LIVESTOCK PRODUCTION SCIENCE

ELSEVIER
Livestock Production Science 40 (1994) 139-155

The Dutch protein evaluation system: the DVE/OEB-system

S. Tamminga a, W.M. Van Straalen b, A.P.J. Subnel c'*, R.G.M. Meijer c, A. Steg a,
C.J.G. W e v e r e, M.C. B l o k e
"Department of Animal Nutrition, Agricultural University, Haagsteeg 4, 6708 PM, Wageningen, Netherlands bCLO-institutefor Animal Nutrition "'De Schothorst", P.O. Box 533, 8200 AM, Lelystad, Netherlands CResearchStation for Cattle, Sheep and Horse Husbandry (PR), Runderweg 6, 8219 PK, Lelystad, Netherlands dResearch Institute for Livestock Feeding and Nutrition (IWO-DLO), P.O. Box 160, 8200 AD, Lelystad, Netherlands eNational Reference Centre of Livestock Production, Runderweg 2, 8219 PK, Lelystad, Netherlands fCentral Bureau for Livestock Feeding, Runderweg 6, 8219 PK, Lelystad, Netherlands
Received 13 August 1993; accepted 1 February 1994

Abstract
In 1991 a new protein evaluation system replaced the Digestible Crude Protein (DCP) system in the Netherlands: the DVE/ OEB-system. The system was mainly developed with the aim to prevent avoidable losses of nitrogen, by feeding according to more exactly defined requirements of dairy cows. A second aim was to predict milk protein production more accurately. Protein requirements for maintenance, milk protein production, growth, mobilisation, metabolic losses in the digestive tract and gestation are expressed in DVE, the sum of digestible feed and microbial true protein available in the small intestine. In the system each feed has a DVE-value composed of the digestible true protein contributed by feed protein escaping rumen degradation ( 1), microbial protein synthesized in the rumen (2) and a correction for endogenous protein losses in the digestive tract (3). Each feed also has a degraded protein balance (OEB) reflecting the difference between the potential microbial protein synthesis based on degraded feed crude protein and that based on energy available for microbial fermentation in the rumen. The framework of the new system is based on what are considered strong elements of other recently developed protein evaluation systems. Additionally new elements are introduced, including undegraded starch (USTA), fermentation products (FP) in ensiled feeds, the role of energy balance in protein supply and the way in which requirements change in the course of lactation. Data within the framework of the system are mainly of Dutch origin. This is particularly true for the regression equations developed to predict the protein values of forages and protein values of a number of by-product ingredients.
Keywords: Dairy cattle; Protein Evaluation; Nitrogen loss; Milk protein

1. Introduction
In the Netherlands dairy rations are formulated according to the energy-standards in a net energy system, the VEM-system (Van Es, 1975, 1978). Until 1991 the Digestible Crude Protein system (DCP, the
*Corresponding author 0301-6226/94/$07.00 1994 Elsevier Science B.V. All rights reserved SSDIO301-6226(94)OOO12-V

difference between CP ( N X 6.25) ingested with the ration and CP excreted in the feces) was used to express protein requirements for dairy cattle as well as the protein value of feeds (CVB, 1990). Protein that really can be utilised is only that part of the ingested CP which is digested in and subsequently absorbed from the small intestine (SI) as amino acids. The DCP-system is a poor predictor of the amount of

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true protein absorbed from the SI, because it does not indicate to which degree the CP in a feedstuff is degraded in the rumen, nor does it take into account microbial protein synthesis in the rumen. These are important limitations, because CP that is degraded in the rumen and lost as ammonia (NH3) and urea can not be utilised by the animal. Because no ruminal Ntransactions are incorporated in the DCP-system, it is hardly possible to improve the efficiency of N-utilisation in dairy cows with this system. In many countries more sophisticated systems describing the digestion and metabolism of N in a more detailed way than the DCP-system does, were introduced in practice (INRA, 1978; ARC, 1984; NKJ-NJF, 1985; NRC, 1985; Ausschuss for Bedarfsnormen, 1986). After their first introduction some of the systems were updated already (Vtrit6 et al., 1987; AFRC, 1992). The objectives of this study were to develop a system that: (a) Describes digestion and metabolism of N in dairy cows in detail, (b) Identifies and quantifies the losses of N due to an inappropriate intake of feed N. Principles of already existing systems were used to develop the framework of the DVE/OEB system, which describes digestion and metabolism of N under Dutch circumstances. Together with information on the digestive behaviour of a wide variety of feeds used in Dutch dairy diets, such a system should make it possible to feed protein more accurately according to the real demands of the dairy cow and prevent unnecessary losses of nitrogen (N).

Table 1 The N-losses (kg N per cow and year) in the Dutch dairy cow producing 6250 kg of milk (kg/year) (Tamminga, 1992) Form Input Losses Output Output Output Output

milk Feed Rumen loss Nucleic acids Undigested Endogenous Maintenance Milk Growth
Total

tissues

faeces

urine 25 15

175 25 25 1 33 3
33 4

50

15 6 24 3
88

2. The prevention of losses of N

For a system which describes and quantifies the losses of N in a ruminant animal, information on the sites and moments in the digestion and metabolic processes at which N losses occur is needed. In Table 1 such an example for the Dutch dairy cow is given (Tamminga, 1992). The annual input of N in the Dutch dairy cow (production level: 6250 kg/year) is around 175 kg. When the roughage part of the basal ration mainly contains grass-products it is slightly higher, when on the other

hand maize-silage is the main part of the basal diet, it is slightly lower. Only about 20% of the 175 kg of ingested N becomes recovered as milk protein or is deposited as protein in lean tissue. Dutch practices of feeding dairy cows result in annual N losses from the rumen of around 25 kg, because degraded CP and energy are usually unbalanced (Demeyer and Tanuninga, 1987; Sniffen and Robinson, 1987; Van Straalen and Tamminga, 1990). Often there is a surplus of rumen degraded protein and breakdown of CP and microbial protein synthesis do not always occur at the same moment. Consequently, a shortage of feed N in the rumen may also occur occasionally. Fecal excretion is also an important loss (Charmley et al., 1988) and amounts to about 50 kg of N a year. Half of this originates from undigested feed CP and microbial CP. The other half is due to metabolic losses (enzymes, epithelial cells and mucus ). Part of the metabolic losses appear in the feces as bacterial CP resulting from microbial activity in the hind gut. Metabolic losses need to be replaced. The process of resynthesis is therefore considered to be associated with N losses and believed to cause an elevated N excretion in the urine (Tamminga, 1992). Table 1 shows that about 65% of the N that is lost, is excreted in the urine, but only a small proportion of the loss is a consequence of the losses in maintenance and growth. These latter losses are difficult to avoid.

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Part of the microbial CP will also be lost in the urine, because it is not true protein (amino acids), but nucleic acids. The animal's tissues and organs do not utilise nucleic acids as well as they utilise amino acids (Armstrong and Hutton, 1975). An important additional urinary N loss is associated with the synthesis of milk protein. Average efficiencies for utilization of absorbed true protein are assumed to be 0.67 for maintenance, 0.50 for growth (NRC, 1985 ) and 0.64 for the synthesis of milk protein (V6rit6 et al., 1987). Reducing these losses succesfully in practice requires first of all a protein evaluation system based on a detailed, preferably dynamic description of the Nmetabolism in the dairy cow. The system should also include a proper description of the protein value of feeds (Van Straalen and Tamminga, 1990), including microbial protein supplied by each feed. Additionally, a correct formulation of the requirements of true protein at every moment in lactation is needed.

Table 2 List of abbreviations AAN NAAN BRE %BRE CASH CFAT CP CPM D DCP DOM DVBE %DVBE DVE DVME DVMFE FOM MREN MREE MVRAS OEB %RRE S STA U UM USTA %USTA VRAS %VRAS :Nitrogen presentin aminoacids :Non amino-acid-N :Undegradedfeed CP :Fraction of undegraded feed CP in total feed CP :Crude inorganic matter :Crude fat :Crude protein :Metaboliccrude protein :Unsolublebut potentiallydegradablefractionin nylon bag incubations :Digested Crude Protein :DigestedOrganicMatter :Digested degraded feed CP :Digestionin smallintestine of the undegraded feed protein :True proteindigestedin the small intestine :Rumen synthesisedmicrobialproteindigested in the small intestine :Endogenousprotein lossesin digestion :FermentableOrganicMatter :Microbialprotein synthesisedin the lumen based on availablenitrogen :Microbialproteinsynthesisedin the rumen basedon availableenergy :Maximumvalue of VRAS :Degradedproteinbalance :Fraction of undegraded feed CP in total feed CP after long term rumen incubation :Soluble fractionin nylonbag incubations :Starch :Undegradablefractionin nylonbag experiments :IndigestedDry Matter :UndegradedStarch :Fraction of Undegraded Starch :Digestiblefractionof Crude Ash :Digestioncoefficientof Crude Ash

3. Modern protein evaluation systems

It was chosen to develop a systembased on the principles already formulated in existing modern protein evaluation systems. The protein value for feeds and the requirements for dairy cows are both expressed as the amount of true protein truly digested in and absorbed from the small intestine of the animal: DVE. Other systems use comparable units viz. MP (AFRC, 1992), PDI (INRA, 1978; V6rit6 et al., 1987), AAT (Madsen, 1985; Hvelplund and Madsen, 1990), AP (NRC, 1985) and AAS (Ausschuss fiir Bedarfsnormen, 1986). It was difficult to decide which of the systems was most appropriate under Dutch circumstances. In literature many comparisons between different protein evaluation systems have been reported (Madsen, 1979, 1980, 1985; Jarrige and Alderman, 1987; Waldo and Glenn, 1984; Orskov and Miller, 1988; Van der Honing and Alderman, 1988), but they all remain rather descriptive. Van Straalen et al. (1993a) evaluated which system could be used best under Dutch circumstances. Their conclusion was, that of the modern systems the PDI-system was the most accurate in predicting milk protein production. An additional advantage of the PDI system is that it well tested and widely used in practice.

The Dutch system was therefore developed with the French PDI-system as a basis. If considered appropriate, elements of other systems were incorporated. For instance for expressing maintenance requirements, the approach in the AP-system (USA) was followed. To get a clear impression of the N-losses in the rumen, the concept of the Scandinavian PBV-value (PBV = protein balance in the rumen) was accepted. A list of abbrevations used in the text is given in Table 2.

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4. The supply of true protein to the small intestine

Different components contribute to the DVE value: ( 1) Undegraded feed CP digested in and absorbed from the small intestine as amino acids: DVBE (2) Microbial CP digested in and absorbed from the small intestine as amino acids: DVME (3) Endogenous losses resulting from digestion: DVMFE The DVE-value of a feedstuff is then formulated as: DVE = DVBE + DVME - DVMFE

mation was available a %BRE value of 35% was assumed. The amount of BRE (g/kg) in individual feeds is estimated as: BRE= 1.11 (%BRE/100) C P (BRE and CP in g/kg; %BRE in %) The factor 1.11 in formula (2) is taken from the French PDI-system. The %BRE-values of roughages were mainly derived from data obtained in Dutch trials (Meijer, 1990; Tamminga et al.; 1991; Bosch et al., 1991; Van Vuuren et al., 1992; De Visser, unpublished). For fresh grasses, grass silages and hays, multiple regression equations, in which chemical composition (CP, DM) and day of harvest (DH) were identified as explaining variables for %BRE, were developed (Van Straalen and Tamminga, 1990; Tamminga et al., 1991; CVB, 1991a): % B R E = a + b l XCP+b2 D H + b 3 D M

(2)

(1)

5. Digestible undegraded feed protein (DVBE)

To quantify the DVBE-value of a feedstuff, information is needed on the total amount of CP as well as on the ratio AAN/total-N. Furthermore the degradation and digestion of the AAN must be estimated.
5.1. Undegraded feed crude protein (BRE)

(3)

The fraction of feed CP escaping degradation in the rumen is estimated from the U, S and D-fraction resulting from incubation of a feed in nylon bags in the rumen (Van Straalen and Tamminga, 1990). The size of the U-fraction is determined after long term ( 10-14 days) rumen incubations. The size of the actually degraded fraction is subsequently estimated from the potentially degradable fraction (D), the rate of degradation (ko) and the rate of passage (kp) (Robinson et al. 1986). It should be pointed out that the size of the potentially degradable fraction is not derived from the estimated asymptote of the degradation curve, as is done in the model of ~3rskov and McDonald (1979), a model often used to describe nylon bag incubation data. Based on international and Dutch data, passage rates (kp) of 4.5% for roughages and 6% for concentrates were adopted. The %BRE-values for feeds were mainly derived from compiled Dutch and international data (Van Straalen and Tamminga, 1990). For feeds on which no data were available the %BRE value was estimated by comparison with feeds of a similar nature and a comparable chemical composition. If no infor-

a, bl, b2, b3: regression coefficients CP (N 6.25): in g/kg DM DH: days of harvest after April 1 st DM: dry matter content (g/kg) R 2 varied from 0.73 for fresh grasses to 0.80 for hays. Values for the regression coefficients are given by CVB (1991a). Because of observed discrepancies between the estimated DVE value and production responses in practice after feeding ensiled feeds, grass silages in particular, for this category of feeds it was additionally assumed that 5% of the S fraction is also washed out of the rumen before being degraded.
5.2. Percentage of amino acids in undegraded feed crude protein

Results of nylon bag incubation studies were used to estimate the percentage of amino acid N (AAN) in undegraded dietary CP. Differences in AAN in residues from nylon bags incubated in the rumen for different lengths of time were found to be small for concentrate ingredients (Vrrit6 et al., 1987; Hvelplund and Hesselholt, 1987). For roughages more variable results

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were reported (Rook et al, 1984; Hvelplund, 1987; Hof et al., 1990; Van Straalen et al., 1993b). Most systems have accepted an AAN/N-ratio of 1.0 (ARC, 1984; NRC, 1985; Vtrit6 et al., 1987). In the Nordic AAT/PBV system, a value of 0.65 was accepted for roughages and 0.85 for concentrates (Madsen, 1985). In the DVE-system, the estimation of the percentage of amino acids in undegraded feed CP is not specified, but taken into account in the estimation of the intestinal digestion of CP.

c, b4, bs: regression coefficients CP (N 6.25): in g/kg DM DH: Days after April 1st Values for the regression coefficients are given by CVB (1991a).

6. Microbial protein digestible in the small intestine (DVME)

To estimate microbial growth, the DVE-system uses the organic matter (OM) available for fermentation by the rumen microbes under anaSrobic conditions as found in the rumen, like in the PDI-system. It is called fermentable organic matter (FOM). FOM is calculated from the apparently digested organic matter (DOM), but DOM is corrected for the amounts of crude fat (CFAT), undegraded feed CP (BRE), undegraded starch (USTA), and end products of fermentation (FP) in ensiled feeds. USTA values are based on rumen escape values determined with nylon bag incubations (Tamminga et al., 1990; Nocek and Tamminga, 1991 ) and if relevant, corrected with a factor 0.75 to account for the observed negative effects of pelleting. To compensate for differences observed between in vivo and in sacco values, it was further assumed that 10% of the starch fraction that is washed out of nylon bags prior to rumen incubation, escapes from rumen degradation (Nocek and Tamminga, 1991). The correction for fermentation end products in ensiled feeds is based on the assumption that from the most important fermentation end products lactic acid and alcohols, rumen microbes can extract the equivalent of about 50% of the energy (ATP) extractable from carbohydrates. Hence, DOM is corrected for 50% of the fermentation end products in ensiled products. The way in which the DOM fraction is corrected is shown in the following equation: FOM = DOM-CFAT-CP (%BRE/100)-STA

5.3. Digestion of undegradedfeed crude protein (%DVBE)

In the DVE system the estimated intestinal digestion of undegraded feed CP is based on results of mobile nylon bags passing the digestive tract (Van Straalen and Tamminga, 1990). Because the intestinal digestion of non-amino acid-N (NPN) is low, and the percentage of amino acids in the undegraded feed CP digested in the small intestine is high (Tamminga and Oldham, 1980), the true digestion of amino acids is given as %DVBE. For feeds of which no direct information was available, %DVBE was estimated on the basis of information of comparable feeds. Feeds with a high cell wall content were given a value of 50%. Products of which information was completely lacking, were given a value of 75%. For roughages insufficient information was available of trials in which the mobile nylon bag technique had been used. The %DVBE is therefore calculated from the %BRE and the indigestible CP fraction after long term rumen incubation (%RRE): %DVBE = 100 ( % B R E - %RRE)/%BRE The amount of DVBE is then calculated as: D V B E = C P (1.11 %BRE/100) (%DVBE/100) (DVBE and CP in g/kg DM; other values in %.) For roughages (grasses, grass silages, hays) %RRE is estimated from the chemical composition and day of harvest (CVB, 1991a): %RRE= c +b4 C P + b 5 D H (5) (4)

(%USTA/100) - 0.50 FP

(7)

(%BRE and %USTA in %; other parameters in g/kg). Values for CP, DOM, CFAT, %BRE, STA and %USTA are given by CVB (1991b) as far as the con-

(6)

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centrates and industrial by-products are concerned. For roughages values are also listed in CVB (1991a). Equations to calculate the FP-content of ensiled feeds were derived from Steg et al., (1990). 6. I. Microbial protein production in the rumen In the French PDI system microbial protein production is based on FOM (Vrrit6 et al., 1989) and is calculated to be 145 g of microbial CP per kg FOM. In the DVE-system a value of 150 g/kg FOM was adopted. This slightly higher value was chosen because, except for maize and milocorn, the PDI system, does not account for undegradable starch. Moreover, the French values were derived from a set of data in which the level of intake was relatively low. From work by e.g. Robinson et al. (1987) it is known that a higher intake stimulates the efficiency of microbial protein synthesis positively. 6.2. Amino acids in microbial crude protein and its true digestion Based on data from Dutch trials (S. Tamminga, unpublished) as well as literature, a value of 0.75 was accepted for the AAN/N ratio in microbial CP and a true digestion of 85% was chosen for AAN. At the same level of FOM, only a minor difference exists between the French and Dutch estimation of digestible microbial protein (DVME). The latter is calculated as: DVME = FOM X 0.150 X 0.75 X 0.85 (all values in g/kg)

accordingly. In rations of average composition, loss of metabolic crude protein (CPM) equals 50 g/kg UDM (CVB, 1991a). UDM intake can be separated in indigestible organic matter and indigestible inorganic matter. In the DVE system the digestible inorganic matter (VRAS) is calculated from the crude inorganic matter (CASH) as: VRAS = %VRAS / 100 X CASH (%VRAS in %; CASH in g/kg DM) Each feedstuff has a maximum amount of VRAS (MVRAS). The introduction of MVRAS prevents overestimation of VRAS, for instance due to a large but variable proportion of sand in an individual batch of a particular feedstuff. More detailed information on MVRAS for different feeds is given in CVB ( 1991a and 1991b). The UDM is calculated as: UDM = 1000-DOM-VRAS (all values in g/kg) Based on a net loss of metabolic protein of 50/kg UDM and an efficiency of resynthesis of 0.67, the requirement of DVE for metabolic protein losses (DVMFE), and used as a correction factor for the protein value of each feedstuff, is calculated as: DVMFE = 0.075 UDM (9)

(lO)

(8)

(11)

8. The degradable protein balance (OEB) 7. Endogenous losses in digestion (DVMFE)

Endogenous CP lost in the digestive process exists of digestive enzymes, bile, desquamated epithelial cells and mucus. Although it originates from the animal itself, its magnitude is considered to depend more on characteristics of the feed than of the animal. The DVE, required to compensate for endogenous losses, called DVMFE, is not restricted to endogenous protein itself but also includes amino acids lost in its resynthesis. DVMFE is considered to be directly related to the undigested DM and the DVE-value of feeds is corrected The OEB value, like the PBV value in the AAT/ PBV system (NKJ-NJF, 1985), shows the (im)balance between microbial protein synthesis potentially possible from available rumen degradable CP (MREN) and that potentially possible from the energy extracted during anaerobic fermentation in the rumen (MREE). It also reflects the difference between PDIN and PDIE in the French system (V~rit6 et al., 1987). When positive, the OEB-value gives the loss of N from the rumen. When OEB is negative, microbial pro-

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tein synthesis may be impaired, because of a shortage of N in the rumen. The optimum OEB-value in a ration is therefore zero or slightly above. Contrary to the PBV value, in the DVE system, the OEB value is recommended not to become negative. The risk of a shortage of N for the microbes is considered too high. In that case the calculated values of DVME may not be achieved. This risk is particularly apparent at higher levels of feed intake, normally found in early lactation, combined with infrequent feeding. OEB = MREN - MREE (all values in g/kg) MREN is calculated as: MREN= C P x ( 1-1.11 x %BRE/100) (MREN and CP in g/kg; %BRE in %). The maximum synthesis of microbial CP based on energy is calculated as: MREE = FOM X 0.15 (FOM in g/kg) (14) (13) (12)

(hair, scurf, skin secretions) divided by the efficiency of utilisation for maintenance: DVE-M = (2.75 X BW 5 +0.2 X BW '6)/0.67) (15) (DVE-M in g/day; B W = body weight in kg)
9. 2. Milk protein production (D VE-P)

Modern protein evaluation systems developed till now, use a constant efficiency factor to express the utilisation of absorbed true protein for milk protein production. The PDI-system uses a value of 0.64. Initially this value was also chosen for the DVE-system. However, production trials performed under Dutch circumstances revealed that this efficiency was variable and dependent on the DVE/NEL-ratio (Subnel and Meijer, 1993; Van Straalen et al, 1993a), and level of production (Subnel and Meijer, 1993). Based on this finding, an equation was derived (Subnel and Meijer, 1993), by which a direct estimation of DVE-requirement for production of milk protein (DVE-P in g/day) can be made: DVE-P = 1.396 X GRP + 0.000195 x GRP 2 (16)

(DVE-P in g/day); GRP: Milk protein production in g/day)

9.3. Growth and mobilisation (DVE-GM)

9. Protein requirements in the DVE-system

Dairy cattle require protein for maintenance, for gestation, for growth and for milk production. In the DVE system the requirements are corrected for protein that is mobilised from or stored in body reserves. As was explained earlier, in the DVE-system the requirement for metabolic losses in the digestive tract is taken into account in the DVE-value of each individual feedstuff. As a result maintenance requirements do not have to account for metabolic losses from the digestive tract and the approach chosen by NRC (1985) was followed.
9.1. Maintenance (DVE-M)

The DVE-requirement for maintenance is calculated as the DVE-requirements for losses in urine and body

Requirements for growth and the amount of protein that becomes available from mobilised body reserves depend on the actttal energy balance of the animal. This balance is calculated as the intake of net energy (VEM) minus net energy required for processes mentioned above. When the balance is positive, energy, part of which is protein, is stored in body reserves. When the balance is negative, i.e. when energy is mobilised, part of that energy will become available as amino acids which can be used for metabolic processes. In the DVE system it is assumed that 10% of the energy in body reserves is present in protein (A.J.H. Van Es, unpublished; Waldo et al., 1991 ). Therefore in body reserves each 6.9 MJ (the equivalent of 1000 VEM), contain about 0.7 MJ in protein, which, under the assumption of 24 MJ/kg of protein, is the equivalant of 29 g of protein. Restoring body reserves in dairy

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cattle, is assumed to be possible with an efficiency of 50% (NRC, 1988). Because of the absence of metabolic losses in digestion and an appropriate amino acid composition, protein from body reserves is expected to be used more efficiently for milk protein production than DVE absorbed from the small intestine (80% versus 64%). In a negative energy balance situation, the mobilised energy is also assumed to be used for milk production with an efficiency of 80% (Van Es, pers. comm.). This means that per 6.9 MJ negative energy balance 36 g of body protein is mobilised, the equivalent of 36 ( 8 0 / 6 4 ) = 4 5 g DVE. For animals in a positive energy balance each 6.9 MJ, will require 57 (29/0.5) g of DVE for restoring body reserves.
9.4. Gestation (DVE-G)

For the DVE-requirements for gestation, the NRC appraoch was followed. From day 141 till day 281 after conception, DVE is required for the foetus and extra maternal tissues. Like in the NRC approach, the efficiency is assumed 50% (NRC, 1985). DVE-G (g/day) = (34.375 exp(8.537- 13.1201) exp~- o.oo262- o~_ 0.00262 D 0.50 (17) (D = days after conception between 141 and 281.)

10. Discussion
10.1. Validation o f modern protein evaluation systems

Validation of new protein evaluation systems has not been performed on a large scale. Some comparisons were either rather theoretical or were made with systems already abandoned for use in practice. Many experiments to evaluate the effect of protein feeding on protein production were carried out in the past, but only a limited number of these were used to validate protein evaluation systems (Jarrige and Alderman, 1987). Differences between predicted and observed milk production varied between systems and experiments (VikMo, 1985; MacRae et al., 1988; Sloan et al., 1988; Garnsworthy, 1989; Broderick et al., 1990; Cody et al., 1990; Robinson et al., 1991; Susmel et al., 1991). Yet it became also apparent from such evaluations that pro-

tein intake estimated according to new systems, usually shows a better relationship with milk protein production than CP or DCP intake does (Thuen and Vik-Mo, 1985; Vik-Mo, 1985; Bricenco et al., 1988). The variation still observed in such comparisons probably results from differences in experimental conditions (design, type of feedstuff), production level and type of animals used (Van Straalen et al., 1993a). It also became apparent that the optimum dietary CP content is variable due to differences in true protein supply and requirement (Waldo and Glenn, 1984; Alderman, 1987). A large data set of Dutch milk production trials was used to make a comparison between the CP, DCP, MP, AAT/PBV, PDI, AAS and the AP-system (Ketelaar, 1986; Van Straalen et al., 1993a). Milk protein production predicted by the different systems was compared with the protein production realised in feeding trials. The accuracy with which milk protein output was estimated was generally improved when the CP- or DCP-system was replaced by a modern system. A large part of the remaining error of prediction could be explained by the protein/energy ratio in the diet. Of the modem systems, the PDI-system was more accurate in predicting the milk protein production than (in following order) the AAT, AAS, AP and MP-systems. The overestimated milk protein production in the AAT and AAS system appeared mainly the result of an assumed (too) high efficiency of utilisation of absorbed protein for milk protein production. The main reason for overestimation in the AP-system was an overestimated AP-supply. In the MP-system not only the MP-supply was overestimated, maintenance requirements appeared underestimated at the same time. The PDI-system was finally chosen as the basis of the DVE/OEB-system, but elements of other systems (NRC, 1985; NKJ-NJF, 1985) were also incorperated in it. Examples are N-losses from the rumen, estimations of maintenance requirements and requirements for gestation. Although the framework was taken from already existing systems, the DVE/OEB-system uses data mainly originating from Dutch research, particularly with regard to the protein values of roughages. Some elements in the recently developed protein evaluation systems are still somewhat uncertain because of a lack of documentation. They are discussed below.

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10.2. Digested undegradedfeed crude protein (DVBE)

A first point of concern is the nylon bag incubation method used to estimate the contribution of undegraded feed CP. This has become the almost universally adopted method. Applying this method for measuring CP degradation in feeds low in CP, fibrous roughages in particular, easily leads to contamination with microbial N of the residues. Using the normal washing procedure this contamination is difficult to wash out of the bags (Kennedy et al., 1984). Microbial contamination is however of more theoretical than of practical importance. Feeds low in CP, even if they are heavily contaminated with microbial N, do not contribute much to the intestinal supply of rumen undegraded true protein. The estimated effective rumen escape is based on the ratio between rate of disappearance through degradation and rate of total disappearance from the rumen, the latter based on a (usually) fixed rate of passage. Rate of passage was shown to vary with level of feed intake (Madsen, 1986) and type of diet (Clark et al., 1992). A high passage rate shifts the site of digestion from the rumen to the lower gut. This is not only the case with protein but also with energy yielding substrates enabling microbial growth. An underestimated rumen escape due to a high rate of passage is therefore likely to be counterbalanced by a decreased ruminal microbial synthesis as well as a decreased intestinal digestion. Different values for the rate of passage (kp) have been adopted in different protein evaluation systems. The Nordic system uses an outflow rate of 8% per hour (Madsen and Hvelplund, 1985). The French PDI system (Vrrit6 et al., 1987) uses 6% for all feeds, despite their claim that 4.5% was more appropiate for roughages. Reviewing data from literature, Tamminga and Ketelaar (1988) concluded that values of 5% for concentrates and 3% for roughages looked appropriate. These latter values seem low compared to what was adopted in most other systems and as a compromise, the DVE-system uses 6% as a mean kp-value for concentrates and 4.5% for roughages. Rate of degradation also varies, not only among feeds, but also with diet composition. Concentrate rich diets usually cause a low ruminal pH, which may reduce the rate of CP degradation (Vik-Mo and Lindberg, 1985; Clark et al., 1992). Because a high pro-

portion of concentrates in the diet usually coincides with or maybe even causes a lower rate of passage (Owens and Goetsch, 1986), effective CP escape may not change. So, although rate of CP degradation in the rumen is a more dynamic process than simulated by nylon bag incubations, intestinal supply of absorbed protein per kg of feed ingested is probably less variable due to compensating mechanisms. Fixed figures for CP escape in individual feeds as used in the DVE and other protein evaluation systems seem therefore justified, at least at present. In the PDI-system, the prediction of the amount of CP (NAN) entering the small intestine was based on multiple regression analysis of duodenal flow data from in vivo experiments and published in international literature. The main body of data originated from trials with sheep. In the equation developed to predict the amount of BRE entering the small intestine, a regression coefficient of I. 11 appeared for the amount of feed CP escaping rumen degradation, based on nylon bag incubation studies. At first sight this suggests that the nylon bag method underestimates the amount of feed CP escaping degradation. Other in vivo observations (Aitchison et al., 1986; Oosting, 1993) do suggest however that nylon bag incubations underestimate rate of degradation, at least that of cell walls. It should be realized, that in a multiple regression equation, regression coefficients are not entirely independant of each other. Explanations are than that the flow of microbial CP into the small intestine is underestimated, that the contribution of endogenous CP is underestimated or that, because part of it is flushed out with the fluid, feed CP leaves the rumen at a faster rate than other feed ingredients (Tammingaet al., 1989). Lack of solid data prevent drawing a firm conclusion, the reason why the regression factor was maintained in the DVE system, rather than to increase passage rates to values which seem physiologically unrealistic. During the development of the DVE system a discrepancy between the DVE-value according to the DVE-system and observed production responses occurred for ensiled roughages. This difference was not apparent in non-fermented (dry or fresh) feeds. The causes for this discrepancy were not clear. A factor of importance may be microbial CP, formed during the ensiling process. This microbial CP, a large part of which can be washed out immediately in nylon bag incubations, may leave the rumen at a fast rate since it

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can easily be washed out of the rumen with the fluid phase and thus partly escape ruminal degradation. For ensiled feeds it was therefore adopted that 5% of the N-fraction that will normally be washed out is escaping degradation in the rumen. Another point of concern is the AAN/total N ratio in feed CP escaping rumen degradation. Nylon bag studies performed to determine the AAN/N ratio in undegraded feed CP, showed little variation for concentrates (Vtrit6 et al., 1987;, Hvelplund and Hesselholt, 1987), but for roughages more variable results were reported. For ryegrass (Hvelplund, 1987) and for grass silage (Rook et al., 1984) it was found that the degradation of CP in the rumen is followed by a slight reduction in the AAN/N ratio in the undegraded CP, compared to the CP in feed. Van Straalen et al. (1993b) on the other hand, found a rise in the AAN/N ratio in the undegraded CP compared to the feed CP. This rise might be due to the fact that NPN is more degradable in the rumen than AAN. Hofet al. (1990) observed an initial rise in AAN/N followed by a decrease. This does suggest that two NPN fractions are present in roughages, one, probably associated with the cell contents, which can easily be washed out and one which is quite resistant against washing out of nylon bags, possibly because this fraction is associated with the cell walls. This also points out the importance of the length of incubation time in such studies. According to Vtrit6 et al. (1987), AAN/N in nylon bag incubation residues was on average 0.85 in all feeds. In the Nordic system it is assumed that of the undegraded feed CP entering the small intestine and originating from concentrates the AAN/N ratio is 0.85, undegraded feed CP originating from roughages is assumed to have an AAN/N ratio of only 0.65. The lower value for roughages was justified based on a comparison of data from nylon bag incubations and data of duodenal flow measurements (Hvelplund and Madsen, 1985). At first sight this assumption seems fair, but it may also be assumed that the major part of non-amino acid-N (NAAN) present in roughages, particularly in silages, is soluble and washed out of nylon bags (Hof et al., 1990; Van Straalen et al., 1993b). Hvelplund and Madsen (1985) derived their value from multiple regression calculations in which the duodenal flow of Non-ammonia-N (DNAN) and that of amino acid N (DAAN) was related to carbohydrates fermented (DCHO) and N escaping degradadion

(UN) in the rumen. DNAN and DAAN estimates were on the same set of data, hence, the ratio between the regression coefficients when estimating DAAN and DNAN from DCHO and UN should give an indication of the AAN/N ratio in microbial and undegraded feed CP respectively. The ratio between the regression coefficients obtained for UN originating from roughage rich (R) and concentrate rich (C) diets was 0.59 and 0.86 respectively. However, the ratio for DCHO originating from R and C diets were 0.74 and 0.57 respectively, suggesting that microbial CP originating from C diets has a much lower AAN/N ratio than those originating from R diets, which is difficult to rationalize. This observation emphasizes the danger of drawing conclusions on individual regression coefficients obtained from multiple regression equations. Despite that part of the feed CP escaping degradation in the rumen is not AAN, most systems assume 100% of the total undegraded feed-CP being AAN. According to Vtrit6 et al. (1987) this is justified because undegraded feed CP really entering the duodenum has a higher AAN/N ratio than residues from nylon bag incubations. This opinion could be challenged, because, when using AAN/N ratios for microbial CP, undegraded concentrate CP and undegraded roughage CP of 0.75; 0.85 and 0.65 respectively, this would suggest an AAN/N ratio in CP flowing into the small intestine of around 0.75. Ratios observed in vivo are usually lower (Hvelplund and Madsen, 1985). This also indicates a high contribution of NPN in endogenous secretions. The NRC (1985) reported data based on regression analysis and found a mean value of 0.80 for the true absorption of AAN entering the duodenum. In the French system the digestion of undegraded feed CP is given per category of feeds, and calculated from the amount of undegraded total CP entering the duodenum and the undigested CP. It varies between 0.6 and 0.95 (INRA, 1978). A good relationship was observed between the calculated digestion and the digestion measured by the mobile nylon bag technique (Vtrit6 et al., 1987). Van Bruchem et al.(1989) estimated an overall true digestion of AAN passing into the proximal duodenum and terminal ileum of 0.85 in sheep fed diets of whole roughage or roughage-mixed with concentrates. In most diets part of the undegraded feed CP in the duodenum originates from roughage. For cows fed a diet

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of pure haylage Hvelplund (1984) found a digestion of 0.61. According to Hvelplund and Madsen (1990), digestion of AAN in this fraction is lower than that of AAN from protein supplements because in roughages the undegraded CP is linked to fibre and thus more resistant to digestion in the small intestine. In the DVE system the intestinal digestion of undegraded feed AAN is based on measurements with the mobile nylon bag method (Van Straalen and Tamminga, 1990). A limitation of this method is that it measures the digestion of the total N rather than of AAN. This approach seems however justified because AAN appears to have a much higher intestinal digestion than NPN (Oldham and Tamminga, 1980). For roughages insufficient information was available from mobile nylon bag experiments.%DVBE was therefore calculated from the undegraded residue after long rumen incubations as given by Eq. (4). This was based on a linear relationship between the size of the truly undegradable fraction (after two weeks of rumen incubation) and the undigested fraction after 16 hours of rumen incubation followed by intestinal digestion in mobile nylon bags (Tamminga and Ketelaar, 1988), observed for concentrate ingredients. Further research is needed to establish a similar relationship for roughages.

10.3. Microbial protein digested in the small intestine (DVME)

In the Nordic AAT/PBV system microbial protein synthesis is related to the carbohydrates digested (DCHO). Microbial protein synthesis in the DVE-system is related to the FOM, like in the French PDIsystem. The differences between the two approaches are small (Hvelplund and Madsen, 1990). When calculating FOM from DOM, theoretically the latter should also be corrected for the OM fermented in the hindgut and for the fecal excretion of endogenous and dietary fat. A further correction should be made for the amount of apparently undigested CP (undegraded feed CP, microbial CP and endogenous CP). It is assumed that the positive and negative influences on FOM as mentioned above largely compensate each other. The full potential of microbial CP yield in the rumen from the energy released during fermentation is only achieved when the microbes leave the rumen at a pas-

sage rate equal to their division rate (Hvelplund and Madsen, 1990). Normally this is not the case and the amount of microbial CP entering the small intestine remains below potential. Clearance of microbes from the rumen is influenced by dilution rate (Van Soest, 1982). A lower passage rate compared to division rate increases the proportion of substrate lost in maintenance of the microbial population (Hvelplund and Madsen, 1990). Factors influencing the ruminal outflow rate of microbes, divide the maintenance costs over a greater or lesser microbial yield (Kennedy et al., 1976; Allen and Harrison, 1979; Harstad and Vik-Mo, 1985; Meyer et al., 1986). The microbial eco-system in the rumen to a large extent depends on cross-feeding between microbial species. It can be argued that cross-feeding circumstances become sub-optimal when maximum feed intake is approached and that net microbial growth, strongly favoured by a rapid passage rate, becomes reduced. Sources of variation in microbial CP flow to the duodenum were reviewed by Hvelplund and Madsen (1990). They include the recycling of NH3 (NRC, 1985; 1988; Nolan and Stachiw, 1979), protozoal predation (Coleman, 1975), the quality of the degraded feed CP (McAllen et al., 1988), and specific requirements for amino acids or peptides (Russell et al., 1983), branched chain volatile fatty acids (Russell and Hespell, 1981) or minerals (Durand and Komisarczuk, 1988). The influence of each single factor is however not yet known (Hvelplund and Madsen, 1990). In most systems microbial yield is only related to the DMI or OM digestion (Nocek and Russell, 1988). Some of the factors influencing microbial yield, are included in the Cornell Net Carbohydrate and Protein System, CNCPS (Fox et al., 1992; Russell et al., 1992; Sniffen et al., 1992). In this system it is assumed (Russell et al., 1992) that dietary NDF-content and the ruminal concentrations of NH 3 and peptides have a significant influence on microbial growth. Different types of microbes use different N-sources for growth. In the CNCPS it is assumed that the amount of carbohydrates or CP digested in the rumen is influenced by their relative rates of degradation and passage. Ruminal passage rates are influenced by DMI, particle size, bulk density and type of feed (forage or concentrates) (Sniffen et al., 1992). Because losses of NH3 from the rumen through pas-

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sage and absorption, are considered inevitable, the PDIsystem (Verit6 et al., 1987) assumes that a maximum of 90% of degraded feed N can be used for microbial protein synthesis; the MP system (AFRC, 1992) assumes that feed N quickly degraded (QDN) can be captured with an efficiency of 80%. The DVE-system, like the AAT/PBV system, assumes that the N from degraded feed CP that can not be used immediately for microbial protein synthesis is compensated by recycling of urea. The maximum efficiency of utilisation of feed N by rumen microbes is therefore assumed to be 100%. Microbial CP is a mixture of protozoal and bacterial CP. Although variable, the protozoal contribution to the total microbial CP is considered relatively small (Harrison and McAllen, 1980). Like N in feed, N in microbes is only partly present as AAN, and the ratio AAN/N seems variable. Variation in the AAN/N ratio in microbes as summarized for the different protein evaluation systems, reflect the variation found in bacterial preparations. As summarized by Hvelplund and Madsen (1990), variation could be due to diet, due to time of isolation compared to feeding and due to feeding regime. Indications for influence of diet are given by Hvelplund (1986). In the French PDI-system the AAN/N ratio in microbial CP is given as 0.8. The ARC (1980) gives a value of 0.81, also calculated by Storm (1982) based on 41 published reports in literature. NRC (1985) has adopted the value of 0.8, the Nordic countries 0.7 (Madsen, 1985) and Germany also 0.7 (Rohr, 1987). In the DVE-system a ratio of 0.75 is assumed. The true digestion of microbial AAN in the French system is 80%. The true digestion of microbial protein was reported 87% by Tas et al. ( 1981 ), 81% by Storm and Orskov (1983), 85% by Storm et al. (1983a) and 85% by Hvelplund (1985). These figures suggest a fairly constant digestion of amino acids in microbial CP. In the DVE-system a value of 0.85 was chosen.

gut is not well documented, but the quantity is assumed to be at least related to the DM (NRC, 1988; Swanson, 1982; Van Bruchem et al., 1987; 1989) or OM (V6rit6 et al., 1987) passing the intestinal tract. Part of the metabolic excretions are re-absorbed from the small intestine. In the hindgut part of it is converted into microbial CP or absorbed as NH3 after microbial degradation. In the opposite direction, transfer of urea from the blood to the hindgut and a subsequent capture by microbes also occurs (ARC, 1984; Tamminga, 1992). Endogenous CP losses have to be replaced by resynthesis. The French PDI-system and the Nordic AAT/ PBV-system includes them in their maintenance requirement, while the NRC system considers them as a seperate requirement. The DVE system was mainly introduced with the aim to control avoidable N losses to the environment. Because these losses can be manipulated by selecting feed ingredients, causing low endogenous losses, it was decided to follow the NRCapproach. Metabolic losses vary with the quality of the diet (Tamminga, 1992). The NRC (1985) suggests a value of 30 g of CP per kg of DM ingested at a digestion of the DM of 67%. This implies a (fecal) CP excretion of 90 g per kg of undigested DM. This approach does not take into account that part of the MFN is in fact undigested microbial CP. In rations of average composition this amount is estimated at about 40 g per kg of undigested DM (CVB, 1991a). These losses have to be corrected for. The total loss of metabolic CP is therefore assumed to be around 50 g per kg of undigested DM. This value is comparable to values found by V6rit6 et al. (1987) who found the following relationship to estimate metabolic CP (CPM) losses: CPM = 0.0289 FOM + 0.216 BRE + 0.060 UDM (FOM, BRE and UDM in g/kg)

10.4. Endogenous losses in digestion (DVMFE)

In reviews (Boekholt, 1976; ARC, 1984; NRC, 1985; Owens, 1987), excretion of metabolic fecal N (MFN) was calculated between 20 and 30 g of CP per kg DM intake. MFN does however not represent true metabolic losses from the gut. The regulation of quantity and composition of endogenous CP losses from the From this equation it appears that with each kg of fecally excreted organic matter, about 60 g of CP is lost. With an average OM/DM ratio in the feces of 85%, this adds up to 51 g of MCP per kg UDM. Assuming an efficiency for synthesis of 67% (NRC, 1985), the DVE-requirement to replace these losses is calculated as 75 g per kg UDM.

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10.5. The degraded protein balance ( OEB)

Compared to the French PDI-system (V6rit6 et al., 1987), the value PDIN is used in a different way in both the AAT/PBV-(NKJ-NJF, 1985) and the DVEsystem. In these systems it is called protein balance in the rumen (PBV or OEB) and reflects the losses of N not captured in the rumen. This was done to visualize the losses of N in dairy cattle diets to those who formulate rations in practice. In the DVE-system the OEB value should not be negative. In the AAT/PBV system a negative value of PBV is allowed. Kristensen et al. (1988) and Hvelplund et al. (1987) showed that in early lactation, values below - 300 g PBV/day reduce production, and hence a value around 0 is recommended. Hvelplund and Madsen (1990) concluded that the PBV value of a ration can be allowed below zero when the AAT supply is above requirement, because the N from the surplus of absorbed amino acids can be recycled to the rumen. This could be of importance in early lactation, but may still result in rumen NH 3 levels low enough to impair feed intake and digestion (Oldham, 1984). In late lactation there is ample N for recycling. Based on results of V6rit6 and Geay (1987), Hvelplund and Madsen (1990) calculated an allowance of around - 10 gr PBV/SFU (Scandinavian Feed Unit) in early lactation and - 15 PBV/SFU in mid-lactation for the Nordic system. V6rit6 et al. (1987) tolerate an even more negative value for dry cows. The (lack of) influence of a negative value of OEB in the DVE system has yet to be demonstrated in production trials, but at present a negative OEB is not recommended for dairy cows.

that the efficiency decreases with increasing ratio of protein and energy (DVE/NEL) in the diet (Webster, 1987; Van Straalen et al., 1993a; Subnel and Meijer, 1993). Subnel and Meijer (1993) showed in addition to the DVE/NEL-ratio the level of production is important. Analysis of a large number of feeding trials at INRA showed that the efficiency varies between 0.58 and 0.69. When PDI-intake decreased, generally protein production decreased and efficiency increased, both in a curvilinear manner. The efficiency-factorof 0.64 used in the French system was taken at a point above which little or no extra production of milk protein due to extra intake of PDI is found (V6rit6 et al., 1987), and is called the optimum efficiency. Initially the factor 0.64 was also chosen in the DVE-system, but based on Dutch production trials a new requirement formula was derived in which the effect of level of production on efficiency is incorperated. This part of the system will be discussed in a seperate paper (Subnel et al., 1993). The new DVE-system was accepted in practice. Wether the chosen parameters in the system are well enough adjusted to the Dutch circumstances is being tested in several production trials. The exact level of OEB at which minimal losses of N from the rumen emerge has theoretically been established at zero. Wether this value can be achieved in pratice without any loss of protein production is being evaluated in seperate production trials.

References
AFRC, 1992. Nutritive requirements of ruminant animals: protein. AFRC Technical Committee on Responses to Nutrients. Report no., 9. Nutr. Abstr. Rev. (Series B), 62: 787-835. Aitchison, E., Gill, M. France, J. and Dhanoa, M.S., 1986. Comparison of methods to describe the kinetics of digestion and passage of fibre in sheep. J. Sci. Food Agric., 37: 1065-1072. Alderman, G., 1987. Comparison of rations calculated in different systems. In: R. Jatrige and G. Alderman (Editors), Feed Evaluation and Protein Requirement Systems for Ruminants. Report EUR 10657EN, ECSC-EEC-EAEC, Brussels, pp., 283-298. Allen, J.D. and Harrison, D.G., 1979. The effect of dietary addition of monensin upon digestion in the stomachs of sheep. Proc. Nutr. Soc., 38: 32A. ARC, 1980. The nutrient requirement of ruminant livestock. Commonwealth Agricultural Bureaux, Slough, England, 351 pp. ARC, 1984. The nutrient requirement of ruminant livestock, Suppl. no.l. Commonwealth Agricultural Bureaux, Slough, England, 45 pp.

10.6. D VE-requirements
Different systems use different protein requirements due to differences in diets, protein values of the feeds, types of animals, production levels and type of trials from which values have been derived (N-balances and feeding trials (INRA, 1978) versus factorial approaches (ARC, 1980 and 1984; NRC, 1985)). For maintenance, the French system uses a requirement of 3.25 g PDI per kg metabolizable body weight (MBW), a value also used in the Nordic system (Madsen, 1985). The NRC gives requirements according to Eq. (16). Although all systems use a constant efficiency of N-utilisation for milk production it is known

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Armstrong, D.G. and Hutton, K., 1975. Fate of nitrogenous compounds entering the small intestine. In: I.W. McDonald and A.C.I. Warner (Editors), Digestion and Metabolism in the Ruminant. University of New England, Armidale, N.S.W., pp. 432447. Ausschuss fur Bedarfsnormen, 1986. Energie und Nahrstoftbedarf landwirtschaftliche Nutztiere Nr 3. Milchkuhe und Aufzuchtrinder, DLG, Frankfurt, 92 pp. Boekholt, H.A., 1976. De stikstofhuishouding van het melkrund en de betekenis van de gluconeogenese uit aminozuren. PhD Thesis, Wageningen Agricultural University, 207 pp. Bosch, M.W., 1991. Influence of stage of maturity of grass silages on digestion processes in dairy cows. 4. Protein digestion and microbial protein synthesis in the rnmen. PhD Thesis, Wageningen Agricultural University, 150 pp. Bricenco, J.V., Van Horn, H.H., Harris, B. and Wilcox, C.J., 1988. Comparison of different methods of expressing dietary protein for lactationm dairy cows. J. Dairy Sci., 71: 1647-1658. Broderick, G.A., Ricker, D.B. and Driver, L.S., 1990. Expeller soybean meal and corn by-products versus solvent soybean meal for lactating dairy cows fed alfalfa silage as sole forage. J. Dairy Sci., 73: 435-462. Charmley, E., Veira, D.M. and Aroeira, L., 1988. Effect of inhibiting plant proteolysis, performance and protein digestion in sheep given alfalfa silage. J. Dairy Sci. 71 (Suppl. 1) : 131 ( Abstr. ). Clark, J.H., Klusmeyer, T.H. and Cameron, M.R., 1992. Microbial Protein synthesis and flows of nitrogen fractions to the duodenum of dairy cows. J. Dairy Sci., 75: 2304-2323. Cody, R.F., Murphy, J.J. and Morgan, D.J., 1990. Effect of supplementary crude protein level and degradability in grass silage based diets on performance of dairy cows, and digestibility and abomasal nitrogen flow in sheep. Anim. Prod., 51: 235-244. Coleman, G.S., 1975. The interrelationship between rumen ciliate protozoa and bacteria. In: I.W. McDonald and A.C.I. Warner (Editors), Digestion and Metabolism in the Ruminant, University of New England, Armidale, N.S.W., pp. 149-164. CVB, 1990. Voedernormen voor landbouwhuisdieren. Voederwaarde veevoeders. CVB-series 5, Lelystad, 48 pp. CVB, 1991a. Eiwitwaardering voor herkauwers: Het DVE-systeem. CVB series 7, Lelystad, 53 pp. CVB, 1991b. Veevoedertabel. Gegevens over chemische samenstelling, verteerbaarheid en voederwaarde van voedermiddelen. CVB, Lelystad. Demeyer, D.I. and Tamminga, S., 1987. Microbial protein yield in the rumen and its prediction. In: R. Jarrige and G. Alderman (Editors), Feed Evaluation and Protein Requirement Systems for Ruminants. Report EUR 10657EN, ECSC-EEC-EAEC, Brussels, pp. 129-144. Durand, M. and Komisarczuk, S., 1988. Influence of major minerals on rumen microbiota. J. Nutr., 118: 249-260. Fox, D.G., Sniffen, C.J., O'Connor, J.D., Russell, J.B. and Van Soest, P.J., 1992. A Net carbohydrate and Protein System for Evaluating Cattle Diets; 11I.Cattle requirements and diet adequacy. J. Anim. Sci., 70: 3578-3596. Garnsworthy, P.C., 1989. The interaction between dietary fibre level and protein degradability in dairy cows. Anim. Prod., 48: 271281.

Harrison, D.G. and McAllan, A.B., 1980. Factors affecting microbial growth yields in the retieuloruraen. In: Y. Ruckebuseh and P. Thivend (Editors), Digestive Physiology and Metabolism in Ruminants. MTP Press Ltd., Lancaster, pp. 205-226. Harstad, O.M. and Vik-Mo, L., 1985. Estimation of microbial and undegraded protein in sheep on silage based diets. Aeta Agric. Scand., 25 (Suppl.): 37--48. Hof, G., Kouwenberg, W.J.A. and Tamminga, S., 1990. The effect of washing procedure on the estimation of the in situ disappearance of amino acids from feed protein. Neth. J. Agric. Sci., 38: 719-724. Hvelplund, T., 1984. Intestinal digestion of protein in dairy cows. Can. J. Anim. Sci., 64(Suppl.): 193-194. Hvelplund, T., 1985. The digestibility of rumen microbial protein and undegraded dietary protein estimated in the small intestine of sheep by the in sacco procedure. Acta Scand., 25(Suppl): 132-144. Hvelplund, T., 1986. The influence of diet on nitrogen and amino acid content of mixed rumen bacteria. Acta Agfic. Scand., 36: 325-331. Hvelplund, T., 1987. Amino acid content of feed and microbial protein and their intestinal digestibility. In: R. Jarrige and G. Alderman (Editors), Feed evaluation and protein requirement systems for ruminants. ECSC-EEC-EAEC. Brussels, pp. 159169. Hvelplund, T. and J. Madsen, 1985. Amino acid passage to the small intestine in dairy cows compared with estimates of microbial protein and undegraded dietary protein from analysis on the feed. Acta Agric. Scand., 25 ( Suppl ): 21-36. Hvelplund, T. and Hesselholt, M., 1987. Digestibility of individual amino acids in rumen microbial protein and undegraded dietary protein in the small intestine of sheep. Acta Agric. Scand., 37: 469-477. Hvelplund, T., Madsen, J., Andersen, P.E., Kristensen, V.F., Andersen, R.H. and Foldager, J., 1987. Testing and implementing the modern systems: Denmark. In: R. Jarrige and G. Alderman (Editors), Feed evaluation and protein requirement systems for ruminants. ECSC-EEC-EAEC. Brussels, pp. 237-248. Hvelplund, T. and Madsen, J., 1990. A Study of the Quantitative nitrogen metabolism in the gastro-intestinal tract and the resultant new protein evaluation system for ruminats. The AAT-PBV system. Thesis. Intstitute of Animal Science. The Royal Veterinary and Agricultural University Copenhagen. INRA, 1978. Alimentation des ruminants. Inst. National de la Rech. Agron. Versailles, 597 pp. Kennedy, P.M., Christopherson, R.J. and Milligan, L.P., 1976. The effect of cold exposure of sheep on digestion, tureen turnover time and efficiency of microbial synthesis. Br. J. Nutr., 36: 221242. Kennedy, P.M., Hazlewood G.P. and Milligan, L.P., 1984. A comparison of methods for the estimation of the proportion of microbial nitrogen in duodenal digesta, and of correction for microbial contamination in nylon bags incubated in the rumen of sheep. Br. J. Nutr., 52: 403-417. Ketelaar, G., 1986. Vergelijking van eiwitwaarderingsystemen op basis van produktiegroepen met hoogproduktief melkvee. MSc thesis, Wageningen Agricultural University, 74 pp.

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R~sum6
Tamminga, S., Straalen, W.M. Van, Subnel, A.P.J., Meijer, R.G.M., Steg, A., Wever, C.J.G. et Blok, M.C., 1994. Le syst~me d'6valuation des prot6ines des ruminants: Le syst~me DVE/OEB. Livest Prod. Sci., 40:139-155 (en anglais). En 1991, un nouveau syst~me d'6valuation du taux prot6ique a remplac6 le syst~me DCP aux Pays-Bas: Le Syst&ne DVE/OEB. Ce syst~me a 6t6 d6velopp6 afin de pr6venir les laertes non-necessaires d'azot6 pour ajuster exactement le r6el besoin de la vache laiti~re et de predire plus exactement la production des prot6ines du lalt. Les besoins en prot6ines pour l'entretien, la production laiti~res, la croissance et la gestation sont exprim6s en DVE, concemant les aliments digestibles et les prot6ines microbiennes dans l'intestin gr~le. Pour chaque mati6re premiere, la valeur de la DVE donne la quantit6 de prot6ines disponsibles consistant en prot6ines by-pass ( 1 ), les prot6ines synth6tis6s par les microorganismes du rumen (2), et une correction pour les prot6ines endog~nes perdues par les processes de digestion (3). Pour chaque mati~re premiere, la balance des prot6ines non d6gradables refl6te la diff6rence entre la synth6se prot6ique microbiennes disponsible bas6e sur la d6gradation des prot6ines et l'6nergie disponsible du6 au proc6d6 de fermentation du rumen. Le syst~me DVE/OEB est bas6 sur ce qui est consid6r6 comme les meilleurs 616ments des syst~mes modemes d'6valuation prot6iques des autres pays. Des nouveaux 616ments sont la f6cule r6sistante ~t fermentation dans le rumen (USTA), les produits de fermentation des silages (FP), la relation entre la balance d'6nergie et la disponibilit6 des prot6ines dans l'intestine et la diff6rence des besoins d6pendant de la production des prot6ines de lait. Le syst~me a 6t6 contruit en utilisant principalement les donn6es des recherches N6erlandaises, particuli~rement pour les valeurs prot6iques fourragi~res et sous-produits.

Zusammenfassung
Tamminga, S., Straalen, W.M. Van, Subnel, A.P.J., Meijer, R.G.M., Steg, A., Wever, C.J.G. und Blok, M.C., 1994. Das Niederl~indische Eiweissbewertungssystem: Das DVE/OEB System. Livest Prod. Sci., 40:139-155 (anf englisch) Das DVE/OEB System wurde 1991 an stelle des bis dahin gebr~uchlichen dcp-Systems eingefiihrt als neues Eiweissbewertungssystem. Dieses System wurde entwickelt, um unn0tige Stickstoffveduste zu vermeiden dutch eine genauere Anpassung der Fiitterung an die wirklichen Bedarfsnotmen und um die Milcheiweissproduktion genauer voraussagen zu kSnnen. Der Eiweissbedarf fiir Erhaltung, Milcheiweissproduktion, Wachtstum and Schwangerschaft wird ausgedriickt in DVE, verdauliches Futter- und Mikrobeneiweiss des Diinndarms. Der DVE-gehalt eines jeden Futtermittels repr~,isentiert das verfiigbare Eiweiss, berechnet ans der Summe aas bestiindigem Eiweiss ( 1 ) and microbiellem Eiweiss (2) vermindert um endogene Eiweissveduste in Verdauugsproze (3). Die unbestiindige Eiweissbilanz (OEB) eines jeden Futtermittels spiegelt den Unterschied zwischen einer m0glichen mikrobiellen Eiweisssynthese, basierent auf abbanbarem Eiweiss und verfiigbarer Energie, und die Fermentationsprozessen im Pansen wider. Das DVE/OEB-System beruht auf was andere L~lnder die festen Elemente der modemem Eiweissbewertungssysteme bezeichnen. Neue Elemente sind best~dige St~rke (USTA), Fermentationsprodukte in Silagen (FP), Energiebilanz bei tier Eiweissversorgung and der EiweissVerwertung in abh~ingigkeit der Milchproduktionsmange. Es wurde aufgebaut aus Daten mit haupts~ichlichem Urspnmg in der holl~ndischen Forschung under besonderer Bem'cksichtigung von Eiweissgehalten in Grundfuttermitteln and einige indusaielle Nebenprodukte.