.Thedailyamount offeed was
adjusted weekly accordingtobodyweight. The feed was mixed with water
(1part feed + 1part water). In addition, water was freely available.
D-xyloseinpigs 103
TABLE 1.Composition of thebasal diet.
Ingredient g/kg
Maize meal 287

Wheat starch 287
Soyabean oil 40
Animal fat 40
Cellulose ("Arbocel B800") 60
Limestone 10
Iodized salt 10
Mineral/vitamin mix * 5
DL-methionine 0.7
Contents (g/kg) Exptl Expt2

Dry matter ** 886 891
Gross energy (MJ/kg)** 17.6 17.8
Crude protein ** 216 223
Calcium *** 8.4 8.4
Phosphorus *** 7.0 7.0
Lysine *** 12.6 12.6
Methionine + cystine *** 6.8 6.8
Threonine *** 8.3 8.3
Tryptophan *** 2.6 2.6
* Provided (mg/kgdiet):magnesium, 400;zinc, 110;copper,25;

manganese,45; iron,80;cobalt,0.5;selenium 0.1;thiamin,2;
riboflavin, 5;nicotinamide,30;pantothenic acid, 12;pyridoxine,3;
cyanocobalamin, 0.04;biotin, 0.1;folic acid, 1;menadione,3;
ascorbicacid, 50;retinol, 3.1;cholecalciferol, 0.045;vitamin E,
40;cholinechloride,1000.
**Analysed ***Calculated
104 Chapter 5
Experiment 1.Theobjectives ofthistrialweretodetermine the effect ofD-

xylose at a dietary inclusion level of 100 g/kg on faecal digestibility of
nitrogen(N)andgrossenergy(GE),retention ofN,andurinaryexcretionof
xylose,Nandenergyinrelationtofeedingfrequency.Thefeeding frequencies
applied were 2 and 4 times/d, respectively. The daily amount of feed was
offered at two, respectively, four equal meals at intervals of 12 and 6 h,
respectively.
The trial involved 12pigs with a mean age of8weeks at the start of the
trial.Thepigswereaccustomed tocagesandbasal diet(Table 1)for 25days
before starting the experimental period. At the start of the experimental
period three groups offour pigs,each ofsimilar average bodyweight, were
formedandfeddietscontainingeitherD-glucoseorD-xylose.Asisillustrated
in Table 2, the experimental period consisted oftwo phases. During both
phases the D-glucose diet (treatment 1)was fed four times/d, whereas the
frequency offeeding ofpigsfed the D-xylose diets (treatments 2and 3)was
changed in phase two.Each ofthe twophases consisted ofa 4d adaptation
and a 5d collection period.
At the start ofthe experimental period, the pigsweighed 25.2 (SD 1.1) kg
and at the end 34.6 (SD 1.2) kg.
TABLE 2. Design of experiment 1.
Treatment n Sugar* Frequency of feeding

Phase 1 Phase 2
1 44 D-glucose
D-glucose 4 times/d
4 times/d 4 times/d
2 44 D-xylose
D-xylose 2times/d
2 times/d 4 times/d
3 44 D-xylose
D-xylose 4 times/d
4times/d 2 times/d
* Included in the diet at a level of 100 g/kg.
Experiment 2.In this trial the effect ofgraded dietary levels (25to 100g/
kg) of D-xylose on N digestibility, N retention, and urinary excretion of
xylose and N in relation to the age of the pigs was examined. Moreover,
specific density and pH of urine, liver and kidney weight, and blood
composition were examined.
This trial involved 16pigs:8 young pigs with an age of 7weeks and 8
older pigs with an age of 15weeks at the start ofthe trial. The pigs were
D-xylose in pigs 105
accustomed tocagesandbasaldiet (Table 1)for 14daysbefore starting the

experimental period.Atthe start ofthe experimental periodtwo treatment
groups each involving 4young and 4 older pigs were formed and fed diets
containing either D-glucose or D-xylose. As is illustrated in Table 3, the
experimental periodconsistedoffour phases,duringwhichtimethepigsof
thetwotreatment groupswerefedconsecutivelyonadietcontaining 25,50,
75and 100gD-glucoseorD-xylose/kg.Thedailyamountoffeedwas offered
at twoequal mealsat intervals of12h.Each ofthefour phases consisted of
a 3dadaptation and a4dcollectionperiod.Atthe start ofthe experimental
period theyoung and older pigsweighed onaverage 17.0(SD 1.3) and 55.3
(SD5.6)kg,respectively.Atthe endofthisperiodthepigsweighed 27.9(SD
1.8) and 74.8 (SD 8.0) kg, respectively.
TABLE 3. Design of experiment 2.
Treatment n Sugar Dietary sugar level (g/kg)

phase 1 phase 2 phase 3 phase 4
1 8 D-glucose 25 50 75 100
2 8 D-xylose 25 50 75 100
faeces and urinewascarried outduringeachfive(Expt 1)and four (Expt 2)
24 h collection periods from the individual animals. The faeces were
collectedatintervalsof12h,andstoredat-20°C.Allfaecesproduced during
eachcollectionperiodwerepooledforeachpigseparately,homogenized and
sampled. Next the samples were freeze-dried. Urine was collected in
containersprovidedwithmerthiolatesodium(Thiomersal,BDH,Chemicals
Ltd, Poole, England) at intervals of approximately 4 h. The portion from
each interval was pooled daily from individual animals.A representative
sample of10%ofthe pooled urine was taken and frozen at -20°C.The five
(Expt 1)and four (Expt 2) day sub-samples ofurine were pooled for each
animal separately, homogenized and sampled. Faeces and urine were kept
at -20 °Cbetween sampling and before analysis.
106 Chapter 5
Analytical methods
Samplesoffeed andfreeze-dried faecesweremilledtopassthrough a 1.0

mmscreen(RetschmillZMI,RetschBV, Ochten,TheNetherlands) before
analysis. All analyses were carried out in duplicate. Dry matter was
determinedbydryingthe samplestoconstantweightat 101°C.Inorganic
matterandnitrogenweredeterminedbystandardmethods(Associationof
OfficialAnalyticalChemists,1975).Grossenergywasdeterminedusingan
IKA-C 4000 adiabatic bomb calorimeter. Concentrations of glucose and
xylose in urine were determined according the procedure described by
Schutteetal.(1991a).
Theresultsofbothtrialswereanalysedbymeans ofanalysisofvariance
(Cochran & Cox, 1957). The computer program Genstat 5 (Reference
Manual,1987,OxfordUniversityPress,NewYork)wasusedtocalculatethe
analysisofvariance.Althoughthetreatmentswereconfoundedbytimeand
age,itwasassumedthat differences areduetothetest sugars orincrease
in dietary sugar. In Expt 1 the treatment factors were type of sugar,
frequency offeedingandphase.Thedifferences inresultsachievedonthe
D-xylosedietsinthe first and secondphase at an equalfeeding frequency
weresmallandstatisticallynotsignificant.Therefore,theresultsobtained
atanequalfeedingfrequency werecombinedinthestatisticalanalysis.In
Expt2thetreatment factors weretypeofsugar,dietarylevelofsugar and
age.Inthisexperimentthesumofsquaresforlevelswaspartitionedintoa
setoforthogonallinear andquadraticpolynomialregression components.
RESULTSAND DISCUSSION
Table 4, which incorporates values of a previous study (Schutte et al.,

1991a), indicates that D-xylose was digested almost completely at the
terminal ileum.Thiswould suggest an almost completeabsorption ofthis
pentose sugar as such. However, administration of D-xylose to pigs was
associatedwithanincreased ilealflowofVFAandadecreased ilealchyme
pH.Bothsymptomspointtoamoreextensivemicrobialactivityinthesmall
intestine. This is further supported by the decrease in apparent ileal
digestibility ofNinpigsfed ontheD-xylosediets.Fromtheresults ofthis
studyiswasconcludedthat atleastpart oftheingestedD-xylosehasbeen
consumedbytheintestinalmicrobes.Thisconclusionissupportedbydata
ofSchiffer etal.(1962),Cookeetal.(1963)and Goldsteinetal.(1970)who
D-xyloseinpigs 107
TABLE4.Apparent ilealdigestibility dataofpigsfedon D-glucose

andD-xylose diets *.
Sugar D-glucose D-xylose

Dietary level (g/kg) 200 100 200
Digestibilities
OM 87.6 a 87.2 a 84.7"
GE 87.6 a 87.5 a 84.5"
N 90.3 a 89.l a 87.2"
D-glucose 99.3
D-xylose 98.7 a 98.6 a
Ileal chyme pH 6.5a 6.2" 6.0C
Ileal flow ofVFA(mg/12h) 1106a 2062" 4888c
* Data from Schutte et al.(199la).

a,b,cWithinarow,meanvalueswith different superscript letters were
reportedanincreasedurinaryxyloseexcretionafterantibioticsinmanwith
small intestinal diverticulosis.
Theextent of microbial degradation of D-xyloseinour studywithpigs
cannotbederived simplyfrom thedifferences inilealflowofVFAbetween
the D-glucose and D-xylosetreatments, because some ofthe VFAwill be
absorbed alreadyinthesmallintestine.Wyngaardenetal.(1957)reported
thatapproximately 15%ofanintravenouslyinfused doseofD-xylosecanbe
recoveredascarbondioxideintheexpiredair.Inapreviousstudy(Schutte
etal.,199la)itwasfoundthatatadietaryinclusionlevelof100g/kg,about
45%ofthe ingested D-xylosewas excreted inthe urine ofpigs.Assuming
that 15%ofthe dosehasbeenmetabolized tocarbondioxide (Wyngaarden
etal.,1957),theremaining40%notaccountedformayhavebeenfermented
bythe intestinal microbes.
Oneofthemainobjectives ofthepresent experiments wasto investigate
whetherurinaryexcretionofxyloseisaffectedbyfeedingfrequency,ageand
dietaryinclusion levelofthispentose sugar inpigs. Theresults ofExpt 1
(Table5)showthat urinary excretion ofxylosewasnotclearlyaffected by
the frequency offeeding diets containing this pentose sugar. The dietary
108 Chapter 5
inclusion level of D-xylose, however, did appear to effect the urinary

excretionofxylose(Expt2,Table6).Inbothyoungandolderpigsapositive
correlationbetweenthedietarylevelandtheurinaryexcretionofxylosewas
found.Thiscorrelationwasmorepronouncedintheyoungthanintheolder
pigs. Taking the results of both ages together it appeared that urinary
excretionofxylosein%ofintakewaslinearly(P<0.01)increasedwhenthe
dietary level ofthis sugar was increased. Similar results were found in a
previoustrialwithileostomizedadultroosters(Schutteetal.,1991b).Wagh
& Waibel (1966) reported that the ME value of D-xylose in chicks was
decreasedwhenthedietarylevelwasincreased.Thisobservation suggests
alsoadosage-dependenturinaryexcretionofD-xylose.Inthepresentstudy
noVFAmeasurementsintheilealchymewereperformed.Thusitcannotbe
simply stated that the observed dosage-dependent urinary excretion of
xyloseisexclusivelyduetodifferencesinmicrobialdegradationofthissugar
at the different dietarylevels.In addition the lowrenal threshold for this
sugarassuggestedbyLoos(1954)mayhaveaffected ourresultsforurinary
excretion.
Fowler&Cooke(1960),Finlayetal.(1964)andHindmarsh(1976)reported
that urinaryxyloseoutputinmandeclineswithage.Thereasonforthisis
unknown,butithasbeenpostulatedthatrenalfunction, and consequently
xylose excretion, is affected bythe ageingprocess (Kendall, 1970).In our
study an age dependent urinary excretion of xylose was not clearly
demonstrated (Table6).
Thelossesofxyloseintotheurinewerereflected intheurinary excretion
ofenergy(Tables5and6),butthedifferences inurinaryexcretionofenergy
betweentheD-glucoseandD-xylosetreatmentscouldnotbefully explained
bythexyloselosses.Calculationshavepointedoutthatwhentheincreases
inurinaryexcretionofenergyoftheD-xylosetreatmentsovertheD-glucose
treatmentswereattributed toD-xylose,thiswouldrepresentabout 50%of
theD-xyloseintakeinExpt1(Table5).Thuscomparedtothelossesofxylose
intheurine(40%),10%oftheenergyexcretedintheurineisnotaccounted
for. In Expt 2(Table6)the extralossesofenergyintotheurine ofpigsfed
ontheD-xylosedietswouldrepresentabout47and52%ofthexyloseintake
inyoungandolderpigs,respectively.Thevaluesfoundforurinaryexcretion
ofxyloseinbothyoungandolderpigsweremuchlower,beingasamean32
and 35%,respectively. It isobviousthat other energy bearing substances
thanxylosehavecontributed totheextralossesofenergyintotheurineof
pigs fed onthe D-xylosediets.This is supported bythe slight increase in
urinary excretion of N when pigs were fed on diets containing 100gD-
xylose/kg(Tables5and 7).Wiseetal.(1954)reportedthat Nretention was
significantly decreased whenpigswere fed ona diet containing 560gD-
D-xylosein pigs 109
TABLE5. Expt1.Influence of feeding frequency of D-xylose diets

onN and energy utilization, and urinary excretion of xylose.
Sugar(100g/kg diet) D-glucose D-xylose SED

Frequency of feeding 4 times/d. 4 times/d.2 times/d. 1-2 2-3
(1) (2) (3) 1-3
DM intake (g/pig/d) 754a 751 a 764a 9.9 4.1
Faecal digestibility (%)

N 92.5 a 92.9 a 93.0a 0.5 0.4
Gross energy 90.3 a 90.5 a nd 0.6 -
Urinary excretion (%of intake)

Glucose + + +
Xylose + 41.0a 39.9 a - 1.8
N 32.2 a 35.9 a 37.4a 2.4 2.3
Energy 2.3 a 7.2b nd 0.6 -
N retained (%of intake) 60.4a 57.0a 55.68 2.3 2.2
ME (MJ/kg DM)
Diets * 16.7a 15.9b nd 0.1
D-glucose ** 15.2 " "
D-xylose *** 7.8 nd
SED =standard error ofdifference ofmeans,

nd =not determined.
Small traces ofglucose (0.1-0.3g/L)and xylose(0.01-0.2g/L)were

found inthe urine ofthese experimental treatments.
CorrectedtozeroNbalancebyusingthefactor 31.4kJ/gretainedN.
Calculated as 98%ofGEvalue.
Derivedfrom hedifferences inMEvaluebetweentheD-glucose and
D-xylosediets.
a,b Within arow,meanvalueswith different superscript letters were
110 Chapter 5
TABLE6.Expt2.Urinary excretion ofxylose andenergy in young

(A)andolder (B)pigs fed onD-glucose (Glue)and D-xylose(Xyl)
diets.
Sugar Dietary DM intake Xylose excreted Energy excreted

level (g/pig/d) murine in urine
(g/kg) (%of intake) (%of intake)
A B A B A B
Glue 25 524 1142 + + 2.76 3.61

50 569 1220 + + 2.92 3.36
75 621 1291 + + 2.75 3.41
100 678 1362 + + 2.96 3.34
Xyl 25 540 1100 18.4 22.6 3.87 4.34

50 604 1254 29.2 34.1 5.04 5.96
75 666 1326 36.6 41.3 5.51 7.60
100 723 1435 43.2 42.5 8.18 9.19
Analysis ofvariance (exclusive DM intake)

between animal stratum
Age 0.41 0.01
Sugar - 0.01
Agex sugar - 0.14
Residual MS 110.5 0.46
+ Small traces (0.01-0.1g/1) ofxylosewerefound inthe urine ofthese

D-xylosein pigs 111
xylose/kg,indicatingalsoagreaterurinaryexcretionofN.Theseinvestigators
believethat the observed reduction inNretention inpigsfed onD-xylose
dietswasduetoanenergydeficiencyandconsequentlygreaterNcatabolism.
ThisissupportedbydataofWagh&Waibel(1966)whofoundthat plasma
uricacidwassignificantlyincreasedwhenchickswerefedondietscontaining
200and400gD-xylose/kg.Further theyreported that feeding these diets
tobirdsresulted in decreased liverweights.Thedata ofWiseet al.(1954)
andWagh&Waibel(1966)arenotstrictlycomparablewiththeresultsofour
study sincelowerlevels ofD-xylosewereincluded inour diets.An energy
deficiency when occuring in our study, was not reflected in liver weights
(Table8).
Wagh&Waibel(1967)reported thatbloodhematocrit,cholesterol, serine
and proline in plasma were increased and plasma glutamic acid was
decreasedwhenbirdswerefedondietscontaining100to400gD-xyloseper
kg.In ordertostudywhether ornotinclusion ofD-xyloseinpigdietswill
TABLE7. Ezpt 2. Apparent faecal digestibility of N, urinary

excretion ofNand Nretained inyoung (A)and older (B) pigs
fed onD-glucose (Glue)and D-xylose (Xyl)diets.
Sugar Dietary N digestibility N excreted N retained

level (%) in urine (%of intake)
(g/kg) (%of intake)
A B A B A B
Glue 50 92.8 93.9 38.1 51.5 54.7 42.4

100 93.2 94.3 40.4 53.7 52.8 40.7
Xyl 50 93.0 94.3 38.5 51.4 54.5 42.9

100 93.1 94.4 41.6 55.9 51.5 38.5
Analysis of variance

Age 0.01 0.01 0.01
Sugar 0.65 0.60 0.65
Age x sugar 0.73 0.95 0.99
Residual MS 1.08 22.81 23.0
112 Chapter 5
changebloodcomposition,bloodsampleswerecollectedfrompigsofExpt2.
These samplesweretaken after termination ofthelastphase(phase4) and
involved the following determinations; leucocytes, hemoglobin (Hb),
erythrocytes, hematocrit, mean corpuscular value, mean corpuscular Hb
concentration, glucose, bilirubin, bilirubinester, cholesterol, triglycerides,
albumine and total protein. The differences in these blood parameters
between pigs fed onthe D-glucosediets and those fed onthe D-xylosewere
small and not significant. Further it is noteworthy that histo-pathological
examination oftheliverandkidneys didnotshowabnormalities inpigs fed
on either D-glucose or D-xylose diets (Expt 2). Specific density of urine
appeared to be slightly increased (Table 8) when pigs were fed on diets
containingD-xylose;thiswillberelatedwiththeincreasedurinary excretion
of energy on this diet. No effect of D-xylose on the pH value of urine was
observed (Table 8).
TABLE 8. Expt 2. Specific density and pH of urine, and liver and

kidneyweight (in%ofbodyweight)inyoung(A)andolder(B)pigs
fed on D-glucose (Glue) and D-xylose (Xyl) diets.
Sugar Urine values * Organ weights

Spec, density pH Liver Kidney
A B A B A B A B
Glue 1.006 1.023 7.1 6.7 1.8 1.5 0.372 0.282
Xyl 1.014 1.026 7.0 6.6 1.9 1.6 0.352 0.301
Analysis of variance

Age 0.01 0.01 0.01 0.01
Sugar 0.06 0.10 0.34 0.96
Age x sugar 0.34 0.75 0.56 0.22
Residual MS 32.11 0.01 0.02 0.01
*Determined during phase 4 only; each value represents 16 individual

observations (one observation/animal/d).
D-xylosein pigs 113
In conclusion it can be stated that the energy value ofD-xyloseis much

lower than that ofD-glucose. From the results ofExpt 1it was calculated
that at a dietary level of 100g/kg,D-xylosehas a ME value ofonly 7.8 MJ/
kg, whichvalueisapproximately 50%ofthat ofD-glucose.Considering the
dataforurinaryexcretionofxylose,itmay beexpectedthatatlower dietary
inclusion levels than 100 g/kg the ME value of this pentose sugar will
increase.ThenetutilizationofD-xyloseinpigsisdifficult toaccessfrom the
presentstudy.Thisbecauseoftheunknownmetabolicpathwayofthis sugar.
Theresultsofapreviousstudy(Schutteetal., 1991a)indicated that at least
partofthexyloseisfermented inthesmallintestine.Itmayevenbepossible
that allxylosehastobefermented beforeitcanbeused asan energy source
for pigs.
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Chesson,A., (1987). Supplementary enzymes to improve the utilization of
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114 Chapter 5
Finlay, J.M., Hogarth, J. &Wightman, K.J.R. (1964).Aclinical evaluation

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Goldstein, F.,Karacadag, S., Wirts, C.W. &Kowlessar, O.D.(1970).
Intraluminal small intestinal utilization ofD-xylose by bacteria.
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Kendall, M.J. (1970).The influence ofage on the xylose absorption test.
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Loos,M. (1954). Studies in the utilization ofpentoses in diabetes.Acta
Miller, M.M. &Lewis, H.B. (1932).Pentose metabolism.l. The rate of
absorption ofD-xylose and the formation ofglycogen in the organism
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Biological Chemistry 98:133-140.
Schiffer,L.M.,Faloon, W.W.&Chodos,R.B.(1962).Malabsorption syndrome
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Schutte, J.B.,Kempen, G.J.M. van, &Hamer, R.J. (1990).Possibilities to
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Schutte, J.B.,Jong, J. de, Polziehn, R. &Verstegen, M.W.A. (1991a).
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D-xyloseinpigs 115
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117
CHAPTER 6
L-ARABINOSE IN PIGS
J.B. Schutte,* J. de Jong,* E.J. van Weerden* and

S. Tamminga**
*TNO-InstituteofAnimal Nutrition and Physiology(ILOB),P.O.Box15,

**Department ofAnimalNutrition,WageningenAgricultural University,
P.O.Box338,6700AH Wageningen,The Netherlands
Accepted for publication in:British Journal of Nutrition

Reproduced bypermission ofthe Nutrition Society.
119
L-ARABINOSE IN PIGS
J.B.SCHUTTE,*J.DEJONG,*E.J.VANWEERDEN*
ANDS.TAMMINGA**
*TNO-Institute ofAnimal Nutrition and Physiology (ILOB), P.O.Box 15,
**Department ofAnimal Nutrition, WageningenAgricultural University,
P.O.Box 338,6700AH Wageningen, The Netherlands
The pentose sugar L-arabinose is one ofthe most abundant components

whichwillbecomefreeinacompletehydrolysisofnonstarch polysaccharides
of feed ingredients of vegetable origin. Two studies were conducted to
investigate the apparent ileal digestibility and urinary excretion of L-
arabinose at dietaryinclusion levelsof50and 100g/kg, and 25,50,75 and
100g/kg,respectively,in pigs.Asa reference D-glucosewasincluded in the
studies. Water intake, ileal flow ofvolatile fatty acids and ileal and faecal
digestibility ofdietary nutrients inpigsfed onthe different diets were also
examined. Castrated pigs were prepared with a post-valvular T-caecum
cannula tomeasure ilealdigestibility. Faecal digestibilitywasmeasured in
non-cannulated pigs.Apparent ileal digestibility ofL-arabinose was found
tobeapproximately 70%.Thepresence ofL-arabinose inthediet increased
ilealflow ofvolatilefatty acidsand lacticacid,suggestingthe occurrenceof
microbialdegradationofL-arabinoseinthepigsmallintestine. L-arabinose
was partly excreted in the urine. The extent of this urinary excretion in
percentage of intake increased linearly (P < 0.01) as the dietary level
increased. In pigs fed on the 25 g L-arabinose/kg diet, 10.9 % of the L-
arabinose consumed appeared in the urine. This level was increased to
14.7%whenpigswerefedadietcontaining 100gL-arabinose/kgdiet.Faecal
digestibility and retention ofnitrogen decreased significantly inpigsfed on
the L-arabinose diets. (Key words:L-arabinose, Digestion, Excretion, Pig)
INTRODUCTION
Traditional pigdiets are mainly composedoffeed ingredients ofvegetable

origin. The carbohydrate fraction ofthese ingredients contains two broad
classesofpolysaccharides,starchand cellwallpolysaccharides;thelast one
120 Chapter 6
may be conveniently referred to as nonstarch polysaccharides. Starch, a

storagecarbohydrate,canbehydrolyzedbypancreaticoc-amylaseandmay
thereforebedigestedinthesmallintestineofpigs,andabsorbedasglucose.
Nonstarchpolysaccharides(NSP)arecomplicatedcompoundsbothfromthe
point ofviewofphysical structure and chemical composition, and include
cellulose,hemi-cellulose,pectinandoligosaccharides.Itiswellrecognized
that NSP are resistant to the digestive enzymes of pigs and pass to the
hindgut where microbial degradation takes place. The end products ofa
microbialdegradationofNSP,lacticacidandvolatilefattyacids,arerapidly
absorbedintothebloodandcanbeutilizedbythepigasanenergysource,
butwithalowerefficiencythane.g.glucose(AgriculturalResearchCouncil,
1981;Just et al. 1983;van Es,1987).
Improving the utilization of NSP may be attained by treatment with
enzymes capable of hydrolysing these carbohydrate fractions to
monosaccharides. However,a completehydrolysis ofthe NSPwill release
not onlyglucose,but alsoothersugars,ofwhichinquantitative terms the
pentosesugarL-arabinoseisoneofthemostimportant(Carre'andBrillout,
1986;Brilloutetal.1988).Thestudiesreportedsofarontheabsorptionand
utilization ofL-arabinose relate to animal species other than pigs.These
studieshaveshownthat L-arabinose isabsorbed from theintestinal tract
in rats (Cori, 1925;Arnal-Peyrot and Adrian, 1974) and chicks (Bogner,
1961;WaghandWaibel,1967),butat alowerratethanglucose.The study
reportedbyArnal-PeyrotandAdrian(1974)showedthatpartoftheingested
arabinose is excreted in the urine. Both, the low absorption rate and the
urinary excretion ofL-arabinose, may have nutritional and physiological
implicationsfortheanimalasindicatedinchicksbyWaghandWaibel(1966)
and Schutte (1990). From these studies, it appears that in chicks the
metabolizableenergyvalueofL-arabinosenotonlywaslowerthanthatof
D-glucose,butalsodecreasedwhenthedietarylevelwasincreased.Moreover,
these investigators found that inclusion of L-arabinose resulted in wet
droppings.
Thetwoexperimentsreportedhereinwereundertakentoobtaininformation
onthe quantitative aspects ofdigestion and utilization ofL-arabinose in
pigs. In the first experiment ileo-cannulated pigs were used in order to
measure the disappearance rate ofL-arabinose at the endofthe terminal
ileumatdietaryinclusionlevelsof50and 100g/kg.Thesecondexperiment
wasperformedwithintactanimalstodeterminetheurinaryexcretionofL-
arabinose at dietary levels of 25, 50,75and 100g/kg. In addition, in
thesetrialstheeffect ofL-arabinose onilealandfaecal digestibilityofthe
dietarynutrientswasinvestigated.D-glucosewasincludedintheexperiments
as a reference.
L-arabinose inpigs 121
Animals and diets
(DutchLandracexDutchYorkshire):onetrial withcannulated pigs(Expt
A)andonewithnon-cannulatedpigs(ExptB).Inbothtrialsthepigswere
individuallyhousedinmetabolismcagesunder a 12hlight -12h twilight
cyclethroughout.Thenutritionallycompletebasaldietusedwasbasedon
maize,wheatstarchandisolatedsoyaprotein.Thecompositionofthebasal
dietanditschemicalcharacteristicsareshowninTables1and2,respectively.
The test sugars (D-glucose and L-arabinose), supplied as anhydrous
monosaccharides,were substituted byweightfor wheat starch.
Inbothtrials the experimental dietswerefed at a dailyrate ofapproxi-
mately 0.9 MJ metabolizable energy (ME)/kg metabolic body weight,
assumingthatL-arabinosehasthesameMEcontentasD-glucose.Thedaily
amount of feed was offered at four equal meals at intervals of 6 h, and
adjusted weeklyaccordingtobodyweight.Thefeed wasmixedwith water
(1part feed + 1 part water).Inaddition, water wasfreely available.
ExptA
Theilealdigestibility ofL-arabinose, and the effect ofthis pentose sugar

onilealdigestibilityofdrymatter(DM),organicmatter(OM),grossenergy
(GE)andnitrogen(N)weredetermined.Moreover,ilealflowofvolatilefatty
acids(VFA)and lacticacid was measured.
Four pigs, 9weeks oldat the start ofthe trial,wereinvolved.Thepigs
were surgically fitted with a post-valvular T-caecum cannula (PVTC)
according tothe procedure described byVanLeeuwen et al. (1988).Post-
operativecareincludedkeepingthepigswarm(25°)andwitholdingfeedfor
24h.Duringthe 3-week post-operative period,the pigs werefed on the
basal diet (Table 1). Theexperimental period involved originally three
122 Chapter 6
TABLE 1.Composition ofthebasal diet (g/kg)
Maize meal 287

Wheat starch 287
Soya-bean oil 40
Animal fat 40
Cellulose* 60
Limestone 10
Iodized salt 3
Mineral mix+ 5
Vitamin mix++ 5
DL-methionine 0.7
* ArbocelB800(Rettenmaier, FRG)
+ Provided(mg/kgdiet):magnesium 400,zinc110,copper25,manganese
45,iron 80,cobalt 0.5,selenium 0.1.
-H-Provided(mg/kgdiet):thiamin 2,riboflavin 5,nicotinamide30,
pantothenic acid 12,pyridoxine 3,cyanocobalamin 0.04,biotin0.1,
folicacid 1,menadione3,ascorbicacid50,retinol3.1,
cholecalciferol 0.045,vitamin E40,cholinechloride 1000.
TABLE2.Chemical composition ofthebasal diet (analysed, g/kg

unless otherwise stated)
Constituent ExptA ExptB
Dry matter 901 894

Ash 53 52
Crude protein (Nx 6.25) 221 223
Crude fibre 59 57
Crude fat 84 83
Gross energy (MJ/kg) 17.8 17.7
Calcium 9.8 9.6
Phosphorus 8.6 8.3
L-arabinose in pigs 123
phases during which time each pigwas fed consecutively a diet containing
100gD-glucose/kg(Glue 100),50 gL-arabinose/kg(Arab 50) and 100g L-
arabinose/kg(Arab 100),witha7dadaptation anda4dcollectionperiod for
each diet. During the first three days ofeach adaptation period, pigs were
gradually changed to the next diet.After termination ofthe third phase, a
fourth phasewasincludedinthetrialinordertoinvestigatewhetherornot
theobservedincreaseinilealdigestaoutputinpigsfed ontheArab 100 diet
returnedtoanormallevelwhentheywerechangedtotheGlue100diet.The
fourth phaseconsistedofan 1dadaptationanda4dcollectionperiod.Atthe
start ofthe experimental period,the pigsweighed onaverage 26.4(SD 1.6)
kg and at the end ofthe fourth phase 51.2 (SD 5.2) kg.
TABLE 3.Experimental design of experiment B.
Treatment Sugar Dietary.3ugar level (g/kg)
Phase 1 Phase 2 Phase 3 Phase 4

(1-11d) (12-22 d) (23-33 d) (34-44 d)
1 D-glucose 25 50 75 100
2 L-arabinose 25 50 75 100
ExptB
The objectives ofthis trial were to determine the urinary excretion of L-

arabinose,and tostudythe effect ofthis sugaronfaecal digestibilityofDM,
OM, GE and N, and N retention. This trial, involving ten 9-week old pigs,
was performed after termination of Expt A. The pigs were accustomed to
cages and basal diet (Table 1)for 3weeks before starting the experimental
period.Thebasaldietwascomposedofthesamebatchesoffeed ingredients
as used in ExptA. At the start ofthe experimental period two groups of5
pigs, each of similar average body weight, were formed and fed diets
containingeitherD-glucoseorL-arabinoseaccordingtoaschemeasoutlined
inTable 3.Asisillustrated inTable 3,the experimental period consisted of
four phases.Each ofthefour phases consisted ofa 7dadaptation and a 4d
collection period. During the first three days ofthe adaptation period, pigs
weregraduallychangedtothenextdietcontainingahigherlevelofthe same
sugar.
At the start ofthe experimental period, the pigsweighed 27.5 (SD 1.5) kg
and the end ofthis period 55.8 (SD 3.0) kg.
124 Chapter 6
Digesta collection
Ilealdigestaduringeach4dcollectionperiodwascollectedfrom the PVTC

cannula quantitatively from individual animals over a 12h period per day
(8.00-20.00hours).Previous studieshad shownthatilealdigesta collection
is almost complete byusing this type ofcannula on the same type ofbasal
diet as in the present trial (Kohler et al. 1991). Previous studies had also
shown that there were nosignificant differences in ileal digestibility when
digesta were collected over a 12h or over a 24 h period per day (E.J. van
Weerden, J. Huisman &P.van Leeuwen, unpublished results).
The digesta were collected continuously in dry ice,weighed daily and
stored at -20°.Atthe end ofthe experiment, the four 12hcollected portions
were pooled for each pig separately, homogenized and sampled. VFA and
lacticaciddeterminations wereperformed inthe digesta as such,the other
measurements in freeze-dried samples. Until analysis all samples were
kept at -20°.
faecesandurinewascarriedoutduringthefour24hcollectionperiods from
the individual animals. The faeces were collected at intervals of 12h, and
storedat -20°.Allfaeces produced duringeachcollectionperiodwere pooled
for eachpigseparately, homogenized and sampled. Then the samples were
freeze-dried.
Urine was collected in containers at intervals of4 h.At the start of each
collection period, the containers were provided with merthiolate sodium
(Thiomersal, BDM, Chemicals, Ltd, Poole, England) as a preservative to
inhibit bacterial activity. Before inclusion, this preservative was dissolved
in an ethanol solution (4 g/100 ml), and added in an amount of 0.4 ml/
container.This amount wasbased onan urine production of200mlper4 h.
The portion of urine from each interval was pooled daily from individual
animals.Arepresentative sample of10%ofthepooledurinewastaken and
frozen at -20°.The 4 d sub-samples of urine were pooled for each animal
separately, homogenized and sampled. Faeces and urine were kept at -20°
between sampling and before analysis.
Analytical methods
Samples offeed and freeze-dried digesta and faeces were milled to pass
through a 1.0 mm screen (Retsch mill ZM1, Retsch B.V.,Ochten, Holland)
beforeanalysis.Allanalyseswerecarriedoutinduplicate.DMwasdetermined
by drying the samples to a constant weight at 101°.Inorganic matter and
Nweredetermined bystandard methods (AssociationofOfficial Analytical
Chemists, 1975),GEwas determined usingan IKA-C4000adiabaticbomb
calorimeter.
Concentrations ofVFAand lactic acid in wet digesta were determined by
amodification ofthegas-liquidchromatographicmethodofImoto&Namioka
(1978). A known portion (about 20 g) of the digesta was centrifuged.
Immediately afterwards the supernatant fraction (5ml)was acidified with
500ulphosphoricacid (850ml/1),3mlofan aqueous solution ofisocapronic
acid(4.0193g/1)wasaddedasaninternalstandard.Distilledwaterwas then
added tothe mixture toobtain a final volume of 10ml.A 1ul sample of the
final solutionwasinjected intothecolumnofthegas-liquid chromatograph.
The gas-liquid chromatograph was fitted with a flame ionization detector
(Packard 419, USA).Aglass column (1850 mm x 2 mm i.d.) packed with
Chromosorb 101 of 80/100 mesh was used. The carrier gas (N2 ) was
saturated with formic acid, and had a flow rate of 25 ml/min. The oven
temperaturewassetat 190°,andtheinletanddetectortemperature at225°.
Standard solutions containing acetic acid, propionic acid, butyric acid,
isobutyricacid,valericacidandisovalericacidwereprepared for gas-liquid
chromatographyinthesamewayasdescribedforisocapronicacid.Calibration
curvesfor these acidswere then made byobtaining the peak-height ratios
ofthe acids:that ofisocapronicacid.Recoveryvaluesbetween 95and 100%
werefound fortheindividualVFAandtheinternal standard.TotalVFAwas
represented as the sum of all six acids. Lactic acid concentrations were
determined enzymatically according toAnonymous (1986).
Concentrations of glucose and arabinose in digesta and urine were
determined as silyl derivatives of monosaccharides by gas-liquid
chromatography (Sweeley et al. 1963).Aknown amount ofwet digesta (1
g) or urine (1ml) was diluted with distilled water (1:10 v/v). The diluted
sample was then deproteinized with potassium ferrocyanate and zinc-
acetateanddesaltedbypassingthroughamixture(1:1w/w)ofanion(Biorad
AG3 x 4 ) and cation (BioradAG50Wx4))exchanger.After centrifugation,
200 ul ofthe supernatant fraction was freeze-dried. To the freeze-dried
sample phenylglucopyranoside (0.4 mg in a 1 ml pyridine solution) was
added as an internal standard. The sample was then derivatized by the
126 Chapter 6
addition of0.6mlhexamethyldisilazane and 0.3ml trimethyl-chlorosilane.

Then the contents were mixed using a Vortex stirrer. After an incubation
period of 30 min at room temperature, the reagents were removed by
evaporationwithN 2 at40°.Theresiduewasthenredissolvedin0.5ml ethyl
acetate. From this sample, 2ulwas analysed using a Hewlett Packard HP
5890, equipped with a flame ionization detector and a Hewlett Packard
3396A integrator. The carbohydrate derivatives were separated with a
chrompack capillary WCOTfused silica columncoated with CP sil5CBof
50mlength.H 2 wasusedascarriergas.Theoventemperature washeld for
3minat 190°,thenraisedattherateof57mintoafinaltemperature of265°,
whichwasheldfor 5min.Thetemperature oftheinjector and detector was
240 and 300°, respectively.
The results of both experiments were analysed by means of analysis of

variance as a randomized blockdesign (Cochran &Cox, 1957). Treatments
were confounded by time and age,but it was assumed that differences are
due to the test sugar or increase in dietary sugar. This will further be
elucidated in the discussion.
In ExptAthe treatment factors were type and dietary level ofsugar.All
four phases ofExptAwere included in the statistical analysis, despite the
differences in the length of the adaptation period between the first three
phases and the fourth phase. This can bejustified because the differences
in results obtained onthe Glue 100diets inthe first and fourth phase were
not statistically significant. Moreover, the ratio between wet ileal digesta
outputandintakeofDMwerealmostsimilarinbothD-glucosephases.This
suggests that an adaptation period of 1 d was sufficient to stabilize the
conditions in the gastrointestinal tract when pigs were changed from the
Arab to the Glue diet.
In Expt B the treatment factors were type ofsugar and the dietary level
ofsugar. The between animal error term was used for testing the effect of
typeofsugarandthewithinanimalerrortermfortestingtheeffect ofsugar
levelsaswellasthetypeofsugarxlevelinteraction.Thesum ofsquares for
levelswaspartitionedintoasetoforthogonallinearandquadraticpolynomial
regression components. All statements of significance are based on a
probability ofP <0.05.
RESULTS
Thepigswerehealthyandconsumedtheirdailyfeedallowancecompletely
for allexperimental treatments.
ExptA
Intake of DM and water, output of digesta, and DM content of digesta

measuredincannulatedpigsonD-glucoseorD-arabinosedietsaregivenin
Table4.Sincetheoutputofdigestawasmeasuredover12h/d,intakeofDM
and water is also presented over a 12 h period. There were significant
differences in DMintake among the treatments. These differences were
causedbythe feeding system applied, sincethis system wascoupled with
liveweightofthepigs.Waterintakeofpigsduringphase 1(Glue100diet)
was on average 2.16 times their daily DM intake. This ratio increased
significantlyby40and78%whenpigswerefedontheArab50(phase2)and
TABLE4. Expt A. Intake (g/12h) of drymatter (DM)and water,

output (g/12h) of wet digesta,and DMcontent (g/kg)of digesta,
measured in cannulated pigs fed on D-glucose (Glue) and
L-arabinose (Arab) diets.
(Meanvaluesoffour pigsper treatment)
Phase 1 2 3 4
Sugar Glue Arab Arab Glue SED*
Dietary level (g/kg) 100 50 100 100 (df9)
Intake ofDM (A) 370a 432 b 496c 528d 5.4

a b c
Intake ofwater (B) 800 1300 1900 1250" 82.9
Ratio B :A 2.16a 3.03b 3.85c 2.37" 0.19
Output ofwet ileal 517 a 874b 1491° 753 ab 142.9
digesta
DM content ileal digesta 120a 94 b 72c 117a 6.1
* Standard error ofdifference betweenmeans.

a,b,cWithinarow,meanvalues withnocommonsuperscript letters were
128 Chapter 6
TABLE 5. EzptA. Apparent ileal digestibility coefficients of dry

matter (DM), organic matter (OM),gross energy (GE), nitrogen
(N), D-glucose and L-arabinose, measured in cannulated pigs
fed on D-glucose (Glue) and L-arabinose (Arab) diets).
Phase 1 2 3 4
Sugar Glue Arab Arab Glue SED* df
Dietary level
(g/kg) 100 50 100 100
DM 83.3 a 81.8 a 79.3 b 83.4a 1.10 9

OM 84.5 a 82.9ab 81.0b 84.9 a 0.93 9
GE 84.9" 83.6ab 81.9" 85.l a 0.94 9
N 88.5 a 88.0a 88.0a 88.6 a 1.51 9
D-glucose 99.4a - - 99.3 a 1.55 3
a
L-arabinose - 70.2 66.8 a - 2.36 3
* Standard error ofdifference between means.

a,b
Within a row,mean values with no common superscript letters were
significantly different (P < 0.05).
Arab 100 (phase 3) diets, respectively. When pigs were changed from the
Arab 100 to the Glue 100 diet (phase 4),the ratio between water and DM
intake decreased significantly to2.37,a value which was almost similar to
that obtained inphase 1.Output ofwetdigesta wasincreased significantly
when pigs were changed successively to the Arab 50 and Arab 100 diet.
However, when pigs were changed from theArab 100to the Glue 100 diet,
ileal output ofdigesta was decreased significantly. The increase in digesta
output in pigs fed on theArab diets was associated with a decrease in DM
content ofthe digesta. However, similar towater intake and ileal output of
digesta,thiswasmorepronounced ontheArab 100dietthan ontheArab50
diet.
Apparent ileal digestibility values for DM, OM, GE, N, D-glucose and L-
arabinose are shown in Table 5.In pigs fed on the Glue 100 diets (phase 1
and 4),similar digestibility coefficients for DM, OM, GE, N and D-glucose
were observed. However, in pigs fed on the Arab diets, lower digestibility
coefficients for DM,OMand GEwere observed;the values ontheArab 100
dietbeing significantly different from those ofpigsfed onthe Glue diets.
There was a tendency for digestibility ofN tobelessin pigsfed ontheArab
diets than in pigsfed onthe Gluediets. Apparent ilealdigestibilityof

D-glucosewasfound tobecloseto 100%.However,ilealdigestibility ofL-
arabinosewasonlyapproximately70%.DigestibilityofL-arabinosewasnot
affected significantly bythe doselevelofL-arabinose.
Data for ilealflowofVFAandlacticacidare giveninTable6.Inpigsfed
ontheArabdiets,theilealflowofVFAwashigher than inpigsfed onthe
Gluediets,beingsignificantinpigsontheArab100diet.Theincreaseinileal
flowofVFAinpigsfedontheArabdietswasmainlycausedbyan increase
inaceticacid.Ilealflowoflacticacidfollowedthesamepatternofresponse
asforVFAwhenpigswerefedtheArabdiets.However,the differences in
ilealflowoflacticacid amongthe treatments werenot significant.
TABLE6. ExptA. Ilealflow ofvolatilefatty acids (VFA)and lactic

acid (mg/12h), measured in cannulated pigs fed on D-glucose
(Glue)and L-arabinose (Arab) diets.
Phase 1 2 3 4
Sugar Glue Arab Arab Glue SED*
Dietary level (g/kg) 100 50 100 100 (df9)
Total VFA 985 a 1233ab 1567b 987 a 166.2

Individual VFA
Acetic acid 713 a 957 a 1297b 823 a 139.6
Propionic acid 108a 44 a 79a 65 s 30.8
Butyric acid 50a 35 a 28 a 34a 12.9
Isobutyric acid 106a 188b 141 ab 41 c 27.8
Valeric acid 8a 9a 22b 24 b 5.0
Isovaleric acid + + + +
(L-) Lactic acid 511 a 632 a 863 a 548 a 176.9
* Standard error ofdifference betweenmeans.

+
Belowthe detection levelof1 mg/100gwet digesta.
a,b
Within a row,meanvalues withnocommonsuperscript letters were
130 Chapter 6
ExptB
The mean values for DM and water intake, output of urine and fresh
faeces, and DM content of faeces in pigs fed on D-glucose (Glue) or L-
arabinose(Arab)dietsaregiveninTable7.Thereweresignificant differences
inDMintake amongthetreatment groups.However,as already stated in
ExptA,thesedifferences werecausedbythefeedingsystemapplied.Water
intakein%ofDMintakeofpigsfedontheGluedietswasnot significantly
affected bythe doselevelofthis sugar.Whenpigswerefed theArabdiets,
waterintakein%ofDMintakeincreasedlinearly(P<0.01)asthelevelof
thissugarwasincreased.Outputofurineandfresh faecesonbothtypesof
sugardietfollowedthesamepatternofresponseasforDMandwaterintake.
On a composite basis the differences in output ofurine and fresh faeces
betweenpigsfedontheGluedietsandthosefedontheArabdietswerenot
significant (P>0.05).DMcontentoffaeceswasnotsignificantly affected by
thetypeand doselevelofthe sugar.
Apparentfaecaldigestibilitycoefficients forDM,OM,GEandNaregiven
inTable8.SimilarresultsforapparentfaecaldigestibilityofDM,OMand
GE were achieved in pigs fed on the Glue and Arab diets. However, the
averagedigestibilityofNwassignificantlylowerinpigsfedontheArabdiets
than thosefed onthe Gluediets.
Resultsforurinaryexcretionofarabinose,energyandN,andretentionof
N are given in Table 9.Arabinose was partly excreted via the urine. The
extentoftheurinaryexcretionofthissugar,as%ofintake,wassignificantly
doserelated (P<0.01).As a result ofthe arabinose losses into the urine,
urinary excretion of energy also increased in pigs fed on the Arab diets.
Urinary excretion of N and retention of N are both affected by age
(McConnelletal.1972;Carretal.1977).Sincetheexperimentaldietswere
fedinsequence,theincreaseinurinary excretionofNand thedecreasein
Nretentionasthedietarysugarlevelswereincreasedisratherduetoanage
effect than to the dose level (Carr et al. 1977). Retention of N, being
approximately 55%ofNintake,washighinthepresent trial as compared
topracticalvalues.ThishighNretentionvaluewillbetheresult ofbotha
wellbalancedhighlydigestibleproteinbasaldietandtherelativelowdaily
feedinglevelapplied(AgriculturalResearchCouncil,1981).Whenpigswere
fedontheArabdiets,less(P<0.05)Nwasretained thenwhenfeeding the
Gluediets.ThisisduetobothalowerNdigestibility andahigher amount
ofNexcreted intheurine inpigsfed ontheArabdiets.
TABLE7. ExptB. Intake (g/24h) of drymatter (DM)and water,

output (g/24h) of faeces andurine, and DMcontent (g/kg) of
faeces,measuredinnon-cannulatedpigsfedonD-glucose(Glue)
andL-arabinose (Arab) diets.
(Meanvaluesoffivepigsper treatment)
Sugar Sugar Intake Ratio Output DM

level DM(A)water(B)B:A urine faeces content
(g/kg) faeces
Glue 25 670 1470 2.19 730 162 426

50 797 1650 2.07 790 186 442
75 917 2040 2.22 1050 215 443
100 1081 2490 2.30 1360 236 440
Arab 25 675 1420 2.10 690 182 393

50 802 1750 2.18 860 192 435
75 926 2230 2.41 1320 227 423
100 1085 2890 2.66 1940 247 417
SED(df32)* 8.8 139 0.16 139 19.9 15.6
SED(df24)** 24.4 176 0.21 167 24.2 17.7
Meanresults per sugar
Glue 866 1912 2.20 982 200 438

Arab 872 2072 2.34 1202 212 417
SED(df8) 23.2 128.4 0.16 115.5 17.1 11.4
* Standard error ofdifference between meanswithin each sugar.

**Standard error ofdifference between means ofboth sugars.
132 Chapter 6
TABLE8. ExptB. Apparent faecal digestibility coefficients of dry

matter(DM),organicmatter(OM),grossenergy(GE)andnitrogen
(N),measuredinnon-cannulatedpigsfedonD-glucose(Glue)and
L-arabinose (Arab) diets.
(Meanvaluesoffive pigsper treatment)
Sugar Sugar
level Digestibilities
(g/kg) DM OM GE N
Glue 25 89.8 91.0 90.9 93.8

50 89.7 91.0 91.2 94.3
75 89.6 91.1 91.1 94.2
100 90.5 91.7 91.5 94.4
Arab 25 89.5 90.4 90.2 93.0

50 89.6 90.9 90.9 93.9
75 89.6 90.9 90.9 93.2
100 90.4 91.3 91.0 93.6
SED(df32)* 0.85 0.61 0.55 0.34
SED(df24)** 1.00 0.99 0.86 0.39
Mean results per sugar
Glue 89.9 91.2 91.2 94.2

Arab ••• 89.8 90.9 90.8 93.4a
SED(df8) 0.68 0.84 0.72 0.26
* Standard error ofdifference between means within each sugar.

**Standard error ofdifference between means ofboth sugars.
a
Meanvalue wassignificantly different from the Gluediets (P<0.05).
TABLE9. ExptB. Urinary excretion (% ofintake) of glucose,

arabinose, energy andnitrogen (N),and retention of N(% of
intake),measuredinnon-cannulatedpigsfedonD-glucose(Glue)
and L-arabinose (Arab) diets.
(Meanvalues offive pigsper treatment)
Sugar Sugar Urinary excretion Retention

level Glucose Arabinose Energy N ofN
(g/kg)
Glue 25 + + 2.8 33.4 60.4

50 + + 2.8 35.7 58.6
75 + + 3.0 38.6 55.6
100 + + 3.2 41.2 53.2
Arab 25 + 10.9 3.3 34.3 58.7

50 + 12.2 3.7 36.8 57.1
75 + 13.7 4.7 41.8 51.4
100 + 14.7 5.2 43.5 50.2
SED(df32)* 1.29 0.20 2.19 2.15
SED(df24)** - 0.29 2.25 2.19
Mean results per sugar
Glue + + 3.0 37.4 57.0

Arab + 12.9 4.2 39.1 54.4a
SED (df 8) - 0.23 1.22 1.15
+ Smalltraces ofglucose (0.2-0.4g/L)and arabinose (0.01-0.2g/L)

werefound inthe urine ofthese experimental treatments.
* Standard error ofdifference between means within each sugar.
**Standard error ofdifference betweenmeans ofboth sugars.
a
Meanvaluewas significantly different from the Gluediets (P<0.05).
DISCUSSION
Indesigningastudyofthiskind,anumberoffactorshavetobetakeninto
account. These factors mainly relate to the experimental design and the
dietaryinclusionlevelsofthetestproduct.Latinsquaresareoften used as
anexperimentaldesigninbalancestudieswithanimals,inwhichthediets
134 Chapter 6
are fed in sequence,the diet sequence being different for each animal. The
advantages ofusingLatin squares arethat variation between animals and
those arisingfrom a common timetrend canbeequilibrated. However, this
is only true when there are no carry-over effects. The results ofa previous
tentative study have shown that carry-over effects ofarabinose cannot be
excludedcompletely.Moreover,theresults ofthat study showed that water
intake and ileal output ofwet digesta ofpigs fed on L-arabinose diets was
increased markedly, especially at dietary inclusion levels above 100 g/kg.
Therefore, in the present experiments the Latin square method was not
followed, whereas themaximum dietary inclusion levelofL-arabinose was
setat 100g/kg.Onedisadvantage offeedingexperimental dietsin sequence
isthat digestibilities may beaffected byan agextreatment interaction. In
the present study, however, no indications were found that our ileal and
faecal digestibility data had been affected by age.This statement for ileal
digestibilityisbasedonthealmostsimilarcoefficients onbothglucose diets
(Table 5).The results for faecal digestibilities (Table 8), showing that the
coefficients within the D-glucoseblockwere almost constant, also indicate
that these data were not confounded by age.
Previous studies have shownthat L-arabinose is not absorbed completely
from thesmallintestineinrats(Cori,1925)andchicks(Bogner, 1961;Wagh
and Waibel, 1967). Their observation is supported in the present study,
indicatinganincompletedigestionofthis sugar at theterminal ileum. This
incompletedigestionwasreflected intheapparent ilealdigestibility values
for DM,OMand GE,showinga decrease in digestibility when including L-
arabinose in the diets. No clear indications were observed that ileal
digestibility ofL-arabinose was doserelated.This finding doesnot support
that ofa previous study with roosters, showingthat ileal digestibility ofL-
arabinose decreased when the dietary level was increased (Schutte et al.
1991a). In this context it seems relevant to comment on the possible
influence of the alimentary tract bacteria on the ileal digestibility of L-
arabinose.Itmaybeassumed that thepresenceofunabsorbed arabinose in
the small intestine ofpoultry and pigswill lead to a microbial attack on
this sugar. In poultry this microbial attack may commence already in the
crop. Unfortunately, no literature data are available on a possible
fermentation ofarabinose in the crop and small intestine ofpoultry and in
thesmallintestine ofpigs.However,itishighlyprobablethatinthe present
studysomemicrobialdegradation ofarabinoseinthesmallintestine ofpigs
had taken place. This statement isbased on the observed increase in ileal
flowofVFAand lacticacidinpigsfedontheL-arabinosediets.This symptom
points to a more extensive microbial activity in the small intestine of pigs
when fed on diets containing L-arabinose. Quantitatively the influence of

the increased microbial activity on the ileal digestibility ofL-arabinose is
difficult to assess. In addition to arabinose, other readily fermentable
components in the diet may also be attacked by an increased intestinal
bacterial activity.However,thishypothesiscouldnotbebackedupfirmly by
the ileal digestibility values for N, since the differences in apparent ileal
digestibleNbetweentheD-glucoseandL-arabinose treatments were small
and not statistically significant.
Administration ofL-arabinose to pigs was associated with an increase in
ilealdigestaflow.Thisfinding isinagreement withtheresultsofa previous
studywithroosters (Schutteetal. 1991a).Thisincreaseinilealdigesta flow
can be explained by the presence of unabsorbed arabinose in the small
intestine which will lead to an inflow ofwater into the intestinal lumen in
order to keep osmolality constant (VanWeerden, 1959;Hof, 1980).
Similar results for apparent faecal digestibility ofDM, OM and GE were
achieved when pigs were fed on diets containing either D-glucose or L-
arabinose. These results indicate that the part ofL-arabinose not digested
in the small intestine was microbially degraded in the hind gut. This was
confirmed bytheabsenceofarabinoseinapooledsampleoffaecesofpigsfed
ontheArabdiets.Consideringthefaecal digestibilityvaluesofN,itislikely
that the presence ofarabinose in the hind gut ofpigs was coupled with an
increased microbial activity. If sufficient substrate is available, this will
result in an increased net microbial protein synthesis. Consequently N
output in the faeces willincrease, thus decreasingfaecal digestibility ofN.
Generally,anincreased Noutput inthefaeces duetoanincreased bacterial
fermentation inthehind gut ofpigsisaccompanied bya reduced urinary N
output, resultingin a nonorsometimes positive overalleffect onN balance
(Partridge et al. 1982; Dierick et al. 1983;Malmhof & Hakansson, 1984;
Morgan &Whittemore, 1988).Contrary tothese findings are the results of
the present study. In addition to a depressed faecal digestibility ofN, also
N losses in urine were slightly increased when pigs were fed on diets
containing L-arabinose.Asa result ofboth, less N was retained in pigs fed
on the L-arabinose diet. Similar results of a depressed N retention were
found in a previous study with the pentose sugar D-xylosewhen fed to pigs
at a dietary inclusion level of 100 g/kg (Schutte et al. 1991b). Why the
pentose sugars L-arabinose and D-xylose tend to decrease efficiency of the
utilization ofabsorbed N in pigs is unknown.
It is generally accepted that D-glucose can be utilized almost completely
inhumanandanimals(Demetrakopoulos, 1978),soonlynegligible amounts
of glucose will be found in the urine. The latter is in agreement with the
136 Chapter 6
finding inthe present study. Onlylittle information isavailable towhich

extent L-arabinose will be excreted in the urine ofmonogastric animals.
Arnal-PeyrotandAdrian (1974)reported anurinarylossof7.5%ingested
arabinosewhenratswerefedadietcontaining60gL-arabinose/kg.Similar
results were obtained in our previous study with roosters at a dietary
inclusion levelof25gL-arabinose/kg (Schutte et al. 1990).The resultsof
that study alsoshowedthat urinary excretion ofarabinosein%ofintake,
increasedlinearlyasthedietarylevelofthissugarincreased.Thesamewas
trueinthepresent study.However,thisdose-dependant urinary excretion
ofL-arabinose couldnotbederivedclearlyfrom thedifferences in urinary
excretion of energy between the D-glucose and L-arabinose treatments.
Calculations indicated that if the increase in urinary excretion ofenergy
overthe D-glucose treatments wereattributed toL-arabinose, this would
representabout23%oftheL-arabinoseintakeatalllevels.The differences
betweenthedeterminedvaluesforurinaryexcretionofarabinose(asamean
12.9%)and those calculated from the urinary excretion ofenergy, relate
mostprobablytothehigherlossesofotherenergybearingcomponentsinthe
urine ofpigsfed onthe L-arabinose diets.Thisissupported bythe higher
losses ofNintheurine inpigsfed onthe L-arabinose diets.
Takentogether,thetwostudies indicatethat thenutritional value ofL-
arabinoseinpigsislowerthanthatofD-glucose.Inaddition,administration
ofL-arabinosetopigsmayresultinadepressedNretention.Quantitatively,
the nutritional value ofL-arabinose forpigsisdifficult toassess from the
present studies.Thisisbecauseoftheunknownmetabolicpathwayofthis
pentosesugar.Fromthepresentstudiesithasonlybecomeclearthatabout
30%ofthe ingested L-arabinose entered the largeintestine whereit was
fermented, whileanother 13%wasexcretedintotheurine.Thepathwayof
theremaining 57%ofingestedL-arabinose isuncertain.Thispart, which
disappeared from the small intestine, may have been utilized as such or
fermented toVFA,orboth ofthese. Ofthese two mechanisms, a possible
microbial fermentation of L-arabinose in the small intestine was
demonstratedinourstudy.Knowledgeaboutapossiblemetabolicpathway
ofthis pentose sugar in the animal body is limited. Segal &Foley (1959)
reported that whengivenanintravenously infused doseofC14labeledL-
arabinosetoman, only0.8%couldberecovered in expired carbondioxide.
Iftheirdataaretransferabletopigs,thiswouldmeanthatL-arabinoseonly
aftermicrobialfermentation canbeusedasanenergysourceforpigs.Based
onthis assumption and taking urinary arabinose losses as 13%,it canbe
calculated that the net energy value ofL-arabinose is at least 40%lower
than that of D-glucose. In this calculation the efficiency ofutilization of
energyvia hindgutfermentation comparedtothatofpraecaecallydigested
glucosewassetat67%.(AgriculturalResearchCouncil,1981;Millleretal.
1989).Basedonthedifferences inenergylossesintheurinebetweentheD-
glucose and L-arabinose treatments, the calculated net energy value for
arabinoseisratheroptimistic.Ontheotherhand,dataofCloseetal.(1989)
suggestthattheefficiencyofutilizationofenergyfromhindgutfermentation
is higher than estimated by Agricultural Research Council (1981) and
Mulleret al.(1989).
ACKNOWLEDGEMENTS
Theauthorswouldliketothank P.VanLeeuwenfor surgical assistance,

and G.B.Derksen andJ. Wiebenga for statistical analysis ofthe data.
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141
GENERAL DISCUSSION
143
GENERAL DISCUSSION
The NSP fraction of feed ingredients of vegetable origin represents the
least digestible component of diets in pigs and poultry. Therefore most
efforts on the use ofenzymes in pig and poultry diets in order to improve
digestibility of feed ingredients have focussed on the NSP fraction. The
benefits of an enzymatic hydrolysis of NSP, however, are not determined
solely by an improvement in digestibility, but also by the potential of the
animal toutilize the products ofhydrolysis.In addition toD-glucose, other
sugars will also be released by complete hydrolysis of NSP, of which in
quantitative terms D-xyloseand L-arabinose are the most important ones.
The literature review (General introduction) indicates that knowledge is
incomplete on the nutritional value of these pentose sugars in pigs and
poultry.Theinvestigationsdescribedinthisthesisweremainlyfocussed on
the digestion of D-xylose and L-arabinose in the various parts of the
gastrointestinal tract, and on renal excretion. Besides, the effect of both
pentosesugarsonsomephysiologicalcharacteristicsinpoultryandpigswas
studied. The results of the investigations were compared with data from
literature. In the present discussion various aspects ofthe results will be
considered in relation to some other effects and properties ofthese sugars
and NSP.
ILEAL DIGESTIBILITY OF XYLOSEAND ARABINOSE
Without an accurate knowledge ofthe extent ofthe removal ofxylose and

arabinose from the precaecal tract, itishardly possible tofollow the fate of
both pentose sugars in the animal organism on a quantitative basis.
Hitherto, most of the reported experiments on intestinal absorption of
xylose and arabinose in monogastric animals have been performed on the
wholeintestinal tract ofrats (Cori, 1925;Miller &Lewis, 1932;)and chicks
(Bogner,1961;Wagh&Waibel,1967a).Theprincipleofthismethodis briefly
asfollows.Theexperimental animalsarefedapredetermined amountofthe
sugarbystomachorcroptube.After agiventime,theanimalsarekilled and
thequantityofsugarremainingintheintestineisdetermined.The difference
between the amount fed and that recovered from the intestinal tract is
considered as the amount ofsugar absorbed bythe animal.This method is
opentoanumberofobjections,chieflyrelatedtothelengthofthe absorption
period. The results reported byBogner (1961)and Wagh &Waibel (1967a)
arebased onan absorption period of30min. Consideringthe data ofWagh
& Waibel (1967a), it appears that not all the sugar administered had
disappearedfromthecropandgizzardduringthistimeperiod.Atadoserate
of15um per gbodyweight, 16.7%ofthe ingested glucosewas still present
in the crop and gizzard. For xylose and arabinose these percentages
amounted to 20.3and 24.5 %,respectively. Miller &Lewis (1932) reported
144 General discussion
that in rats absorption coefficients of xylose tended to increase as the

absorption period was prolonged. Mean values of 29, 39 and 46 mg of
absorbed xylose per h per 100 g of body weight were obtained during
absorption periods of 1, 2and 3h, respectively.
Inourstudiestheilealfistula techniquewasused tomeasure absorption
of the sugars. This would permit of basing the results on a much longer
absorption period, and to study possible side effects from the feeding ofD-
xylose and L-arabinose to pigs and poultry in more detail. As a longer
absorption period couldlead toan increased attack by intestinal microbes,
this type of experiments does not distinguish between sugar removal by
absorption and by fermentation. Both probably occur in all sections of the
precaecal tract, but to greatly varying extents. The ratio between removal
byabsorption andfermentation dependsonthevelocityofabsorption ofthe
sugars and on the precaecal activity of microorganisms. From literature
data it iswellknown that glucose is absorbed at a higher rate than xylose,
the latter being superior to arabinose (General introduction). Thus, when
precaecal fermentation ofthese sugars occurs,the sequence is likely to be
of arabinose >xylose > glucose. This sequence ofmicrobial fermentation,
however, could notbeconfirmed inthepresent studies with pigs (Chapters
4and 6).Atequaldietarydoselevelsof100g/kg,ilealflowof VFAincreased
more in pigs fed on the xylose diet than in those fed on the arabinose diet.
Whenilealflowof VFAfortheglucosetreatment was setat 100%,the value
for the xylose treatment was 186%and for the arabinose treatment 159%.
These results suggest that the precaecal microbes have a preference for
xylose over arabinose. While it is questionable whether both trials are
comparable,inthis case it canbejustified sincethe same type ofbasal diet
was used inboth trials and theilealflow of VFAonthe glucose treatments
were similar. Some evidence of a discrimination between sugars by the
bacteriainthealimentarytractisprovidedbyLuckey(1987).Thisinvestigator
reviewed the capability of the human alimentary tract bacteria to utilize
simple sugars. His study indicated that there are considerable differences
betweenbacterialspeciesintheircapacitytoutilizevarioussugars.Glucose
appeared to be utilized by all of the identified intestinal bacteria. In
contrast, xylose as well as arabinose were not used by Lactobacillus
acidophilus, and arabinose also not by Bacteroides fragilis. Thus, it
would seem that the extent ofmicrobial degradation ofsugars depends not
onlyonthedensity,but alsoonthecompositionoftheintestinalbacteria. In
this context it seems relevant to consider also the differences in ileal
digestibilityofthesugarsbetweenpigsandadultroosters(Chapters2,4and
6).In both animal species,D-xylosewas digested almost completely at the
terminal ileum. Considering the ileal digestibility data of L-arabinose, it
appears that in pigs fed this pentose sugar at 50 and 100g/kgdiet, 30 and
33%,respectively,oftheingested arabinose appeared intheilealchyme. In
adult roosters, however, at these dietary inclusions, only 5 and 25 %,
respectively, ofthe ingested arabinose was recovered in the ileal chyme.

Theseresultssuggestthatprecaecalmicrobesinroostersareabletoutilize
moreL-arabinose than thoseinpigs.However, care should betaken with
this conclusion.This conclusion has toberestricted sincefermentation of
arabinoseinfowlsmayalreadycommenceinthecrop.Whiletheroleofthe
cropasastorageorganinthebirdiswellestablished,itsdigestive function
is not well understood. Bolton (1965) demonstrated that as a sugar
disappearedfromthecrop,theamountoflacticacid,aceticacidandethanol
inthecropincreased.Bayeretal.(1978)fedbroilerchicksonhighandlow
fibre diets and noted that lactic and acetic acid were the predominant
organicacids in the crop.Medl&Scharrel (1978)demonstrated an active
trans-epithelial movementofsodiumacrossthechicken'scrop,suggesting
that nutrientscanbeabsorbedfrom thecrop.Thishypothesisissupported
by data of Soedarmo et al. (1961), who showed that about 7% of an
administered doseofC14-labeledglucoseintotheligatedcropdisappeared
from this organ.Theynoted alsothat whengivenC14labeledxylose,this
sugar did not leave the ligated crop in appreciable amounts. Thus, both
absorptionofsugarsandmicrobialfermentation mayoccurinthechicken's
crop,buttoarelativelysmallextentonly.Onthebasisofcertainassumptions
itispossibletoestimatefromthepresentstudiestheamountsofxyloseand
arabinose which have been removed from the precaecal tract by either
absorption or fermentation. This will be discussed further in the next
section.
FATE OF THE PRECAECALLY DIGESTED XYLOSE AND

ARABINOSE
(1)D-xylose
Thexyloseabsorptiontestiswidelyusedinthediagnosisofmalabsorption.
The test isbased onthe assumption that xyloseisnotmetabolized toany
significant extent in the body.Thus,this would mean that the amountof
xyloseexcretedintheurineafter oraladministrationreflectstheamountof
xyloseabsorbedinthegastro-intestinaltract.Nodirectevidenceisavailable
tostate whether conversiontoglucoseisafate ofD-xylosein monogastric
animals (Segal & Foley, 1959). There is conclusive evidence, however,
indicatingthatsomeoftheD-xyloseabsorbedismetabolizedintheorganism.
After intravenousadministration ofD-1-C14xylosetoman,about 14to16
%couldberecovered as carbon dioxide in expired air (Wyngaarden et al.,
1957;Segal&Foley,1959).SimilarresultswerereportedbyWeser&Laster
(1968)whofoundthatabout11%ofanintraperitonealinjectionofD-xylose
-1-C14toguineapigswasrecoveredascarbondioxideinexpiredair.Amuch
lower value was reported in chicks by Wagh & Waibel (1967b). These
investigatorsfound thatonly4%ofasubcutaneousdoseof C14-labeledD-
xylose was recovered as carbon dioxide in expired air. In addition to

catabolism of D-xylose to carbon dioxide, a proportion of both orally and
intravenously administered D-xylosehas been shown tobe converted into
a four-carbon polyol (D-threitol) in the human organism (Pitkanen &
Svinhufvud, 1965).Their data suggest further that D-threitol is excreted
into the urine with no or very little tubular reabsorption. According to
experimental data of Weser &Laster (1968), D-xylose catabolism occurs
mainly in the kidney and liver.The roleofthe liver in the conversion ofD-
xylosetoD-threitol hasbeenstudiedmoreextensivelybyPitkanen (1977).
Urinary excretion of D-xylose and D-threitol was studied after oral
administration of25gxylosetoman withportal livercirrhosis,with active
fattyliverdiseaseandwithnoliverdisease,respectively.Thisstudy pointed
out that 15%ofthe doseofD-xylose excreted in the urine was recovered as
D-threitolinmanwithoutliverdiseaseorwithfattyliverdisease. D-threitol
excretion was decreased to8%incirrhotic patients. Based onthese results
the author concluded that a substantial proportion ofthe conversion ofD-
xyloseto D-threitol conversion occursin the liver. In order to test whether
conversion ofD-xylosetoD-threitol isofsignificant importance inpigs and
poultry, additional analyses were carried out in the urine ofadult roosters
and pigs involved inthe studies reported in Chapters 2and 5.In the urine
ofboth pigs and adult roosters fed on D-glucosediets, D-threitol could not
bedetected. Considerable amounts ofD-threitol were found in the urine of
pigs and adult cocksfed on diets containing D-xylose (Table 1).The extent
ofthe urinary excretion ofD-threitol agreeswellwith the data of Pitkanen
(1977).The presence ofD-threitol in the urine may alsopartly explain the
discrepancy between the renal excretion ofD-xylose and the energy losses
into the urine observed in our studies.
TABLE 1.Urinary excretionofD-xylose andD-threitol (%of intake)

in pigs and adult roosters fed on diets containing 100g D-xylose/
k g (mean values ± SD of 4 pigs and 3 adult roosters )
Animal Urinary excretion of: D-threitol/

species D-xylose D-threitol D-xylose ratio *
Pigs 37.2 ±7.6 7.9+1.4 0.17

Adult roosters 20.2 ± 2.0 4.6 ±0.5 0.19
*Not corrected for the differences in molecular weight.
The metabolic path for the conversion ofD-xylose toD-threitol isnot fully
known. There is evidence that D-xylose metabolism isinitiated by glucose
dehydrogenase or by a more specific pentose dehydrogenase ( Weser &
Laster, 1968).AsproposedbyPitkanen(1977),further stepsinthe conversion
mayoccurthroughtheglucuronatecycleenzymesforwhichD-xylonateand
D-erythrulose are intermediates. A similar direct oxidation pathway of
galactose through galactose dehydrogenase and beta-L-hydroxy-acid
dehydrogenasehasalsobeendemonstrated inman(Cuatrecasas & Segal,
1966;Segal &Cuatrecasas, 1968).
Inourstudiesitwasfoundthatinbothadultroostersandpigstheurinary
excretion ofD-xylosein %ofintake was significantly increased when the
dietarylevelofthis sugarwasincreased from 25gto 100g/kg(Chapters2
and 5).Ontheassumption that atthesedoselevels 10%oftheingestedD-
xyloseiscatabolized tocarbondioxide,andthat D-threitolrepresents15%
oftherenalD-xyloseexcretion,theproportionofD-xylosefermented prior
to the caecum can be calculated. The results of these calculations are
summarized inTable2.In consideringthe data presented inthis table, it
should be realized that the calculations relate to a certain physiological
status of the animals. Both renal excretion and precaecal microbial
degradationofD-xylosemaybeaffectedbyage(Hindmarsh,1976).Thedata
(Table2)arebasedonpigswithaliveweightof20to80kg.Withinthisperiod
renalexcretionofxylosewasnotaffected toanyextentbyage(Chapter5).
Renalexcretionofxyloseinpigsaged4weekswasfoundtobesimilartothat
inolderpigs(Schutteetal.,unpublishedresults).Theresultsofthat study
alsoshowedthatinyoungpigsD-xylosewasnotdigestedcompletelyat the
terminalileum.Whenfedthesepigsonadietcontaining100gD-xylose/kg,
approximately 10%oftheingestedxylosewasrecoveredintheilealchyme.
TABLE2.EstimatedproportionofD-xylosemicrobially fermented
in theprecaecal tract ofpigs and fowls.
Animal Dietary Fate ofprecaecally digested D-xylose(%ofintake)
species inclusion Excreted in urine Catabolized Microbially
(g/kg) xylose* D-threitol toC02 degraded
Pigs 25 20 4 10 66
50 32 6 10 52
75 39 7 10 44
100 43 8 10 39
Fowls 25 7 1 10 82
50 13 2 10 75
75 17 3 10 70
100 20 4 10 66
*Datafrom thestudiespresentedinChapter 5(pigs)andChapter2(adult

roosters).
As stated before, in older pigs this recovery value was nearly zero.Most
probably these differences inileal apparent digestibility coefficients of D-
xylosebetweenyoungandolderpigsareduetolessmicrobialactivityinthe
precaecal tract oftheyoungpigs.
It is highly likely that there are differences in precaecal microbial
degradation ofD-xylosebetween chicksand adult roosters.The observed
slight increase inceacalweight ofchicksfed ondiets containingD-xylose
mayprovidesomeevidenceforthishypothesis(Chapters 1and3).Thefew
data available on the precaecal fermentation ofD-xylosein monogastric
animalsareconflicting.Usingtheevertedsactechnique,Heneghan(1963)
investigated the absorption of D-xylose from the mid small intestine in
conventionalandgermfree rats andmice.Intheabsenceofmicrobialflora
therewasatwofoldincreaseintheabsorptionofD-xylose.Thisobservation
wasbasedonanabsorptionperiodofonehour,atimewhenboth germfree
andcontrolanimalshadabsorbed lessthanhalfoftheadministered dose.
Tennant et al. (1970) analysed the intestinal tract of germfree and
conventionalrats6hoursafteradministrationofD-xylose.Attheendofthe
testperiod, 11.3% oftheD-xyloseremainedinthestomachofthe germfree
rats compared to 3.8% in conventional rats. There was no difference
betweengermfreeandconventionalgroupsintheamountofD-xylosewhich
remained in the small intestine. Further studies are in progress at our
Institute with germfree pigs and chicksin order to clarify the role ofthe
intestinal microbial flora onxylose digestion.
(2)L-arabinose
The literature on the metabolic pathway of L-arabinose is very scarce.

Data ofSegal &Foley (1959) suggest that metabolism of L-arabinose to
carbondioxideisofnosignificance. Whengiven anintravenously infused
dose ofC14-L-arabinose to man, only 0.8%could be recovered as carbon
dioxide in expired air.A slightly higher value was reported in chicksby
Wagh&Waibel(1967b).Theseinvestigatorsobservedthatofasubcutaneous
doseofC14labeledL-arabinose,4.6% couldberecoveredintheexpiredair
ascarbondioxide.Inspectionofourchromatographybandsforurineofpigs
fed on L-arabinose diets, showed that there was an unidentified peak,
representing about 50% ofthearabinose peak.Theseresults suggest that
L-arabinose isexcretedintheurinepartlyinaform otherthan arabinose.
Thisissupportedbythehigherlossesofurinaryenergythanexpectedfrom
the renal excretion of arabinose (Chapter 6). Few literature data are
availableonthecatabolisme ofL-arabinose toproductsother than carbon
dioxide.The existence ofan enzyme converting L-arabinose toL-arabitol
hasbeensuggested sincethissubstancehasbeenisolatedfrom pentosuric
urineofman (Tbuster& Harewell, 1958).Inaddition somearabinosemay

beoxidizedtoL-arabonicacidaswasdemonstrated inbacteria (Weimberg
&Doudoroff, 1955).Additionalanalysesperformed intheurineofpigsfed
ondietscontainingL-arabinose(Chapter6)revealedthepresenceofbothL-
arabitol and L-arabonic acid. Further studies are in progress to quantify
bothmetabolites intheurine ofpigsandfowls fed on L-arabinose diets.
On the assumption that 4%ofthe ingested arabinose is catabolized to
carbondioxideandthatarabinosemetabolitesrepresent50%oftherenalL-
arabinoseexcretion,microbialdegradationofL-arabinoseinthe precaecal
tractofpigsandadultroosterswascalculated(Table3).Inthis calculation
the incomplete ileal digestion ofarabinose was taken into account. Since
ilealdigestionofarabinoseinpigswasonlymeasuredatdietaryinclusions
of50and 100g/kg,calculationscouldnotbemadefordoselevelsof25and
75g/kg.ItshouldbenotedthatthedatainTable3havetobeconsideredwith
some reservation, as they relate to a certain physiological status of the
animals;inpigstotheliveweightperiodof25to55kgandinfowltoadult
roosters.Noexperimentaldataareavailabletoindicatewhetherbothrenal
excretionandmicrobialdegradationofthispentosesugarareaffectedbyage
oftheanimal.Theresultsofatrialontheeffect ofthefrequency offeeding
diets containing D-xyloseonthe renal excretion ofthis pentose sugar are
discussed in Chapter 5.The attack ofprecaecal microbes onboth pentose
sugarsmaybeexpected tobegreaterinanimals continuouslyfed ondiets
containingthesepentosesugars.Consequently,renalexcretionofD-xylose
andarabinosewoulddecrease.Thishypothesiswasnotbesupportedinthe
trialwithD-xylosebecauserenalexcretionofxylosewassimilarinanimals
fed D-xylose diets twice or four times a day.Asimilar study showed that
renal excretion of L-arabinose was also not affected distinctly when
arabinosedietswerefedtwiceorfourtimesaday(Schutteetal.,unpublished
results).Studiesarenowbeingundertaken withgermfree pigsand chicks
toinvestigatetheroleoftheintestinalmicrobialflorainarabinosedigestion.
TABLE 3. Estimated proportion of L-arabinose microbially

fermented in the precaecal tract of pigs and fowls.
Animal Dietary Ileal Fate ofprecaecally digested L-arabinose (%ofintake)

species inclusion digested Excreted inurine Catabolized Microbially
(g/kg) (% )* arabin.* metabolites toC02 degraded**
Hgs 25 ND 11 5 4
50 70 12 6 4 48
75 ND 14 7 4
100 67 15 7 4 41
Fowls 25 95 9 4 4 78
50 94 11 5 4 74
75 80 14 7 4 55
100 75 17 8 4 46
* Data from studies presented in Chapter 6 (pigs) and Chapter 2 (adult

roosters).
**Preceacally fermented. The amount ofL-arabinose not digested at the
terminal ileum can be considered to be fermented in the hind gut.
ENERGETIC VALUE OF D-XYLOSEAND L-ARABINOSE
From stoichiometry, itcanbecalculated that the content ofgrossenergy is

2.82MJ/moleforD-glucoseand 2.34MJ/moleforD-xyloseand L-arabinose.
Thismeansthatallthreesugarshaveasimilargrossenergycontentof15.65
M J / k g substance.Severalsystemshavebeendevisedtoexpressthe energy
value offeed ingredients or diets in terms ofthe energy which is useful to
theanimal.Inthisrespectthetermmetabolizableenergy(ME)iscommonly
used, especially in poultry. In this system the losses ofenergy in faeces and
urine are taken into account.The MEvalue ofD-glucose canbe considered
tobesimilar tothe GE content, because this sugar isabsorbed and utilized
almost completely in monogastric animals. (Demetrakopoulos & Amos,
1978). The results of our studies have shown that the ME value of both
pentose sugars was not only lower than that of D-glucose but also dose
dependent. The latter statement was based on the increase in urinary
excretion ofbothpentose sugarswhen the dietaryinclusion ofthese sugars
was increased. Inpoultry,this dose dependent ME value was confirmed by
direct measurements of the ME value of both pentose sugars at dietary
levelsof50and 100g/kg(Chapter 1).Basedonthesemeasurements,theME
value ofD-xylosefor poultrywas estimated tobe 11.1MJ/kgat 50g/kg diet
and 8.5 MJ /kg at 100 g/kg diet. Corresponding values for L-arabinose at
thesedietarylevelswere9.6and5.7MJ/kg.Theseestimated MEvalues for

both pentose sugars have tobeconsidered with some caution because they
concern a far extrapolation to 100%.Anincrease in the dietary level of the
pentose sugarsmayalsohaveanindirectnegative effect ontheMEofother
dietary energybearingcomponents. Inpigs,the directmeasurement ofthe
ME value of both pentose sugars in our studies were made for a dietary
inclusionof100g/kgonly.Basedonthisdoselevel,theMEvalueofD-xylose
andL-arabinoseforpigswasestimatedtobe7.8and8.1MJ/kg,respectively,
(Chapter 5;Schutteetal.,unpublished results).However,asalready stated
for poultry, alsothese ME values have tobe considered with some caution.
The results referred to above indicate that it is not possible to give a fixed
ME value for D-xylose or L-arabinose because these values are dose
dependent.Apartfrom this,theuseoftheMEvalueinpredictingthe energy
value ofboth pentose sugars is questionable. From both pathways of the
pentose sugars after ingestion,metabolism per seand fermentation, losses
in energy can arise.Asa result from these lossesin energy,the ME content
ofthese sugars overestimates the energy that is actually available to the
animals as compared to D-glucose. It is well recognized that the ME ofD-
glucose is utilized with a great efficiency in terms ofproducing adenosine
triphosphate (ATP).According toVan Es (1974),the energetic efficiency of
utilization ofthe ME ofD-glucose is approximately 80%,resulting in a net
energy (NE) value ofapproximately 12.5 MJ/kg. However, the ME of both
pentose sugarsismainlyintheform ofenergyfrom afermentation process,
which process is coupled with considerable losses of energy. These losses
arisefrom theheat offermentation, the production ofmethane and the less
efficient utilizationoftheabsorbedendproducts(organicacids)as compared
to D-glucose. The overall utilization of fermented ME is still under
discussion.The present estimations about the utilization offermented ME
relativetothatoftheMEofD-glucosevarybetween50and 70%(Agricultural
Research Council, 1981;Just etal., 1983;VanEs, 1987;Mulleret al.,1989).
No data are available to which extent the part of the absorbed pentose
sugars not excreted into the urine, can be used by the animal. This
uncertainty together with the discrepancies on the efficiency of utilization
offermented ME,means that the NE valueofbothpentose sugars can only
be estimated roughly. On the assumption that metabolism ofboth pentose
sugars is analogous to that proposed in the previous section (Tables 2 and
3),thefollowingequationwasusedtoestimatetheNEvalueofboth pentose
sugars.
NE (in MJ/kg) =Ax 0.80 + 0.80 (B x 0.60 )
Where for the particular pentose sugar concerned:

A =the amount (in MJ/kg) ofthe ME which was calculated to arise from
a metabolism ofboth pentose sugars per se .The energetic efficiency
ofutilization ofthis fraction was assumed to be equal to that of
absorbed D-glucose (=80%)
B=the amount (inMJ/kg)oftheMEwhichwascalculated toarisefrom the

fermentation process. The relative utilization ofthis fraction as com-
pared to D-glucose was assumed to be 60%(mean value of literature
data)
According to this method the NE value of D-xylose and L-arabinose at a

dietary inclusion of 100 g/kg for pigs would be 4.2 and 4.0 MJ/kg.
Corresponding values for poultry at this dose level are 4.4 and 2.8 MJ/kg,
respectively. Notwithstanding the uncertainties involved in these
estimations,there isnodoubt that the energyvalue ofboth pentose sugars
isconsiderably lowerthan that ofD-glucose inpigs and poultry.An overall
estimate ofthe energy value ofboth pentose sugars of25to 35%ofthat of
D-glucose is probably not far from the real value. The study with chicks
described in Chapter 3, in which carcass energy was measured, suggests
indeed that the energy ofD-xylose and L-arabinose can onlybe utilized to
a limited extent by the animal.
ASSOCIATED EFFECTS OF D-XYLOSEAND L-ARABINOSE
Like galactose, D-xylose can potentially cause cataracts on the eyes

(Kinoshita, 1974).Noreliable data are available astowhether L-arabinose
alsoinducescataract formation. Thecataractogenicpotential ofD-xylosein
rats was first reported by Darby and Day (1939, 1940). Indeed, xylose
cataracts have occurred only in young Wistar albino rats when fed on a
regular diet enriched with 350 g or more D-xylose/kg. The cataracts
appeared onthe4thand5thdietarydayandprogresseduptothe 8thor9th
day(VanHeynigen, 1959).Despite the discontinuation ofD-xylose feeding,
the lenses became completely transparent and opacities did not reoccur
(VanHeynigen, 1967).Olderratsdidnotdevelopcataractswhenfedondiets
containing 250 to 500 g D-xylose/kg (Lerman & Heggeness, 1961; Van
Heynigen, 1969).Similarly,cataractswerenotobservedinadult ratsfedon
diets containing 50to 150gD-xylose/kg for a period ofa year (Loos, 1954).
There is only one report (Wise et al., 1954) available on the occurrence of
cataracts in pigs. These authors produced cataracts in young piglets by
feeding D-xylose at a level of 560 g/kg diet, but not when fed on a diet
containing approximately 200 g of this sugar/kg. Additionally, other

detrimentaleffects(anorexia,vomitingandseverediarrhoea)werereported
in young piglets bythese investigators when fed these high dietary dose
levelsofD-xylose.Theseeffectswerenotobservedinourexperimentswhen
pigswerefed ondietscontainingat amaximumthelowestdietarylevelof
D-xylosefed byWiseet al. (1954).The sameholdstrue for L-arabinose at
dietarylevelsupto100g/kg,beingthemaximumdietaryinclusioninvolved
inourexperiments.Noinformationisavailableaboutdetrimentaleffectsof
this pentose sugar at higher doselevels.
Consideringthedata of Wagh&Waibel(1967b),othersideeffects from
the feeding ofD-xyloseorL-arabinose alsomainlymayoccurwhenfed at
relatively high dose levels. These authors reported that in chicks,blood
hematocrit,cholesterol,serineandprolineinplasmaincreased significantly
when feeding these pentose sugars at dietary levels of200and 400g/kg.
However,bloodcompositionwasnotaffected significantlywhenfedatalevel
of100g/kgdiet.Thelatterresultsareinagreementwiththeresultsofour
blood chemistry studies in pigs (Chapter 5; Schutte et al., unpublished
results).A decreased liver weight ofchicks fed on diets containing high
levelsofeitherpentosesugar(400g/kgdiet),andatendencyforthisdecrease
at dietary levels of200g/kg,was reported bythe same authors (Wagh&
Waibel,1966,1967b).Theseobservationstogetherwithourresultsinchicks
(Chapters 1and3)indicatethatbelowdietarylevelsof200g/kgneitherD-
xylosenor L-arabinose willaffect liver weight.
Whendietarysugarsarenotabsorbedcompletelyinthesmallintestine,
therewillbeconsiderablewaterretentioninthelumen(Cunninghametal.,
1963; Kidder et al., 1968; Gray, 1983; De Groot, 1987). Consequently,
diarrhoeamayoccurwhenfeedingsugarswithanincompleteabsorptionin
excessivequantities(Darly&Day,1939;Boothetal.,1953;Wiseetal.,1954;
Walker &Faichney, 1964;Bar, 1985;De Groot, 1987).The results ofour
experiments with D-xylose and L-arabinose showed an increased water
intake,and adecreased drymatter contentofilealdigestaandoffaeces in
pigs(Chapters4and6).Similarly,inchickswaterintakewasincreasedand
drymatter content ofexcretawasdecreased whenfed ondiets containing
either D-xyloseorL-arabinose (Chapters 1and 3).These disturbances in
ileal digesta, faeces and excreta consistency are usually due to increased
amounts of osmotically active compounds in the caudal parts of the
gastrointestinal tract(Hof, 1980).Theextraosmoticloadmaypartlybethe
resultoftheunabsorbed sugarsandpartlyoftheacidsproducedastheend
products ofbacterial fermentation ofthe former (DeGroot, 1987).Due to
osmoticallyattractedwaterinthelumen,anincreaseinthesizeandweight
ofthecaecumisafurther resultofincreased amountsofosmoticallyactive
compoundsintheintestinaltract(Leegwateretal.,1974;Walker,1978;Bar,
1985). Caecal enlargement is considered to be an adaptive phenomenon
ratherthanatoxiceffectsincenosignsoftissuedegenerationwereobserved
from an increased intestinal weight (WHO, 1974). In our chick studies,

caecalenlargement wasmainly observed whenL-arabinose wasfed. This
couldbeexpectedbecause,unlikeD-xylose,partoftheingested L-arabinose
enters thehind gutformicrobial fermentation.
Carbohydrates which stimulate fermentation in the hind gut usually
haveanegativeeffect onproteindigestibilityduetoincreasedfaecallossof
N. Generally, an increase in N excretion in the faeces due to increased
fermentation is accompanied bya reduction inurine Nexcretion without
effect onNbalance(Partridge et al., 1982;Dierick etal., 1983;Malmlof&
Hakansson,1984;Morgan&Whittemore,1988).OurstudiesonL-arabinose
inpigsdonotsupportthesefindings(Chapter6).Inadditiontoadepressed
faecalNdigestibility,Nlossesinurinewereincreasedslightlyinpigsfedon
dietscontainingL-arabinose.Consequently,thesepigsretainedlessN.This
reducedNretentionwasobservedinallL-arabinosetreatmentsregardless
the dose administered. In chicks both treatments with 25 and 75 g L-
arabinose/kgdietcausedasimilardecreaseinproteinutilization (Chapter
3).AsimilarpatternofdecreaseinproteinutilizationwasobtainedwhenD-
xylosewasfedtochicksandpigs(Chapters3and 5).However,considering
the data of our results with pigs (Chapter 5), this decrease in protein
utilization from D-xyloseseemstobeexclusively the result ofan increase
inurinary Nexcretion.Thefact that, in contrast toL-arabinose, faecal N
excretion isnot affected byD-xylosecanbe explained bythe absence ofa
microbialdegradation ofxyloseinthehind gut.Takingtheresultsofboth
pentose sugars on protein utilization together, it can be concluded that
feeding either D-xylose or L-arabinose to pigs and poultry results in an
increased N excretion via the urine. The mechanism responsible for this
phenomenon isnotyetclear.
IthasbeenreportedthatfeedingD-xylosereducedvoluntaryfeedintake
in birds (Baker, 1977). This was confirmed in our studies with chicks
showingthatevenatalowdietarylevelof25g/kg,feedintakewasdepressed
(Chapters1and3).Thiseffectwasnotobservedwhenbirdswerefedondiets
containingL-arabinose(Chapters1and3).Ourobservationsareinagreement
withthosereportedbyKare&Medway(1959).Theseauthorsreported an
almost totalrejection ofwater solutionscontainingD-xylosewhen offered
tobirdstogetherwithuntreated water.Insimilartestsperformed bythese
investigators,othersugarssuchaslactose,galactose,raffinoseandarabinose
hadonlyminoreffectsonwaterconsumption.Afterstudyingmanyvariables
including sweetness, viscosity, refractive index.concentration, osmotic
pressure and density,Kare& Medway(1959)concluded that therewasno
distinct basisfor thebirds'selection other than an absolute specificity for
the sugars involved. No reliable data are available on whether the pig
discriminates between glucose,xyloseand arabinose.
CONCLUSIONS
It appears that D-xylose and L-arabinose, in spite of their identical
molecular size,haveadifferent modeoftransportinthesmallintestineof
monogastricanimals.ThepentosesugarL-arabinoseisabsorbedatalower
rate than D-xylose,whereas absorption velocity of D-xyloseis slightly
lowerthanthatofD-glucose.Bothpentosesugarsarepartlyexcretedinthe
urine.Theextentofthisurinaryexcretioninpercentageofintakeincreases
as the dietary inclusion of either D-xylose or L-arabinose is increased.
However, at equal dietary dose levels, more xylose than arabinose is
excretedintheurine.Thereisnodirectevidenceavailabletostatewhether
conversion toglucoseisafate ofD-xyloseand L-arabinose in monogastric
animals. It appears that some D-xylose may be catabolized to carbon
dioxide,but this pathway seems tobe ofno significance for L-arabinose.
Therefore it is concluded that the energy value of both pentose sugars
mainly depends ontheir degree offermentation. Taking into account the
losses in energy arising from this fermentation process,it was estimated
thattheenergyvalueofthetwopentosesugarsisapproximately25to35%
ofthat ofD-glucose.
FeedingofD-xyloseorL-arabinosetopigsandpoultrycausesaseriesof
physiological changes. In some cases, these changes are similar to those
observed in rats fed lactitol, lactose, xylitol and sorbitol, all ofwhich are
knowntobepoorlyabsorbed inthe small intestine.Unlikewell digestible
sugarsand starches,substantial amountsoftheseproductsare fermented
microbially. Consequently, this will result in an increased production of
volatilefattyacidsandlacticacid,increasedwaterretentionintheintestinal
contentsandthefaeces,anddistentionandincreasedweightofthecaecum.
Changes inducedbyD-xylosearelesspronounced than thoseproducedby
L-arabinose.Thelattercanbeexplainedbythedifferenceinilealdigestibility
betweenbothpentosesugars;D-xyloseisdigestedalmostcompletelyatthe
terminal ileum,whilearabinose isalsopartly digested inthe hind gut.
Thereareindicationsthatbothpentosesugarsaffect protein utilization.
Both faeces N and urine N increase when L-arabinose is fed to pigs.
However,inpigsfedondietscontainingD-xyloseonlyanincreaseinurine
Nwas observed. When D-xyloseisincluded in chick diets,voluntary feed
intakeisdepressed.ThisphenomenonisnotobservedwhenL-arabinoseis
added tochickdiets.
In conclusion it can be stated that not onlythe nutritional value ofD-
xyloseandL-arabinoseforpigsandpoultryislow,butthesesugarsalsomay
induceunwantednutritiveproblemstogetherwithwetdroppingsinchicks.
Considering these aspects, the benefits of a dietary application of NSP
degradingenzymeswhichwillreleasemainlythesesugars(e.g.hemicellulose)
areverydoubtful.Inthisstatementapossibleimprovementofthedigestibility
ofdietaryNandfatasaresultfromanenzymaticaldegradationofsuchNSP
isnottakeninto account.
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161
SUMMARY
163
SUMMARY
Plant carbohydrates arethemajor energycomponentofpigsand poultry

diets. These carbohydrates contain twobroad classes ofpolysaccharides,
starchandthepolysaccharidesoftheplantcellwallwhichareconveniently
referred to as nonstarch polysaccharides (NSP). Starch, a storage
carbohydrateinplantssuchascerealgrains,isana-linkedglucansusceptible
tohydrolysisbypancreaticcc-amylase.Thus,itcanbedigestedinthesmall
intestine of monogastric animals, and absorbed as glucose. Nonstarch
polysaccharides, which comprise the remainder of the dietary plant
polysaccharides, are all resistant toa-amylase. They areprincipallycell
wall structures and include a mixture of substances such as cellulose,
hemicellulose,pectinandoligosaccharideswhichcontainhexoseandpentose
sugars and uronic acids. In some feed ingredients including cereal by-
products,soyabeans,sunflower seed,groundnuts,rapeseedandlupins,the
carbohydrate fraction exists mainly as NSP. In spite of the absence of
indigenous NSP degrading enzymes in the small intestine of pigs and
poultry, some NSP is utilized by these animal species via microbial
degradationinthehindgut.Volatilefatty acidsandlacticacidformed asa
resultofthemicrobialdegradationcanbeabsorbedandutilizedasasource
ofenergy.However,inmicrobialconversionandthesubsequent utilization
ofthese organicacidsapproximately 30to50%ofthe energyislost.
ThereisconclusiveevidencethatthedigestibilityofNSPcanbeimproved
bytreatment withenzymeswhichhydrolysetheNSPtomonosaccharides.
Thebenefits ofahydrolysisofNSP,however,arenotdetermined solelyby
an improvement indigestibility,but alsobythepotential ofthe animal to
utilize theproducts ofhydrolysis.Inaddition toglucose,other sugarswill
also be released by complete hydrolysis ofNSP in normal practical feed
compositions. In quantitative terms the pentose sugars D-xylose and L-
arabinose arethemostimportant ones.Areviewoftherelevant literature
(GeneralIntroduction)revealedthatknowledgeaboutthenutritionalvalue
ofthesesugarsinpigsandpoultryisincomplete.Mostoftheliterature data
are concerned with the rate ofabsorption ofbothpentose sugars from the
intestinaltract insmalllaboratory animalsandman.Itappearsthat both
pentose sugars can be absorbed from the intestinal tract of monogastric
animals,but at alowerrate thanglucose.Moreover,absorptionvelocityof
L-arabinoseappearstobelowerthanthatofD-xylose.Theliteraturereview
alsoindicated that bothpentose sugars arepartly excreted inthe urine.
The present studies were designed to obtain quantitative data on the
digestion and utilization ofD-xyloseand L-arabinose inpoultry andpigs.
Inaddition,somephysiologicaleffects ofthepentosesugarsinbothanimal
species were examined. Six studies were performed, three with poultry
(Chapters 1,2and 3)and three withpigs(Chapters4,5and6).
164 Summary
InChapter 1the effects ofgraded levelsfrom 25to 150g/kgofdietary D-

xyloseand L-arabinose onchickperformance andthemetabolizable energy
(ME) value of both pentose sugars were studied. Weight gain and feed
conversion efficiency were decreased linearly when the dietary level of
either D-xyloseor L-arabinose was increased. The same was true for daily
feedintakeontheD-xylosetreatments.Waterintakewaslinearly increased
as the dietary level ofboth pentose sugars increased, and as a result dry
matter content ofthe droppings decreased. Caecal length and weight were
markedly increased byfeeding L-arabinose and intermediately by feeding
D-xylose. The concentration of glucose in the blood was not affected by
feedingeitherD-xyloseorL-arabinose.TheMEvalueofbothpentose sugars
was considerably lower than that of glucose and negatively related with
dose.
In Chapter 2, results are reported of a study with ileostomized adult
roostersinwhichtheilealdigestibilityandurinaryexcretionofboth pentose
sugars at dietaryinclusions rangingfrom 25to 100g/kg,were determined.
IlealdigestibilityofD-xylosewasfoundtobecloseto100%.Ileal digestibility
ofL-arabinose wasdoserelated andvariedbetween 95(at25g/kgdiet) and
75% (at 100 g/kg diet). Both pentose sugars were partly excreted in the
urine. The extent of urinary excretion in percentage of intake increased
linearly as the dietary level increased. When fed 25gD-xylose per kg diet,
about 7%ofthe ingested xyloseappeared in the urine.This level increased
to 20% when roosters were fed a diet containing 100 g D-xylose/kg.
Corresponding values for L-arabinose in urine at these dietary inclusion
levels were approximately 9 and 17%.
In Chapter 3the results ofa study are described in which the utilization
ofboth pentose sugars at dietary inclusion of 25,50 and 75g/kg in chicks
was examined. Although the experimental diets were balanced for
metabolizable energy content, the performance of birds fed the pentose
sugarswasinferior tothosefedthereference diet.Basedoncarcass analysis
itwas concluded that bothpentose sugars canprovide onlysomeenergy to
chicks. Protein utilization tended to decrease when either D-xylose or L-
arabinose was included in the diets.
In Chapter 4ilealdigestibility and urinary excretion ofD-xyloseand the
associated effects ofthis pentose sugar on ileal and faecal digestibility of
DM,OM,GEand Nwereinvestigated inpigs.IlealdigestibilityofD-xylose
wasfoundtobecloseto 100%.ThepresenceofD-xyloseinthediet increased
the ileal flow ofvolatile fatty acids, suggesting the occurrence of microbial
degradationofD-xyloseinthesmallintestine.Inpigsfedondiets containing
100 and 200 g D-xylose /kg, 44.5 and 52.6%,respectively, of the D-xylose
intake appeared in the urine. Ileal and faecal digestibility ofDM,OM, GE
and N as well as N retention, were not affected in pigs fed on the 100gD-
Summary 165
xylose/kgdiet.Atahigherinclusionlevelof200gD-xylose/kgdiet,digestibility
ofall these parameters and retention ofN was decreased significantly.
In Chapter 5,urinary excretion ofxylosein relation to age,frequency of
feeding and dietary inclusion ofD-xylose were examined in pigs. Urinary
excretion ofxylose was not affected significantly by age and frequency of
feeding, but certainly bydose level.In pigsfed ona diet containing 25gD-
xylose/kg,about 20%ofthe D-xyloseconsumed appeared inthe urine. This
level increased toabout 43%when pigswere fed on a diet containing 100 g
D-xylose/kg.Retention ofNwas slightly decreased when pigswere fed 100
gD-xylose/kgdiet.Liverandkidneyweight,pHofurineandbloodcomposition
were not significantly affected by the inclusion ofD-xylose in the diets.
In Chapter 6 the results of a study are reported in which the ileal
digestibility and urinary excretion ofL-arabinose were investigated. Ileal
digestibility ofL-arabinose was approximately 70%.Administration of L-
arabinose to pigs was associated with an increase in ileal flow of volatile
fatty acids, suggesting the occurrence of microbial degradation of this
pentose sugar inthe smallintestine.Inpigsfed ona dietcontaining 25gL-
arabinose/kg, about 11%ofthe arabinose consumed appeared inthe urine.
This level increased to about 15%when pigs were fed on a diet containing
100 g L-arabinose/kg. Faecal digestibility and retention of N decreased
significantly in pigs fed on the L-arabinose diets.
IntheGeneralDiscussion,theresultsoftheseexperimentswerecombined
withdataofliteratureonthefateofbothpentosesugarsinpoultryandpigs.
There isnodirect evidence availabletostatewhether conversiontoglucose
isafateofD-xyloseandL-arabinoseinmonogastricanimals.Itappears that
someD-xylosemaybecatabolizedtocarbondioxide,butthispathway seems
to be ofno importance for L-arabinose. Therefore, it is concluded that the
energy value of both pentose sugars depends mainly on their degree of
fermentation. Their energy value was estimated to be 25 to 35%ofthat of
D-glucose. In addition to the low energy value, both pentose sugars may
induceunwanted nutritiveproblemstogetherwithwetdroppingsinchicks.
Therefore, the benefits ofa dietary application ofNSP degrading enzymes
which will release mainly these sugars (e.g.hemicellulose) are considered
to be very doubtful. In this statement a possible improvement of the
digestibilityofdietaryNandfatasaresultfromanenzymatical degradation
ofsuch NSP is not taken into account.
167
SAMENVATTING
169
SAMENVATTING
Varkensenpluimvee zijnvoorhun energievoorzieninggrotendeels aange-

wezen opdekoolhydraten uit plantaardige grondstofFen. De koolhydraten
kunnen globaal in twee groepen worden ingedeeld; zetmeel en de niet
zetmeel-koolhydraten. De laatste groep wordt gemakshalve afgekort tot
NSP. Deze afkorting is afgeleid van de engelse benaming voor de niet
zetmeel-koolhydraten(nonstarchpolysaccharides).Voorzoverdekoolhydraat
fraktie uitzetmeelbestaat (b.v.ingranen)levertdeverteringbijvarkens en
pluimvee veelal geen bijzondere problemen op.Beide diersoorten beschik-
kennl.overeenenzymsysteemdathetzetmeelkanafbreken totglucose;een
suikerdiezeergoedwordtbenut doorvarkensenpluimvee.Deniet-zetmeel
koolhydraten (NSP) zijn gecompliceerde verbindingen, zowel uit het oog-
punt van hun fysische structuur als hun chemische samenstelling. Tbt de
NSPwordengerekenddekoolhydraatkomponentencellulose,hemicellulose,
pectine en oligosacchariden. In sommige veevoedergrondstoffen zoals
sojaschroot, zonnebloemzaadschroot, raapzaad en graan bijprodukten be-
staat de koolhydraatfraktie vrijwel geheel uit NSP. Varkens en pluimvee
beschikken niet over enzymen die de NSP frakties kunnen afbreken. Als
gevolghiervankomendezefrakties vrijwel onverteerd indeblindeen dikke
darm terecht, waar zemicrobieel afgebroken kunnen worden. De eindpro-
duktenvanditfermentatieproces(vluchtigevetzurenenmelkzuur)worden
doorhetdiergeresorbeerdinhetbloedenvervolgensbenutalsenergiebron.
In vergelijking met een enzymatische vertering treden bij een microbiele
fermentatie van voeder-componenten echter aanzienlijke verliezen aan
energie op. De schattingen omtrent de hoogte van deze verliezen lopen
uiteen van 30 tot 50%.
Verbetering van de vertering van NSP is in principe mogelijk door het
opnemenvan enzymen in devoeders diedeNSPfrakties kunnen afbreken.
Dit wordt bevestigd door literatuur gegevens en eigen onderzoekingen.
Betreffende de kennis van de toepassingsmogelijkheden van enzymen,
vooral gericht op de Nederlandse situatie, zijn er nog vele leemten. Deze
leemten inkennis doenzichmet name voorophet gebiedvan de benutting
van deprodukten dievrijkomen bijeenenzymatische afbraak van NSP.Bij
een volledige hydrolyse van NSP worden naast glucose nl. tevens
monosacchariden vrijgemaakt dienormaliter niet v66rkomen in de dunne
darmbijvarkensenpluimvee.Debelangrijkste inditopzichtzijndepentose
suikersD-xyloseenL-arabinose.Deliteratuur (AlgemeneInleiding)betref-
fende het metabolisme en de benutting van deze pentose suikers door
varkens en pluimvee is beperkt en verwijst meestal naar inzichten geba-
170 Samenvatting
seerd ophumaan-medisch onderzoek. Volgens de literatuur kunnen beide

strikers geresorbeerd worden uit de dunne darm, maar met een lagere
efficientie dan glucose. Daarnaast blijkt D-xylose slechter te worden
geresorbeerd dan L-arabinose. Uit het literatuur onderzoek kwam even-
eens naar voren dat beide pentose suikers voor een deel weer worden
uitgescheiden via de urine.
Het doelvan deinditproefschrift beschreven experimenten was tweele-
dig.Enerzijds washet gericht ophet verkrijgen van inzicht in de vertering
en benutting van de betreffende pentose suikers bij varkens en pluimvee.
Anderzijds werd onderzochtinhoeverrebijhet opnemenvanbeide suikers
in het rantsoen, fysiologische veranderingen bij deze twee diersoorten
optraden. Beide pentose suikers werden onderzocht in doseringen varie-
rend tussen de 25 en 150g/kgrantsoen.Als referentie suiker werd steeds
D-glucosemeegenomen. In totaal werden zes studies uitgevoerd waarvan
driemetpluimvee(Hoofdstukken 1,2en3)endriemetvarkens (Hoofdstuk-
ken 4, 5en 6).Voorhet meten van de ileale verteerbaarheid werd gebruik
gemaaktvanvarkensenvolwassenhanendiewarenvoorzienvaneen darm
canule aangebracht aan het eind van het ileum.
In hoofdstuk 1 worden de resultaten van een onderzoek behandeld,
waarin het effect van oplopende doseringen (25-150 g/kg)aan D-xylose en
L-arabinose in het rantsoen op een aantal parameters bij kuikens werd
bestudeerd. Het opnemen van D-xylose ofL-arabinose in devoeders resul-
teerde in een verslechtering van de groei en voederconversie en een
verhogingvandewaterconsumptie.Ditlaatsteleiddetoteendalingvanhet
drogestofgehalte indeexcreta. DoorD-xylosewerd verder de voeropname
negatiefbeinvloed.Dematevanvoornoemdenegatieve effecten nam toebij
het oplopen van de dosering aan de pentose suikers in het rantsoen. De
omzetbare energie (OE)waarde van beide pentose suikers was niet alleen
aanzienlijk lager danvan D-glucose dochtevens negatief gecorreleerd met
de dosering. Het glucose gehalte in het bloed werd niet beinvloed door de
aanwezigheid van D-xyloseofL-arabinose inhetrantsoen. Er werden geen
aanwijzigingen verkregen dat onder invloed van de pentose suikers be-
paaldeveranderingen indeorganenoptraden.Ditlaatstemet uitzondering
van de blinde darm gewichten, welke met name bij het verstrekken van
rantsoenen met L-arabinose sterk toenamen.
Ineenvolgendonderzoek(Hoofdstuk 2)werddeilealeverteerbaarheid en
de renale uitscheiding aan D-xylose en L-arabinose bij volwassen hanen
bepaald.Aan het einde van de dunne darm werd vrijwel geen xylose meer
aangetroffen, hetgeen inhoudt dat deze pentose suiker vrijwel volledig
ileaal was verteerd. De ileale verteerbaarheid van L-arabinose was dosis
afhankelijk. Opgenomen in een dosering van 25 g/kg rantsoen bleek L-
Samenvatting 171
arabinose voor ongeveer 95%ileaal te zijn verteerd. Dit percentage liep

terug tot ongeveer 75%bij een dosering van 100 g/kg rantsoen. Een
belangrijk deelvandeoraal verstrekte pentose strikers werd viade urine
weeruitgescheiden.Dematevandezeuitscheidingwassterkpositiefgecor-
releerd metdedosering. D-xylose werd bijdoseringen van25en 100g/kg
rantsoen voor respektievelijk 7en20%uitgescheiden viadeurine. Decor-
responderende waarden voor L-arabinose bij deze doseringen waren 9en
17%.
Inhetlaatste onderzoekmetpluimvee(Hoofdstuk3)laghetaccentophet
bestuderen vandebenutting vandetwee pentose suikers door kuikens bij
doseringenvan25,50en75g/kgrantsoen.Devoederswerdenopisocalorische
(OE)basis verstrekt aandedieren. Desondanks werden slechtere groeien
voederconversie resultaten gemeten opderantsoenen met D-xylose ofL-
arabinose. Gebaseerd opkarkas analyses werden aanwijzingen verkregen
dat de energie van beide pentose suikers slechts in beperkte mate kan
worden benut door kuikens. Daarnaast bleken beide pentose suikers de
eiwit retentie negatief te beinvloeden.
In hoofdstuk 4 worden studies beschreven welke in hoofdzaak waren
gerichtophetbestuderenvandeilealeverteerbaarheidenderenaleexcretie
van D-xylose bijvarkens. D-xylose bleek vrijwel volledig ileaal te worden
verteerd. Deaanwezigheid vanD-xyloseinhetrantsoen ginggepaard met
een sterke verhoging vandepassage aanvluchtige vetzuren aanhet eind
van de dunne darm. Dit verschijnsel werd in verband gebracht meteen
gedeeltelijke fermentatie vanD-xylose in de dunne darm. Opgenomen in
doseringenvan 100en200g/kgrantsoen,werdrespektievelijk 44,5en52,6%
van deopgenomen hoeveelheid D-xylose uitgescheiden viadeurine. Ileale
enfaecale verteerbaarheid van dedrogestof,organischestof,bruto energie
en eiwit alsmede deeiwit-retentie werden niet duidelijk beinvloed doorD-
xylose bijeendosering van 100g/kg rantsoen. Opgenomen in een hogere
dosering van 200 g/kg rantsoen, werden de verteringscoefficienten van
voornoemde parameters endeeiwit-retentie sterk negatief beinvloed.
Inhetvolgendeonderzoekmetvarkens(Hoofdstuk 5)werddeinvloedvan
enkele faktoren die mogelijk de renale uitscheiding aan xylose kunnen
beinvloeden, onderzocht. De mate van de uitscheiding aan xylose via de
urine wassterk positief gecorreleerd metdedosering. Opgenomen ineen
doseringvan 25g/kgrantsoen werd 20%van deopgenomen hoeveelheid D-
xyloseuitgescheiden indeurine.Ditpercentage liepoptot43%als varkens
een rantsoen kregen verstrekt met 100g D-xylose/kg. De leeftijd vande
dieren endefrequentie vandevoederverstrekking hadden geen duidelijke
invloed opderenale uitscheiding aanxylose.Lever- enniergewichten, pH
van de urine en de samensteling van het bloed werden niet significant
172 Samenvatting
beinvloed door D-xylose.

In hoofdstuk 6 tenslotte zijn de resultaten van een onderzoek met L-
arabinose bijvarkens beschreven. Opileaal niveau bleek L-arabinose voor
ongeveer 70% te worden verteerd. Verstrekking van rantsoenen met L-
arabinose ging gepaard met een verhoging van de passage aan vluchtige
vetzurenaanheteindvandedunnedarm.Ditverschijnsel werdin verband
gebracht met een gedeeltelijke fermentatie van L-arabinose in de dunne
darm.DerenaleuitscheidingaanL-arabinosewaspositiefgecorreleerdmet
dedosering.Bijeendoseringvan 25g/kgrantsoen werdvan de opgenomen
hoeveelheid L-arabinose ongeveer 11%via de urine uitgescheiden. Bij een
dosering van 100g/kgrantsoen bedroeg dit 15%. Faecale verteerbaarheid
vanheteiwitendeeiwit-retentiewerdendoorL-arabinose negatiefbeinvloed.
De resultaten van de hier beschreven proeven werden tesamen met
gegevens uit de literatuur nader geevalueerd (Algemene Discussie). De
literatuur levert geen direkt bewijs dat D-xylose en L-arabinose
gemetaboliseerd kunnen worden tot glucose. Een andere mogelijke route
van beide suikers, omzetting tot C02, lijkt alleen in beperkte mate van
toepassing te zijn op D-xylose. Op basis van deze twee uitgangsstellingen
werd geconcludeerd dat de energiewaarde van beide pentose suikers voor
pluimvee en varkens in hoofdzaak wordt bepaald door de mate van hun
fermentatie inhetmaag-darmkanaal.Eenglobaleberekeningleerdedat de
energiewaarde vanbeidepentose suikershooguitgesteld kanwordenop25
a 35%vandievanD-glucose.Ditbetekent datbeidepentose suikers slechts
in geringemate eenbijdrage kunnen leveren aan deenergievoorzieningbij
varkens en pluimvee. Gezien ditlaatste enmedegeletophet feit dat beide
pentose suikers aanleidingkunnen geven tothet optredenvan ongewenste
neveneffekten, lijkt eenvolledige enzymatische afbraak vanNSP frakties,
waarbij in hoofdzaak xylose en arabinose vrijkomen (b.v. hemicellulose),
voormonogastrische landbouw-huisdieren geenbijzondere voordelen op te
leveren.Hierbijisgeenrekeninggehoudenmeteeneventueelpositief effekt
vaneenenzymatischeafbraak vandezeNSPfrakties opde verteerbaarheid
van eiwit en vet.
173
CURRICULUM VTTAE
Johannes Bernardus Schutte werd geboren op 21 mei 1936 ter gemeente

Odoorn. Achtereenvolgens behaalde hij het diploma van de lagere en
middelbare landbouwschool, de MULO en de praktijkschool voor de
pluimveeteelt. In 1976 slaagde hij cum laude voor het diploma Register
Ingenieur. In 1988 werd aan hem vrijstelling verleend voor het doctoraal
examen aan deLandbouwuniversiteit Wageningen.Van 1958tot 1984 was
hij verbonden aan het ILOB (Instituut voor Landbouwkundig Onderzoek
van Biochemische produkten).Sinds 1mei 1984ishet ILOBopgenomen in
de TNO organisatie onder de naam TNO Instituut voor Diervoeding en
Fysiologie. Zijn huidige funktie is wetenschappelijk medewerker en hoofd
van de sectie voedingsonderzoek pluimvee en varkens.

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NUTRITIONAL VALUE AND PHYSIOLOGICAL

EFFECTS OF D-XYLOSE AND L-ARABINOSE

0000 0456 7893

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10. Desuggestiedateenlinolzuur-rijk eigezonderisdaneennormaal ei,berust

11. Mettuinieren alshobbylooptmenzeldenpersoonlijk gezichtsverlies op;de

12. Debevolkinginhet westenvan onsland dient zichmeerbewust te worden

Proefschrift J.B. Schutte.

Gaarne maak ikvan dezegelegenheid gebruik omalientebedanken die direkt

Chapter 1. Nutritional implications and metabolizable energy 23

Chapter 2. Ileal digestibility and urinary excretion ofD-xylose 39

Chapter 3. Nutritional value ofD-xyloseand L-arabinose for 57

Part B.Studies with pigs 77

Chapter 4. Nutritional implications ofD-xyloseinpigs 79

Chapter 5. Urinary excretion ofD-xyloseinpigsasaffected 99

Chapter 6. Nutritional implications ofL-arabinose inpigs 117

General discussion 141

Curriculum Vitae 173

NATURE OFNONSTARCH POLYSACCHARIDES

The nonstarch polysaccharides (NSP) are complicated compounds, both

Cellulose Hemi- Pectin Oligo- Starch+

FIGURE 1. Classification of carbohydrates in vegetable feeds

important constituent ofthe NSP in legume seeds.The raffinose family is

TABLE1.Contents ofcarbohydrates, starch (+sugars) andNSP in

Ingredient Ibtal Starch NSP

1)Valuesderived from figures givenbyAnonymous(1988).

HYDROLYSIS PRODUCTS OF NSP

From aliteraturereview,Rexen(1981)and Chesson (1987)concluded that

NSP fraction Hexoses Pentoses Other

Hemicellulose D-glucose D-xylose

Pectins D-galactose L-arabinose Uronic

*Mainlyasgalacturonic acid(=derived form ofD-galactose ofwhich the

In addition to glucose, the pentose sugars xylose and arabinose are in

TABLE3.Composition of NSP isolated from various feed in-

Glucose Xylose Arabi- Galac- Mannose Uronic Lignin

Maize 24.2 24.0 19.1 4.1 0.8 6.0 5.4

ABSORPTION AND UTILIZATION OF D-XYLOSE AND L-

intestinal mucosa by simple diffusion was initially reported (Cori, 1925;

FIGURE2.Structural formulae of D-glucose, D-xylose and L-

D-glucose D-xyl()se L-arabinose

Themetabolicpathways ofbothpentose sugarsisstill under discussion.

infusion of L-arabinose in man. Hiatt (1957) observed in mice a slight

AIM AND OUTLINE OF THE STUDY

Carre\ B.&Leclercq,B.(1985).Digestionofpolysaccharides, protein and

delanature et duniveau desfibres brutes sur la digestibilityil^aleet

Journal ofNutrition, 62: 563-577.

Schiffer, L.M.,Faloon,W.W.&Chodos,R.B.(1962).Malabsorption syndrome

STUDIES WITH POULTRY

NUTRITIONAL IMPLICATIONS AND

TNO-InstituteofAnimalNutrition and Physiology(ILOB),

Published inPoultry Science69: 1724-1730(1990)