* Thierry Gidenne, Rachel Bellier

Transcription

1 Livestock Production Science 63 (000) locte/ livprodsci Use of digestile fire in replcement to ville crohydrtes Effect on digestion, rte of pssge nd cecl fermenttion pttern during the growth of the rit * Thierry Gidenne, Rchel Bellier Institut Ntionl de l Recherche Agronomique, Sttion de Recherches Cunicoles, BP 7, 336 Cstnet-Tolosn, Frnce Received 4 Septemer 998; received in revised form 0 April 999; ccepted 0 My 999 Astrct The dietry replcement of ville crohydrtes (minly strch) y digestile fire ws ssessed on rit growth nd digestion. Two diets supplying hemicellulose (diet HC) or mix of hemicellulose nd pectins (HC P) were compred to control diet hving higher strch content, ut similr content in low digested fire (lignocellulose ADF 5 55 g/ kg). Three groups of 7 rits were llocted t wening (8 d) to ech diet. Nine nimls of ech group were cnnulted to follow the cecl microil ctivity from 45 to 70 d old. Compred to control, rits fed HC or HC P incresed feed intke ( 0%), ut only HC P resulted in higher weight gin nd in similr feed conversion (men.00), prticulrly during the postwening period (4 6 wk old). Trnsit of liquids ws similr mong diets, either in the ceco colic segment or in the whole trct. HC P diet gve short ceco colic trnsit for prticles (3. h), nd lso the highest cecl SCFA level (6. mm/l) nd pool (3.8 mm), while HC nd control diet led to similr cecl fermenttion (men nd.5, respectively for SCFA level nd pool). SCFA concentrtion incresed ccording to the men intke of hemicellulose pectins, suggesting tht microil ctivity nd fire degrdtion were more ffected y the nture of the cell wll thn y the ceco colic retention time. 000 Elsevier Science B.V. All rights reserved. Keywords: Rit digestion; Cecl fermenttion; Strch; Hemicellulose; Pectins; Dietry fire. Introduction hindgut flor nd to ensure digestive (motility nd trnsit) nd metolic functions of the host (Mthers, For monogstric species nd prticulrly for the 99; Crre, 99; Edwrds nd Prrett, 996). growing rit, supplying dietry fire corresponds Digestive consequences of fire re determined s to two min roles: to provide nutrients for the well y the quntity of dietry fire consumed s y the degrdility of cell wll components in reltion *Corresponding uthor Tel.: ; fx: to their physico chemicl structure Previous studies on growing rit hve ddressed E-mil ddress: gidenne@toulouse.inr.fr (T. Gidenne) the prolem of incresing the intke of low digestile / 00/ $ see front mtter 000 Elsevier Science B.V. All rights reserved. PII: S030-66(99)003-

2 4 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 fire to prevent digestive troules (minly dirrhoe), Tle ut ssocited to negtive effect on growth per- Composition of the experimentl diets formnce (Mıtre ˆ et l., 990). Conversely, dietry Diet Control HC HC P strch could lso ply sustntil role in interction Ingredients (g/kg diet) with fire, s it is incompletely hydrolysed in the Whet strw smll intestine of the wenling rit (Bls et l., Whet rn ; Bls et l., 994) cusing ltertion of the Sugr Beet pulp Sunflower mel ceco colic digestion ssocited with digestive dis- Soy en mel turnces. Whet Therefore, the reduction of strch supply should Minerls nd vitmins e explored without excessive reduction in the digestile energy nd without modifying the supply Chemicl composition (g/kg DM) Orgnic mtter in low digested fire frction. Becuse the ceco Strch colic fermenttion is reltively short (7 h; Giden- Crude protein (N 3 6.5) ne, 997), the use of rpidly fermented fire (i.e., Gross Energy (MJ/ kg DM) digestile fire) replcing ville crohydrtes Wter insolule cell-wll could e of interest to supply energy to the host vi Neutrl-detergent fire Acid-detergent fire cecl microil ctivity, without ltertion of rte of Acid-detergent lignin pssge. The level of degrdtion for hemicellulose Hemicellulose frctions (30 45%) nd for pectins (70%) in the Uronic cids c rit clsses these cell wll constituents in the Pectins digestile fire compred to low-digested fires Contined: (g/ kg, respectively for C, HC; HC P diets) such s lignocellulose (Gidenne et l., 998). The clcium cronte nd iclcium phosphte (.0, 7.5, 8.0), present study ssessed two wys of strch replce- DL-methionine (0.5,.5,.5), L-lysine (.5,.0, 0.0) slt (7.0, 7.0, 6.5); nd 5.0 g/kg diet of vitmin premix (retinol 540 mg/kg, ment either minly through supply of hemicellulose choleclcipherol 5 mg/ kg, -tocopherol 3.3 mg/ kg, roenidine or with mix of hemicellulose nd pectins. Further- 3. mg/ kg). more, we imed to develop new pproch to otin Hemicellulose frction estimted y difference etween NDF more comprehensive informtion on cecl digesc nd ADF. tion, tht included simultneous in-vivo mesureingredients Wter insolule pectins: clculted from tulted dt on ment of rte of pssge, nutrient digestion nd cecl (IO7, 993). microil ctivity (CMA), over sustntil period of the rit growth (6 0 wk old). ADF), minly provided y one source (whet strw). Mterils nd methods to void mjor chnge in the type of cellulose or lignin. The rtio digestile protein/ digestile energy.. Diets nd feeding ws mintined constnt mong the diets, y djusting the incorportion of soy en mel nd whet. Three experimentl diets were prepred (Tle ) in pelleted form with rw mteril clssiclly used.. Animls, housing nd experimentl protocol in rit feed formultion, to otin vrition in the level of digestile fire: control diet, nd two Three groups of 7 New Zelnd white rits diets where portion of strch ws sustituted either were llocted (fter controlling the effect of litter y hemicellulose (evluted s NDF-ADF) supplied origin) t wening (8 d old) to one of the exminly y whet rn (HC diet), or y perimentl diets. They were housed in individul hemicellulose pectins through supply in eet metolism cges ( cm) nd sumitted to pulp (HC P diet). In ddition, the three diets hve h light (7:00 to 9:00) nd h drk schedule. similr level of low-digestile fire (lignocellulose, After seven-dy dpttion period to the diet, nine

3 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) rits of ech group were surgiclly fitted with whole trct ws clculted ccording to MRT 5 Mt i i/ T-cnnul in the cecum s descried y Bellier et M i, where ti ws the time tht hs elpsed etween l. (995). Smpling of cecl digest nd diges- mrker dministrtion nd the ith defection nd M i tiility mesurements were performed fter one ws the quntity of mrker excreted in feces. MRT week recovery period. Apprent fecl digestiility includes the miniml trnsit time (TTm), which ws coefficients were clculted from individul nd the time tht hs elpsed etween dosing nd the first dily totl collection during six consecutive dys (4 mrker ppernce in the feces. TTm reflects the to 48 d of ge). retention time of digest without dely in the Smples of cecl digest were individully col- mixing comprtments, so it represents the rte of lected in vivo vi the cnnul t 45, 50, 60 nd 70 pssge in the tuulr segment of the trct, i.e., dys of ge. Rits were plced in specil minly in the smll intestine nd lso in the distl hmmock for 0 min, nd digest drined y grvity colon (Gidenne, 994). in smpling tues (Gidenne nd Bellier, 99). Cecl volume ws determined in-vivo ccording Smpling time ws fixed t the end of the cecot- to the following mrker dilution procedure, on six rophy period (:00) when inter-individul vri- 5 rits per group. The concentrtion of chromiumtions re lower (Bellier et l., 995). After smpling, EDTA ws determined in smples of cecl digest the cecl mteril ws immeditely sujected to the collected,, 4 nd 6 h fter the introduction of the determintion of ph. Portions of cecl digest mrker dose (etween :30 nd :00). A liner smple (5 to 0 g fresh mtter) were plced in tues djustment of the decrese in mrker concentrtion contining (%, v/ v) H3PO4 or HSO4 storge ws performed. The cecl volume ws then osolution ( nd ml/ tue), respectively for further tined y extrpolting to zero time the curve of nlyses of short chin ftty cids (SCFAs) nd trcer concentrtion ginst time (Leng nd mmonicl nitrogen (NH 3), nd stored t 58C. Hornicke, 976), nd expressed in g of fresh mtter. 5 A solution of chromium-edta nd cell wll The cecl pool of SCFAs ws clculted y multipprticles lelled with 4 cerium (Gidenne, 994) lying the SCFA concentrtion (mm/ l) nd the liquid were used s liquid- nd solid-phse mrkers. Cell content of the cecum (in ml, otined from cecl wll prticles were prepred from ech diet, nd volume nd dry mtter determintion). corresponded to the NDF residue fter sieving on 0.05 mm screen. Digest men retention time of.3. Anlyticl methods solid nd liquid phse ws mesured on six cnnulted nimls per group, y following the kinetics of Dry mtter ws determined on feeds nd feces y the fecl excretion of the two mrkers, using n heting for 4 h t 038C. Orgnic mtter (OM) ws utomtic fecl smpler (Automtic Smpler; API, determined y shing t 5508C for 5 h. Mesure- Cstnet, Frnce). First, rte of pssge etween ments of fire frctions (neutrl detergent fire: cecum nd rectum ws recorded during five-dy NDF, cid detergent fire: ADF, cid detergent period (from 48 to 5 d old), following the dminis- lignin: ADL) were mde ccording to Vn Soest et l. trtion (t :30) of the two mrkers in the cecl (99) using n mylolytic pre-tretment with cnnul. Then, rte of pssge in the whole trct ws thermostle mylse, nd wter insolule cell wlls otined on the sme cnnulted rits, during were otined ccording to Crre nd Brillouet six-dy period (from 53 to 57 d old) following the (989). Nitrogen ws determined y DUMAS comorl dministrtion (t 8:30) of mrkers. A dose of ustion method using Leco pprtus (Model FPlelled prticle corresponded to 00 mg DM 48, Leco Corp., St. Joseph, MI, USA), nd con- (ctivity 5 kbq) nd ws given orlly using verted to crude protein using the fctor 6.5. Gross modified syringe (-ml model). The mrker con- energy ws mesured y n ditic clorimeter centrtions were determined in dried feces using (PARR Instrument, Moline, IL, USA). Ammoni gmm spectrometer (Pckrd Instrument, Model concentrtion ws determined spectrophotometriclly 5530, Downersgrove, IL, USA). t 660 nm ccording to Verdow et l. (977). In The digest men retention time (MRT) in the feeds, uronic cids were determined colorimetriclly

4 44 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 y the m-phenylphenol method (Blumenkrntz nd 3. Results Asoe-Hnsen, 973). Strch ws hydrolysed enzymticlly nd the resultnt glucose ws mesured According to the ojectives of the formultion, the y using the hexokinse (EC.7..)-glucose-6- three experimentl diets were similr in their levels phosphte dehydrogense (NAD) (EC...49) sys- of lignocellulose (menly 55g ADF/ kg, Tle ). tem (Boehringer Mnnheim). SCFAs were nlysed They differed minly y their level of strch (high y gs liquid chromtogrphy (CP9000, Chrom- for control) in lnce with the hemicellulose or pck, Middelurg, The Netherlnds) on semi-cpil- pectin level (high for HC nd HC P), lthough the lry column (Bellier, 994). totl fire level (NDF or WICW) differed slightly (0%) mong diets..4. Sttisticl nlyses Rits hving n normlly low feed intke (3 SD under men) during the mesurements were discrded from the sttisticl nlyses. A two wys nlysis of vrince ws done using the generl liner model GLM procedure (Sttisticl Anlysis System, 988), with the effect of cnnultion nd diets nd interction etween diet nd cnnultion s min effects for the prmeters of intke nd growth. Digestiility nd rte of pssge dt were exmined y one-wy nlysis of vrince (ANOVA). The dt on the cecl content composition (Tle 6 nd Fig. ) were exmined to evlute the effect of the ge, diet nd ge 3 diet interctions. The effect of the individul ws controlled in this model, to tke into ccount tht the mesurements (four sm- ples t four successive ges) were repeted on the sme nimls. The significnce of the effect of the diet ws thus clculted using the men squre of the intr-individul vritions s n error term (split-plot design). Lest-squre mens comprisons were per- formed using the PDIFF test of GLM. Dt from unhelthy nimls or nimls hving n norml SCFA pttern (i.e., with propionte/ utyrte rtio over ) were not included in the nlysis. The in vivo SCFA pttern from 6 to 0 weeks of ge ws sttisticlly nlysed fter pooling the results from the three diets, s no interction etween the diet nd the ge of the rit ws found. In ddition, the effect of ge on CMA ws nlysed ccording to the GLM procedure (liner model). The prtil correltion coefficients etween the prmeters of the cecl fermenttion nd the digestion of the nutrients (over the period 6 0 wk old) were otined through multivrite nlysis of vrince (tking into ccount the effect of the diet). 3.. Effect of the cnnultion nd of the diet on intke nd growth Among the six groups of rits, only one niml receiving HC diet died ecuse of n cute dirrhoe (t 48 d of ge). Following the implnttion of the cnnul the rits recovered norml feed intke within four or five dys, so tht from 8 to 45 d of ge the dily feed intke ws only 0% lower for cnnulted nimls. This resulted in lower live weight gin (30%) for cnnulted nimls compred to control (Tle ). From 45 to 70 dys old, the dily feed intke nd the weight gin did not differ significntly mong cnnulted or control group. In ddition, interction etween the effect of the diet nd the group of rits ws not significnt whtever the prmeter. Dietry tretments ffected the intke s well s the weight gin, prticulrly during the two weeks fter wening. The feed intke ws significntly higher ( 0%) for HC nd HC P groups from four to six weeks of ge, ut resulted in sig- nificntly higher weight gin only for HC P group (Tle ). Consequently the feed conversion ws not significntly different (P 5 0.9) mong the three groups for the period 4 6 wk (men.00), ut it ws significntly lower (P ) for the control group compred to HC P during the 6 0 weeks old period (respectively,.73 vs. 3.06). 3.. Nutrient intke nd digestiility Food intke vritions comined with differences in the dietry chemicl composition produced exgg- erted difference in nutrient intke. Thus, lthough

5 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) Tle Effect of cecl cnnultion nd of dietry tretment on feed intke nd growth Effect of the cnnultion Effect of diets Sttisticl difference n 5 3 n 5 4 n 5 6 n 5 5 n 5 6 Cnnultion Control Cnnultion Control HC HC P RMSE Cnnultion Diet 3 Diet Live weight (g) At 8 dys old , At 45 dys old , Intke (g/d/kg LW) 8 to 45 dys old , to 70 dys old Weight gin (g/d) 8 to 45 dys old , 0.00, to 70 dys old Implnttion of cecl cnnul t five weeks old. Numer of rits. RMSE 5 Root men squre error., : Mens, etween diets, hving common superscript did not differ significntly t the level P the dietry ADF level ws similr, the ADF intke versely the strch intke is reduced y round 0% incresed y 0% for HC P group (Tle 3). for these two groups. Compred to control group, the intke of hemicellul- Orgnic mtter digestiility did not differ sigose incresed y four units ( 40%) for HC group, nificntly from control when strch is replced while the intke of pectin tripled for HC P. Re- minly y hemicellulose nd pectin (HC P group), Tle 3 Effect of dietry tretment on the intke of nutrients (totl nd digestile), from 45 to 70 dys old Dily intke Control HC HC P Sttisticl nlysis (g/d/kg LW, ir dry sis) n 5 8 n 5 7 n 5 7 RMSE Rw intke Totl nutrients Orgnic mtter Crude protein Strch , 0.00 NDF , 0.00 ADF Hemicellulose 8.0 c , 0.00 Pectins c , 0.00 Digestile nutrients Orgnic mtter Crude protein NDF , 0.00 ADF Hemicellulose , 0.00, : See Tle. See Tle. P. F

6 46 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 Tle 4 Effect of dietry tretment on the totl trct pprent digestiility Control HC HC P Sttisticl nlysis n 5 8 n 5 7 n 5 7 RMSE Orgnic mtter , 0.00 Crude protein Energy , 0.00 Neutrl-detergent fire , 0.00 Acid-detergent fire Hemicellulose , 0.00 Nutritive vlue (ir dry sis) Digestile protein, DP (g/ kg) Digestile energy, DE (MJ/ kg) DP/ DE (g/ MJ).7.8.9, : See Tle. RMSE: Root men squre error. See Tle. P. F lthough there ws unit decrese (Tle 4). In The minimum trnsit time ws identicl etween contrst, the replcement of strch y hemicellulose solid nd liquid phse, thus only men vlues for the led to 6 unit decrese in OM digestiility. From 6 two phses were given in the Tle 5. TTm for the to 0 wk old, the intke in digestile OM ws similr whole trct ws menly 4.4 h from which 50% mong diets (Tle 3) nd logiclly led to similr ccounted for ceco rectl trnsit. TTm remined weight gin (Tle ). Crude protein digestiility or unffected y the dietry tretments. intke in digestile protein were not significntly Contrry to the liquid phse, the MRT of the solid ffected y dietry tretments. In ccordnce with phse ws ffected y the diets, prticulrly in the the ims of the dietry formultion the rtio DP/ DE ceco colic segment when it decresed (P ) ws similr mong the diets. Whtever the diet, the y 50% for HC P group compred to control. hemicellulosic frction ws digested t higher rte Although not sttisticlly significnt (high interincompred to lignocellulose, reching level of 44% dividul vritions) MRT for the totl trct tended to for HC P group. Compred to control diet, the e lower (P ) for HC P group. TTm ws NDF digestiility ws improved with the ddition of not significntly different mong the three groups, pectin, while it decresed with the ddition of indicting therefore tht the chnge in totl nd hemicellulose (minly from whet rn). This re- ceco rectl MRT occurred in the mixing comprtsulted in high quntity of fire degrded (g/ d) for ments, nd minly in the cecum nd proximl HC P group whtever the fire frction (Tle 3), colon. wheres fire degrdtion ws similr for control nd HC groups Cecl fermenttion nd microil ctivity 3.3. Rte of pssge Whtever the vrile, no significnt interctions were found etween the effect of ge nd the effect The rte of pssge in the whole trct of the of the diet (Tle 6), so these two effects will e digest solid phse ws menly 5.5 h, from which presented seprtely herefter. t lest the two-thirds ws ttriutle to the ceco colic segment (Tle 5). The MRT of the liquid Effect of ge phse ws three-fold higher (men 48 h), nd more The in-vivo totl SCFA concentrtions in the thn 80% originted from the ceco colic segment. cecum ugmented linerly (P ) y 0% from

7 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) Tle 5 Men retention time (MRT) of solid nd liquid phse of digest in the whole trct nd ceco colic segment of the growing rit, ccording to dietry tretment Control HC HC P Sttisticl nlysis n 5 5 n 5 4 n 5 5 RMSE Live weight (g) Feed intke (g/ d) MRT: whole trct (h) Solid phse Liquid phse MRT: cecum rectum (h) Solid phse Liquid phse TTm: miniml trnsit time Whole trct (h) Cecum rectum (h) , : See Tle. Men vlues for the period dys of ge. P. F Tle 6 In-vivo mesurement from 6 to 0 weeks old, of ph, mmoni nd short-chin ftty cids (SCFAs) concentrtion (mmol/ l) in the cecl content of rits ccording to dietry tretment Effect of the diet Sttisticl nlysis n 5 9 n 5 4 n 5 6 Diet 3 Control HC HC P RMSE Diet Age 3 ge Totl SCFAs Acette Propionte Butyrte , Minor SCFAs Propionte:utyrte Ammoni ph Mens from smples covering the period 45 to 70 dys old., : See Tle. 3 Comprison of in vivo smpling ccording to ge (t 45, 50, 60 nd 70 dys old). 4 Liner effect of the ge (P, 0.05) 5 Isoutyrte vlerte isovlerte. 45 d to 70 d of ge (Fig. ), ecuse of significnt 0.00) with ge from 0.7 (t 45 d) to 0.36 (t 70 d). liner increse (P ) in cette (from 48.4 to Ammoni or ph were not significntly ffected y 5.5 mmol/l) nd utyrte (from 4.3 to 8.9 mmol/l) the ge (Tle 6). levels. From 45 to 70 d of ge, we registered two-fold increse of the utyrte molr proportion Effect of the diet (Fig. ), ssocited with slight decrese of the Compred to control, the totl SCFA levels in cette molr proportion (5%). Consequently the HC P group were 8 units higher (P ) s the rtio propionte: utyrte declined linerly (P, level of cette incresed y 0% (P ). On the

8 48 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 Fig.. In vivo cecl SCFA pttern ccording to the ge of the rits. Vlues re mens of the three diets., : Mens hving common superscript did not differ significntly t the level P () Liner effect of the ge (P, 0.05). other hnd, levels of utyrte propionte or mmoni prmeters (SCFA levels or molr proportion), exremined unffected y dietry tretments. In return, cept the level of mmoni tht ws positively relted the cette molr proportion ws significntly lower to crude protein digestiility (r , P 5 0.0). (P ) for control compred to HC nd HC P A reltively low numer of nimls ws involved group (respectively, 8.6% %), while in the in-vivo determintion of the cecl volume the utyrte molr proportion tended to e higher nd SCFA pool (Tle 7), ecuse the mrker (P ) for control (.%) compred to the two dilution procedure is reltively difficult to mnge other groups (men 9.%). nd time consuming. Therefore, only lrge differ- The reltionship etween cecl fermenttion nd ences mong tretments could ppered s signifithe digestion of the min nutrients (OM, CP, NDF, cnt. At 70 d of ge, the digest content of the ADF nd NDF-ADF) hs een ssessed y clcult- cecum corresponds menly to 55g of fresh mtter ing the prtil correltion coefficients (r) etween tht represented 5% of the live weight. The cecl ph, NH3 or SCFA levels nd the digestiility content reched 7g for HC P tht ws 80% over coefficients or the dily quntity of nutrients di- the vlue for HC group. Therefore the SCFA pool gested. Digestiility or nutrients intke were not ws significntly higher for HC P(65%) comsignificntly correlted with ny of the fermenttion pred to the two other groups.

9 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) Tle 7 In-vivo mesurement of the digest content nd of SCFA pool in the cecum of growing rits (7 wk old) ccording to dietry tretment Control HC HC P Sttisticl nlysis n 5 4 n 5 3 n 5 5 RMSE Live weight (g) Cecl digest content (g) SCFAs (mm/ l) SCFA pool (mm) , : See Tle. SCFAs (mm/l) 3 liquid cecl content (ml). P. F 4. Discussion higher thn in hrd feces, Luick et l. 99) tht re ingested vi cecotrophy prctice. Consequently 4.. In-vivo pproch of the digestive processes of portion of the liquid mrker ws recycled nd the growing rit contriuted to increse liquid retention time, s oserved previously in ceco colic segment y Although cecl cnnultion led to temporry Gidenne nd Jehl (996) using the sme method. loss in feed intke nd live weight, no interction Leng et l. (977) mentioned shorter MRT for with the effect of the diets ws detected for intke or liquids (men 6.4 h) etween cecum nd rectum, growth, s oserved in previous experiments using 4 ut they used C -PEG (polyethylene glycol) tht is the sme methodology (Bellier nd Gidenne, 996; incompletely recovered in excret. Rte of pssge Gidenne et l., 998). Such technique ppered of prticles in the ceco colic segment (MRT prticulrly convenient to ssess in-vivo the hindgut TTm) ws menly 8.4 h tht ws in the rnge of digestive processes, s vrious mesurements could previous results otined y mthemticl modelise performed over complete growth period: digest tion (Gidenne, 994; Gidenne nd Perez, 996). composition, cecl volume (using mrker dilution As expected, prticles nd liquids hve strictly the technique) or ceco rectl rte of pssge. Further- sme retention time in the distl colon (corremore, our study demonstrted tht it ws possile to sponding to the ceco rectl TTm, Gidenne, 994) comine SCFA nlysis in digest nd digestive s digest re lredy ggregted into fecl pellets volume mesurements, to evlute in-vivo the pro- (Gidenne nd Jehl, 996). Similr conclusions could duction of fermenttion end-products. lso e drwn for digest retention in the smll The ssocition of doule lelling technique intestine, which ws ssessed y the difference (simultneous mesurements for liquid nd solid etween totl nd ceco rectl TTm (menly. h). phse) to tht of cecl cnnultion let us to provide The oro cecl retention time ws estimted, for vlule new informtions out digest rte of liquid nd prticles, y difference etween totl nd pssge in different segment of the rit trct, such ceco rectl MRT mesured consecutively on the s oro cecl trnsit of liquids. In the whole trct, sme nimls. A differentil trnsit seemed to occur liquid retention time reched 48 h, in greement with etween liquid nd solid phse, s we found Luick et l. (99). The ceco colic segments shorter rte of pssge for prticles (men 4.9 h) thn (cecum proximl colon) were the min site of for liquid (men 7.5 h). This differentil trnsit could liquid retention (menly 39 h, otined y difference e ttriuted to the retention in the stomch, since etween ceco rectl MRT nd TTm). The retro- liquids nd prticles moved without differentil grde trnsport of fluid from the proximl colon to trnsit in the smll intestine. This higher gstric the cecum (occurring when hrd feces re pro- retention time for liquids would e relted to the duced) is minly responsile of this phenomenon retention in the fundus of soft feces hving high (Bjornhg, 98). It lso explins the high con- concentrtion in liquid mrker. centrtion of Cr-EDTA in soft feces (up to 00-fold The in-vivo mesurement of the rit cecl

10 50 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 volume ws first ssyed y Leng nd Hornicke HC P). However, slight effect of the overll (976) using the mrker dilution method nd de- dietry fire level could not e excluded on feed scried diurnl vritions ccording to cecotrophy intke, s NDF vried y 0% mong diets. In prctice. Therefore we pplied this procedure fter return, only supply in hemicellulose nd pectin the cecotrophy period (t :30) to reduce potentil resulted in higher weight gin, corresponding lso to vritions. Our dt indicted tht the cecl digest digestiility similr to tht of high strch diet. This content rnged from 4 to 6% of the live weight. confirmed the previous study of Gidenne nd Jehl Similr vlues were reported for seven-week-old (996) reporting the high vilility for the growing rits in the review of Gidenne (997), thus rit of mix of hemicellulose nd pectins in diets vlidting this procedure. with higher content of lignocellulose. The growing Very few studies hve ssessed in-vivo cecl rit compensted the replcement of strch y fermenttive ctivity (CFA) over complete growth higher fire digestion (over 40% for hemicellulosic period, ecuse of difficulties to otin individul frction) comined with higher fire intke, thus consecutive mesurements over severl weeks. Dur- leding to n incresed quntity of fire degrded ing the growth, the fermenttion pttern evolved to nd to high cecl fermenttive ctivity. In conpttern similr to the dult, corresponding minly to trst, the hemicellulose rich diet ws of low di liner increse in utyrte nd decrese in cette gestiility, even for the fire frctions, ut it ws not molr proportion, s previously descried (Bellier et ssocited with reduction of the cecl SCFA l., 995; Gidenne et l., 998). Our pproch ws levels. Although there ws no significnt individul lso of interest to control tht dietry effects re- reltionship etween the min nutrients intke nd mined constnt ccording to ge. We confirmed our the cecl fermenttion ctivity, we would point out previous results (Gidenne et l., 998), indicting tht the men SCFA cecl concentrtion incresed ny significnt interction etween ge nd diet, ccording to the men intke of digestile fire from 6 to 0 wk of ge. During this period, the (hemicellulose pectins). In prllel, the reduction cecl SCFA concentrtion incresed linerly y of the ceco colic retention time did not ffect the 0% (reching 65 mm/ l). A similr increse ws fermenttion processes. Contrry to ruminnts, milso oserved y Gidenne et l. (998) with diets croil ctivity nd fire degrdtion in the rit hving similr NDF/ strch rtio (rnging from 0.9 seemed not lrgely ffected y the ceco colic rte to.). With diets hving higher NDF/strch rtio of pssge, ut would e minly controlled y the (over.3) Jehl nd Gidenne (996) did not mention nture of fire. The cecl retention time is too short such rise in cecl SCFAs with ge, proly to llow cteril degrdtion of cell wll tht re ecuse the SCFA concentrtions (over 65 mm/ l) not esily ccessile (such s cellulose), ut is lredy levelled off t six weeks of ge. For instnce, sufficiently long to llow sustntil degrdtion of erlier stilistion of cecl SCFA level (t five the more esily digestile fire frctions (e.g., pectins weeks old) ws oserved y Cndu et l. (978) or even hemicellulose). with diet rich in digestile fire (such s hemicel- Studies on digestive trnsit recover for the rit lulose or pectins) nd with low strch content. n dditionl interest in terms of preventing excessive retention time tht fvours dysfunction or 4.. Replcement of redily ville ltertion of the cecl microil ctivity nd thus crohydrtes y digestile fire digestive troules (Gidenne, 997). Compred to less digestile fire frction, pectins re reputed to slower The replcement of ville crohydrtes (es- the trnsit in monogstric (Hillmnn et l., 983; sentilly strch) y fire frctions of high digestiili- Potkins et l., 99). In rit diets, pectins re often ty (such s pectins or hemicellulose), without supplied through incorportion of eet pulp, tht re chnges in the supply (level nd nture) of lignocel- lso suspected to slower trnsit (predisposing to lulose (low digested), led to similr increse development of pthogenic flor), since Auvergne ( 0%) in the voluntry feed intke either for low et l. (987) oserved tht lucerne replcement with (diet HC) or for high proportion of pectins (diet eet pulp resulted in slower oro ilel rte of pssge

11 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) of liquids. Inversely, our results pointed out tht crohydrtes, ut without chnges in lignocellulose incorportion of 0% of eet pulp did not modify the content, fvoured cecl microil ctivity without trnsit of liquids either etween proximl or distl slowing the digestive trnsit. This ppered prprt of the trct. This grees with the study of Frg ticulrly eneficil to the rit growth during the et l. (99) reporting tht eet pulp, in sustitution postwening period. to lucerne, did not lter the trnsit of liquids in the whole trct. Compred to strchy diet, the rte of pssge of prticles ws similrly stimulted with hemicellulose or with mix of hemicellulose nd Acknowledgements pectins. The effect ws logiclly pronounced in the The uthors thnk Andre Lpnouse nd Muriel ceco colic segment responsile for the two-thirds Segur for their technicl ssistnce (Sttion de of the totl retention time, nd it ws relted with Recherches Cunicoles, INRA, Toulouse, Frnce). NDF intke. However, Gidenne nd Jehl (996) did This study ws prtly supported y fellowship not mention stimultion of the ceco colic trnsit grnt from Rlston Purin Frnce (Sint Quentin En for rits hving high fire intke (. 30 g NDF/ Yvelines, Frnce). d), proly ecuse ceco colic trnsit ws lredy t miniml level (men 7 h). Animls fed pectin-rich diet presented high cecl volume, s lredy descried for rit pig or rt References (Grci et l., 993; Kesting et l., 990; Brunsgrd Auvergne, A., Bouyssou, T., Piret, M., Bouillier-Oudot, M., et l., 995). This ws here ssocited to high level Ruckeusch, Y., Cndu, M., 987. Nture de l liment, finesse of SCFA, therefore incresing the cecl SCFA pool. de mouture et donnees ntomo fonctionnelles du tue This increment in cecl fermenttive ctivity would digestif proximl. Reprod. Nutr. Devel. 7, contriute to explin tht pectin-rich feeds were s Bellier, R., 994. Controle ˆ nutritionnel de l ctivite fermentire cecle. These ` efficiently utilised s strchy feed for rit growth de Doctort, Institut Ntionl Polytechnique, Ecole Ntionle Superieure d Agronomie de Toulouse, (similr feed conversion on the overll growth Toulouse. period). This effect ws prticulrly pronounced Bellier, R., Gidenne, T., 996. Consequences of reduced fire during the postwening period when young rit intke on digestion, rte of pssge nd cecl microil hd higher growth rte with feeds contining mix ctivity in the young rit. Br. J. Nutr. 75, of hemicellulose nd pectin. Such fvourle effect Bellier, R., Gidenne, T., Verny, M., Colin, M., 995. In vivo study of circdin vritions of the cecl fermenttion pttern of strch replcement y digestile fire for the in the post-wened nd dult rits. J. Anim. Sci. 73, 8 postwened rit ws lso reported y Jehl nd 35. Gidenne (996) nd ws ssocited with etter Bjornhg, G., 98. The retrogrde trnsport of fluid in the helth sttus in reltion with stimultion of the proximl colon of rits. Swed. J. Agric. Res., Bls, E., Fernndez Crmon, J., Cerver, C., 988. Effect of cecl microil ctivity (increse of the cecl digestive ctivity nd strch intke on mylse ctivity nd iomss nd SCFA level). In contrst, the supply of pncretic juice of rit. In: Proceedings of the 4th Congress hemicellulose (HC diet) with feed contining of the World Rit Science Assocition, WRSA, Budpest, pp. ingredients of lmost one origin (the whet) gve similr growth nd cecl fermenttive ctivity thn Bls, E., Cerver, C., Fernndez Crmon, J., 994. Effect of two strchy diet. The lck of diversity in the otnicl diets with vried strch nd fire levels on the performnces of 4 7 weeks old rits. W. Rit Sci., 7. origin of fire might e unfvourle to the CMA, s Blumenkrntz, N., Asoe-Hnsen, G., 973. New method for lredy oserved recently (Gidenne et l., 998). quntittive determintion of uronic cids. Anl. Biochem. 54, In conclusion, the dietry lnce etween low nd highly digestile fire needs to e determined Brunsgrd, G., Eggum, B.O., Sndstrom, B., 995. Gstrointesti- with precision for the growing rit, to fvour nl growth in rts s influenced y indigestile polyscchrides nd dpttion period. Comp. Bioch. Phys. A, cecl digestion without dverse effects for the Cndu, M., Delpon, G., Fiormonti, J., 978. Influence de l growth. The dietry incorportion of digestile nture des glucides memrnires sur le developpement cell wll constituents in replcement to ville ntomofonctionnel du trctus digestif du lpin. In: Proceed-

12 5 T. Gidenne, R. Bellier / Livestock Production Science 63 (000) 4 5 ings of the nde Journees de l Recherche Cunicole, INRA, effects of pectin, cellulose nd lignin on stool PH, trnsit-time Toulouse, pp...4. nd weight. Br. J. Nutr. 50, Crre, B., Brillouet, J.M., 989. Determintion of wter insolule IO7, 993. L Bnque de Donnees de l Alimenttion Animle, cell-wlls in feeds: interlortory study. J. Assoc. Off. Agric. V.5, Assocition Frnçise de Zootechnie, Pris. Chem. 7, Jehl, N., Gidenne, T., 996. Replcement of strch y digestile Crre, B., 99. Fctors ffecting the digestiility of non-strch fire in the feed for the growing rit.. Consequences for crohydrtes in monogstric nimls. Poultry Sci. 6, 57 microil ctivity in the cecum nd on incidence of digestive 69. disorders. Anim. Feed Sci. Technol. 6, Edwrds, C.A., Prrett, A.M., 996. Plnt cell wll polysc- Kesting, U., Schnel, E., Boldun, G., 990. Degree nd effects chrides, gums nd hydrocolloids: nutritionl spects. In: of fermenttions in the hind gut of the pig.. Ilel flow-rtes, Elisson, A.-C. (Ed.), Crohydrtes in Food, pp fermentility nd colon cpcity. Arch. Anim. Nutr. 0, 933 Frg, M.J., Perez De Ayl, P., Crno, R., 99. Effect of 94. type of fier on the rte of pssge nd on the contriution of Leng, E., Hornicke, H., 976. Diurnl vritions in the rits soft feces to the nutrient intke of finishing rits. J. Anim. cecl volume in vivo with C4-PEG. Z. Vet. Med. 3, Sci. 69, Leng, E., Cluss, W., Hornicke, H., 977. Colon pssge time in Grci, G., Glvez, J.F., De Bls, J.C., 993. Effect of sustitution rits in reltion to the formtion of cecotrophes. Z. Vet. of sugreet pulp for rley in diets for finishing rits on Med. 4, growth performnce nd on energy nd nitrogen efficiency. J. Luick, B.R., Ayers, A., Cheeke, P.R., 99. Rte of pssge of Anim. Sci. 7, Blck Locust lef nd lflf mel in the rit gut. J. Appl. Gidenne, T., Bellier, R., 99. Etude in vivo de l ctivite Rit Res. 5, 94 9, Proceedings of the 5th congress of fermentire cecle chez le lpin. Mise u point et vlidtion the World Rit Science Assocition, 5 30 July, Corvllis, d une nouvelle technique de cnultion cecle. Reprod. Nutr. OR. Dev. 3, Mıtre, ˆ I., Les, F., Arveux, P., Bourdillon, A., Duperry, J., Gidenne, T., 994. Effets d une reduction de l teneur en fires Sint Cst, Y., 990. Tux de lignocellulose, ADF de Vnlimentires sur le trnsit digestif du lpin. Comprison et Soest, et performnces de croissnce du lpin de chir. In: vlidtion de modeles ` d justement des cinetiques d excretion Proceedings of the 5th Journees de l Recherche Cunicole, Vol. fecle des mrqueurs. Reprod. Nutr. Dev. 34, , ITAVI, Pris, pp Gidenne, T., Jehl, N., 996. Replcement of strch y digestile Mthers, J.C., 99. Digestion of non-strch polyscchrides y fire in the feed for the growing rit.. Consequences for non-ruminnt omnivores. Proc. Nutr. Soc. 50, 6 7. digestiility nd rte of pssge. Anim. Feed Sci. Technol. 6, Potkins, Z.V., Lwrence, T.L.J., Thomlinson, J.R., 99. Effects of structurl nd non-structurl polyscchrides in the diet of the Gidenne, T., Perez, J.M., 996. Dietry cellulose in the growing growing pig on gstric emptying rte of pssge of digest to rit.. Consequences on digestion nd rte of pssge. Ann. the terminl ileum nd through the totl gstrointestinl trct. Zootech. 45, Br. J. Nutr. 65, Gidenne, T., 997. Ceco colic digestion in the growing rit: Sttisticl Anlysis System, 988. SAS/ STAT Guide For Personl impct of nutritionl fctors nd relted disturnces. Livest. Computers, Version 6.03, SAS Institute, Cry, NC. Prod. Sci. 5, Vn Soest, P.J., Roertson, J.B., Lewis, B.A., 99. Methods for Gidenne, T., Crno, R., Grci, J., De Bls, C., 998. Fire dietry fier, neutrl detergent fier, nd non strch polyscdigestion in the rit. In: De Bls, C., Wisemn, J. (Eds.), The chrides in reltion to niml nutrition. J. Diry Sci. 74, Nutrition of the Rit, CABI Pulishing, pp , Ch Gidenne, T., Bellier, R., Vn Eys, J., 998. Effect of the dietry Verdow, H., Vn Echteld, C.J.A., Dekkers, E.M.J., 977. Ammoni fire origin on the digestion nd on the cecl fermenttion determintion sed on indophenol formtion with sodium pttern of the growing rit. Anim. Sci. 66, slicylte. Wter Res., Hillmnn, L., Peters, S., Fisher, A., Pomre, E.W., 983. Differing