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Apparent digestibility (ileal and overall) of nutrients as

evaluated with PVTC-cannulated or ileo-rectal anastomised pigs

fed diets containing two indigestible markers

a ,1 c d e e

Y.-L. Yin

, J.D.G. McEvoy , H. Schulze , U. Hennig , W.-B. Souffrant ,

a,b ,

_*

K.J. McCracken

Department of Agricultural and Environmental Science, The Queen’s University of Belfast, Newforge Lane, Belfast BT9 5PX, Northern Ireland, UK

Department of Agriculture for Northern Ireland, Newforge Lane, Belfast BT9 5PX, UK

Veterinary Sciences, Department of Agriculture for Northern Ireland, Stoney Road, Belfast BT4 3SD, Northern Ireland, UK

Finnfeeds International Ltd., Marlborough, Wiltshire SN8 1AA, UK

¨ ¨

Forschungsbereich Ernahrungsphysiologie‘‘Oskar Kellner’’, Forschungsinstitut f ur die Biologie Landwirtschaftlicher Nutztiere, Rostock, Germany

Received 28 July 1998; received in revised form 3 February 1999; accepted 15 June 1999

Abstract

The techniques of post-valve ‘‘T’’ caecal cannulation (PVTC) and ileo-rectal anastomosis (IRA) and the indigestible markers Cr O and TiO for determination of ileal digestibility of nutrients were studied in this experiment. The results2 3 2

showed that the marker Cr O₂ ₃ gave consistently lower ileal recovery and ileal apparent digestibility of nutrients. The average value of ileal recovery of Cr O measured with the IRA method was only 87%. A nearly 100% of ileal recovery of2 3

TiO measured with the IRA method suggests that the use of TiO as a marker may be a better choice for determination of₂ ₂ ileal digestibility of nutrients when partial-sampling ileal digesta. The reason for the lower ileal recovery of Cr O may be2 3

due to the chemical analytical problems of Cr in excreta. The low recoveries of markers (TiO 67.8% and Cr O 60.6%) and₂ ₂ ₃ the high values for ileal digestibility of nutrients in the high fibre content diet based on wheat bran (WB) measured by the PVTC total collection method indicated that a quantitative collection of digesta had not been achieved and hence that it is necessary to use a marker for the PVTC method. However, the results from the present experiment showed that there is a high ileal recovery of TiO (95–100%) for the low and medium fibre diets based on wheat (W), recombined wheat (flour₂ plus bran plus middlings, WR) and wheat middlings (WM) measured by the PVTC method. The ileal digestibility of nutrients measured by PVTC (TiO ) agreed well with the IRA method for all of the for measuring the ileal digestibility of₂ nutrients in pigs.  2000 Elsevier Science B.V. All rights reserved.

Keywords: Ileal digestibility; Indigestible markers; PVTC-cannulation; Ileo-rectal anastomised pigs

*Corresponding author. Corresponding address: Agricultural and Environmental Science Division, Agriculture and Food Science Centre, Newforge Lane, Belfast BT9 5PX, Northern Ireland, UK. Tel.: 144-1232-255-368; fax: 144-1232-662-007.

E-mail address: [email protected] (K.J. McCracken)

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1. Introduction opposite the ileo-caecal valve. Once the cannula is opened the valve protrudes into the cannula and Since the 1970s, many studies have been per- digesta can be sampled as it leaves the ileum. Recent formed to estimate ileal digestibility using different studies showed that PVTC has a number of advan-methods for digesta collection. These have been tages over the simple ‘‘T’’cannulation methods (van

reviewed by Sauer and Ozimek (1986), Fuller (1991) Leeuwen et al., 1991). Kohler et al. (1991) compared and Yin and McCracken (1996). The methods three different cannulation techniques and reported frequently applied for quantitative collection are re- higher ileal Cr O recovery and digestibility values2 3 entrant cannulation, end-to-end ileorectal anas- with the re-entrant techniques than with the PVTC or tomosis (IRA) or the steered ileo-caecal valve simple ileal ‘‘T’’ cannula. However, with all three (SICV) technique (Mroz et al., 1996). These meth- methods Cr O recovery was less than 100%. The2 3 ods are generally regarded as more accurate for authors concluded that the results showed less vari-measuring ileal digestibility of nutrients, since they ation with the PVTC cannula compared with the involve the total collection of ileal digesta and it is simple ileal ‘‘T’’ cannula. Therefore on this basis, unnecessary to use a marker, such as chromic oxide. and in view of the simplicity of handling, the PVTC However, the re-entrant cannulation technique poses cannula appears to be a good alternative to the other problems in that it interrupts the transport of the methods. The results from Yin et al. (1999) indicated normal migrating myoeletric complex which is that the post-valvular ‘‘T’’-caecum cannula method necessary for normal digesta passage (Laplace, 1980; is a more precise and less labour intensive technique Low, 1982; Sauer and Ozimek, 1986; Fuller, 1991). for ileal digestibility studies than the simple ileal

This method is also generally associated with block- ‘‘T’’ cannula. Furthermore Kohler et al. (1991) and age of the cannula by coarsely ground or fibrous Yin et al. (2000) suggested that the PVTC method is diets which prevents its use with many feed com- almost a quantitative collection method. However ponents. With the IRA method, normal functioning this aspect requires further investigation.

of the large intestine is obviously not maintained and The low recovery of markers in faeces reported by this may have an influence on the water and mineral Mueller (1956), Moore (1957, 1959), Ishikawa

absorption and on growth (Kohler et al., 1991). (1966), Ishikawa and Sugimura (1973), Moughan et However, Hennig et al. (1986, 1992) and Yin et al. al. (1991), Greer (1992) and McClean (1993) is a (1993) reported that the loss of the function of the major cause for concern. Barnicoat (1945), Moore large intestine can be compensated by supplying the (1957, 1959), McClean (1993) and Yin et al. (2000) animal with large amounts of water, minerals and suggested that this may be due to retention and / or vitamins and Laplace et al. (1994) showed that the absorption in the large intestine.

end-to end procedure was preferable to the end-to- Although chromic oxide is frequently used as a ¨

side operation used by Kohler et al. (1991). There is marker in pig digestibility studies, some authors have ample evidence, therefore, that the IRA method can found that recoveries are variable, less than 100% be used for measurement of ileal digestibility of dry and that this material may have carcinogenic prop-matter, crude protein, energy and amino acids no erties (Mueller, 1956; Moore, 1957; Peddie et al., simultaneous information on faecal digestibility is 1982; Jagger et al., 1992; Monaghan, 1996). Data obtained. The SICV method appears to have some from ileal digestibility studies have indicated that advantages but has not yet been widely tested and no chromic oxide may move out of phase with digesta,

direct comparisons with the IRA method are avail- particularly when fibrous diets are fed (Kohler et al., able since the use of the latter is prohibited in the 1990). The problems encountered with Cr O led to2 3

Netherlands. its replacement by some workers with other markers,

Methods frequently applied for partial-sampling e.g., titanium dioxide and acid-insoluble ash (Jagger involving the use of a marker include the simple ileal et al., 1992; McCarthy et al., 1974; Moughan et al., ‘‘T’’ and post-valve ‘‘T’’ caecal cannulae (PVTC). 1991; Njaa, 1961; van Leeuwen et al., 1996). Unlike the simple ileal ‘‘T’’ cannula method, with Titanium dioxide has been used successfully as an the PVTC technique the caecum is partially removed indicator in digestibility studies with rats, chickens and replaced by the PVTC cannula, which is placed and pigs (Njaa, 1961; Peddie et al., 1982; Jagger et

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al., 1992). However, there is little information about crude protein (CP), energy and amino acids (AAs) the use of TiO as a marker for determination of the2 were calculated for the two experiments. Ileal re-ileal digestibility of nutrients in pigs. covery of TiO2 and Cr O2 3 and IAD of DM, CP, The present studies afforded the opportunity to (1) gross energy and amino acids measured with TiO2 compare the use of Cr O and TiO as indigestible2 3 2 and Cr O2 3 were compared by Student’s t-test. In markers, (2) compare the PVTC with the IRA order to compare three methods (PVTC TiO2 method, (3) evaluate the PVTC method for quantita- marker, PVTC Cr O marker and PVTC total collec-2 3 tive collection of digesta, using wheat and its by- tion), the 144 data values corresponding to 12 pigs3 product-based diets with a range of fibre content. 4 periods33 methods were analysed using a split plot model with diets as the main plot factor and methods as the sub-plot factors. The analysis was

2. Materials and methods carried out using the REML procedure of the Genstat

5 Committee (1993). Because of the differences in Two separate studies were conducted. In experi- location, pig breed and statistical design no statistical ment 1, 12 Large White Landrace boars, approxi- comparison of the PVTC and IRA techniques was mately 26 kg, were fitted with post-valve ‘‘T’’ caecal considered valid.

cannulae (PVTC) according to the procedures of van Leeuwen et al. (1991). The pigs were randomly allocated, in a four-period crossover design to eight

3. Results

diets based on wheat (W), wheat plus middlings (WM), wheat plus bran (WB) or recombined wheat

3.1. Ileal recoveries of TiO and Cr O2 2 3 (flour plus bran plus middlings, WR) without or with

xylanase addition. Each of the four main diets was

The ileal TiO2 and Cr O2 3 recoveries for the mixed as a single batch and then divided for enzyme

experimental diets W, WR,WM and WB and the addition. The diets, which contained both TiO and2 _{average values are given in Table 1. With both} Cr O as indigestible markers, were cold-pelleted (32 3 _{markers the recoveries measured with the PVTC} mm die) and fed dry. This experiment was conducted

method for diet WB were significantly (P,0.001) at The Queen’s University of Belfast. After a 5-d

lower than for the other diets, values for TiO and2 adaptation period, faeces were collected for 7 d and

Cr O being, respectively, 31 and 24 units less than2 3 ileal digesta were collected for 12 h (08:00–20:00 h)

those for diet W. The ileal recovery of TiO for all of2 on the next two days.

the tested diets measured with the IRA method and In experiment 2, 30 German Landrace barrows,

for diets W, WR and WM measured with the PVTC approximately 15 kg, were subjected to an ileo-rectal

method were close to 100%, whereas recoveries of anastomosis by the procedure described by Hennig et

Cr O were 11 units lower (P2 3 ,0.001), for both the al. (1986). The experiment was conducted at

Ros-PVTC and IRA methods. tock. The six diets, based on W, WM and WB, of

As a consequence of the lower Cr O2 3 recovery, identical composition to those used in Belfast, were

the average values for IAD of DM, CP, energy and fed to five pigs per diet. Ileal digesta were collected

AAs in the four diets measured with TiO2 were for 5 d after a 7-d adaptation period. Samples of ileal

significantly higher than with Cr O (Table 2).2 3 digesta were pooled within period and pig for each

dietary treatment and freeze–dried. TiO was mea-2

sured according to Leone (1973). Cr O was mea-2 3 3.2. Ileal apparent digestibility of nutrients sured using the method of Saha and Gilbreath (1991) measured with markers (PVTC) and total by atomic absorption spectrophotometry. Further collection (PVTC and IRA) methods details of the diet compositions and methods of

analysis are given in a previous paper (Yin et al., As illustrated in Table 3, apart from CP

diges-2000). tibility (P,0.05), all diet3method interactions for

Ileal recoveries of TiO2 and Cr O , and ileal2 3 DM, CP and energy were highly significant (P, apparent digestibility (IAD) of dry matter (DM), 0.001). Apart from the numerically lower DM

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di-Table 1

Ileal recovery (%) of TiO and Cr O2 2 3

Diet W WR WM WB S.E.D. P Mean

a a a b

PVTC TiO2 99.9 94.9 95.5 67.8 4.92 ,0.001 89.7

a a a b

Cr O2 3 84.9 87.4 82.8 60.6 4.69 ,0.001 79.0

IRA TiO2 98.8 ND 101.1 95.2 2.70 NS 98.4

Cr O2 3 84.3 ND 88.7 86.8 2.39 NS 86.8

PVTC: Measured with PVTC total collection method. IRA: Measured with IRA method.

a,b

Values in the same row without a common superscript are significantly different (P,0.05). ND: Not determined.

Table 2 _{(TiO ) and IRA methods for W, WR and WM but the}

Comparison of average values of ileal apparent digestibility of _{high fibre WB diet resulted in higher estimates of} nutrients in diets W, WR, WM and WB measured with PVTC

apparent digestibility by the total collection method. cannula and either TiO or Cr O2 2 3

TiO2 Cr O2 3 S.E.D. P

DM 0.702 0.671 0.0095 ,0.001 _{4. Discussion}

CP 0.772 0.748 0.0081 ,0.001

Energy 0.710 0.680 0.0084 ,0.001

4.1. Comparison of Cr O and TiO as markers2 3 2 Threonine 0.747 0.72 0.0092 ,0.001

for determination of ileal apparent digestibility of

Valine 0.748 0.722 0.0074 ,0.001

Cystine 0.785 0.760 0.0083 ,0.001 nutrients

Methionine 0.888 0.880 0.0039 ,0.05

Isoleucine 0.810 0.790 0.0042 ,0.001 _{Low faecal recoveries of Cr O and TiO (ranging} 2 3 2

Leucine 0.820 0.801 0.0052 ,0.001

from 71 to 85%) in pigs have been often reported in Tyrosine 0.831 0.814 0.0048 ,0.001

the literature (Mueller, 1956; Moore, 1957, 1959; Phenylalanine 0.923 0.812 0.0199 ,0.001

Lysine 0.819 0.800 0.0054 ,0.001 Ishikawa, 1966; Ishikawa and Sugimura, 1973; Histidine 0.850 0.834 0.0055 ,0.001 _{Moughan et al., 1991; Greer, 1992; McClean, 1993;} Arginine 0.883 0.870 0.0042 ,0.001 _{Yin et al., 2000). The present results show that the} Aspartic acid 0.756 0.730 0.0091 ,0.001

ileal recoveries of TiO and Cr O are much higher2 2 3 Serine 0.812 0.792 0.0073 ,0.001

than those at the faecal level. For example, the ileal Glutamic acid 0.892 0.881 0.0036 ,0.001

Proline 0.787 0.763 0.0061 ,0.001 recoveries of TiO2 and Cr O2 3, measured with the Glycine 0.721 0.690 0.0086 ,0.001 _{IRA method, were 12.6 and 5.3 percentage units} Alanine 0.719 0.688 0.0054 ,0.001 _{higher, respectively, than the overall (faecal)} re-Total AAs 0.817 0.798 0.0061 ,0.001

coveries of TiO and Cr O observed by Yin et al.2 2 3 (2000). The nearly 100% ileal recovery of TiO for2 gestibility for diet WM the results for PVTC using the low fibre diets W, WR and the medium fibre diet TiO2 and for IRA were very similar, and, in line WM obtained with the PVTC and IRA pigs indicates with the expected effects of fibre, values declined that the disappearance of TiO2 observed by Greer from diet W to diet WB. Similar to the results for the (1992); McClean (1993) and Yin et al. (2000) did digestibility of DM, CP and energy, ileal apparent not occur at the ileal level. This suggests that the digestibilities of amino acids were also comparable lower recovery of marker in faeces is due to marker between the PVTC (TiO ) and IRA techniques,2 retention, or absorption or both in the large intestine. except for valine, phenylalanine and proline. Values The reasons for retention and absorption in the large for the DM, CP, gross energy and AA with PVTC intestine of pigs may be a combination of the long using Cr O were lower than for the other methods.2 3 retention time (36 h) of digesta (Kidder and Man-PVTC by total collection agreed well with the Man-PVTC ners, 1978), the complex and capacious structure of

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Table 3

Comparison of ileal apparent digestibility of DM, CP, gross energy and selected AAs measured either with markers (TiO or Cr O ) using2 2 3

PVTC-cannula or by total collection (PVTC or IRA)

Diet TiO2 Cr O2 3 PVTC S.E.D. Diet3method IRA

DM W 0.773 0.741 0.772 0.763

WR 0.766 0.752 0.783 ND

WM 0.673 0.632 0.688 0.0155 P,0.001 0.624

WB 0.594 0.556 0.732 0.597

CP W 0.807 0.779 0.807 0.811

WR 0.804 0.790 0.808 ND

WM 0.757 0.727 0.769 0.0311 P,0.05 0.747

WB 0.720 0.693 0.808 0.745

Gross energy W 0.787 0.757 0.777 0.784

WR 0.771 0.759 0.792 ND

WM 0.683 0.644 0.698 0.0142 P,0.001 0.656

WB 0.596 0.558 0.724 0.601

Threonine W 0.77 0.737 0.768 0.764

WR 0.767 0.753 0.785 ND

WM 0.743 0.71 0.756 0.0128 P,0.001 0.732

WB 0.708 0.68 0.794 0.702

Valine W 0.766 0.733 0.764 0.805

WR 0.777 0.763 0.793 ND

WM 0.749 0.717 0.761 0.0123 P,0.001 0.769

WB 0.701 0.673 0.797 0.756

Isoleucine W 0.831 0.808 0.83 0.843

WR 0.837 0.826 0.848 ND

WM 0.803 0.778 0.813 0.0093 P,0.001 0.815

WB 0.768 0.746 0.841 0.8

Leucine W 0.849 0.827 0.849 0.851

WR 0.841 0.832 0.853 ND

WM 0.807 0.783 0.817 0.0121 P,0.001 0.82

WB 0.78 0.76 0.85 0.807

Phenylalanine W 0.935 0.83 0.85 0.876

WR 0.935 0.846 0.865 ND

WM 0.918 0.797 0.828 0.0068 P,0.001 0.851

WB 0.904 0.777 0.861 0.834

Lysine W 0.843 0.811 0.834 0.844

WR 0.832 0.822 0.845 ND

WM 0.82 0.797 0.829 0.0093 P,0.001 0.833

WB 0.786 0.767 0.855 0.807

Histidine W 0.865 0.845 0.864 0.869

WR 0.863 0.855 0.872 ND

WM 0.847 0.827 0.854 0.0075 P,0.001 0.865

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Table 3. Continued

Diet TiO2 Cr O2 3 PVTC S.E.D. Diet3method IRA

Tyrosine W 0.849 0.828 0.849 0.851

WR 0.853 0.844 0.863 ND

WM 0.833 0.813 0.841 0.0082 P,0.001 0.858

WB 0.789 0.771 0.856 0.834

Arginine W 0.887 0.871 0.887 0.908

WR 0.882 0.875 0.89 ND

WM 0.886 0.871 0.89 0.0062 P,0.001 0.901

WB 0.876 0.865 0.917 0.895

Aspartic acid W 0.777 0.746 0.776 0.799

WR 0.779 0.765 0.796 ND

WM 0.751 0.72 0.764 0.012 P,0.001 0.769

WB 0.716 0.689 0.807 0.75

Serine W 0.834 0.81 0.833 0.836

WR 0.837 0.827 0.849 ND

WM 0.8 0.775 0.81 0.009 P,0.001 0.801

WB 0.779 0.757 0.85 0.786

Glutamic acids W 0.909 0.897 0.908 0.912

WR 0.908 0.902 0.915 ND

WM 0.879 0.863 0.885 0.0066 P,0.001 0.873

WB 0.873 0.861 0.913 0.879

Proline W 0.768 0.731 0.83 0.891

WR 0.881 0.874 0.732 ND

WM 0.763 0.733 0.763 0.0247 P,0.001 0.858

WB 0.709 0.683 0.77 0.854

Glycine W 0.749 0.712 0.745 0.784

WR 0.742 0.726 0.761 ND

WM 0.717 0.679 0.73 0.0157 P,0.001 0.74

WB 0.675 0.644 0.782 0.732

Alanine W 0.737 0.7 0.735 0.77

WR 0.75 0.734 0.769 ND

WM 0.719 0.683 0.732 0.0144 P,0.001 0.741

WB 0.666 0.635 0.773 0.714

Total AAs W 0.833 0.81 0.832 0.856

WR 0.851 0.846 0.856 ND

WM 0.808 0.784 0.817 0.0091 P,0.001 0.825

WB 0.783 0.764 0.854 0.814

IRA not included in statistical analysis, since the experimental design was different from the PVTC experiment. ND: Not determined.

the large intestine (Barnicoat, 1945; Moore, 1957; and Monaghan (1996) the present results highlight 1959) and the high specific gravity of TiO2 and the problems of use of Cr O2 3 as a marker. Mean Cr O (Lide and Frederikse, 1995).2 3 values of the ileal recovery of Cr O measured with2 3 In common with the studies of McCarthy et al. the IRA method was only 87% in this study. (1974), Moughan et al. (1991), Jagger et al. (1992) Consequently the ileal digestibility of DM, CP,

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energy and total AA measured with Cr O2 3 were Cr O2 3, TiO2 is insoluble in water, HCl, HNO3 or respectively 4.6, 3.2, 4.4 and 2.4% lower than those dilute H SO (Merck Index, 1989) and analysis of2 4 measured with TiO (Table 2). Jagger et al. (1992)2 TiO2 is a relatively simple and straightforward found that the ileal digestibilities of N and AAs procedure. High recoveries (95–100%) in samples of measured with Cr O2 3 were significantly lower (N, feeds and excreta of poultry and pigs have been 6.7% and the average value of 11 AAs, 6.1%) than reported (Njaa, 1961; Leone, 1973; Peddie et al., those measured with TiO , when both markers were2 1982; Jagger et al., 1992).

added to a barley and wheat based diet fed to

growing and finishing pigs. Furthermore, the results 4.2. Comparison of methods for measurement of of Jagger et al. (1992) and of this experiment show ileal apparent digestibility of nutrients

that the standard errors associated with apparent ileal

digestibility values calculated using each marker The excellent marker recoveries with TiO for the2 were lower with TiO2 than with Cr O2 3. This is W, WR and WM with the PVTC method and for the somewhat surprising in that the level of incorpora- W, WM and WB diets with the IRA method indicate tion of Cr O was higher than for TiO2 3 2.At these low that, under these conditions, digesta collection was levels it would be possible for problems to arise due fully representative of the material reaching the to lack of homogeneity of mixing. However, the low terminal ileum. This would be expected with the IRA standard errors obtained with TiO2 would suggest method where a complete 5-d collection was made that diet mixing was not a problem. but it is noteworthy that similar results occurred with The primary reason for the low recovery and the much shorter period of collection (24 h) using the variable results from Cr O2 3 may be related to PVTC technique. However, the result for the high-analytical problems arising mainly from interference fibre diet WB using the PVTC technique demon-by other minerals present in the sample (Williams et strates that quantitative collection of digesta was not al., 1962; Saha and Gilbreath, 1991). For example, achieved. This result therefore supports the reports of

Williams et al. (1962) found a 12% decrease in Cr Kohler et al. (1991) and van Leeuwen et al. (1991) recovery when phosphate (PO ) was added to the4 of variable recovery with the PVTC technique solution of Cr. Saha and Gilbreath (1991) also dependant on diet composition and confirms the showed significant effects on recovery when varying necessity of using an appropriate indigestible marker. the concentration of Ca, P or Mg in diets and faeces Although a direct statistical comparison of the of pigs. These minerals occur in high concentrations PVTC and IRA techniques is not valid due to the in the digesta or faeces of pigs. This possibility is separate locations in which the techniques were used presently under investigation in our laboratory. In an and the different sources of pigs, the results given in initial study, three levels of Cr O corresponding to2 3 Table 3 for PVTC (TiO ) and IRA show close2 5, 10, 20 ppm were added to the faeces of pigs which numerical agreement. This, taken with the excellent had not received Cr O in the diet. The recoveries2 3 marker recoveries (Table 1), suggests that both were 88.0, 87.7, 83.3%, respectively. This is similar methods were giving reliable estimates of ileal to the mean recovery values reported by Saha and digestibility. Furthermore, the good agreement be-Gilbreath (1991) and the mean ileal recovery of tween IRA and PVTC (TiO ) for diet WB, where2 Cr O measured in this study.2 3 PVTC collection was not quantitative, suggests that In addition, although Cr O is insoluble in water,2 3 the digesta recovered was similar in composition to it is slightly soluble in acids and alkalis (Merck that by-passing the cannula.

Index, 1989) and therefore it might be soluble to a As discussed in the Introduction, the main advan-certain extent in the acid or alkaline digestive juices tage of the end-to-end IRA technique is that it is a of the gastrointestinal tract of pigs (Moore, 1959) or quantitative method for collecting ileal digesta and, the acid solution used during Cr analysis. Saha and therefore, it is not necessary to use any marker for Gilbreath (1991) suggested that the soluble Cr might calculating digestibility. However, compared with be lost due to the formation of volatile Cr com- the PVTC method, the main disadvantages are: (1) pounds at the sample digestion stage. Compared with relatively complex surgical procedures and

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post-beans in diets for growing pigs. M.Sc. Thesis. The Queen’s operative animal care. (2) Longer time for recovery

University of Belfast, Belfast. from the surgery. (3) Need for greater care to

Hennig, U., Noel, R., Herrmann, U., Wunsche, J., Mehnert, E., minimise skin irritation and discomfort due to the _{1986. Nutrition–physiologic studies in pigs with ileo-rectal} outpouring of digesta around the hind legs. (4) The anastomoses. 1. Operation methods, biochemical and

mor-phological findings. Arch. Anim. Nutr. 36, 585–596. need for additional water, minerals and vitamins to

Hennig, U., Wunsche, J., Souffrant, W.B., Borgmann, E., offset the loss of the large intestine. Compared with

Kreienbring, F., 1992. Methodenverleich zur Bestimmung der the IRA method, the surgery for the PVTC cannula- _{PrSzSkalen NShrstoffverdaulichkeit und AminosSureresorption}_{ˇ ˇ} _ˇ _ˇ tion is much easier to perform and less severe on the _{zwischen Schweinen mit ileorectal-anastomose und ileozSkal-}ˇ

¨ ¨

Bruckenkanule. Arch. Anim. Nutr. 42, 197–211. animals than the IRA procedure and the PVTC

Ishikawa, S., 1966. Reliability of polyethyleneglycol as an in-cannulation technique causes less discomfort to the

dicator for digestion studies with swine. 1. Rate of passage of animal. Furthermore this method permits simulta- _{polyethyleneglycol through the digestive tract. Agric. Biol.} neous measurement of ileal and overall digestibility _{Chem. 30, 278–284.}

in contrast to the IRA method. Taking all of these Ishikawa, S., Sugimura, K., 1973. Movement of polyethylalcohol through the digestive tract as a digestion indicator with swine. aspects into account it is concluded that the PVTC

Agric. Biol. Chem. 37, 203–206. method is a better method for measuring ileal

Jagger, S., Wiseman, J., Cole, D.J.A., Craigon, J., 1992. Evalua-digestibility in most situations. It would, however, be _{tion of inert markers for the determination of ileal and faecal} of considerable interest to make a direct comparison digestibility values in the pig. Br. J. Nutr. 68, 729–739. of the PVTC and SICV techniques. Kidder, D.E., Manners, M.J., 1978. Digestion in the Pig, Kingston

Press, Bath. ¨

Kohler, T., Huisman, J., Den Hartog, L.A., Mosenthin, R., 1990. Comparison of different digesta collection methods to

de-5. Conclusions termine the apparent digestibilities of nutrients at the terminal

ileum in pigs. J. Sci. Food Agric. 53, 465–475. ¨

Kohler, T., Huisman, J., Den Hartog, L.A., Mosenthin, R., The results highlighted problems with poor

re-Verstegen, M.W.A., 1991. Digestibility measurements in pigs covery of Cr O marker resulting in low estimates of2 3 _{using post valve T-caecum cannulation or end to side} ileo-ileal digestibility, while the TiO marker appears to2 _{rectal anastomosis. J. Anim. Physiol. Anim. Nutr. 66, 278–286.} be satisfactory as shown by good agreement with the Laplace, J., 1980. Stomach and small intestine motility in the pig: IRA (total collection) method. PVTC total collection Electromyography in nutritional studies. In: Low, A.G., Par-tridge, I.G. (Eds.), Current Concepts of Digestion and Absorp-proved is satisfactory for measurement of ileal

tion in Pigs. Technical Bulletin 3, p. 24. National Institute for digestibility with low- or medium-fibre diets but not

Research in Dairying, Reading, U.K.

with a high-fibre diet. Considering its simplicity, _{Laplace, J.P., Souffrant, W.B., Hennig, U., Chabeauti, E., Fevrier,} acceptability on animal welfare grounds and the C., 1994. Measurement of precaecal dietary protein and plant good agreement with in measurements of digestibili- cell digestion in pigs; comparison of four surgical procedures

for ileo-rectal anastomosis. Livest. Prod. Sci. 40, 313–328. ty made with the IRA technique, the PVTC

tech-Leone, J.L., 1973. Collaborative study of the quantitative de-nique, in combination with TiO2 as a marker, is _{terminations of titanium dioxide in cheese. J. Assoc. Anal.} considered to be a good method for determination of _{Chem. 56, 535–537.}

ileal digestibility of nutrients for pigs. Lide, D.R., Frederikse, H.P.R., 1995. CRC Handbook of Chemis-try and Physics, CRC Press, Boca Raton, FL.

Low, A.G., 1982. Digestibility and availability of amino acids from feedstuff for pigs. A review. Livest. Prod. Sci. 9, 511–

References _518.

McCarthy, J.F., Aherne, F.X., Okai, D.B., 1974. Use of HCl-Barnicoat, C.R., 1945. Nutrient digestion in animals. NZ J. Sci. insoluble ash as an index for determining apparent digestibility

Technol. 27, 202–205. with pigs. Can. J. Anim. Sci. 54, 107–109.

Fuller, M.F., 1991. Methodologies for the measurement of diges- McClean, D., 1993. Effects of processing of raw materials on tion. In: Verstegen, M.W.A., Huisman, J., den Hartog, L.A. digestibility of diets for weaned pigs. Ph.D. Thesis. The (Eds.), Vth International Symposium on Digestive Physiology Queen’s University of Belfast, Belfast.

in Pigs, Pudoc, Wageningen, pp. 273–288. Merck Index, 11th ed., Budavari, S. (Ed.), Encyclopedia of Genstat 5 Committee, 1993. Genstat 5 Manual, Clarendon Press, Chemicals, Drugs and Biologicals, Merck, Rahway, NJ, 1989. Oxford, Release 3 Reference. Monaghan, C., 1996. Effects of feed enzymes on ileal and faecal Greer, H.E., 1992. Relative nutritive value of soya, peas and field digestibility of selected cereal and protein sources for growing

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pigs. M.Phil. Thesis. The Queen’s University of Belfast, Van Leeuwen, P., van Kleef, D.J., van Kempen, G.J.M., Huisman, Belfast. J., Verstegen, M.W.A., 1991. The postvalve T-caecum cannula-Moore, J.H., 1957. Diurnal variations in the composition of the tion technique in pigs applicated to determine the digestibility faeces of pigs on diets containing chromium oxide. Br. J. Nutr. of amino acids in maize, groundnut and sunflower meal. J.

11, 273–288. Anim. Physiol. Anim. Nutr. 65, 183–193.

Moore, J.H., 1959. The use of indicators in digestibility studies. Van Leeuwen, P., Veldan, A., Boisen, S., Deuring, K., Derksen, Agric. Prog. 34, 48–63. G.B., Verstegen, M.W.A., Schaafsma, G., 1996. Apparent ileal Moughan, P.J., Smith, W.C., Schrama, J., Smits, C., 1991. dry matter and crude protein digestibility of rations fed to pigs Chromic oxide and insoluble ash as faecal markers in di- and determined with the use of chromic oxide (Cr O ) and2 3

gestibility studies with young pigs. NZ J. Agric. Res. 34, acid-insoluble ash as digestive markers. Br. J. Nutr. 76, 551–

85–88. 562.

Mroz, Z., Bakker, G.C.M., Jongbloed, A.M., Dekker, R.A., Williams, C.H., David, D.J., Iismaa, O., 1962. The determination Jongbloed, R., 1996. Apparent digestibility of nutrients in diets of chromic oxide in faeces samples by atomic absorption with different energy density, as estimated by direct and spectrophotometry. J. Agric. Sci. 59, 381–385.

marker methods for pigs with and without ileo-caecal cannulas. Yin, Y.-L., McEvoy, J., Schulze, H., McCracken, K.J., 1999. J. Anim. Sci. 74, 403–412. Studies on effects of cannulation method, and alternative Mueller, W.J., 1956. Feasibility of the chromic oxide and the different indigestible markers, diet composition and the effects lignin indicator methods for metabolism experiments with of feed enzyme supplementation in barley-based diets on ileal chickens. J. Nutr. 58, 29–36. and overall apparent digestibility in growing pigs. Anim. Sci. Njaa, L.R., 1961. Determination of protein digestibility with in press.

titanium dioxide as indicator substance. Acta Agric. Scand. 11, Yin, Y.-L., McEvoy, J., Schulze, H., Hennig, U., Souffrant, W.B., 227–241. McCracken, K.J., 2000. Apparent digestibility (ileal and over-Peddie, J., Dewar, W.A., Gilbert, A.B., Waddington, D., 1982. The all) of nutrients and endogenous nitrogen losses in growing use of titanium dioxide for determining apparent digestibility in pigs fed wheat (var. Soissons) or its by-products without or mature domestic fowls. J. Agric. Sci. 99, 233–236. with xylanase supplementation. Livest. Prod. Sci. 62, 119–132. Saha, D.C., Gilbreath, R.L., 1991. Analytical recovery of Yin, L Y.-L., McCracken, K.J., 1996. Methodological aspects of in chromium from diet and faeces determined by colorimetry and vivo measurement of ileal amino acid digestibility in pigs – a atomic absorption spectrophotometry. J. Sci. Food Agric. 55, review. Asian–Australasian J. Anim. Sci. 9, 495–502. 433–446. Yin, Y.-L., Huang, R.L., Zhong, H.Y., 1993. Comparison of ileo-Sauer, W.C., Ozimek, L., 1986. Digestibility of amino acids in rectal anastomosis and conventional method for the measure-swine: results and their practical application. A review. Livest. ments of ileal digestibility of protein sources and mixed diets in Prod. Sci. 15, 367–369. pigs. Anim. Feed Sci. Technol. 45, 233–298.

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Table 1

Ileal recovery (%) of TiO and Cr O2 2 3

Diet W WR WM WB S.E.D. P Mean

a a a b

PVTC TiO2 99.9 94.9 95.5 67.8 4.92 ,0.001 89.7

a a a b

Cr O2 3 84.9 87.4 82.8 60.6 4.69 ,0.001 79.0

IRA TiO2 98.8 ND 101.1 95.2 2.70 NS 98.4

Cr O2 3 84.3 ND 88.7 86.8 2.39 NS 86.8

PVTC: Measured with PVTC total collection method. IRA: Measured with IRA method.

a,b

Values in the same row without a common superscript are significantly different (P,0.05). ND: Not determined.

Table 2

_{(TiO ) and IRA methods for W, WR and WM but the}

Comparison of average values of ileal apparent digestibility of

_{high fibre WB diet resulted in higher estimates of}

nutrients in diets W, WR, WM and WB measured with PVTC

apparent digestibility by the total collection method.

cannula and either TiO or Cr O2 2 3

TiO2 Cr O2 3 S.E.D. P

DM 0.702 0.671 0.0095 ,0.001

_{4. Discussion}

CP 0.772 0.748 0.0081 ,0.001

Energy 0.710 0.680 0.0084 ,0.001

4.1. Comparison of Cr O and TiO as markers

2 3 2

Threonine 0.747 0.72 0.0092 ,0.001

for determination of ileal apparent digestibility of

Valine 0.748 0.722 0.0074 ,0.001

Cystine 0.785 0.760 0.0083 ,0.001

nutrients

Methionine 0.888 0.880 0.0039 ,0.05

Isoleucine 0.810 0.790 0.0042 ,0.001

_{Low faecal recoveries of Cr O and TiO (ranging}

2 3 2

Leucine 0.820 0.801 0.0052 ,0.001

from 71 to 85%) in pigs have been often reported in

Tyrosine 0.831 0.814 0.0048 ,0.001

the literature (Mueller, 1956; Moore, 1957, 1959;

Phenylalanine 0.923 0.812 0.0199 ,0.001

Lysine 0.819 0.800 0.0054 ,0.001

Ishikawa, 1966; Ishikawa and Sugimura, 1973;

Histidine 0.850 0.834 0.0055 ,0.001

_{Moughan et al., 1991; Greer, 1992; McClean, 1993;}

Arginine 0.883 0.870 0.0042 ,0.001

_{Yin et al., 2000). The present results show that the}

Aspartic acid 0.756 0.730 0.0091 ,0.001

ileal recoveries of TiO and Cr O are much higher

2 2 3

Serine 0.812 0.792 0.0073 ,0.001

than those at the faecal level. For example, the ileal

Glutamic acid 0.892 0.881 0.0036 ,0.001

Proline 0.787 0.763 0.0061 ,0.001

recoveries of TiO

and Cr O

2 3,

measured with the

Glycine 0.721 0.690 0.0086 ,0.001

_{IRA method, were 12.6 and 5.3 percentage units}

Alanine 0.719 0.688 0.0054 ,0.001

_{higher, respectively, than the overall (faecal)}

re-Total AAs 0.817 0.798 0.0061 ,0.001

coveries of TiO and Cr O observed by Yin et al.

2 2 3

(2000). The nearly 100% ileal recovery of TiO for

gestibility for diet WM the results for PVTC using

the low fibre diets W, WR and the medium fibre diet

TiO

and for IRA were very similar, and, in line

WM obtained with the PVTC and IRA pigs indicates

with the expected effects of fibre, values declined

that the disappearance of TiO

observed by Greer

from diet W to diet WB. Similar to the results for the

(1992); McClean (1993) and Yin et al. (2000) did

digestibility of DM, CP and energy, ileal apparent

not occur at the ileal level. This suggests that the

digestibilities of amino acids were also comparable

lower recovery of marker in faeces is due to marker

between the PVTC (TiO ) and IRA techniques,

retention, or absorption or both in the large intestine.

except for valine, phenylalanine and proline. Values

The reasons for retention and absorption in the large

for the DM, CP, gross energy and AA with PVTC

intestine of pigs may be a combination of the long

using Cr O were lower than for the other methods.

2 3

retention time (36 h) of digesta (Kidder and

Man-PVTC by total collection agreed well with the Man-PVTC

ners, 1978), the complex and capacious structure of

(2)

Table 3

Comparison of ileal apparent digestibility of DM, CP, gross energy and selected AAs measured either with markers (TiO or Cr O ) using2 2 3

PVTC-cannula or by total collection (PVTC or IRA)

Diet TiO2 Cr O2 3 PVTC S.E.D. Diet3method IRA

DM W 0.773 0.741 0.772 0.763

WR 0.766 0.752 0.783 ND

WM 0.673 0.632 0.688 0.0155 P,0.001 0.624

WB 0.594 0.556 0.732 0.597

CP W 0.807 0.779 0.807 0.811

WR 0.804 0.790 0.808 ND

WM 0.757 0.727 0.769 0.0311 P,0.05 0.747

WB 0.720 0.693 0.808 0.745

Gross energy W 0.787 0.757 0.777 0.784

WR 0.771 0.759 0.792 ND

WM 0.683 0.644 0.698 0.0142 P,0.001 0.656

WB 0.596 0.558 0.724 0.601

Threonine W 0.77 0.737 0.768 0.764

WR 0.767 0.753 0.785 ND

WM 0.743 0.71 0.756 0.0128 P,0.001 0.732

WB 0.708 0.68 0.794 0.702

Valine W 0.766 0.733 0.764 0.805

WR 0.777 0.763 0.793 ND

WM 0.749 0.717 0.761 0.0123 P,0.001 0.769

WB 0.701 0.673 0.797 0.756

Isoleucine W 0.831 0.808 0.83 0.843

WR 0.837 0.826 0.848 ND

WM 0.803 0.778 0.813 0.0093 P,0.001 0.815

WB 0.768 0.746 0.841 0.8

Leucine W 0.849 0.827 0.849 0.851

WR 0.841 0.832 0.853 ND

WM 0.807 0.783 0.817 0.0121 P,0.001 0.82

WB 0.78 0.76 0.85 0.807

Phenylalanine W 0.935 0.83 0.85 0.876

WR 0.935 0.846 0.865 ND

WM 0.918 0.797 0.828 0.0068 P,0.001 0.851

WB 0.904 0.777 0.861 0.834

Lysine W 0.843 0.811 0.834 0.844

WR 0.832 0.822 0.845 ND

WM 0.82 0.797 0.829 0.0093 P,0.001 0.833

WB 0.786 0.767 0.855 0.807

Histidine W 0.865 0.845 0.864 0.869

WR 0.863 0.855 0.872 ND

WM 0.847 0.827 0.854 0.0075 P,0.001 0.865

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Table 3. Continued

Diet TiO2 Cr O2 3 PVTC S.E.D. Diet3method IRA

Tyrosine W 0.849 0.828 0.849 0.851

WR 0.853 0.844 0.863 ND

WM 0.833 0.813 0.841 0.0082 P,0.001 0.858

WB 0.789 0.771 0.856 0.834

Arginine W 0.887 0.871 0.887 0.908

WR 0.882 0.875 0.89 ND

WM 0.886 0.871 0.89 0.0062 P,0.001 0.901

WB 0.876 0.865 0.917 0.895

Aspartic acid W 0.777 0.746 0.776 0.799

WR 0.779 0.765 0.796 ND

WM 0.751 0.72 0.764 0.012 P,0.001 0.769

WB 0.716 0.689 0.807 0.75

Serine W 0.834 0.81 0.833 0.836

WR 0.837 0.827 0.849 ND

WM 0.8 0.775 0.81 0.009 P,0.001 0.801

WB 0.779 0.757 0.85 0.786

Glutamic acids W 0.909 0.897 0.908 0.912

WR 0.908 0.902 0.915 ND

WM 0.879 0.863 0.885 0.0066 P,0.001 0.873

WB 0.873 0.861 0.913 0.879

Proline W 0.768 0.731 0.83 0.891

WR 0.881 0.874 0.732 ND

WM 0.763 0.733 0.763 0.0247 P,0.001 0.858

WB 0.709 0.683 0.77 0.854

Glycine W 0.749 0.712 0.745 0.784

WR 0.742 0.726 0.761 ND

WM 0.717 0.679 0.73 0.0157 P,0.001 0.74

WB 0.675 0.644 0.782 0.732

Alanine W 0.737 0.7 0.735 0.77

WR 0.75 0.734 0.769 ND

WM 0.719 0.683 0.732 0.0144 P,0.001 0.741

WB 0.666 0.635 0.773 0.714

Total AAs W 0.833 0.81 0.832 0.856

WR 0.851 0.846 0.856 ND

WM 0.808 0.784 0.817 0.0091 P,0.001 0.825

WB 0.783 0.764 0.854 0.814

IRA not included in statistical analysis, since the experimental design was different from the PVTC experiment. ND: Not determined.

the large intestine (Barnicoat, 1945; Moore, 1957;

and Monaghan (1996) the present results highlight

1959) and the high specific gravity of TiO

and

the problems of use of Cr O

2 3

as a marker. Mean

Cr O (Lide and Frederikse, 1995).

2 3

values of the ileal recovery of Cr O measured with

2 3

In common with the studies of McCarthy et al.

the IRA method was only 87% in this study.

(1974), Moughan et al. (1991), Jagger et al. (1992)

Consequently the ileal digestibility of DM, CP,

(4)

energy and total AA measured with Cr O

2 3

were

Cr O

2 3,

TiO

is insoluble in water, HCl, HNO

or

respectively 4.6, 3.2, 4.4 and 2.4% lower than those

dilute H SO (Merck Index, 1989) and analysis of

2 4

measured with TiO (Table 2). Jagger et al. (1992)

TiO

is a relatively simple and straightforward

found that the ileal digestibilities of N and AAs

procedure. High recoveries (95–100%) in samples of

measured with Cr O

2 3

were significantly lower (N,

feeds and excreta of poultry and pigs have been

6.7% and the average value of 11 AAs, 6.1%) than

reported (Njaa, 1961; Leone, 1973; Peddie et al.,

those measured with TiO , when both markers were

1982; Jagger et al., 1992).

added to a barley and wheat based diet fed to

growing and finishing pigs. Furthermore, the results

4.2. Comparison of methods for measurement of

of Jagger et al. (1992) and of this experiment show

ileal apparent digestibility of nutrients

that the standard errors associated with apparent ileal

digestibility values calculated using each marker

The excellent marker recoveries with TiO for the

were lower with TiO

than with Cr O

2 3.

This is

W, WR and WM with the PVTC method and for the

somewhat surprising in that the level of incorpora-

W, WM and WB diets with the IRA method indicate

tion of Cr O was higher than for TiO

2 3 2.

At these low

that, under these conditions, digesta collection was

levels it would be possible for problems to arise due

fully representative of the material reaching the

to lack of homogeneity of mixing. However, the low

terminal ileum. This would be expected with the IRA

standard errors obtained with TiO

would suggest

method where a complete 5-d collection was made

that diet mixing was not a problem.

but it is noteworthy that similar results occurred with

The primary reason for the low recovery and

the much shorter period of collection (24 h) using the

variable results from Cr O

2 3

may be related to

PVTC technique. However, the result for the

high-analytical problems arising mainly from interference

fibre diet WB using the PVTC technique

demon-by other minerals present in the sample (Williams et

strates that quantitative collection of digesta was not

al., 1962; Saha and Gilbreath, 1991). For example,

achieved. This result therefore supports the reports of

¨

Williams et al. (1962) found a 12% decrease in Cr

Kohler et al. (1991) and van Leeuwen et al. (1991)

recovery when phosphate (PO ) was added to the

of variable recovery with the PVTC technique

solution of Cr. Saha and Gilbreath (1991) also

dependant on diet composition and confirms the

showed significant effects on recovery when varying

necessity of using an appropriate indigestible marker.

the concentration of Ca, P or Mg in diets and faeces

Although a direct statistical comparison of the

of pigs. These minerals occur in high concentrations

PVTC and IRA techniques is not valid due to the

in the digesta or faeces of pigs. This possibility is

separate locations in which the techniques were used

presently under investigation in our laboratory. In an

and the different sources of pigs, the results given in

initial study, three levels of Cr O corresponding to

2 3

Table 3 for PVTC (TiO ) and IRA show close

5, 10, 20 ppm were added to the faeces of pigs which

numerical agreement. This, taken with the excellent

had not received Cr O in the diet. The recoveries

2 3

marker recoveries (Table 1), suggests that both

were 88.0, 87.7, 83.3%, respectively. This is similar

methods were giving reliable estimates of ileal

to the mean recovery values reported by Saha and

digestibility. Furthermore, the good agreement

be-Gilbreath (1991) and the mean ileal recovery of

tween IRA and PVTC (TiO ) for diet WB, where

Cr O measured in this study.

2 3

PVTC collection was not quantitative, suggests that

In addition, although Cr O is insoluble in water,

2 3

the digesta recovered was similar in composition to

it is slightly soluble in acids and alkalis (Merck

that by-passing the cannula.

Index, 1989) and therefore it might be soluble to a

As discussed in the Introduction, the main

advan-certain extent in the acid or alkaline digestive juices

tage of the end-to-end IRA technique is that it is a

of the gastrointestinal tract of pigs (Moore, 1959) or

quantitative method for collecting ileal digesta and,

the acid solution used during Cr analysis. Saha and

therefore, it is not necessary to use any marker for

Gilbreath (1991) suggested that the soluble Cr might

calculating digestibility. However, compared with

be lost due to the formation of volatile Cr com-

the PVTC method, the main disadvantages are: (1)

pounds at the sample digestion stage. Compared with

relatively complex surgical procedures and

(5)

post-beans in diets for growing pigs. M.Sc. Thesis. The Queen’s

operative animal care. (2) Longer time for recovery

University of Belfast, Belfast.

from the surgery. (3) Need for greater care to

Hennig, U., Noel, R., Herrmann, U., Wunsche, J., Mehnert, E.,

minimise skin irritation and discomfort due to the

_{1986. Nutrition–physiologic studies in pigs with ileo-rectal}

outpouring of digesta around the hind legs. (4) The

anastomoses. 1. Operation methods, biochemical and

mor-phological findings. Arch. Anim. Nutr. 36, 585–596.

need for additional water, minerals and vitamins to

Hennig, U., Wunsche, J., Souffrant, W.B., Borgmann, E.,

offset the loss of the large intestine. Compared with

Kreienbring, F., 1992. Methodenverleich zur Bestimmung der

the IRA method, the surgery for the PVTC cannula-

_{PrSzSkalen NShrstoffverdaulichkeit und AminosSureresorption}_{ˇ ˇ} _ˇ _ˇ

tion is much easier to perform and less severe on the

_{zwischen Schweinen mit ileorectal-anastomose und ileozSkal-}ˇ

¨ ¨

Bruckenkanule. Arch. Anim. Nutr. 42, 197–211.

animals than the IRA procedure and the PVTC

Ishikawa, S., 1966. Reliability of polyethyleneglycol as an

in-cannulation technique causes less discomfort to the

dicator for digestion studies with swine. 1. Rate of passage of

animal. Furthermore this method permits simulta-

_{polyethyleneglycol through the digestive tract. Agric. Biol.}

neous measurement of ileal and overall digestibility

_{Chem. 30, 278–284.}

in contrast to the IRA method. Taking all of these

Ishikawa, S., Sugimura, K., 1973. Movement of polyethylalcohol through the digestive tract as a digestion indicator with swine.

aspects into account it is concluded that the PVTC

Agric. Biol. Chem. 37, 203–206.

method is a better method for measuring ileal

Jagger, S., Wiseman, J., Cole, D.J.A., Craigon, J., 1992.

Evalua-digestibility in most situations. It would, however, be

_{tion of inert markers for the determination of ileal and faecal}

of considerable interest to make a direct comparison

digestibility values in the pig. Br. J. Nutr. 68, 729–739.

of the PVTC and SICV techniques.

Kidder, D.E., Manners, M.J., 1978. Digestion in the Pig, Kingston

Press, Bath. ¨

Kohler, T., Huisman, J., Den Hartog, L.A., Mosenthin, R., 1990. Comparison of different digesta collection methods to

de-5. Conclusions

termine the apparent digestibilities of nutrients at the terminal ileum in pigs. J. Sci. Food Agric. 53, 465–475.

Kohler, T., Huisman, J., Den Hartog, L.A., Mosenthin, R.,

The results highlighted problems with poor

re-Verstegen, M.W.A., 1991. Digestibility measurements in pigs

covery of Cr O marker resulting in low estimates of

2 3 _{using post valve T-caecum cannulation or end to side}

ileo-ileal digestibility, while the TiO marker appears to

2 _{rectal anastomosis. J. Anim. Physiol. Anim. Nutr. 66, 278–286.}

be satisfactory as shown by good agreement with the

Laplace, J., 1980. Stomach and small intestine motility in the pig:

IRA (total collection) method. PVTC total collection

Electromyography in nutritional studies. In: Low, A.G., Par-tridge, I.G. (Eds.), Current Concepts of Digestion and

Absorp-proved is satisfactory for measurement of ileal

tion in Pigs. Technical Bulletin 3, p. 24. National Institute for

digestibility with low- or medium-fibre diets but not

Research in Dairying, Reading, U.K.

with a high-fibre diet. Considering its simplicity,

_{Laplace, J.P., Souffrant, W.B., Hennig, U., Chabeauti, E., Fevrier,}

acceptability on animal welfare grounds and the

C., 1994. Measurement of precaecal dietary protein and plant

good agreement with in measurements of digestibili-

cell digestion in pigs; comparison of four surgical procedures

for ileo-rectal anastomosis. Livest. Prod. Sci. 40, 313–328.

ty made with the IRA technique, the PVTC

tech-Leone, J.L., 1973. Collaborative study of the quantitative

de-nique, in combination with TiO

as a marker, is

_{terminations of titanium dioxide in cheese. J. Assoc. Anal.}

considered to be a good method for determination of

_{Chem. 56, 535–537.}

ileal digestibility of nutrients for pigs.

Lide, D.R., Frederikse, H.P.R., 1995. CRC Handbook of Chemis-try and Physics, CRC Press, Boca Raton, FL.

Low, A.G., 1982. Digestibility and availability of amino acids from feedstuff for pigs. A review. Livest. Prod. Sci. 9, 511–

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Technol. 27, 202–205. with pigs. Can. J. Anim. Sci. 54, 107–109.

(6)

pigs. M.Phil. Thesis. The Queen’s University of Belfast, Van Leeuwen, P., van Kleef, D.J., van Kempen, G.J.M., Huisman,

Belfast. J., Verstegen, M.W.A., 1991. The postvalve T-caecum

cannula-Moore, J.H., 1957. Diurnal variations in the composition of the tion technique in pigs applicated to determine the digestibility faeces of pigs on diets containing chromium oxide. Br. J. Nutr. of amino acids in maize, groundnut and sunflower meal. J.

11, 273–288. Anim. Physiol. Anim. Nutr. 65, 183–193.

gestibility studies with young pigs. NZ J. Agric. Res. 34, acid-insoluble ash as digestive markers. Br. J. Nutr. 76, 551–

85–88. 562.

titanium dioxide as indicator substance. Acta Agric. Scand. 11, Yin, Y.-L., McEvoy, J., Schulze, H., Hennig, U., Souffrant, W.B.,

227–241. McCracken, K.J., 2000. Apparent digestibility (ileal and

over-Peddie, J., Dewar, W.A., Gilbert, A.B., Waddington, D., 1982. The all) of nutrients and endogenous nitrogen losses in growing use of titanium dioxide for determining apparent digestibility in pigs fed wheat (var. Soissons) or its by-products without or mature domestic fowls. J. Agric. Sci. 99, 233–236. with xylanase supplementation. Livest. Prod. Sci. 62, 119–132. Saha, D.C., Gilbreath, R.L., 1991. Analytical recovery of Yin, L Y.-L., McCracken, K.J., 1996. Methodological aspects of in chromium from diet and faeces determined by colorimetry and vivo measurement of ileal amino acid digestibility in pigs – a atomic absorption spectrophotometry. J. Sci. Food Agric. 55, review. Asian–Australasian J. Anim. Sci. 9, 495–502.

433–446. Yin, Y.-L., Huang, R.L., Zhong, H.Y., 1993. Comparison of

ileo-Sauer, W.C., Ozimek, L., 1986. Digestibility of amino acids in rectal anastomosis and conventional method for the measure-swine: results and their practical application. A review. Livest. ments of ileal digestibility of protein sources and mixed diets in