A net carbohydrate and protein system for evaluating cattle diets: II.

Carbohydrate
and protein availability

C. J. Sniffen, J. D. O'Connor, P. J. Van Soest, D. G. Fox and J. B. Russell

J Anim Sci 1992. 70:3562-3577.

The online version of this article, along with updated information and services, is located on
the World Wide Web at:
http://jas.fass.org

www.asas.org

Downloaded from jas.fass.org by on September 2, 2008.

Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
 A Net Carbohydrate and Protein System for
Evaluating Cattle Diets: 11. Carbohydrate
and Protein Availability
C. J. Sniffen*J, J. D. O'Connor*p2, P. J. Van
D. G , Fox*, and J. B. Russell*#+
*Department of Animal Science, Cornell University, Ithaca, NY 14853 and
W.S. Dairy Forage Research Center, ARS, USDA, Madison, WI 53706 and
U.S. Plant, Soil, and Nutrition Laboratory, Ithaca, NY 14853
ABSTRACT: The Cornell Net Carbohydrate and
Protein System (CNCPS) has a submodel that
predicts rates of feedstuff degradation in the
rumen, the passage of undegraded feed to the
lower gut, and the amount of ME and protein that
is available to the animal. In the CNCPS, struc-
tural carbohydrate (SC) and nonstructural carbo-
hydrate (NSC) are estimated from sequential NDF
analyses of the feed. Data from the literature are
used to predict fractional rates of SC and NSC
degradation. Crude protein is partitioned into five
fractions. Fraction A is NPN, which is trichloroa-
cetic (TCA) acid-soluble N. Unavailable or protein
bound to cell wall (Fraction C) is derived from acid
detergent insoluble nitrogen (ADIP), and slowly
degraded true protein (Fraction B3) is neutral
Key Words: Cattle, Nutrition, Models, Feed Composition Tables, Metabolizable Energy,
Metabolizable Rotein
Introduction
The Weende system for proximate analysis and
the TDN system have been used for about a
century as the basis for predicting the energy and
protein available from feedstuffs (Morrison, 1956).
Net energy systems were developed to adjust for
methane, urinary, and heat increment losses
(NRC, 1976, 19781. The NE systems have worked
well under standard feeding conditions, but the
'Present address: W. H. Miner Agric. Res. Inst., Chezy, NY
12921.
'Present address: P. 0. Box 2077, Cowallis, OR 97339.
Received May 24, 1991.
Accepted July 13, 1992.
3562
Downloaded from jas.fass.org by on September 2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
Soest*,
detergent insoluble nitrogen (NDIP) minus Frac-
tion C. Rapidly degraded true protein (Fraction B11
is TCA-precipitable protein from the buffer-soluble
protein minus NPN. True protein with an inter-
mediate degradation rate (Fraction B2) is the
remaining N. Protein degradation rates are esti-
mated by a n in vitro procedure that uses Strep-
tomyces griseus protease, and a curve-peeling
technique is used to identify rates for each
fraction. The amount of carbohydrate or N that is
digested in the rumen is determined by the
relative rates of degradation and passage. Rumi-
nal passage rates are a function of DMI, particle
size, bulk density, and the type of feed that is
consumed (e.g., forage vs cereal grain).
J. Anim. Sci. 1992. 70:3562-3577
tabular value of NE for a particular feed is
typically computed from TDN and represents the
average expected value based on a group of feeds
rather than the NE that will be derived by a
particular group of cattle eating that feed. Be-
cause the feeding conditions of cattle are variable
and often unique, accurate NE values usually are
not available N a n Soest et al., 1984). Crude and
digestible protein determinations do not com-
pletely account for the dynamics of ruminal
fermentation and the potential loss of nitrogen as
ammonia. The Cornell Net Carbohydrate and
Protein System (CNCPS) has equations that esti-
mate the fermentation and passage of feed carbo-
hydrate and protein fractibns. This information
can be used as a basis for predicting ME and
protein absorption.
 PREDICTING FEED ENERGY AND PROTEIN VALUES
Fermentation, Passage, and Absorption
Van Soest et al. (1984)summarized the effects of
variations in intake. feed composition, ruminal
digestion rates, and passage rates on the NE and
protein values of feeds and developed discount
factors to adjust for these effects. The Van Soest
discount system works well under a variety of
feeding conditions. Static adjustments, however,
cannot fully accommodate the dynamics of rumi-
nal fermentation (Russell et al., 1992). New protein
systems (NRC, 1985,1989)have attempted to define
the amount of protein escaping ruminal degrada-
tion, but these systems have 1) a single N pool; 2)
used TDN or OM rather than ruminally degraded
carbohydrates to control microbial protein synthe-
sis; 3) not partitioned ruminal microorganisms
according to carbohydrate and N utilization; 4)not
accommodated the effect of growth rate, amino
acid availability, or pH on the efficiency of
microbial growth; and 5)not integrated the inverse
relationship between rates of carbohydrate and
protein fermentation.
In a companion paper (Russell et al., 1992) we
have demonstrated that the CNCPS equations
could be used to predict the growth of ruminal
bacteria accurately so long as 1) carbohydrates
were partitioned into structural (SC) and nonstruc-
tural (NSC) components, 2) degradation rates of
these components could be estimated, 3) N availa-
bility could be described in terms of ammonia and
peptides, and 4) changes in ruminal pH could be
accommodated by variations in diet composition.
In this paper we provide 1)tables of feed composi-
tion that partition feedstuffs into carbohydrate
and protein fractions, 2) methods for calculating
the amount of N and carbohydrate that can be
used to drive ruminal microbial growth, and 3)
mechanistic equations for predicting metaboliza-
ble energy and protein values that are based on
ruminal rates of digestion and passage, microbial
growth kinetics, and postruminal digestibilities.
Feed Fractions
The CNCPS assumes that feedstuffs are com- the soluble protein in common feedstuffs is
posed of protein, carbohydrate, fat, ash, and
water. Protein and carbohydrate DM are further
subdivided by chemical composition, physical
characteristics, ruminal degradation, and postru-
minal digestibility characteristics (Table 1). The
values presented in Table 1 are from NRC (1982, of the total soluble protein (approximately 5%) and
1984,19891,Krishnamoorthy et al. (19821,Van Soest
et al. (19841,Abdalla et al. (1988a,b),and estimates
by the authors based on the above and on our
unpublished data. Fractions of protein, carbohy-
drate, ash, fat, and water in a feedstuff can be
Downloaded from jas.fass.org
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3563
derived from the following standard laboratory
chemical analyses:
1. DM of the feed (AOAC, 1980);
2. NDF and lignin (Van Soest et al., 1991);
3. Total nitrogen as assayed by macro- or micro-
Kjeldahl (AOAC, 1980);
4. Soluble protein as determined by the proce-
dure of Krishnamoorthy et al. (1983);
5. Nitrogen that is insoluble in neutral deter-
gent (without sodium sulfite) and acid deter-
gent (Van Soest et al., 1991);
6. Ash (AOAC, 1980);
7. Solvent-soluble fat (AOAC, 1980); and
8. Computation of NSC from the determinations
of NDF, protein, fat, and ash or directly (Van
Soest et al., 1991): 100 - NNDF - NDF protein)
+ fat + ashl.
Feed Protein Fractions
Each feed is further described by the proportion
of NDF that is effective in meeting fiber require-
ments because of particle size, based on the values
of Mertens (1985).
Feed protein is partitioned into three fractions:
nonprotein nitrogen (NPN), true protein, and un-
available nitrogen (Van Soest et al., 1981). These
have been described as Fractions A (NPN),B (true
protein), and C (bound true protein), respectively
(Pichard and Van Soest, 1977). True protein is
further fractionated into three subfractions (Bl,
B2, and B3) based on their inherent rates of
ruminal degradation (Van Soest et al., 1981;
Krishnamoorthy et al., 1983). Roe et al. (1990)
presented a summary of recommended procedures
to determine protein fractions. Fractions A and B1
are soluble in buffer (Roe et al., 1990) and B1 is
determined as the TCA-precipitable fraction (Van
Soest et al., 1981; Krishnamoorthy et al., 1983).
Nonprotein N (ammonia, peptides, amino acids) is
rapidly converted to ammonia in the rumen.
Essentially all the soluble protein in silages and
cut forages is in the form of NPN (Pichard, 1977;
Pichard and Van Soest, 1977). The NPN content of
presented in Table 1.
Fraction B is subdivided to estimate rates of
ruminal degradation. Fraction B1 is rapidly
degraded in the rumen (Van Soest et al., 1981). In
harvested forages Fraction Bl is a small fraction
concentrates can contain twice as much Fraction
B1 as forages do (Pichard, 1977;Krishnamoorthy et
al., 1982). Most of the soluble protein in fresh
pastures is Fraction B1 (Van Soest, 1982). In the
CNCPS
2, 2008. B1 is degraded in the rumen.
Fraction
by on September
3564 SNIFFEN ET AL.
Table 1. Carbohydrate and protein fractions, fat, and ash in common feedsa

NDF, Lignin, NSP, Starch, CP, Soluble, ADFIP, NPN, NDFIP, Fat, Ash,
Yo of O
h of % of % of % of Yo of Yo of o/o of Yo of Yo of O h of
Feedstuff DM NDF DM NSC DM CP CP SP CP DM DM
Grains
Barley heavy 19.0 10.5 4 90 13.0 17.0 6.4 29.4 8.0 2.1 2.6
Beet pulp 54.0 3.7 30 90 9.7 26.5 11.0 97.4 52.6 .e 4.4
Canola meal 18.2 19.4 90 42.3 32.4 6.4 85.0 10.6 1.2 7.8
Corn dry grain 9.0 11.0 2 90 10.1 11.0 5.0 70.0 15.0 4.3 1.6
Corn HM grain 9.0 11.0 2 90 10.1 40.0 5.3 100.0 15.9 4.3 1.6
Corn flaked grain 9.0 11.0 2 90 10.1 8.0 5.0 80 15.0 4.3 1.6
Corn dry ear 28.0 7.1 2 90 9.0 16.0 7.8 70.0 17.8 3.7 1.9
Corn HM Ear 28.0 7.1 2 90 9.0 30.0 8.3 100.0 18.7 3.7 1.9
Corn hominy 55.0 3.6 90 11.5 18.0 3.0 80.0 7.8 7.7 3.1
Milo ground 23.0 5.0 90 12.5 12.0 4.8 30.0 10.0 3.1 2.1
Milo HM or flake 23.0 5.0 100 12.5 30.0 5.1 100.0 10.1 3.3 2.2
Molasses beet 0 0 0 8.7 100.0 0 100.0 0 .2 11.3
Molasses cane 0 0 0 5.8 100.0 0 100.0 0 .1 13.1
Oats 489 g/L 32.0 9.4 4 90 13.3 53.1 5 .O 18.5 11.0 5.4 3.4
Oats 412 g/L 42.0 9.4 4 90 13.1 53.1 5.0 18.5 11.0 4.9 4.6
Soybean hulls 67.0 3.0 14 0 12.1 17.9 14.0 60.8 20.0 2.1 5.1
Wheat ground 16.0 6.3 2 90 16.0 30.0 2.0 25.0 4.0 2.0 1.9
Wheat middlings 37.0 5.9 2 90 18.4 40.0 2.6 30.0 4.0 4.9 5.2
Protein concentrates
Alfalfa meal 45.0 24.4 7 90 18.9 28.0 17.1 100.0 25.0 3.0 10.6
Bloodmeal 0 0 0 91.7 4.9 1.2 4.9 0 2.4 10.2
Brewers grain dry 46.0 13.0 100 25.4 4.1 12.0 70.7 40.4 6.5 4.8
Brewers grain wet 42.0 11.9 100 29.2 8.0 10.0 50.0 38.0 6.5 4.8
Corn distill. dry 44.0 9.1 100 29.5 22.0 20.0 77.3 63.1 10.3 5.1
Corn distill. wet 40.0 9.1 100 29.5 25.0 12.0 65.6 54.8 9.9 4.8
Corn gluten feed 45.0 2.1 100 25.6 49.0 2.1 100.0 7.8 2.4 7.5
Corn gluten meal 14.0 7.1 100 65.9 4.2 2.0 71.4 11.0 2.4 1.8
Cottonseed whole 44.0 22.7 90 23.0 40.0 6.0 2.0 6.0 20.0 4.8
Cottonseed meal 29.0 25.0 90 44.8 20.0 7.6 40.0 10.0 1.3 6.3
Feathermeal 10.0 10.0 90 88.6 3.8 2.4 4.9 50.0 3.2 3.8
Fishmeal 2 .o 0 90 66.6 12.0 .9 0 1.o 5.1 25.4
Linseed meal 25.0 24.0 25 0 38.3 20.0 2.4 50.0 10.0 1.5 8.5
Soybean meal 44 14.0 3.0 29 0 49.0 20.0 2.0 55.0 5.0 1.5 7.3
Soybean meal 49 8.0 3.0 29 0 55.1 20.0 2.0 55.0 5.0 1.0 6.5
Soybeans raw 13.4 10.0 19 0 42.8 44.2 2.9 22.6 4.4 18.8 5.8
Soybeans heated 13.4 10.0 19 0 42.8 5.7 7.3 100.0 23.6 18.8 5.8
Sunflower meal 40.0 30.0 0 25.9 30.0 4.8 36.7 7.7 1.2 6.3
Urea 0 0 0 281.0 100.0 0 100.0 0 0 0
Hay crop forages, northb
Alfalfa hay prebl 39.0 16.6 7 10 21.7 30.0 10.0 98.0 15.0 3.0 10.0
Hay earlybl 42.0 16.9 7 10 19.0 30.0 10.0 96.0 17.8 3.2 9.0
Si1 earlybl 42.0 16.9 7 10 10.0 50.0 15.0 100.0 26.7 3.2 9.0
Hay midbl 46.0 18.9 7 10 17.0 28.0 14.0 98.0 25.2 2.6 9.0
Si1 midbl 46.0 18.9 7 10 17.0 45.0 18.0 100.0 32.0 2.6 9.0
Hay mature 55.0 22.2 7 10 12.0 25.0 20.0 92.0 35.6 1.6 8.0
Pasture spg 33.0 8.0 8 28.0 46.0 2.2 2.2 10.0 2.7 10.0
Pasture sum 38.0 8.5 8 24.0 46.0 3.0 2.2 12.0 2.7 9.8
Grass hay late veg 55.0 5.5 1 6 16.0 25.0 5.7 96.0 31.0 2.6 7.2
Hay midbl 67.0 7.5 1 6 9.1 25.0 6.1 96.0 31.0 2.6 6.3
Hay mature 72.0 12.5 1 6 7.0 25.0 6.5 96.0 31.0 2.6 6.0
Pasture spg 50.0 6.0 1 5 24.0 41.0 2.0 2.4 14.5 3.7 10.7
Pasture s u m 55.0 7.0 1 5 15.0 42.0 2.2 4.8 24.0 3.7 9.0
Pasture fall 48.0 6.5 10 5 22.0 43.0 2.0 2.3 18.4 3.7 10.0
Hay crop forages, southb
Alfalfa hay prebl 37.0 18.9 7 10 27.0 30.0 10.0 96.0 15.0 4.0 10.2
Hay earlybl 40.0 20.0 7 10 25.0 30.0 10.0 96.0 17.8 3.0 9.6
Hay midbl 44.0 22.7 7 10 22.0 28.0 14.0 96.0 25.2 2.6 9.1
Hay mature 58.0 24.8 7 10 14.0 25.0 20.0 92.0 35.0 2.0 8.9

Downloaded from jas.fass.org by on September 2, 2008.

Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
 PREDICTING FEED ENERGY AND PROTEIN VALUES 3565
Table 1 (continued). Carbohydrate and protein fractions, fat, and ash in common feedsa

NDF, Lignin, NSP, Starch, CP, Soluble, ADFIP, NPN, NDFIP, Fat, Ash,
% of % of % of % of % of 96 of % of Yo of % of % of Yo of
Feedstuff DM NDF DM NSC DM CP CP SP CP DM DM
Bermuda hay late veg 70.0 8.e 1 6 10.0 25.9 8.9 25.4 34.2 2.5 8.0
Fescue, K31, hay 63.0 6.3 1 6 16.4 25.9 8.9 25.4 34.2 8.1 9.0
Hay full bloom 67.0 7.5 1 6 12.1 25.9 8.9 25.4 34.2 5.3 8.0
Hay mature 70.0 10.0 1 6 9.2 25.9 8.9 25.4 34.2 4.3 7.0
Grain crop forages, northb
Corn silage 45% gr. 41.0 7.3 100 9.0 45.0 7.9 100.0 16.4 3.2 5.0
Corn silage 35Y0 gr. 46.0 8.7 100 8.6 50.0 8.0 100.0 16.0 2.8 7.0
Corn silage 259'0 gr. 52.0 9.6 100 8.3 55.0 8.5 100.0 16.0 2.1 8.0
Grain crop forages, southb
Corn silage 40% gr. 45.0 9.0 100 9.2 45.0 7.9 100.0 16.4 3.1 4.0
Corn silage 25% gr. 55.0 10.9 100 8.1 50.0 8.0 100.0 16.0 2.1 7.0
'NSP is nonstructural polysaccharides (pectin, galactins, fructans, betaglucans, etc.), NSC is nonstructural carbohydrates,
ADFIP is acid detergent insoluble protein, SP is soluble protein, NDFIP is neutral detergent insoluble protein. See Van Soest and
Fox (1992)for a more complete feed library containing these chemical composition values.
bDifferences between north and south are primarily a function of degree days and growing season temperatures. The north
values are typical of northwestern, corn belt, lake and northeastern states in the United States, and values for north grasses are
representative of cool-season grasses.

Unavailable or bound protein, Fraction C, is the thy et al., 1982). Prolamin proteins, such as zein
protein that is insoluble in the acid detergent (acid protein in corn, are found in Fraction B3 (Van
detergent insoluble protein, ADIP) fraction Soest et al., 1981). In the CNCPS, a high percen-
(Pichard and Van Soest, 1977).Fraction C contains tage of Fraction B3 escapes degradation in the
protein associated with lignin, tannin-protein com- rumen.
plexes, and Maillard products that are highly Buffer insoluble protein minus the protein in-
resistant to microbial and mammalian enzymes soluble in neutral detergent is used to estimate
(Krishnamoorthy et al., 1982, 1983). Fraction C Fraction B2. Some Fraction B2 is fermented in the
cannot be degraded by ruminal bacteria and does rumen and some escapes to the lower gut. The fate
not provide amino acids postruminally (Krish- of Fraction B2 depends on the relative rates of
namoorthy et al., 1982). The Fraction C content of digestion and passage. Fraction B2 is typified by
feedstuffs has been measured by several research- the glutelin protein found in small grains (Van
ers (Goering and Adams, 1973; Pichard and Van Soest et al., 19811.
Soest, 1977; Waldo and Goering, 1979; Krish- The following equations can be used to calcu-
namoorthy et al., 1982; Muscato et al., 1983; Van late the five protein fractions contained in the jth
Soest and Sniffen, 1984). At least five common feedstuff from the values given in Table 1:
feeds may contain important amounts of protein in
the bound or indigestible form: 1) haycrop silages,
2) dehydrated alfalfa, 3) citrus pulp, 41 corn
distillers grains, and 5 ) brewers dried grains
(Waldo and Goering, 1979). Table 1 contains
estimates of the Fraction C content of common
feedstuffs.
Fraction B3 is insoluble in neutral detergent but
soluble in acid detergent (neutral detergent insolu- where CPj(%DM)= percentage of crude protein of
ble protein [NDIPI minus ADIP; Goering and Van the jth feedstuff; NPNj(%CP) = percentage of
Soest, 1970; Krishnamoorthy et al., 1982). Fraction crude protein of the jth feedstuff that is non-protein
B3 is slowly degraded in the rumen because it is nitrogen x 0.25; SOLPj(%CP)= percentage of the
associated with the cell wall (Pichard, 1977; Van crude protein of the jth feedstuff that is soluble
Soest et al., 1981; Krishnamoorthy et al., 1983). The protein; NDIPj(%DMl = percentage of the jth
NDIP and ADIP contents of common feedstuffs feedstuff that is neutral detergent insoluble pro-
have been measured (Krishnamoorthy et al., 1982; tein; ADIPj(%DM)= percentage of the jth feedstuff
Muscato et al., 1983). Protein supplements contain that is acid detergent insoluble protein; PAj(%CP)
a small amount of Fraction B3, but forages, = percentage of crude protein in the jth feedstuff
fermented grains, and byproduct feeds contain that is non-protein nitrogen; PBIj(%CP) = percen-
significant amounts of Fraction B3 Downloaded
(Krishnamoor- tage
from jas.fass.org of crude
by on September protein in the jth feedstuff that is
2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3566
rapidly degraded protein; PB2 (%CPl = percen-
h
tage of crude protein in the jt feedstuff that is
intermediately degraded protein; PB3j(%CPl =
percentage of crude protein in the jth feedstuff that
is slowly degraded protein; and PCj(%CP) =
percentage of crude protein in the jth feedstuff that
is bound protein.
Feed Carbohydrate Fractions
Given the crude protein, fat, and ash content of
a feedstuff, expressed as a percentage of DM, the
total carbohydrate (CHO)content in the feedstuff
can be estimated, calculated by difference (100 -
CP - fat - ash). Carbohydrates can then be
classified according to degradation rate, as with
protein (Fraction A is fast and is sugars; Fraction
B1 is intermediate and is starch; Fraction B2 is
slow and is available cell wall; and Fraction C is
unavailable cell wall). These fractions are com-
puted from feed content of NSC, SC, and indigesti-
ble fiber (C). The fraction CHO C is lignin x 2.4
(Smith et al., 1972; Mertens, 19731, the material
remaining after 72 h of digestion in vitro (Mertens,
1973). The lignin content of forages ranges from 5
to 25% of the plant cell wall; legumes have a
higher content than do grasses (Van Soest, 1982).
The NDF component includes cellulose, hemicellu- where CPj(%DMl = percentage of crude protein of
lose, and lignin (Goering and Van Soest, 1970). the jth feedstuff; CHOj(%DM) = percentage of
Available SC (Fraction B2) can be determined by
subtracting the fraction CHO C from ash-free NDF
that has been corrected for associated protein.
Fraction B2 is slowly fermented in the rumen by
bacteria that require ammonia as their sole
nitrogen source (Russell et al., 1992).
Nonstructural carbohydrates contain sugars
(Fraction A) and starch and pectin (Fraction Bl).
The NSC fraction represents the carbohydrates
that are soluble in neutral detergents and it can be
estimated as 100 minus protein, NDF corrected for
protein, lipids, and ash in a feedstuff. The NSC can
also be measured directly (MacGregor et al., 1983).
The calculated NSC is usually in close agreement
with direct measurements, but feeds containing
large amounts of pectin will have a somewhat
lower true NSC. Given the proportion of starch
and pectin in the NSC, the Fraction A (sugars and
organic acids) can be determined by difference.
The sugar content of most ruminant diets is
normally quite low unless sugar byproduct feeds
or fresh, lush grasses are fed (Van Soest, 1982;
Sniffen et al., 1983). Starch is a major component
of cereal grains and is a major component of
forages. Sugars are fermented rapidly by ruminal
microorganisms. Starches from ensiled and
processed grains are rapidly digested in the
rumen, but dried grains can contain a significant
Downloaded from jas.fass.orgples
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
SNIFFEN ET AL.
amount of insoluble starch, which is digested
slowly. Pectins are unimportant in grasses and
cereals, but legume forages, seed products, citrus
pulp, and beet pulp contain significant amounts of
pectin (Van Soest, 1982). Pectins are rapidly
fermented in the rumen (Van Soest, 1982). All NSC
fractions are fermented by ruminal bacteria that
can utilize either ammonia or peptide as a
nitrogen source (Russell et al., 1992).
Using the chemical analyses described above,
equations used to calculate carbohydrate composi-
tion (Table 1) of the jth feedstuff are listed below:
CHOj(%DM)= 100 - CPj(%DM)- FATj(%DMl
- ASHj(YoDM1
CCj(YoCH0) = 1OO*(NDFj(%DMl* . O 1 +LIG-
NINj(%NDFl*2.4)/CHOj(O/oDM)
CB2j(%CHOl = 1OO*((NDFj(%DMl -
NDIPj(%CP)* .O1 * CPj(%DMl
- NDFj(%DM).O 1 * LIGNINj(%
NDFI 2.4l/CHOj(%DM))
CNSCj(%CH-= 100 - B2j(%CHO)- Cj(%CHO)
0)
CBj(%CHOl = STARCHj(%NSC)* (100
- B2j(%CHO) - Cj(%CH0))/1OO
CAj(%CHO)= (100 - STARCHj(%NSC)l~(lOO
- B2j(%CHO)- Cj(%CH0))/1OO
carbohydrate of the jth feedstuff; FATj(%DM) =
percentage of fat of the jth feedstuff; ASHj(%DM)
= percentage of ash of the jth feedstuff;
NDFj(%DMl = percentage of the jth feedstuff that
is neutral detergent fiber; NDIPj(%DM)= percen-
tage of neutral detergent insoluble protein of the
jth feedstuff; LIGNIN~WONDF) = percentage of
lignin of the jth feedstuffs NDF; STARCHj(YoNSC1
= percentage of starch in the nonstructural
carbohydrate of the jth feedstuff; SUGARj(%NSC)
= percentage of sugar in the nonstructural carbo-
hydrate of the jth feedstuff; CA(%CHO)= percen-
tage of carbohydrate of the feedstuff that isjh
sugar; CBlj(%CHO)= percentage of carbohydrate
of the jth feedstuff that is starch + NSP;
CB2j(%CHO)= percentage of carbohydrate of the
jth feedstuff that is available fiber; and CCj(%CHO)
= percentage of carbohydrate in the jth feedstuff
that is unavailable fiber.
Carbohydrate and protein fractions in feeds are
then summed to determine the intake of each
fraction.
Ruminal Kinetics
Sophisticated, dynamic models of the rumen
that incorporate many of the mechanistic princi-
of rumen
by on September 2, 2008. function have been developed
 PREDICTING FEED ENERGY AND PROTEIN VALUES
(Baldwin et al., 1977; Mertens and Ely, 1979; France
et al., 1982; Gill et al., 1984) and some of the
principles used in these earlier models have been
incorporated into this system. Most models of
ruminant digestion assume that nutrient ingestion
occurs on a steady-state, continuous basis and
that digestion rates follow fist-order kinetics
(Waldo et al., 1972; Mertens, 1973; Mertens and
Ely, 1979; Van Soest et al., 1981; Krishnamoorthy
et al., 1983; Ewing and Johnson, 1987). Based on
the model of Waldo et al. (19721, rumen digestibil-
ity (RD) is defined as the specific rate of rumen
digestion (Kdl divided by the specific rate of
disappearance due to digestion and passage (Kd +
Kpl: RD = Kd/(Kd + Kp). Ruminal escape (RE) is
defined as RE = Kp/(Kd + Kp).
Ruminal Degradation and Passage Rates
Ruminal passage rate constants (Kp) are shown
in Tables 2 and 3, based on the work of Hartnell
and Satter, 1979; Colucci et al., 1982; and Erdman
et al., 1987. Recent research (Welch, 1986) has
shown that particle size, density, and hydration
rate can affect the passage rates of feeds, and
these effects were incorporated into the passage
rate estimates, based on judgments made by the
authors. Passage rates are also influenced by the
level of intake, and adjustments for intakes above
maintenance are presented in Tables 2 and 3.
These rates assume additivity across feeds in a
particular diet. The use of variable ruminal pas-
sage rates for each ingredient in the diet provides
a method for estimating variations in their rumi-
nal digestibility. As rates of passage increase, the
extent of ruminal digestion and energy availability
are reduced (Van Soest et al., 1984).
The model computes Kp for each feed ingredient
as described by Chalupa et al. (1991):
Kp [forages] = .388 + (.002.DMI/BW.75)
+ (.00024forage O!O in DM21
Kp [concentratesl = -.424 + (1.45 Kp[foragel1 e REPB2j = DIET
where DMI is in grams and BW.75 is in kilograms.
The Kp is adjusted (AD for particle size using the
effective NDF (ENDF) values in Tables 2 and 3:
fif[forages] = lOO/[ENDF + 701
Af [concentratesl = lOO/(ENDF + 90).
Protein Degradation and Passage Rates
Potentially degradable true protein, the protein
B fractions, range in degradability from .04 to
26Oo/o/h across most common feeds (Tables 4 to 61.
Residual protein corrected for bacterial
Downloadedprotein
from jas.fass.org by on September 2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3567
can be measured by enzymatic or in situ digestion
(Nocek, 1985; Nocek and Russell, 1988). After the
residual N is plotted as a natural logarithm vs
time (Pichard and Van Soest, 19771, curve-peeling
techniques can be used to estimate the digestion
rate of each fraction. However, peptides arising
from protein degradation are only utilized by
ruminal microorganisms at a rate of .07 g of
peptide per gram of microorganism per hour
(Russell et al., 19921. When the degradation of the
protein is rapid, peptides accumulate and small
amounts of peptides as well as protein can escape.
The following equations calculate the amounts
of protein fractions that are ruminally degraded:
RDPAj = DIET PAj
RDPBlj = DIET PBIj.(Kdlj/(Kdlj + KPjN
RDPB2j = DIET PB2j * (Kdzj/&dzj + Kpj1)
RDPB3j = DIET PB3j.(Kdaj/(Kd3j + KpjN
RDPEPj = RDPBlj + RDPB2j + RDPB3j
where Kdlj = rate of ruminal digestion of the
rapidly degraded protein fraction of the jthfeed-
stuff, h-l; Kd2j = rate of ruminal digestion of the
intermediately degraded protein fraction of the jth
feedstuff, h-l; Kd3j = rate of ruminal digestion of
the slowly degraded protein fraction of the jth
feedstuff, h-1; Kpj = rate of passage from the
rumen of the jth feedstuff, h-l; RDPAj = amount of
ruminally degraded NPN in the jth feedstuff, g/d;
RDPBIj = amount of ruminally degraded B1 true
protein, in the jth feedstuff, g/d; RDPB2j = amount
of ruminally degraded B2 true protein, in the jth
feedstuff, g/d; RDPB3i = amount of ruminally
degraded B3 true protein in the jth feedstuff, g/d;
and RDPEPj = amount of rumen degraded pep-
tides from the jth feedstuff, g/d.
The undegraded protein is passed to the small
intestine. The following equations calculate the
amount of each protein fraction that escapes
ruminal degradation:
REPBlj = DIET PBIj*(Kpj/(KdIj+ KPj))
PB2j*(Kpj/(Kdzj+ KPj1)
REPB3j = DIET PB3j*(Kpj/(Kd3j+ Kpjl1
REPCj = DIET PCj
where
REPBlj = anount of ruminally escaped B1 true
protein, in the jth feedstuff, g/d;
REPB2j = amount of ruminally escaped B2
true protein, in the jth feedstuff, g/d;
REPB3j = amount of ruminally escaped B3
true protein in the jth feedstuff, g/d;
and
REPCj = amount of rumen escaped bound C
protein from the jth feedstuff, g/d.
3568
Carbohydrate Degradation and Passage Rates
As with protein, the model assumes that all feed
carbohydrate must disappear through passage or
digestion, and that the kinetics are first-order.
Because sugars, the Fraction CHO A, are fer-
mented very rapidly in the rumen, 300%/h (Sniffen
et al., 19831, virtually none of the sugar escapes
ruminal degradation. More slowly fermented but
potentially available carbohydrates, the Fractions
CHO B1 and B2, are digested a t rates varying from
2 to 50%/h, and some of this material may escape
ruminal degradation. Tables 4, 5, and 6 list logical
ranges for rates of ruminal fermentation of the
carbohydrate fractions for common grains, protein
supplements, and forages. Carbohydrate rates,
like protein rates, can be obtained using in situ
procedures (Nocek, 1985; Nocek and Russell, 1988).
It is not possible at this time to obtain reliable
estimates of digestion rates for carbohydrates by
enzymatic procedures.
Digestion rates for the Fraction CHO B1 are
typically 3 to 8%/h. Pectins are rapidly digested,
but certain types of starch can be degraded slowly.
Uncooked starches ranked in the order of decreas-
ing rate of digestion are wheat, barley, oats, corn,
sorghum, and legume (Van Soest, 1982).
The Fraction CHO B2 of legumes is generally
degraded at a faster rate than in grasses (Van
Soest, 1982; Varga and Hoover, 1983). The Fraction
CHO B2 of mature grains has a mean digestion
rate of 5.l%/h (Smith et al., 19721, and feeds with
50 to 60% CP (corn gluten meal, soybean meal, and
peanut meal) had rate constants of 4.8 to 5.4%/h
(Varga and Hoover, 1983). Protein sources with 25
to 30% CP (distillers grains, corn gluten feed, and
brewers grains) had more rapid rates, 6.5 to 7.2%/
h (Varga and Hoover, 19831. Corn and corn
byproducts (hominy, corn gluten feed, and corn
gluten meal) all had lower rates than wheat
products [middlings and bran) or barley feeds
(barley and brewers grains) (Varga and Hoover,
1983).
The following equations are used to calculate
the amounts of each of the carbohydrate fractions
of the jth feedstuff that are ruminally digested:
RDCAj = DIET CAj'(Kd4j/(Kd4j + KPjN
RDCBlj = DIET CBlj.(Kd5j/KdSj + Kpj))
RDCBPj = DIET CBSj.(Kdsj/(Kdej+ Kpj))
where K&j = rate of ruminal sugar digestion of
the jth feedstuff, h-l; Kdsj = rate of ruminal starch
digestion of the jth feedstuff, h-l; K&j = rate of
ruminal available fiber digestion of the jth feed-
stuff, h-l; RDCAj = amount of ruminally degraded
sugar from the jth feedstuff, g/d; RDCBlj =
amount of ruminally degraded starch fromfrom
Downloaded thejas.fass.org
jth
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
SNIFFEN ET AL.
feedstuff, g/d; and RDCB2j = amount of ruminally
degraded available fiber from the jth feedstuff, g/d.
The following equations are used to calculate
the amounts of each of the carbohydrate fractions
of the jth feedstuff that escape the rumen:
RECAj = DIET CAj.(Kpj/(Kd4j + KPjN
RECBlj = DIET CBlj*(Kpj/(Kd5j+ KpjN
RECB2j = DIET CBzj.(Kpj/(Kdej + Kpj))
RECCj = DIET CCj
where RECAj = amount of ruminally escaped
sugar from the jth feedstuff, g/d; RECBIj =
amount of ruminally escaped starch from the jth
feedstuff, g/d; RECB2j = amount of ruminally
escaped available fiber from the jth feedstuff, g/d;
and RECCj = amount of ruminally escaped un-
available fiber from the jth feedstuff, g/d.
Microbial Flow to the Small Intestine
All microorganisms that pass from the rumen
are assumed to be bacteria, and microbial tur-
nover (starvation and predation) is accounted by
variations in the yield coefficient (Russell et al.,
1992). Based on the microbial composition given in
a companion paper, bacterial fractions escaping
the rumen are computed as follows:
REBTPj = .60..625. BACTj
REBCWj = .25 .625. BACTj
REBNAj = .15 * .625 *BACTj
REBCHOj = .21.BACTj
REBFATj = .07*BACTj
REBASHj = .044*BACTj
where BACTj 7 amount of bacterial total protein
passed to the intestines by the jth feedstuff, g/d;
REBTPj = amount of bacterial true protein passed
to the intestines by the jth feedstuff, g/d; REBCWi
= amount of bacterial cell wall protein passed to
the intestines by the jth feedstuff, g/d; REBNAj =
amount of bacterial nucleic acids passed to the
intestines by the jth feedstuff, g/d; REBCHOj =
amount of bacterial carbohydrate passed to the
intestines by the jth feedstuff, g/d; REBFATj =
amount of bacterial fat passed to the intestines by
the jth feedstuff, g/d; and REBASHj = amount of
bacterial ash passed to the intestines by the jth
feedstuff, g/d.
Intestinal Protein Absorption
The absorption of bacterial N (true protein,
nucleic acid, and cell wall protein) and escaped
protein (B1,
by on September B2, B3, and Cl is calculated by
2, 2008.
 PREDICTING FEED
multiplying each fraction by its respective digesti-
bility. The model uses intestinal true digestibilities
of 100, 100, 100, and 8O%, respectively, for the
peptide, B1, B2, and B3 protein fractions, based on
data summarized by Van Soest (1982).The ADIP or
C protein fraction is completely unavailable for
digestion and does not contribute to absorbable
amino acids (NRC, 1985). Because feed protein
appearing in the feces (insoluble NDIP and ADIP
protein that is either keratin, Maillard products, or
bound to lignin) is resistant to peptic digestion, it
seems that feces contain little true protein (Van
Soest, 1982).
The availability of microbial N has not been
directly determined, but Storm et al. (1983) and Tas
et al. (1981) estimated that bacterial true protein
was 85 and 87% digestible, respectively. The
ENERGY AND PROTEIN VALUES 3569
CNCPS assumes that the total tract digestibility of
bacterial true protein is 100Y0. The literature is
inconsistent relative to the digestibility and path-
ways of nucleic acid utilization by cattle. The true
digestibilities of ruminal bacterial RNA and DNA
have been reported to be 89 and 8O%, respectively
(Storm et al., 19831 and 87 and 819'0, respectively
(Smith and McAllan, 1971). In the CNCPS,nucleic
acid is entirely digested postruminally, but the
absorbed nucleic acid is excreted in the urine
(Smith, 1969). Bacterial cell wall protein is not
released by proteolytic enzymes in the abomasum
or small intestine and seems to have little nutri-
tional value (Mason and White, 1971; Mason and
Palmer, 1971).Therefore, bacterial cell wall protein
is completely unavailable for digestion and does
not contribute to the absorbed amino acid pool.

Table 2. Ruminal passage rate constants for concentrates (%/hJa

Level of maintenanceb Effective NDFC

Ingredient lx 2x 3x % of NDF
Lightweight concentrates
Dried brewers grains 2.0 2.5 3.0 18
Wheat middlings 2.0 2.5 3.0 2
Soybean mill feed 1.0 2.0 3.0 33
Citrus pulp 1 .o 2.0 2.5 33
Beet pulp 1 .o 2.0 2.5 33
Wheat bran 2.0 2.5 3.0 33
Whole cottonseed 1.5 2.0 2.5 100
Whole soybeans 1.5 2.0 2.5 100
Dehy alfalfa 2.0 2.5 3.0 6
Corn cobs, ground 2.0 3.0 3.0 56
Intermediate-weight concentrates
Ground barley 2.5 3.0 3.5 34
Ground wheat 2.5 3.0 3.5 34
Ground oats 2.5 3.0 4.0 34
Fish meal 2.5 3.5 4.0 9
Hominy feed 2.5 3.0 3.5 9
Distillers, w/sol 3.0 3.5 4.0 4
Corn and cobmeal 2.5 3.0 3.5 56
Blood meal 2.5 3.5 4.0 9
Heavyweight concentrates
Whole dry corn 2.5 4.0 6.5 100
Corn meal 3.0 4.0 6.0 48
Cracked corn 3.5 4.O 5.0 60
High-moisture corn
Whole 3.5 4.0 5.0 100
Coarsely rolled 2.5 3.0 4.0 70
Intermediately rolled 2.0 2.5 3.0 60
Finely rolled 3.0 4.0 5.0 48
Soybean meal 3.5 4.0 5.0 23
Cottonseed meal 3.5 4.0 5.0 36
Corn gluten meal 3.0 4.0 6.0 36
Corn gluten feed 3.0 4.0 6.0 36
Peanut meal 3.5 4.0 5.0 36
Meat and bonemeal 3.0 4.0 6.0 8
&Valuesare based on data of Hartnell and Satter (19791,Colucci et al. (19821,and Erdman et al. (1987). Assumed NDF intake is >
1 YO of body weight with 75% of NDF intake coming from forage NDF. If NDF intake, as a percentage of body weight, is e 1%, then
increase passage rates by approximately 20%.
bComputed as total ration DM consumed/ration DM needed for maintenance.
CProportionof NDF that is effective in meeting fiber requirements.
Downloaded from jas.fass.org by on September 2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3570 SNIFFEN ET AL.
Table 3. Ruminal passage rate constants for forages (%/hIa

Effective
Level of maintenanceb NDFC
Ingredient 1x 2x 3x Yo of NDF
Legumes
High-quality, 18 to 21% CP
Long 2.5 3.0 4.0 92
20% > 2.54 cm length 3.0 3.5 4.5 82
.635 cm length 4.0 5.0 6.0 67
Average quality, c 18% CP
Long 2.0 2.5 3.0 92
20% > 2.54 cm length 2.5 3.0 3.5 82
3 3 5 cm length 3.0 3.5 4.0 67
Grasses
Long 2.0 2.5 3.0 98
20% > 2.54 cm length 2.0 3.0 4.0 88
.635 cm length 3.0 3.5 4.5 73
Corn silage
Mature, > 50% grain
Normal chop 2.0 2.5 3.0 71
Fine chop 4.0 5.0 6.0 61
Intermediate, 30 to 50% grain
Normal chop 1.5 2.0 2.5 81
Fine chop 3.0 4.0 5.0 71
Immature, c 30% grain
Normal chop 1.o 1.5 2.0 81
Fine chop 2.0 3.0 4.0 71
&Valuesare based on data of Hartnell and Satter (19791, Colucci et al. (19821, and Erdman et al.
(91871. AssumedNDF intake is > 1% of body weight with 75% of NDF intake coming from forage
NDF. If NDF intake, as a percentage of body weight, is < 146, then increase passage rates by
ap roximately 20%.
%Computed as total ration DM consumedhation DM needed for maintenance.
Choportion of N D F that is effective in meeting fiber requirements.

Equations for calculating digested protein from 1981; Siciliano-Jones and Murphy, 1989; Swingle et
feed and bacterial sources are listed below: al., 1990; Zinn, 1990). Only small amounts of starch
are recovered in the feces (Van Soest, 1982)if grain
DIGPBlj = REPBlj is adequately processed. Postruminal true digesti-
DIGPBBj = REPB2j bilities are given in Table 7. The small intestine
DIGPB3j = .80 * REPB3j lacks the enzymes to digest cellulose and hemicel-
DIGFPj = DIGPBlj +
DIGPB2j + DIGPBSj lulose (MacRae and Armstrong, 1969; Beaver et al.,
DIGBTPj = REBTPj 1972; Thompson et al., 19721, but cellulose and
DIGBNAj = REBNAj hemicellulose can be fermented by bacteria in the
DIGPj = DIGFPj + DIGBTPj + DIGBNAj large intestines. Lower tract digestion of cellulose
and hemicellulose ranged from 18.5 to 49.5% and
where DIGPBlj = digestible B1 protein from the 2.5 to 46%, respectively (Hoover, 1978). Therefore,
jth feedstuff, g/d; DIGPB2i = digestible B2 protein a n average postruminal true digestibility of 20% is
from the jth feedstuff, g/d; DIGPB3j = digestible used. The model assumes a postruminal true
B3 protein from the jth feedstuff, g/d; DIGFPj = digestibility of 95% for bacterial carbohydrate
digestible feed protein from the jth feedstuff, g/d; (Van Soest, 1982).
DIGBTPj = digestible bacterial true protein The equations for calculating digested carbohy-
produced from the jth feedstuff, g/d; DIGBNAj = drate due to the jth feedstuff are listed below:
digestible bacterial nucleic acids produced from
the jth feedstuff, g/d; and DIGPj = digestible VFAj = RDCAj + RDCBlj + RDCB2j
protein from the jth feedstuff, g/d. DIGFG = RECAj + std&*RECBLj+ .20*RECB2j
DIGBG = .95*REBCHOj
DIGCj = VFAj + DIGFCj + DIGBCj
Intestinal Carbohydrate Absorption
where stdig = postruminal starch digestibility, g /
Postruminal starch digestibility seems to be in g; DIGFCj = intestinally digested feed carbohy-
the range of 60 to 100% (Karr et al., 1966; Tucker et drate from the jth feedstuff, g/d; VFAj = ruminally
al., 1968; Waldo, 1973; Hoover, 1978; Russell et al., digested carbohydrate from the jth feedstuff, g/d;
Downloaded from jas.fass.org by on September 2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
 PREDICTING FEED ENERGY AND PROTEIN VALUES 3571
DIGBCj = digested bacterial carbohydrate Ruminants readily hydrolyze and absorb
produced from the jth feedstuff, g/d; and DIGCj = triglycerides, and pos truminal true digestibility
digestible carbohydrate from the jn feedstuff, g/d. approaches 100% (Van Soest, 1982). The CNCPS
assumes that bacterial fat has a postruminal true
digestibility of 95% Wan Soest, 19821. Equations
Intestinal Fat Absorption for calculating digestible fat from feed and bac-
terial sources are listed below:
Ruminal bacteria hydrogenate fat, but they do
not ferment it. Therefore, the model assumes that
all dietary fat passes to the small intestine. The DIGFFj = .95*REFATj
following equation is used to calculate ruminally DIGBFj = .95 REBFATj
escaped fat from the jth feedstuff: DIGFj = DIGFFj + DIGBFj

REFATj DIET FATj where DIGFFj = digestible feed fat from the jth
feedstuff, g/d; DIGBFj = digestible bacteria1 fat
where REFATj = amount of ruminally escaped fat from the jth feedstuff, g/d; and DIGFj = digestible
from the jth feedstuff, g/d. fat from the jth feedstuff, g/d.

Table 4. Digestion rate constants [%AI) for grainsa

Carbohydrate Protein
Ingredient A B1 B2 B1 B2 B3
Corn
Dry, whole shell corn
Whole 75-150 5-10 3-5 120-150 3-5 .06-.07
Cracked corn 100-200 10-20 5-7 140-160 4-6 .08-. 10
Cornmeal 200-300 20-30 7-0 150-175 6-9 .09-.12
High-moisture corn
> 35% moisture
Whole 150-200 10-15 5-7 140-160 4-6 .09-.12
Coarsely rolled 200-300 15-20 6-8 200-250 9-10 .lo-.20
Intermediate rolled 300-400 20-30 6-8 200-250 10-11 .15-.25
Finely rolled 300-400 30-40 8-10 200-250 11-12 .20-.30
30 to 35% Moisture
Whole 100-150 10-15 4-6 125-150 4-7 .08-.09
Coarsely rolled 150-250 15-20 6-8 125-150 8-9 .OO-.15
Intermediate rolled 250-350 20-30 6-8 125-250 9-10 .lo-.20
Finely rolled 250-350 30-40 8-10 125-250 10-11 .15-.25
25 to 30% Moisture
Whole 75-125 10-15 4-6 120-150 3-5 .07-.08
Coarsely rolled 125-175 15-20 6-8 120-150 6-7 .09-.10
Intermediate rolled 250-350 20-30 6-8 120-150 8-9 .lo-.15
Finely rolled 250-350 30-40 8-10 120-150 9-10 .lo-.20
.e 25% Moisture
Whole 75-125 10-15 3-5 120-140 3-5 .06-.07
Coarsely rolled 150-200 15-20 6-8 120-150 5-6 .07-.08
Intermediate rolled 200-300 20-30 6-8 120-150 7-8 .08-.10
Finely rolled 250-350 30-40 6-8 200-300 8-9 .09-.15
Steam-flaked corn 150-200 20-30 6-8 120-150 5-6 .07-.08
Sorghum
Dry,rolled 100-200 5-15 4-5 120-150 6-8 .09-. 15
Steam-flaked 200-300 15-20 6-8 150-170 8-10 .10-.20
Oats
Ground 250-350 30-40 4-6 300-350 12-15 .20-.50
Barley
Ground, dry 250-350 20-30 4-6 250-350 12-15 .20-.50
Ground, wet 250-350 30-40 4-6 250-350 13-16 .25-.55
Wheat
Dry,rolled 250-350 35-45 8-10 250-350 12-15 .20-.50
Steam-flaked 250-350 40-50 10-14 300-400 14-16 .25-.55
'Based on data of Waldo et al. (10721, Mertens (19731, Mertens and Ely (19791, Ewing and Johnson (19891, Van Soest et al. (10811,
Krishnamoorthy et al. (19831, Hoover (19831,Downloaded
and Smith etjas.fass.org
from al. (1972).by on September 2, 2008.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3572 SNIFFEN ET AI,.
Table 5 . Digestion rate constants (%h)
for proteinaceous feedsa

Carbohydrate Protein
Ingredient A B1 B2 B1 B2 B3
Soybean
Whole, raw 250-350 25-35 2 4 150-250 6-10 .10-.30
Whole, heated 250-350 3545 4-6 100-200 5-6 .15-.20
Meal, solvent 250-350 40-50 4-8 200-260 9-12 .lo-.30
Meal, expeller 250-350 35-45 4-8 150-250 6-8 .15-.20
Canola, solvent 250-350 4050 4-8 200-260 11-13 .10-.30
Peanut, solvent 250-350 40-50 4-8 200-260 12-14 .lo-.30
Cottonseed
Solvent 250-350 30-40 4-8 120-200 6-10 .lo-.20
Expeller 250-350 25-35 4-8 100-150 6-8 .lo-.15
Whole, linted 250-350 20-30 3-5 150-200 10-12 .20-.30
Whole, delinted 250-350 20-30 1-2 100-200 6-10 .20-.30
Corn gluten meal 250-350 40-60 4-6 100-200 2 4 .05-. 10
Corn gluten feed 250-350 40-60 6-8 100-200 2 4 .05-. 10
Corn distillers w/sol 250-350 15-20 6-8 100-200 3-4 .05-.15
Wheat middlings 250-350 60-85 10-15 200-300 5-6 .08-.15
Animal meals
Fishmeal 0 0 0 100-200 5-6 .08-. 15
Meat and bonemeal 0 0 0 100-200 5-6 .08-.15
Bloodmeal 0 0 0 50-100 2 4 .05-.08
Feathermeal 0 0 0 100-1 50 34 .05-.10
Brewers grain 250-350 3540 4-8 100-200 8-8 .lo-.20
Alfalfa meal, dehy 25-50 3540 8-10 100-200 7-9 .lo-.20
Whey 250-350 0 0 300400 0 0
&Based on Waldo et al. (19721, Mertens (19731, Mertens and Ely (1979), Ewing and Johnson (19891, Van Soest et al. (19811,
Krishnamoorthy et al. (19831, Hoover (19831, and Smith et al. (19721.

Table 6. Digestion rate constants (Wh) for foragesa

Carbohydrate Protein
Ingredient A B1 B2 B1 B2 B3
Corn silage
> 40% DM
Coarsely chopped 200-300 10-20 3-6 150-250 8-9 .08-.10
Finely chopped 250-350 20-30 4-8 250-350 10-12 .lo-.20
3 0 4 0 % DM
Coarsely chopped 200-300 15-25 4-8 200-300 9-10 .lo-.20
Finely chopped 250-350 25-30 8-10 250-350 10-1 1 .15-.25
< 30% DM
Coarsely chopped 200-300 25-35 4-8 250-350 10-1 1 .15-.25
Finely chopped 250-350 3540 8-10 250-350 10-12 .20-.30
Legumes
Hay 200-300 25-35 3-0 100-200 8-10 1.0-1.5
Silage
Coarsely chopped 200-300 3040 4-7 100-200 10-12 1.5-2.0
Finely chopped 250-350 3545 5-9 100-200 12-14 1.5-2.0
Grasses
Hay 200-300 25-35 2-4 120-150 10-12 .08-.10
Silage
Coarsely chopped 200-300 3540 3-5 200-250 12-14 1.0-1.2
Finely chopped 200-300 4045 4-6 250-300 13-15 1.1-1.3
'Based on Waldo et al. (19721, MertensDownloaded
(19731, Mertens and Ely
from jas.fass.org by (19791, Ewing
on September and Johnson (19891, Van Soest et al. (19811,
2, 2008.
Krishnamoorthy et al.©(19831,
Copyright HooverSociety
1992 American (19831, and Smith
of Animal etAll
Science. al.rights
(19721.
reserved. For personal use only. No other uses without permission.
 PREDICTING FEED ENERGY AND PROTEIN VALUES 3573
Fecal Losses Table 7. Postruminal starch digestibilities (%)a

Direct chemical analysis shows the absence of % Entering

plant cell contents in ruminant feces; the only Feed intestines
exceptions are starch and heat-damaged proteins, corn
both of which can be measured (Van Soest, 1982). Whole corn 50-80
There is little evidence of potentially digestible Dry, rolled 65-75
feed protein in feces; feed protein appearing in the Cracked 70-80
feces is insoluble and is either NDIP or ADIP Cum meal 80-90
High moisture, whole 80-90
protein (Van Soest, 1982). The following equations High moisture, ground 85-95
calculate undigested feed residues appearing in Steam-flaked 92-97
the feces from NDIP, ADIP, starch, fiber, fat, and Sorghum
ash fractions, based on data summarized by Van Dry, rolled 60-70
Soest (19821: Dry, ground 70-80
Steam-flaked 90-95
FEPBSj = (1 - .80l*REPB3j *Based on Swingle et al. (1990) and Zinn (1990), using dairy
FEPCj = REPCj and beef cattle.
FEFPj = FEPB3j + FEPCj
FECBlj = (1 - stdig1.RECBlj
FECBZj = (1 - .201*RECBOj FEBFj = amount of bacterial fat in feces from the
FECCj = RECCj jth feedstuff, g/d; FEBASHj = amount of bacterial
FEFCj = FECBlj + FECB2j + FECCj ash in feces from the jth feedstuff, g/d; and
FEFAj = DIET ASHj FEBACTj = amount of bacteria in feces from the
jth feedstuff, g/d.
where FEPBBj = amount of feed B3 protein Endogenous substances consist of calcium and
fraction in feces from the jth feedstuff, g/d; FEPCj magnesium salts of fatty acids, bile salts, sloughed
= amount of feed C protein fraction in feces from animal cells, mucus, and keratinized tissue Wan
the jth feedstuff, g/d; FEFPj = amount of feed Soest, 1982). According to Lucas et al. (19611,
protein in feces from the jth feedstuff, g/d; FECBlj endogenous protein, carbohydrate, and ash are:
= amount of feed starch in feces from the jth
feedstuff, g/d; FECB2j 7 amount of feed available FEENGPj = .0387.DIET PROT;
fiber in feces from the jth feedstuff, g/d; FECCj = FEENGFj = .017*DIETFAT;
amount of feed unavailable fiber in feces from the FEENGAj = .OIIQ*DIETASH
jth feedstuff, g/d; FEFCj = amount of feed carbohy-
drate in feces from the jth feedstuff, g/d; and FEFAj where FDj = feed DM consumed; FEENGPj =
= amount of feed ash in feces from the jth amount of endogenous protein in feces from the jth
feedstuff, g/d. feedstuff, g/d; FEENGFj = amount of endogenous
A variety of metabolic materials are excreted in fat in feces from the jth feedstuff, g/d; and
the feces, including microbial matter and en- FEENGAj = amount of endogenous ash in feces
dogenous substances. Microbial matter in from the jth feedstuff, g/d.
ruminant feces is largely indigestible cell walls of Total fecal DM is calculated by summing p r o
ruminal bacteria and bacterial cells produced in tein, carbohydrate, fat, and ash DM contributions
the lower tract (Van Soest, 1982). Microbial matter from undigested feed residues, microbial matter,
appearing in the feces is composed of indigestible and endogenous matter:
bacterial cell walls, bacterial carbohydrate, fat,
and ash (Van Soest, 19821: FEPROTj = FEFPj + FEBCPj + FEENGPj
FECHOj = FEFq + FEBCj
FEBCWj REBCWj FEFATj FEBFj + FEENGFj
FEBCPj = FEBCWj FEASHj = FEFAj + FEBASHj + FEENGAj
FEBCj = (1 - .951*REBCHOj IDMj = FEPROTj + FECHOj + FEFATj +
FEBFj = (1 - .95).REBFATj FEASHj
FEBASHj REBASHj
FEBACTj = FEBCPj + FEBCj + FEBFj + FEBASHj where FEPROTj = amount of fecal protein from
the jth feedstuff, g/d; FECHOj = amount of
where FEBCWj = amount of fecal bacterial cell carbohydrate in feces from the jth feedstuff, g/d;
wall protein from the jth feedstuf'f, g/d; FEBCPj = FEFATi = amount of fat in feces from the jth
amount of fecal bacterial protein from the jth feedstuff, g/d; FEASHj = amount of ash in feces
feedstuff, g/d; FEBq = amount of bacterial from the jth feedstuff, g/d; and IDMj = amount of
carbohydrate in feces from the jthDownloaded
feedstuff, by on September 2,DM
from jas.fass.orgindigestible
g/d; 2008.in feces from the jth feedstuff, g/d.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3574
Total Digestible Nutrients
True TDN can be calculated by subtracting
fecal losses from the potentially digestible nutrient
intake. Apparent TDN is potentially digestible
nutrient intake minus indigestible bacterial and
feed components appearing in the feces:
TDNAPPj = (DIET PROTj - FEPROTj) + (DIET
CHOj - FECHOj)+ 2.25 * DIET FATj -
FEFATj)
where TDNAPPj = apparent TDN from the jth
feedstuff, g/d.
Metabolizable and Net Energy
Values of Feeds
The CNCPS computes ME and NE values from
TDN rather than from the absorbed end products
of total tract digestion. It is aggregated at the level
that is the first critical step beyond present NRC
systems in improving accuracy of feed energy and
microbial yield values used to evaluate and formu-
late diets. That step is to compute carbohydrate
fraction pool sizes and microbial yield sensitive to
intake level, ruminal fermentation and passage
rates, ruminal microbial growth, bacterial compo-
sition, postruminal digestibilities, and endogenous
nutrient contributions. The determination of NE
values directly from individual absorbed VFA,
starch, and fat requires a much more complex
submodel of the rumen. Based on our preliminary
unpublished observations, the more complex sub-
models of the rumen we have evaluated did not
seem to improve the prediction of feed energy and
microbial yield compared to the CNCPS. However,
we are currently evaluating the addition of deter-
minations for pH and pool sizes of end products of
ruminal fermentation to the present CNCPS struc-
ture.
The ME values for each feed are based on the
assumption that 1 kg of TDN is equal to 4.409 Mcal
of DE and 1 Mcal of DE is equal to .82 Mcal of ME
(NRC, 1976):
MEaj = .001 .TDNAPPj*4.409*.82
MECi = ME,i/(FDi * .001)
i-1
MEC = MEIADMI * .OO1)
where MEaj = metabolizable energy available
from the jth feedstuff, Mcal/d; MECj = metaboliza-
ble energy concentration of the jth feedstuff, Mcal/
kg; ME1 = metabolizable energy supplied by the
diet, Mcal/d; and MEC = metabolizable energy
concentration of the diet, Mcal/kg. Downloaded from jas.fass.org(DIP)
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
SNIFFEN ET AL..
NE, and NE, values are computed from ME,
based on NRC (1984) equations to adjust for
differences between forages and concentrates in
efficiency of use of ME. The term NE1 is calculated
based on NRC (1989).
NEgaj = 1.42.MECj - .174.MECf + .0122.MECB -
1.65)
NEmaj = (1.37.MECj - .138*MECj]+ .0105*MEC?-
1.12)
NElaj = MECje.65
NEga = (1.42.MEC - .174.MEC2 + .0122.MEC3-
1.65)
NEma = (1.37-MEC- .138*MEC2+ .0105.MEC3-
1.121
NEla = MECje.65
where NEgaj = net energy for gain content of the
jth feedstuff, Mcal/kg; NEmd = net energy for
maintenance content of the ith feedstuff, Mcal/kg;
NE1,j = net energy for lactation content of the jth
feedstuff, Mcal/kg; NEga = net energy for gain
content of the diet, Mcal/kg; NEma = net energy
for maintenance content of the diet, Mcallkg; and
NE1a = net energy for lactation content of the diet,
Mcal/kg.
Metabolizable Protein
Metabolizable protein (MP) is digested feed and
bacterial protein minus bacterial nucleic acids.
The model generates a variable MP estimate for
each ingredient based on protein composition,
ruminal protein digestion rates, passage rates,
bacterial yield, bacterial composition, and postru-
minal digestibilities of feed and bacterial protein
fractions. Total feed MP is the s u m of each feed
MP:
MPai = DIGPi - DIGBNAi
i-1
where MPaj = metabolizable protein from the jth
feedstuff, g/d, and MP, = metabolizable protein
available in the diet, g/d.
Differences Between the Cornell Net
Carbohydrate and Protein System and
the National Research Council System
The CNCPS differs from the NRC (1985, 1989) in
many respects. In the NRC system, protein is
divided into two fractions, degraded intake protein
and 2,undegraded
by on September 2008. intake protein (UIP),
 PREDICTING FEED ENERGY AND PROTEIN VALUES
whereas the CNCPS divides protein into five
fractions L4, B1, B2, B3, and C). Average, static
tabular values for DIP and UIP are used by the
NRC (1985, 19891, whereas the CNCPS generates
MP values that are based on the various protein
fractions, ruminal digestion, and passage rates.
The NRC uses a constant, intestinal true digesti-
bility of 80% for UIP, whereas the CNCPS applies
true digestibilities of 100, 100, 80, and 0% for B1,
B2, B3, and C protein fractions. In the NRC, bound
protein contributes to absorbed protein, but the
CNCPS assumes that bound protein does not
contribute to absorbed protein. The use of a single
DIP value in the NRC does not accommodate
differences in ruminal bacterial N utilization
(ammonia vs amino N). The CNCPS accounts
ammonia, peptides, and protein as separate pools
and the ruminal bacteria are partitioned accord-
ing to N utilization.
The NRC protein systems (1985, 1989) use empir-
ical regression equations that relate NE values to
static TDN and microbial crude protein produc-
tion to daily TDN intake. The amount of TDN is a
poor indicator of ruminal fermentation when
rations are high in fat or oil (Satter and Roffler,
1975). Ruminal bacteria only grow on ruminally
degraded carbohydrate and do not use fat as an
energy source (Nocek and Russell, 19881; therefore,
NRC systems may overestimate microbial growth.
Another disadvantage of empirically based TDN is
the large negative intercept that may underesti-
mate microbial protein contributions at low TDN
intakes. The CNCPS uses mechanistic equations
that predict a variable TDN and microbial protein
yield from fermentable SC and NSC carbohydrate
intake, rates of fermentation, the availability of
amino N, and pH.
Associative Effects of Feeds
Validations with our unpublished experimental
and case study data indicate that the digestion
and passage rates presented in Tables 2 through 6
are appropriate for use with typical mixed diets.
However, we have found that users may need to
adjust these rates, as well as the microbial growth
rates, based on their knowledge of factors that
may result in unusually low or high digestion and
passage rates. The ruminal digestion rates assume
a normal pH, and the submodel of the rumen
adjusts microbial protein yield in high grain
rations in which pH can be expected to be 6.0 to
6.2. However, it does not automatically adjust
microbial growth on the SC fraction, which is very
pH-sensitive. In an evaluation of data from experi-
ments with 0 to 10% forage, we concluded that the
use of a growth rate of 0 for the SC bacteria was
Downloaded from jas.fass.org by ondigestion,
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
3575
appropriate for this type of diet. The associate
effects of N sources are accounted for primarily by
determining the ammonia and peptide require-
ments of the SC and NSC pools. By adding a
calculation to estimate the branched-chain VFA
requirement for SC bacteria as described in the
submodel of the rumen (Russell et al., 19921, the
positive effects of using true protein to provide
ammonia for SC bacteria in high-forage diets can
be accounted for.
Implications
The Cornel1 Net Carbohydrate and Protein
System uses standard chemical analyses to esti-
mate the pool sizes of biologically significant
carbohydrate and protein fractions and microbial
growth on these fractions. Because ruminal and
lower gut digestion is a function of fermentation
and passage rates, total digestible nutrients and
metabolizable protein can be predicted in a more
mechanistic fashion for field application in specific
feeding situations. This system provides the begin-
ning structure necessary to predict more ac-
curately ruminally degraded carbohydrate and
nitrogen fractions and absorbed amino acids from
microbial and feed protein. Further research
should result in field-usable models of the rumen
with lower levels of aggregation.
Literature Cited
Abdalla, H. O., D. C. Fox, and R. R. Seaney. 1988a. Protein
distribution in four cool-season grass varieties alone or in
combination with trefoil. J. Anim. Sci. 66:2325.
Abdalla, H. O., D. C. Fox,and R. R. Seaney. 1988b. Variation in
protein and fiber fractions in pasture during the grazing
season. J. Anim.Sci. 68:2883.
AOAC. 1980. Official Methods of Analysis (13th Ed.). Association
of Official Analytical Chemists, Washington, DC.
Baldwin, R. L., L. J. Koong, and M. J. Ulyatt. 1977. A dynamic
model of ruminant digestion for evaluation of factors af-
fecting nutritive value. Agric. Systems 2:255.
Beaver, D. E.,J. F. Coelho da Silva, J.D.H. Prescott, and D. G.
Armstrong. 1972. The effect in sheep of physical form and
stage of growth on the sites of digestion of a dried grass. 1.
Sites of digestion of organic matter, energy and carbohy-
drate. Br. J. Nutr. 28:347.
Chalupa, W., C. J. Sniffen, D. G. Fox, and P. J. Van Soest. 1991.
Model generated protein degradation nutrition informa-
tion. In: Proc. Cornel1 Nutr. Conf. p 44. Ithaca, NY.
Colucci, P. E., L. E. Chase, and P. J. Van Soest. 1982. Feed
intake, apparent diet digestibility, and rate of particulate
passage in dairy cattle. J. Dairy Sci. 65:1445.
Erdman, R. A,, J. H. Vandersall, E. Russek-Cohen, and G.
Switalski. 1987. Simultaneous measures of rates of ruminal
digestion and passage of feeds for prediction of ruminal
nitrogen and dry matter digestion in lactating dairy cows.
J. Anim. Sci. 04:585.
Ewing, D. L., and D. E. Johnson. 1987. Corn particle starch
September 2,passage
2008. and size reduction in beef steers: A
3576 SNIFFEN
dynamic model. J. Anim. Sci. 64:1194.
France, J. J., H. M. Thornley, and D. E. Beever. 1982. A mathe-
matical model of the rumen. J. Agric. Sci. (Camb.1 88:343.
Gill, M., J.H.M. Thornley, J. L. Black, J. D. Oldham, and D. E.
Beever. 1984. Simulation of the metabolism of absorbed
energy-yielding nutrients in young sheep. Br. J. Nutr. 52:
621.
Goering, H. K., and R. S. Adams. 1973. Frequency of heat-
damaged protein in hay, haycrop silage and corn silage. J.
Anim. Sci. 37:295 (Abstr.).
Goering, H. K., and P. J. Van Soest. 1970. Forage fiber analyses
(apparatus, reagents, procedures, and some applications).
Agric. Handbook 379. ARS, USDA, Washington, DC.
Hartnell, G. F., and L. D. Satter. 1970. Determination of rumen
fill, retention time and ruminal turnover rates of ingesta at
different stages of lactation in dairy cows. J. Anim. Sci. 48:
381.
Hoover, W. H. 1978. Digestion and absorption in the hindgut of
ruminants. J. Anim. Sci. 46:1789.
Karr, M. R.,C. 0.Little, and G. E. Mitchell, Jr. 1966. Starch
disappearance from different segments of the digestive
tract of steers. J. Anim. Sci. 25:652.
Krishnamoorthy, U.C., T. V. Muscato, C. J. Sniffen. and P. J.
Van Soest. 1082. Nitrogen fractions in selected feedstuffs. J.
Dairy Sci. 65:217.
Krishnamoorthy, U. C., C. J. Sniffen, M. D. Stem, and P. J. Van
Soest. 1983. Evaluation of a mathematical model of digesta
and in-vitro simulation of rumen proteolysis to estimate the
rumen undegraded nitrogen content of feedstuffs. Br. J.
Nutr. 50:555.
Lucas, H. L., Jr., W.W.G. Smart, Jr., M. A. Cipolloni, and H. D.
Gross. 1061. Relations between digestibility and composi-
tion of feeds and foods. S-45 Report, North Carolina State
College.
MacGregor, C. A,, M. R.Stokes, W. H. Hoover, H. A. Leonard, L.
L. Junkins, Jr., C. J. Sniffen, and R. W. Mailman. 1983.
Effect of dietary concentration of total nonstructural carbo-
hydrates on energy and nitrogen metabolism and milk
production of dairy cows. J. Dairy Sci. 66:39.
MacRae, J. C., and D. G. Armstrong. 1969. Studies on intestinal
digestion in the sheep. 2. Digestion of some carbohydrate
constituents in cereal and hay-cereal. Br. J. Nutr. 23:377.
Mason, V. C., and R.Palmer. 1071. Studies on the digestibility
and utilization of the nitrogen of irradiated rumen bacteria
by rats. J. Agric. Sci. (Camb.) 76:567.
Mason, V. C., and F. White. 1071. The digestion of bacterial
mucopeptide constituents in sheep. I. The metabolism of 2,6
diaminopimelic acid. J. Agric. Sci. (CambJ 7791.
Mertens, D. R. 1073. Application of theoretical mathematical
models to cell wall digestion and forage intake in
ruminants. Ph.D. Dissertation. Cornell Univ., Ithaca. NY.
Mertens, D. R. 1985. Effect of fiber on feed quality for dairy
cows. 46th Minnesota Nutr. Cod. p 209. St. Paul, MN.
Mertens, D. R.,and L. 0. Ely. 1979. A dynamic model of fiber
digestion and passage in the ruminant for evaluating
forage quality. J. Anim. Sci. 49:1085.
Morrison, F. B. 1956. Feeds and Feeding Wnd EdJ. Morrison
Publishing Co., Clinton, I k
Muscato, T.V., C. J. Sniffen, U. C. Krishnamoorthy, and P. J.
Van Soest. 1083. Amino acid content of noncell and cell
wall fractions in feedstuffs. J. Dairy Sci. 66:2108.
Nocek, J. E. 1985. Evaluation of specific variable affecting in
situ estimates of ruminal dry matter and protein digestion.
J. Anim. Sci. 60:1347.
Nocek, J. E., and J. B. Russell. 1088. Protein and energy as an
integrated system. Relationship of ruminal protein and car-
bohydrate availability to microbial protein synthesis and
milk production. J. Dairy Sci. 71:2070.
NFlC. 1076. Nutrient Requirements of Beef Cattle (5th Ed.). Na-
tional Academy Press, Washington, DC. Downloaded from jas.fass.org by onNY.
Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
--
NRC. 1978. Nutrient Requirements of Dairy Cattle (5th Ed.).
National Academy Press, Washington, DC.
NRC. 1981. Nutritional Energetics of Domestic Animals and
Glossary of Energy Terms (2nd Ed.). National Academy
Press, Washington, DC.
NRC. 1982. United States-Canadian Tables of Feed Composi-
tion. National Academy Press, Washington, DC.
NRC. 1984. Nutrient Requirements of Beef Cattle (6th Ed.). Na-
tional Academy Press, Washington, DC.
NRC. 1985. Ruminant Nitrogen Usage. National Academy
Press, Washington, DC.
NRC. 1989. Nutrient Requirements of Dairy Cattle (6th Ed.).
National Academy Press, Washington, DC.
Pichard, D. G. 1977. Forage nutritive value. Continuous and
batch in vitro fermentations and nitrogen solubility. Ph.D.
Dissertation. Cornell Univ., Ithaca, NY.
Pichard, D. G., and P. J. Van Soest. 1977. Protein solubility of
ruminant feeds. Proc. Cornell Nutr. Conf. p 91. Ithaca, NY.
Roe, M. B., C. J. Sniffen, and L. E. Chase. 1990. Techniques for
measuring protein fractions in feedstuffs. Proc. Cornell
Nutr. Conf. p 81. Ithaca, NY.
Russell, J. B., J. D. O’Connor, D. G. Fox, P. J. Van Soest, and C.
J. Sniffen. 1092. A net carbohydrate and protein system for
evaluating cattle diets I. Ruminal fermentation. J. Anim.
sci. 70:355 1.
Russell, J. R.,A. W. Young,and N. A. Jorgensen. 1981. Effect of
dietary corn starch intake on ruminal, small intestinal and
large intestinal starch digestion in cattle. J. Anim. Sci. 52:
1170.
Satter, L. D., and R. E. Roffler. 1975. Nitrogen requirement and
utilization in dairy cattle. J. Dairy Sci. 581219.
Siciliano-Jones, J., and M. R. Murphy. 1980. Nutrient digestion
in the large intestine as influenced by forage to concen-
trate ratio and forage physical form. J. Dairy Sci. 72:471.
Smith, L. W., H. K. Goering, and C. H. Gordon. 1972. Relation-
ships of forage compositions with rates of cell wall diges-
tion and indigestibility of cell walls. J. Dairy Sci. 55:1140.
Smith, R.H. 1969. Nitrogen metabolism and the rumen. J. Dairy
Res. 36:313.
Smith, R. H., and .4. B. McAllan. 1971. Nucleic acid metabolism
in the ruminant. 3. Amounts of nucleic acids and total and
ammonia nitrogen in digesta from the rumen, duodenum
and ileum of calves. Br. J. Nutr. 25:181.
Sniffen, C. J., J. B. Russell, and P. J. Van Soest. 1983. The
influence of carbon source, nitrogen source and growth
factors in rumen microbial growth. Proc. Cornell Nutr.
Conf. p 26. Ithaca, NY.
Storm, E., D. S. Brown, and E. R. Orskov. 1983. The nutritive
value of rumenmicroorganisms in ruminants. 3. The diges-
tion of microbial and nucleic acids in, and losses of e n
dogenous nitrogen from, the small intestine of sheep. Br. J.
Nutr. 50:470.
Swingle, R.S., J. Moore, M. Moore, and T.Eck. 1990. Utilization
of starch from processed sorghum grain. I n Proc. South-
west Nutr. and Management Conf. p 52. Tempe, AZ.
Tas,M. V., R.A. Evans, and R.F.E. Axford. 1981. The digestion of
amino acids in the small intestine of sheep. Br. J. Nutr. 45:
167.
Thompson, D. J., D. E. Beever, J. F. Coelho da Silva, and D. G.
Armstrong. 1972. The effect in sheep of physical form on
the sites of digestion of a dried lucerne diet. Br. J. Nutr. 28:
31.
Tucker, R. E., G. E. Mitchell, Jr., andC. 0. Little. 1088. Ruminal
and postruminal starch digestion in sheep. J. Anim. Sci. 27:
824.
Van Soest, P. J. 1982. Nutritional Ecology of the Ruminant. O&B
Books, Cornallis, OR.
Van Soest, P. J., and D. G. Fox. 1092. Discounts for net energy
and protein. In: Proc. Cornell Nutr. Conf. (In press). Ithaca,
September 2, 2008.
 PREDICTING FEED ENERGY AND PROTEIN VALUES
Van Soest, P. J., D. G. Fox, D. R. Mertens, and C. J. Sniffen. 1984.
Discounts for net energy and protein-fourth revision. Proc.
Cornell Nutr. Conf. p 121. Ithaca, NY.
Van Soest, P. J.,J. B. Robertson, and B. A. Lewis. 1991. Methods
for dietary fiber, neutral detergent fiber, and nonstarch
polysaccharides in relation to animal nutrition. J. Dairy
Sci. 74:3583.
Van Soest, P. J., and C. J. Sniffen. 1984. Nitrogen fractions in
NDF and ADF. Proc. Dist. Feed Conf. 39:73.
Van Soest, P. J., C. J. Sniffen, D. R. Mertens, D. G. Fox, P. H.
Robinson, and U. C. Krishnamoorthy. 1981.A net protein
system for cattle: The rumen submodel for nitrogen. In: F.
N. Owens (Ed.) Protein Requirements for Cattle: hoceed-
ings of a n International Symposium. MP-109. p 265. Div. of
Agric., Oklahoma State Univ., Stillwater.
3577
Varga, G. A., and W. H. Hoover. 1983. Rate and extent of
neutral detergent fiber degradation of feedstuffs in situ. J.
Dairy Sci. 66:2109.
Waldo, D. R. 1973. Extent and partition of cereal grain starch
digestion in ruminants. J. h i m . Sci. 37:1082.
Waldo, D. R.,and H. K. Goering. 1979. Insolubility of proteins in
ruminant feeds by four methods. J. Anim. Sci. 49:1560.
Waldo, D. R.,L. W. Smith, and E. L. Cox. 1972. Model of
cellulose disappearance from the rumen. J. Dairy Sci. 55:
125.
Welch, J. G. 1986. Physical parameters of fiber affecting pas-
sage from the rumen. J. Dairy Sci. 692750.
Zinn, R. A. 1990. Optimizing the value of steamflaked corn in
diets for feedlot cattle. In: Proc. Southwest Nutr. and
Management Conf. p 36. Tempe, AZ.

Downloaded from jas.fass.org by on September 2, 2008.

Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.
Citations This article has been cited by 51 HighWire-hosted articles:
http://jas.fass.org#otherarticles

Downloaded from jas.fass.org by on September 2, 2008.

Copyright © 1992 American Society of Animal Science. All rights reserved. For personal use only. No other uses without permission.