An Analysis of The Nutritive Value of Heat Processed Legume Seeds For Animal Production Using The DVE-OEB Model A Review

Animal Feed Science and Technology
99 (2002) 141–176
An analysis of the nutritive value of heat processed

legume seeds for animal production using the
DVE/OEB model: a review
P. Yu a,∗ , J.O. Goelema b , B.J. Leury c , S. Tamminga b , A.R. Egan c
a Department of Animal and Poultry Science, University of Saskatchewan, 6D34 Agricultural Building, 51
Campus Drive, Saskatoon, SK, Canada S7N 5A8
b Department of Animal Nutrition, Wageningen Agricultural University, Zodiac, Marijkeweg 40, 6709 PG,
Wageningen, The Netherlands
c Department of Animal Production, Institute of Land and Food Resources,
University of Melbourne, Parkville, Vic. 3052, Australia

Received 15 March 2000; received in revised form 20 February 2002; accepted 12 April 2002
Abstract
Recently obtained information on structural and compositional effects of processing of legume
seeds is reviewed, in relation to legume seed characteristics affecting digestive behavior and nutrient
utilization. The emphasis is on (1) manipulation of digestive behavior by heat processing methods,
particularly on protein and/or starch components of legume seeds, (2) prediction of changes of
potential nutrient supply to ruminants from legume seeds after heat processing, using the newly
developed DVE/OEB model, and (3) obtaining information on optimal processing conditions of
legume seeds as intestinal protein source in term of protein DVE and OEB values (to achieve target
values for potential high net absorbable protein in the small intestine while holding any N loss in the
rumen to a low level), using the protein evaluation DVE/OEB model. The information described in
this paper may give better insight in the mechanisms involved and the changes occurring upon pro-
cessing of legume seeds. A focus of the review is on evaluation of the models and new approaches to
establishment of a protein evaluation system that more accurately accounts for digestive processes
in the ruminant on a quantitative basis.
© 2002 Elsevier Science B.V. All rights reserved.
Keywords: Digestive behavior; Legume seeds; Optimal seed processing; The DVE/OEB model
∗ Corresponding author. Tel.: +1-306-966-4150; fax: +1-306-966-4151.
E-mail address: yupe@sask.usask.ca (P. Yu).
0377-8401/02/$ – see front matter © 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 7 - 8 4 0 1 ( 0 2 ) 0 0 1 1 4 - 1
142 P. Yu et al. / Animal Feed Science and Technology 99 (2002) 141–176
Nomenclature
ABCP truly absorbed rumen undegradable protein in the small intestine (g/kg DM)
AMP truly absorbed microbial protein in the small intestine (g/kg DM)
BCP rumen undegradable protein (g/kg DM)
%BCP percent rumen undegradable protein (% of CP)
BSt rumen undegradable starch (g/kg DM)
%BSt percent rumen undegradable starch (% of starch)
dASH digestibility of ash (%)
dBCP digestibility of rumen undegradable protein in the small intestine (%)
dTCP total tract digestibility of crude protein (%)
D insoluble but potentially degradable fraction in sacco (%)
DOM digested organic matter (g/kg DM)
DVE truly digested protein in the small intestine (g/kg DM)
ED effective degradability of CP (% of CP) or starch (% of starch)
ENDP endogenous protein secretion in the small intestine (g/kg DM)
E MP microbial protein synthesized in the rumen based on available energy
(g/kg DM)
FOM organic matter fermented in the rumen (g/kg DM)
Kd the rate of degradation of the D fraction (%/h)
Kp ruminal passage rate (%/h)
N MP microbial protein synthesized in the rumen based on available nitrogen
(g/kg DM)
OEB degraded protein balance (g/kg DM)
S soluble fraction in sacco (%) which is assumed to be degraded rapidly
and completely
T0 lag time (h) in which no degradation of D takes place
TPSI true protein supplied to the small intestine (g/kg DM)
U undegradable fraction in sacco (%)
UASH undigested ash (g/kg DM)
UDM undigested dry matter (g/kg DM)
UOM undigested organic matter (g/kg DM)
1. Introduction
Dairy cows require large quantities of amino acids (AA) and glucose supplied by the diet
to maintain high milk production (Ferguson, 1975). This can only be assured if sufficient true
protein and glucose are available to be absorbed from the small intestine. Therefore, dairy
cows with a high milk production need not only N available for rumen microbial protein
synthesis, but also digestible dietary protein and non-structural carbohydrates bypassing the
rumen (Tamminga, 1979; Grummer and Clark, 1982; Satter, 1986; Chalupa and Sniffen,
1991; Nocek and Tamminga, 1991).
It is easy to understand that it would be advantageous to enhance the rate of degrada-
tion of structural carbohydrates in the rumen, but to optimize the extent to which proteins
P. Yu et al. / Animal Feed Science and Technology 99 (2002) 141–176 143
and non-structural carbohydrates are degraded in the rumen (Nocek and Tamminga, 1991;
Goelema, 1999). Rumen degradation of the latter two feed components should be restricted
to what is necessary for maintaining efficient microbial activity and growth. Digestion in
the small intestine of feed protein and non-structural carbohydrates has the advantage that
less protein and energy is lost (Goelema and Tamminga, 1994). In achieving such circum-
stances, the balance of site of digestion and synchrony of fermentation rate and continuing
availability of the nutrients essential for microbial growth is of critical importance.
Legume seeds have attracted attention as components of feedstuffs for dairy cows as
protein supplements in recent years mainly because: (1) legume seeds usually have a par-
ticularly high protein and/or starch contents; and (2) a diversity of legume species, with
high seed yields, are well suited to the various ecological and climatic conditions in many
countries (Cerning-Beroard and Filiatre, 1977; Ensminger and Olentine, 1978). However,
the use of legume seeds in dairy cows is limited and the utilization is inefficient under certain
conditions. An important reason is that the soluble or rapidly degradable protein content
of many legume seeds is too high, the values approach 90% (Van Straalen and Tamminga,
1990), and the rate of degradation is also very high (Yu, 1995). These cause an imbalance
between feed protein breakdown and microbial protein synthesis, resulting in unnecessary
N loss from the rumen. Often there is an overall surplus of rumen degraded protein. This
is because microbial growth rate and protein synthesis cannot be increased at the same
time as NH3 concentration is maximizing, and subsequently after early rapid degradation,
a shortage of feed N in the rumen will occur (Egan, 1986; Tamminga et al., 1994).
To achieve a higher milk production, it is advantageous under certain conditions to have
more starch escape degradation in the rumen (Nocek and Tamminga, 1991). Starch escaping
ruminal degradation will not only prevent fermentation losses in the rumen, it (if digested
in the intestines) will also supply the animal with more glucose, an important precursor for
lactose. This may result in a sparing of AA (from oxidation, or as precursors for gluco-
neogensis) which can be used for other purposes in the production of milk protein. Most
starches in legume seeds are degraded very rapidly (Goelema, 1999; Yu, 1999). As a result
little starch escapes in the digesta flow from the rumen, and VFAs and lactic acid are being
generated rapidly with a concomitant decrease in pH, to which cell wall degrading bacteria
are sensitive. Rapid degradation usually means that a large proportion of VFA is propionate,
an important precursor for lactose synthesis, which in large amounts may depress the con-
centration of milk fat. Generally, starch digested post-ruminally is used more efficiently for
milk synthesis than that digested in the rumen (Nocek and Tamminga, 1991).
If legume seeds are to be more efficiently utilized in ruminant diets as intestinal protein
source, particularly in the diet of high yielding dairy cows, under certain circumstance, the
rate and extent of degradation in the rumen must be reduced without diminishing the extent
of their intestinal digestion.
Recently obtained information on structural and compositional effects of processing of
legume seeds is reviewed, in relation to feed characteristics affecting digestive behavior and
nutrient utilization. The emphasis is on manipulation of digestive behavior of legume seeds
by processing methods, particularly on protein, prediction of changes of potential nutrient
supply to ruminants from legume seeds after heat processing, using the newly developed
DVE/OEB model, and obtaining information on optimal processing conditions of legume
seeds as intestinal protein sources in term of protein DVE and OEB value (to achieve target
values for potential high net absorbable protein in the small intestine while holding any N
loss in the rumen to a low level), using the DVE/OEB model. Besides, rumen starch digestive
behavior and characteristics of starch-containing legume seeds were also evaluated.
2. Several important aspects of ruminant digestion and nutrient utilization of

legume seeds
The major nutrient constituents of organic matter in legume seeds are protein and/or
starch. The chemical and physical properties within and among them largely determine
their fate in the digestive tract of ruminants.
2.1. Protein of legume seeds
In legume seeds, 85–100% of protein is in albumins (water soluble) and globulins (salt
soluble), none in prolamines (alcohol soluble) and 0–15% in glutelins (dilute alkali soluble)
(McDonald et al., 1988; Van Straalen and Tamminga, 1990).
Protein degradation of legume seeds in the rumen involves two steps, as described by
Goelema (1999), being hydrolysis of the peptide bond by protease and peptidases and
decarboxylation and/or deamination of AA. The first step results in release of peptides and
AA, while the end products of the second step are VFAs and branched chain fatty acids,
CO2 and NH3 (Blackburn, 1965; Hungate, 1984). Proteolysis in the rumen results from the
proteolytic enzymes from bacteria, protozoa and fungi. Bacterial proteases in the rumen are
mainly cysteine endoproteases, but metallo- and serine proteases are also present. Protozoa
also exhibit proteolytic activity, cysteine and aspartate proteases, whereas rumen fungi
have an extracellular metalloprotease (Hungate, 1966; Hungate, 1984; Wallace, 1991). As
a result of proteolytic digestion, polypeptides are released, these are then broken down
into dipeptides and AA. Some of the AA are incorporated into microbial protein, but the
main part of the AA-N is converted to NH3 by deamination (Bryant, 1970; Russell and
Hespell, 1981; Wallace, 1991). Part of the NH3 is reincorporated into microbial protein,
but much is lost by diffusion across the rumen wall. The processes of N transaction in the
rumen are described quantitatively by Nolan (1974) and Egan et al. (1985). For protein
breakdown to occur, microorganisms and their proteolytic enzymes must gain access to
their substrate. Soluble proteins are more susceptible to degradation than are insoluble
proteins, but exceptions to this rule do exist. Cross-linking, particularly through disulphide
bonds, has a major influence on protein degradability. A reduction in the rate or extent of
degradation of protein of legume seeds in the rumen will only be beneficial to the animal
when the protein escaping degradation in the rumen is digested and AA and peptides are
absorbed from the small intestine.
2.2. Starch of legume seeds
Starch, as a main component in non-structural carbohydrates of legume seeds, plays

an important role in production diets for ruminants. Goelema (1999) reviewed that the
mechanism of starch degradation by rumen microorganisms has been little studied, but the
physical characteristics of starch bodies affect enzyme accessibility. Isolated enzymes from
various strains of rumen bacteria have proved to be similar to amylases from other bacterial
and mammalian sources. Despite this, no accurate prediction of starch degradation could be
obtained by using purified enzymes instead of freeze-dried, cell free preparations of rumen
fluid (Cone, 1991). Part of the amylolytic activity of bacteria is cell bound (Cone, 1991), but
this has not been studied extensively. Although not an essential or consistently represented
contributor to starch utilization by ruminants, the protozoa may play a role as a stabilizing
agent. Engulfment of starch granules by protozoa limits the amount of starch available
for the usually rapid bacterial fermentation. This helps to prevent lowering of rumen pH
which, in severe cases may become detrimental to cell wall degrading bacteria (Chesson and
Forsberg, 1988) or worse, inhibit normal rumen contraction and overall digestive function.
Amylolytic activity in the rumen is stimulated by a low pH as well as by the presence of
starch (Tamminga et al., 1989).
2.3. Optimization of rumen fermentation
For optimum microbial protein synthesis, the ratio of rumen available proteins to ru-
men available carbohydrates is a critical determinant. Czerkawski (1986) reported that
around 25 g N/kg available carbohydrates should be appropriate and sufficient for achiev-
ing optimum microbial growth. Tamminga et al. (1990) reported that optimum ratio of
carbohydrates (g/kg DM) and proteins (g/kg DM) is approximately 5:1 (approximately
33 g N/kg carbohydrates). Legume seeds usually have the high ratio of available N to
available carbohydrates, for example lupin is about 112 g N/kg carbohydrates, faba bean
is about 55 g N/kg carbohydrates (Yu, 1999), which indicates a potential N loss in the
rumen under certain conditions. Not only must the ratio of available carbohydrates to avail-
able proteins be optimum, but also degradation of carbohydrate and protein should be
synchronized in order to achieve efficient microbial growth and little N loss in the rumen
(Tamminga et al., 1990).
In order to optimize rumen fermentation, large discontinuous inputs of soluble or rapidly
degradable carbohydrates should be avoided, because they may give rise to an excessive
VFA production resulting in a low pH. This in turn will slow down the microbial degrada-
tion of structural carbohydrates. Likewise, discontinuous input of large amounts of soluble
or rapidly degradable protein should be avoided, because they may result in excessive
NH3 production followed by high levels of urea excretion, so that N utilization is reduced
(Chalupa, 1974; Beever and Thomson, 1977).
To optimize rumen fermentation, several possible approaches could improve degradative
behavior of substrates in the rumen, such as changing the process by microbial manipulation,
changing the process by feeding strategies (selecting feed ingredients, meal frequency,
balancing the forage and concentrate quality in terms of protein supply, and synchrony with
energy supply, etc.), and/or changing the process by processing feed (changing characteristic
of ingredients). Feed processing, such as treatments related to heat, can be used to manipulate
site of digestion as well as degradation characteristics in the rumen and is therefore a helpful
tool in optimizing ruminant production under certain circumstance (Broderick et al., 1991;
Goelema and Tamminga, 1994; Goelema, 1999). The paper focuses on manipulation of
digestive behavior and characteristics of legume seeds by heat processing and prediction of
potential nutrient supply of legume seeds as intestinal protein sources for ruminants by the
model.
3. The modern evaluation system for describing digestive behavior of

legume seeds in ruminants
3.1. Rumen degradation model
Each feed has its own rumen degradation characteristics. Different feed components also
have different rumen degradation characteristics, which are becoming critical in modern
feed formulation.
Various models to describe rumen degradation characteristics of feed components are
possible. In the choice of variables used to describe the model, biological or mathemat-
ical considerations may prevail. Although mathematical background models, such as the
Gompertz equation (Sauvant et al., 1985) may be more accurate in empirically fitting to
the data, their biological interpretation is often difficult. This is why biologically orientated
models are more popular (Tamminga et al., 1990). Among the biologically orientated mod-
els, the most widely used model is a first order kinetics degradation model introduced first
by Ørskov and McDonald (1979). Methods used to solve such a model include non-linear
iterative least square regression, least square regression of logarithmic-transformed residues
with or without correction for an estimated or measured ruminally undegraded residue, and
curve peeling (Tamminga et al., 1990).
The important rumen degradative characteristics described by the first-order kinetics
degradation model (Ørskov and McDonald, 1979), modified by Robinson et al. (1986) and
Tamminga et al. (1990) are S (%), U (%), D (%), Kd (%/h), T0 (h) and ED (%). Because
each component of legume seed has its own pattern of rumen degradation characteristics,
to describe its degradation accurately the models used for different components (such as
protein, starch, etc.) are different.
The degradation model of protein is:
R(t) = U + (100 − S − U ) × e−Kd ×(t−T0 )
where R(t) is the residue (in %) of the amount of incubated material after t h of rumen
incubation (Tamminga et al., 1990).
Rumen degradation of starch of legume seeds is described well by assuming two fractions
(S and D) in the first-order kinetics degradation model, one soluble fraction of S which can
be washed out without rumen incubation and one insoluble fraction of D which is degraded
exponentially. The degradation model of starch is:
R(t) = (100 − S) × e−Kd ×t
where R(t) is the residue (in %) of the amount of incubated material after t h of rumen
incubation and assuming T0 = 0; U = 0 (Tamminga et al., 1994).
The above different models were employed to describe rumen degradative behavior and
characteristics of protein and starch separately in legume seeds.
3.2. A modern protein evaluation system: the DVE/OEB model
More recently evaluation systems based on protein truly or apparently absorbed from in-
testine have been developed such as ARC (1984), NKJ-NJF (1985), PDI (INRA, 1978), AP
(NRC, 1985). Based on the principles already formulated in existing modern protein evalu-
ation systems as mentioned above, a new modern protein evaluation system: the DVE/OEB
model (Tamminga et al., 1994) has been developed. This new system considers the strong
elements of other recently developed protein evaluation systems and it also introduces new
elements, such as the role of energy balance in protein supply.
In the DVE/OEB system, each feed has a protein DVE value, which stands for true ab-
sorbable protein in the small intestine, composed of the digestible true protein contributed
by digestible feed true protein escaping rumen degradation (ABCP), digestible true micro-
bial protein (AMP) synthesized in the rumen and a correction for endogenous protein losses
(ENDP) in the digestive tract. Each feed also has a protein OEB value, which stands for
rumen degraded protein balance. The OEB value shows the (im)balance between microbial
protein synthesis potentially possible from available rumen degradable protein (N MP),
and that potentially possible from the energy extracted during anaerobic fermentation in the
rumen (E MP). When OEB is positive, it indicates the potential loss of N of a feed from the
rumen. When negative, microbial protein synthesis may be potentially impaired because of
a shortage of N of a feed in the rumen (Tamminga et al., 1994).
To describe quantitatively aspects of how the heat treatments affect legume seeds degrada-
tion and digestion in detail, proven evaluation methods, in sacco and mobile bag techniques,
and the newly developed protein evaluation DVE/OEB model (Tamminga et al., 1994) were
employed. Using the DVE/OEB model could provide estimates of rumen degradation char-
acteristics (S, D, U, Kd , T0 , ED), BCP, BSt, FOM, E MP, N MP, TPSI, AMP, ENDP, ABCP,
DVE and OEB values. The most important is that this DVE/OEB model can give the quanti-
tative aspects of protein potential degradation and digestion of both raw and heat-processed
legume seeds in detail. It can predict the changes of potential nutrient supply to ruminants
from legume seeds after heat processing. It can also provide information for the decision for
the optimal treatment conditions of legume seeds in term of protein DVE and OEB values,
under certain circumstances.
The detailed concepts and formulas of the DVE/OEB model were provided by Tamminga
et al. (1994). Here, a brief explanation and points were given as follows in order to understand
how to use it to calculate and predict potential protein nutrient supply of legume seeds
affected by heat processing.
(1) The BCP values were calculated as: BCP (g/kg DM) = 1.11 × CP (g/kg DM) ×
%BCP/100, where BCP was calculated using a Kp value of 6%/h; the factor 1.11 in
the formula was the regression coefficient of in vivo on in sacco degradation data.
(2) The BSt were calculated as: BSt (g/kg DM) = starch (g/kg DM) × %BSt/100, where
BSt was calculated using a Kp value of 6%/h. For the factor 0.1 in the formula and
assuming that for starch 10% of S escape rumen fermentation (Tamminga et al., 1994).
(3) The FOM in the rumen was calculated as: FOM (g/kg DM) = DOM (g/kg DM) −
CFat (g/kg DM) − BCP (g/kg DM) − BSt (g/kg DM) − FP (g/kg DM), where FP:
fermentation products for conserved forages, not for legume seeds.
(4) Subsequently, E MP was estimated as: E MP (g/kg DM) = 0.15 × FOM (g/kg DM),
where the factor 0.15 means that per kg FOM, 150 g of microbial protein is assumed
to be synthesized.
(5) The TPSI value was calculated as: TPSI (g/kg DM) = BCP (g/kg DM) + 0.75 × MP
(g/kg DM), where factor 0.75 means that 75% of microbial N is present in AA, the
remaining part of N in nucleic acids.
(6) The previous BCP and TPSI did not give exact enough information on the amount of
AA absorbable from the small intestine. A correction is needed for protein losses due
to incomplete digestion and resulting from endogenous excretion.
(7) True digestibility of microbial protein is assumed to be 85% (Egan et al., 1985) and,
therefore, the amount of AMP can be estimated as: AMP (g/kg DM) = 0.85 × 0.75 ×
0.15 × FOM (g/kg DM).
(8) The ABCP value is calculated as: ABCP (g/kg DM) = dBCP (%)/100 × BCP (g/kg
DM).
(9) In DVE/OEB model, ENDP in the intestine is related to the amount of DM excreted in
the feaces. According to the DVE/OEB model, 75 g of absorbed protein per kg DM in
feacal excretion is required to compensate for endogenous losses. Therefore, ENDP is
estimated as: ENDP (g/kg DM) = 75×UDM (g/kg DM), where UDM (g/kg DM) =
UOM (g/kg DM) + UASH (g/kg DM); UOM (g/kg DM) = OM (g/kg DM) − DOM
(g/kg DM); UASH (g/kg DM) = ASH (g/kg DM) − ASH (g/kg DM) × dASH (%),
dASH are 50% (CVB, 1996).
(10) The DVE value was estimated as: DVE (g/kg DM) = ABCP (g/kg DM) + AMP
(g/kg DM) − ENDP (g/kg DM).
(11) The OEB value is balance between potentially microbial protein synthesis from ru-
men degradable CP and that potentially from the energy extracted during anaero-
bic fermentation in the rumen. Therefore, the OEB value was estimated as: OEB
(g/kg DM) = N MP (g/kg DM) − E MP (g/kg DM), where N MP (g/kg DM) =
CP (g/kg DM) − BCP (g/kg DM) = CP (g/kg DM) − 1.11 × %BCP/100; E MP
(g/kg DM) = 0.15 × FOM (g/kg DM).
4. Heat processing methods for legume seeds
Some chemical properties of legume seeds reduce the biological availability and di-
gestibility of some nutrients thus lowering the feeding value. Various heat processing tech-
niques have therefore been developed in an attempt to improve utilization of these nutrients.
Heat treatments of legume seeds can inactivate ANFs (Van der Poel et al., 1990), it also ef-
fective in reducing the solubility of the protein and/or starch in the rumen and increasing the
amounts of protein and/or starch entering the small intestine for absorption and digestion.
It is feasible, environmentally safe and relatively cheap compared to other methods (such
as plant breeding and chemical treatments), and does not pose any health hazards to ani-
mals or consumers (Goelema, 1999). In particular heat treatment may have advantages for
practical application and for economical and ecological reasons. Dry roasting, microniza-
tion, pressure toasting, extrusion, expander treatment, pelleting are the methods that have
been found to be used for heat processing legume seeds for ruminants in publications.
The methodological description of those heat processing was reviewed by Goelema (1999)
as follows:
4.1. Dry roasting
Dry roasting is a dry heat treatment, in which heat is transferred by conduction, convection
and radiation. Heat can originate from gas burners or from electrical heaters. Moisture levels
should be adjusted before the treatment. Processing temperature can be up to 200 ◦ C, while
residence time is unlimited and may exceed several hours (Plegge et al., 1985; Baiden,
1997; Yu, 1999; Yu et al., 1999; Yu et al., 2001a,b,c).
4.2. Micronization
Micronization involves internal heating of the legume seeds by gas fired infrared heaters,
to temperatures between 125 and 135 ◦ C over a residence time of 40–90 s. The infrared rays
cause the molecules of the material to vibrate in accordance with the resonance to their own
frequency resulting in rapid internal heating and a loss in moisture content (Melcion and
van der Poel, 1993).
4.3. Pressure toasting
Pressure toasting can be carried in pressurized barrels. There is a positive relation be-
tween steam pressure and the temperature in the pressurized barrels. Processing time and
temperature can be varied (Van der Poel et al., 1990; Melcion and van der Poel, 1993; Yu,
1995; Goelema, 1999; Yu et al., 2000a).
4.4. Extrusion
Extruding consists of passing dry whole seeds through tapered screw(s) in a fixed metal
barrel. The screws convey the ground material from the feed end of the barrel to the die(s),
cooking it by heat due to friction and shear, and additional heat applied to the barrel (Melcion
and van der Poel, 1993). As the material exits the die, steam derived from the moisture
content is flashed due to pressure differential expanding the feed material. The combination
of temperature, pressure, moisture and shear, followed by expansion when the material
leaves the barrel, changes the properties of the material, including its digestive behavior in
the rumen. Processing time in extruders varies from 30 to 50 s, while temperatures range
from 80 to 200 ◦ C (Cros et al., 1991, 1992; Kibelolaud et al., 1993; Goelema, 1999).
4.5. Expander treatment
Expanders are similar to single screw extruders, but have an annular discharge valve,
instead of a die. An electrically or hydraulically adjustable cone is used to increase the
pressure during operation up to 3800–4000 kPa (Pipa and Frank, 1989). Steam can be used
for heating the barrel wall, as well as for injection in the feed mash to increase processing
temperature. Residence time in the expander varies from 5 to 15 s, while temperatures are
ranging from 80 to 140 ◦ C. Shear action during expander treatment is much less than during
extrusion (Goelema et al., 1996; Goelema, 1999).
4.6. Pelleting
Pelleting involves the compression of a conditioned feed mash through a die. Residence
time in the die does usually not exceed 15 s. Conditioning can be carried out by the addition
of water and/or steam, which influences the amount of friction between feed particles, the
barrel wall and in the die. The pelleted mashes are usually pre-conditioned at temperature
ranging from 65 to 90 ◦ C.
5. Digestive characteristics and behaviors of legume seeds as ruminant feedstuffs
5.1. Legume seeds and common and alternative names
The leguminosae is a large family of plants with about 12,000 recognized species. Four
tribes within the family are of importance since they include all the common peas and beans:
(1) the Hedysareae contains the groundnut; (2) the Vicieae contains the genera Vicia, Cicer,
Pisum, Lens and Lathyrus; (3) the Genisteae contains the genus Lupinus; (4) the Phaseoleae
contains the genera Phaseolus, Dolichos and Glycine (McDonald et al., 1988). Table 1 gives
the botanical name and common and alternative names for legume seeds.
5.2. Nutrient values of legume seeds
Legume seeds are primarily used as food for human consumption. They are also used as
feedstuffs in animal nutrition. Among them, lupin seeds, peas, faba beans and soybeans are
four major commercially important legume seeds in crop rotations due to their wide use
as feedstuffs (Batterham and Egan, 1986). They represent the range of legume seeds. They
have high protein and energy contents, are highly digestible and are palatable. Among them,
they also have differences. Lupin seeds and soybeans contain hardly any starch, but faba
bean and peas contain >40% starch content (Petterson and Mackintosh, 1994; Goelema,
1999; Yu, 1999). The AA profile of legume seeds generally complements that of cereal
grains and only small amounts of limiting AA, usually lysine and sometimes methionine,
need to be added or drawn from alternative protein sources to give a balanced diet (Petterson
and Mackintosh, 1994). Table 2 gives the nutrient levels for four major legume seeds used
as animal intestinal protein sources.
5.3. Rumen degradation characteristics of raw legume seeds
5.3.1. Rumen protein degradation of raw legume seeds

In practical dairy nutrition, the use of legume seeds is limited and their utilization is
inefficient. An important reason is that for legume seeds (lupin, faba beans, peas and soy-
beans) the soluble protein or rapidly degradable protein content is a high proportion of total
Table 1
Botanical name and common and alternative names for legume seeds
Botanical name Common name and alternative name
Cajanus cajan Pigeonpea, Congo pea, Cajan, red gram, Congo pea, dhal arhar
Canavalia ensiformis Jack bean, sword bean
Cicer arietinum Chickpea (desi or Kabuli type), garbanzo, Bengal gram, Egyptian pea, gram,
chana, shihu
Cyamopsis tetragonoloba Guar, cluster bean
Lablab purpureus Lablab, Dolichos lablab, hyacinth bean, bonavist(a), Indian (butter) bean
Lathyrus sativus Grass pea, Chickling vetcch
Lathyrus cicera Flat pod pea vine
Lathyrus ochrus Cyprus vetch
Lens culinaris Lentil, masurdahl, adas, mercimek
Lupinus angustifolius Narrow-leafed lupin, blue pupin (Europe), sweet lupin, Australian sweet lupin,
NZ blue (bitter)
Lupinus albus White lupin, white lupin(e), Lupini bean, Albus pupin, L. graesus, L. termis
Lupinus cosentinii Sandplain lupin, WA blue lupin
Lupinus luteus Yellow lupin
Marotyloma uniforum Horse gam
Medicago sativa Lucerne, alfafa
Phaseolus vulgaris Navy bean, Borlotti bean, Great Northern, Culinary bean, butter bean, pinto
bean, dry bean, haricot bean, kidney bean
Phaseolus lunatus Lima bean, butter bean
Phaseolus polyanthus Year bean
Phaseolus coccineus Runner bean
Phaseolus acutifolius Tepary bean
Pisum sativum Marrowfat pea, small blue pea, field pea, dry pea, arveja, muttar
Psophocarpus tetragonoloba Winged bean, Goa bean, four-angled bean, Manila bean, Princess pea
Trigonella foenumgraecum Fenugreek
Vicia faba Faba bean, field bean, horse bean, tick bean, baakla, feverole, fulmedames
Vicia narbonensis Narbon bean, bakla balcigi, ful iblis, koca fig, mohren
Vicia sativa Vetch, Chickling pea, khesari
Vicia articulata One-flowered vetch
Vicia villosa Woolly pod vetch
Vigna radiata Green mung bean, green gram, golden gram, moong
Vigna mungo Black mung bean, black gram, urd, matpe, mungo bean, mash
Vigna unguiculata Cowpea, crowder pea, black-eyed pea, lobtya
Vigna angularis Adzki bean, V. angulans, haricot konde
Vigna aconitifolia Moth bean, mat bean
Voandzeia subterranea Bambara groundnut
Source: Petterson and Mackintosh (1994).
protein. Van Straalen and Tamminga (1990) stated that the protein fraction in these legume
seeds consists of 85–100% albumins and globulins, which are highly soluble and easily
degradable (up to 75%) in the rumen. A high rumen degradation rate causes an imbalance
between seed protein breakdown and rate of microbial protein synthesis, resulting in un-
necessary N loss from rumen (Van Straalen and Tamminga, 1990; Tamminga et al., 1990).
High degradability of legume seeds is also contributed to by the high ratio of available N
to available carbohydrates (Yu, 1999).
Table 2
Nutrient values of four major legume seeds (lupin seeds, faba beans, peas and soybeans, %)
Nutrient Lupin seeds Faba beans Peas Soybeans
Crude protein 35.0 30.6 26.1 39.4

Total lysine 1.51 1.50 0.16 0.28
Total sulphur amino acids 0.71 0.53 0.57 0.13
Total tryptophan 0.42 0.16 0.21 0.52
Crude fat 5.6 1.2 0.9 19.8
ME, cattle (MJ/kg) 12.0 13.4 11.7 13.7
ME, sheep (MJ/kg) 12.4 11.5 12.0 13.7
Total calcium 0.21 0.12 0.09 0.40
Total phosphorus 0.33 0.44 0.39 0.60
Cruder fiber 14.4 8.2 5.4 6.7
ADF 19.7 11.2 7.8 9.0
NDF 24.9 15.2 11.9 12.6
Lignin 0.9 0.3 0.2 1.05
Source: Petterson and Mackintosh (1994) and Goelema (1999).
Table 3 presents the rumen protein degradation characteristics (S, D, U, Kd , T0 , BCP) of

those four major legume seeds, raw lupin seeds, faba beans, peas and soybeans, as evaluated
in different laboratories. They provided evidence that protein degradabilities of lupin seeds,
faba beans, peas and soybeans are very high around 62–95, 83–90, 79–88, and 62–98%,
respectively. There would thus be little feed BCP for digestion in the small intestine.
5.3.2. Rumen starch degradation of raw legume seeds

In reports on use of legume seeds as feedstuffs for dairy cows, little attention is paid to
the starch digestive behavior. Information in the literature on rumen starch degradation of
legume seeds is scarce. In Table 4, rumen starch degradation characteristics of lupin seeds,
faba beans and peas from our laboratories are presented. Table 4 shows not only is protein
degradability high in legume seeds, but also the starch degradability. The studies indicate
that starch in lupin seeds, faba beans and peas has a high Kd (12.5, 5.0–12.2 and 5–14%/h,
respectively), and very high rumen degradabilities (88, 71–88 and 68–78%, respectively).
Therefore, in order to utilize these kinds of legume seeds in a more efficient manner and
prevent imbalance between nutrients breakdown and microbial synthesis in ruminants, the
high rate of degradation of protein and/or starch should be reduced.
6. Manipulation of digestive behavior and characteristics of protein and/or

starch of legume seeds by heat treatments
6.1. Effect of heat treatments on protein digestive behavior of legume seeds
Many attempts have been made to protect protein in these grain legumes against rapid
rumen degradation by treatments involving application of heat with or without moisture
(Cros et al., 1991, 1992; Aguilera et al., 1992; Robinson and McNiven, 1993; Singh et al.,
1995; Yu, 1995; Goelema et al., 1996, 1999; Yu et al., 1999; Goelema, 1999; Yu et al.,
2000b, 2001a). The objective of heat treatment is to reduce rumen degradation of protein
Table 5
Effect of heat treatment on denaturation and degradation of protein
Temperature (◦ C) Effect of heating
50 Increase hydration, some loss of crystalline structure

70–80 Disulfide splitting, loss of tertiary structure
80–90 Loss of secondary structure disulfides
90–100 Intermolecular disulfides formed
100–150 Lysine and serine loss, isopeptide formation
150–200 Peptidization and more isopeptide formation
200–250 Pyrolysis of all amino acid residues
Sources: Finley (1989) and Goelema (1999).
and increase BCP without adversely affecting total tract protein digestion. In other words,
heat treatments may provide the means of shifting protein digestion to the small intestine
to provide more dietary AA to the animal. In this way, under certain circumstances, it will
benefit to animals.
The mechanism of altering the degradative behavior of protein, as described by Goelema
(1999), with heat treatments involves principally denaturation (which is a disorganization
of the overall molecular shape of a protein); unfolding or uncoiling of a coiled or pleated
structure, or the separation of the protein into its subunits, which may then unfold or uncoil
(Holum, 1982). Any temperature change in the environment of the protein which can influ-
ence the non-covalent interactions involved in the structure may lead to an alteration of the
quaternary, tertiary and secondary structure. More extreme forms of processing may lead to
the destruction of the primary structure, often called ‘degradation’ of protein (Finley, 1989).
Depending on the processing conditions several processes may occur (Table 5), as summa-
rized by Goelema (1999), although it is obvious that not only temperature during treatment
plays a role, but also factors such as residence time and moisture level. Reactions of the
Maillard type are very common (Hurrell and Finot, 1985). These include non-enzymatic
browning, involving condensation reactions between primary amines and reducing sug-
ars, followed by isomerization and polymerization of the products formed. Epsilon amino
groups of lysyl residues are the most reactive amines in proteins. As products formed by
isomerization and polymerization reactions are not digestible, the Maillard reactions has
been perceived by nutritionists as being only detrimented to the nutritional value of pro-
teins. However, Finot et al. (1977) showed reaction intermediates preceding isomerization
to be fully available to rats. If properly controlled, non-enzymatic browning might be an
effective means of protecting soybeans from ruminal proteolysis without reducing protein
digestibility. Cleale et al. (1987a,b,c) reported that controlled, non-enzymatic browning may
be effective for reducing ruminal degradation of soybeans, increase flow of soybean N to the
intestine; improved efficiency of soybean protein utilization by ruminants. Other reactions
also occur (Goelema, 1999) including the formation of isopeptide cross-links between lysine
residues and asparagine and glutamine. Additionally, methionine, cystine and tryptophan
may be involved (Broderick et al., 1991). Heat treatment results in structure stabilization
and cross linkages to carbohydrates, which protects them from ruminal hydrolysis or at
least slows down their rate of degradation. Such linkages are beneficial in the rumen, unless
they are not reversible in the intestine (Goelema and Tamminga, 1994; Goelema, 1999).
Generally, common heat treatments applied to grain legumes are pressure (steam) toasting
(Yu, 1995; Goelema, 1999; Yu et al., 2000a), extrusion (Cros et al., 1991, 1992; Kibelolaud
et al., 1993), autoclaving (Aguilera et al., 1992) and roasting (Robinson and McNiven, 1993;
Singh et al., 1995; Yu, 1999) (Table 6). The rate and extent of rumen degradation of legume
seeds depend on the treatments applied. The extent of intestinal digestion also depends on
the type and degree of treatments (Cros et al., 1991 and 1992; Goelema, 1999). The type of
grain legumes and proportion in the diet can also influence digestion site.
In the publications, only a few studies (Yu, 1995, 1999; Goelema, 1999; Yu et al., 2000a)
did systematic research on legume seeds, at various conditions, quantitatively evaluated
changes of nutritional values and determined the optimum heat processing conditions in
term of protein DVE and OEB values. The studies (Table 6) show that pressure toasting
(Yu, 1995; Goelema et al., 1996, 1999; Goelema, 1999; Yu et al., 2000a) increased BCP of
lupins, horse beans, peas and soybeans from 21 to 51, 17 to 53, 21 to 54 and 28 to 43%, re-
spectively. This was mainly because pressure toasting reduced S, increased D and decreased
Kd . The effects increased with pressure temperature and time (Table 6). The digestibility of
BCP in the small intestines and total tract digestion of CP were not significantly affected by
pressure toasting (Goelema, 1999; Yu et al., 2000a). Yu (1999) did systematic research on
effect of dry roasting (which is as a common heating method used on farm basis in Australia)
on white lupin seeds and whole faba beans. The results also show that dry roasting could be
used a method to protect protein of legume seed degradation in the rumen and shift more
legume protein into intestines (Table 6). Cros et al. (1991) reported that extrusion of whole
lupin seeds at 120, 150 and 195 ◦ C decreased N degradability value: 83.9, 72.9 and 53.0
versus 95.3% (raw), assuming Kp of 6%/h. He also reported the total tract N disappearance
was high for each treatment, all at approximately 98%. Such results indicated that extru-
sion can shift protein digestion from rumen to the intestines without adversely affecting the
total N digestibility (Table 6). However, the effect of even higher extrusion temperatures
on rumen degradation and overall N disappearance from the digestive tract is unknown,
as optimum conditions were not determined. Kibelolaud et al. (1993) undertook a similar
study to that of Cros et al. (1991) and found that extrusion did improve nutritional value
of whole lupin seeds for the ruminant in terms of BCP and total tract digestion of protein.
Again no optimal treatment was found in the study.
However, undesirable effects of heat treatments through increasing rumen protein degra-
dation have also been reported. Goelema, et al. (1999) reported that pelleting did not re-
duced but increased rumen degradation of protein in mixture of peas, lupins and faba beans
(Table 6). In terms of the rumen degradation characteristics identified before, pelleting sig-
nificantly increased Kd (3.6 versus 10%/h) and decreased BCP (64.7 versus 28.0%). This
study thus indicated that pelleting would be expected to shift the site of digestion to the
rumen rather than to the intestines.
6.2. Effect of heat treatments on starch digestive behavior of legume seeds
Less research has been done on the digestive behavior of starch as affected by heat
treatments and there is little reported work particularly on legume seeds. Recent data
(Table 4) shows that starch from peas and faba beans is highly soluble and easily degrad-
able. The purpose of heat treatments of starch in legume seeds is to reduce the degradation
rate in the rumen and through this to manipulate the amount degraded in the rumen
and digested in the small intestine, which may benefit to animals under certain
circumstances.
The mechanism of altering the degradative behavior of starch with heat treatment involves
three processes: swelling, gelatinization and retrogradation (Nocek and Tamminga, 1991).
Goelema reported that swelling results from the exposure of starch to water when gradually
heated (55 ◦ C). At low temperature (<60–80 ◦ C), swelling is reversible after cooling and
drying. Above that temperature irreversible gelatinization takes place. Starch granules lose
their crystallinity (French, 1973). Retrogradation of starch is the reassociation of starch
molecules separated after gelatinization, in which hydrogen bonding between amylose and
amylopectin is reestablished. Retrograded starch does not completely regain the native
starch character, but changes can be reversed to some degree by reheating (Rooney and
Pfugfelder, 1986).
Few systematic studies were found on rumen starch degradation characteristics of legume
seeds affected by heat treatments. The data presented in Table 7 were from our laboratory.
The effects of heat treatment on starch degradability of legume seeds are very much depen-
dent on the treatment type and the conditions applied during processing. Yu et al. (1999),
Goelema (1999) and Yu et al. (2000a) found that both pressure toasting and dry roasting
decreased starch degradation of faba beans and increased BSt (Table 7). The effects of
increasing BSt or reducing ED of starch increased with increasing time and temperature.
Also experiments done by Goelema et al. (1999) showed that compared with the control
sample of peas, pressure toasting increased BCP from 32.3 to 50.5% mainly due to pressure
toasting decreasing S from 52.2 to 11.1% and increased D from 47.8 to 88.9%. But pressure
toasting did not affect Kd much. All above methods decreased rumen starch degradation
and increased the rumen BSt. In contrast, Goelema et al. (1999) found that pelleting of
mixture of legume seeds, peas, faba beans and lupins did not increased BSt but reduced BSt
(Table 7).
Therefore, changes in starch degradability in the rumen in response to heat treat-
ments appear to differ among feeds. Such variability in effects of heat treatment may be
related to differences in the crystallinity of the starch source and/or in the association
between starch and the protein matrix surrounding the starch granules (Theurer, 1986).
Differences in degradative behavior of starches can also be caused by several other fac-
tors such as their amylose and amylopectin content, crystallinity, particle size and the
presence of enzyme inhibitors (Cone and Wolters, 1990), as reported by
Goelema (1999). Such factors may contribute to different characteristic effects of heat
treatments on the digestive behavior of starch in legume seeds when compared with cereal
grains.
6.3. Physicochemical characteristics of starches between legumes and cereals grains
In cereal grains, the effect of heat treatments is usually to increase rumen degradability
of starch (Theurer et al., 1999). However, in legume seeds, heat treatments except pelleting
decreased rumen degradability of starch (Yu, 1995; Goelema, 1999; Yu, 1999). This may be
contributed to different physicochemical characteristics of starches between legume seeds
and cereals grains.
The study by Naivikul and D’Appolonia (1979) showed that the starch content in various
legumes was considerably lower than in cereals such as wheat (35–43 versus 70–80%).
Legume starch granules were oblong and did not exhibit as wide a variation in size as in
wheat, which has a mixture of large, intermediate and small spherical granules
(Kulpd, 1973; Naivikul and D’Appolonia, 1979). Birefringence of the legume starch gran-
ules was similar among the various legumes, but differred from the cereals. Legume starches
had a higher resistance to swelling and rupture than cereal starch. The pastes in legume
starches were relatively stable and that the granules did not rupture, which was not the case
with cereal starches. The amylose content for the legume starches was slightly lower than
that for cereal starches. Acetylated legume starch amylose had a lower molecular weight
than did the acetylated cereal starch amylose (Naivikul and D’Appolonia, 1979). Legumes
starches exhibited a highly hydrated fine fiber fraction (presumably from the cell walls en-
closing the starch granules) and also the high content of insoluble protein. Legume starch
granules were irregularly swollen with an elongated hilum (Schoch and Maywald, 1968).
Madhusudhan and Tharanathan (1996) did structural studies of linear and branched frac-
tions of legume and cereal starches by using beta-amylase and pullulanase to explain at a
molecular level the differences in the digestibility of legume (chickpea, Bengal gram) and ce-
real (finger millet, ragi) starches. The authors found that legume starches were less digestible
(approximately 45%) with glucoamylase than the cereal starches (>70%). Of the various
fractionation methods employed, concanavalin A precipitation gave pure amylopectin (Ap).
The crude amylose on subfractionation with hot 1-butanol yielded pure amylose (Am) and an
intermediate fraction (Ax). The content of “true” Am and Ax was higher in legume (33 and
6.5%) than in cereal starches (22 and 6%, respectively). The molecular weight of legume
starch fractions was much higher than those of cereal starch fractions. Beta-amylolysis
and debranching studies indicated legume-Am to contain sparsely distributed side-chain
branches, whereas cereal-Am was more linear. The legume-Ap was comprised of very long
B (dp 70), long B (dp 56), long A/short B (dp 25), and short A (dp 17) chains; whereas
cereal-Ap had long B (dp 48), short B (dp 25), and short A (dp 17) chains. The structure of
Ax lay between those of Ap and Am.
Above studies clearly show that physicochemical characteristics of legume and cereal
starches are quite different. This may partly explain the different degradative behavior of
legume and cereal starches due to structural differences.
The study by Hoover and Vasanthan (1994) on effect of heat-moisture treatment (at
100 ◦ C/16 h at moisture contents between 10 and 30%) on the structure and physicochemical
properties of cereal (wheat, oats) and legumes (lentil) found that the heat treatment did not
change granule size and shape. However, in cereal starches, granules were less compactly
packed after heat treatment. The X-ray diffraction patterns of cereal starches remained
unchanged, while those of legume starches became more cereal-like. In all starches, the
swelling factor and amylose leaching decreased. Heat treatment induced complex formation
between amylose and native lipids. Differential scanning calorimetry of the heat-treated
samples showed a broadening of the gelatinization-temperature range and a shifting of the
endothermal transition towards higher temperatures. The gelatinization enthalpy of cereal
and legume starches remained unchanged on the heat treatment. Heat treatment increased
the 95 ◦ C viscosity of cereal starches, but decreased those of legume starches. The results
indicated that the extent of starch-chain associations within the amorphous regions and the
degree of crystalline order were altered during heat-moisture treatment. The magnitude of
these changes were found to be dependent upon the moisture content during heat treatment
and on the starch source.
Theurer et al. (1999) reported that increased moisture, temperature and pressure consis-
tently increases in vitro enzymatic starch hydrolysis of grains and the proportion of starch
digested in the rumen and the total starch digestibility by cattle. These increased grain di-
gestibilities are caused by disruption of the protein matrix surrounding the starch granules
in the grain endosperm and disorganization of the starch granules.
The above information provide different inside physicochemical characteristics of starches
between legumes and cereal grains and how they are affected by heat processing.
7. The prediction of changes of potential nutrient supply to ruminants from legume

seeds affected by heat processing, using the newly developed DVE/OEB model
A number of publications (such as Annexstad et al., 1987; Cros et al., 1991, 1992;
Benchaar et al., 1994) indicated that heat treatment does reduce ED and increase BCP, but
none of them provide details how heat processing affected microbial protein synthesis and
potential nutrient supply and none of them were optimal heating conditions identified or
studied. Heating above the optimal temperature may overprotect the protein so that the pro-
tein is neither fermented in the rumen nor digested in the small intestine (Stern et al., 1985).
Yu (1995, 1999), Yu et al. (2000a) and Goelema (1999) have attempted to use the newly
developed DVE/OEB model to predict potential microbial protein synthesis and potential
nutrient supply to ruminants from four major legume seeds as indicated before affected by
pressure toasting (Table 8) and dry roasting (Table 9) and to determine optimal processing
conditions of pressure toasting and dry roasting of faba beans, lupin seeds, peas and soybeans
in term of the protein DVE and OEB values, according to protein evaluation DVE/OEB
model. These studies systematically evaluated the effects of heat treatments on rumen
degradation and intestinal digestion characteristics of raw and heated legumes seeds at
various conditions to give the quantitiative aspects of how the heat treatments affected
protein degradation and digestion in detail and provided information for ration formula and
the decision for the optimal treatment conditions in dairy industry. The items assessed in
those experiments were not only rumen degradation characteristics in terms of S, D, U, Kd ,
T0 , ED but also BCP, BSt, FOM, ABCP, E MP, N MP, TPSI, AMP, ENDP, DVE and OEB
values. The most important values are potential protein DVE and OEB values, which were
changed by heat processing for each legume seed.
Tables 8 and 9 show the prediction of potential nutrient supply to ruminants from raw
and heated lupin seeds, faba beans, peas and soybeans by using the DVE/OEB model. Input
data from in sacco and mobile bag techniques were from dairy cows. Consequently, the
data generated by the model, though of significance in the dairy cows, are best regarded as
characteristics of the test materials.
These studies indicated that pressure toasting (Table 8) significantly increased the pre-
dicted BCP (76–185; 47–148; 58–149; 121–189 g/kg DM for lupin seeds, faba beans, peas
and soybeans, respectively), increased BSt (94–174; 158–248 g/kg DM for faba beans and
peas; lupin seeds and soybeans contain little starch and no BSt was reported), increased
true protein value of TPSI supplied to the small intestine (164–260; 121–200; 136–206;
186–246 g/kg DM for lupin seeds, faba beans, peas and soybeans, respectively), increased
truly intestinal absorbed rumen undegradable protein value of ABCP (71–176; 40–139;
54–145; 102–178 g/kg DM for lupin seeds, faba beans, peas and soybean, respectively),
and increased truly digested intestinal protein value of DVE (133–232; 89–170; 115–189;
152–222 g/kg DM for lupin seeds, faba beans, peas and soybean, respectively). Pressure
toasting (Table 8) decreased rumen fermented OM value of FOM, microbial protein value
of E MP synthesized in the rumen based on available energy, truly intestinal absorbed
rumen synthesized microbial protein value of AMP, microbial protein value of N MP syn-
thesized in the rumen based on available N and degraded protein balance value of OEB
with increasing temperature and time (Table 8).
Though pressure toasting (Table 8) reduced microbial protein synthesis due to a reduction
in FOM and a reduction in rumen protein degradation, the total truly digested intestinal
protein value of DVE did not decrease but increased markedly. This was due to the fact that
absorbable BCP value was increased more than enough to compensate for the computed
decrease in microbial protein production. Therefore, the net absorbable DVE value in the
animal was substantially increased (DVE = ABCP + AMP-ENDP). The largest increase
of the DVE value for all legume seeds investigated was found at 136 ◦ C/15 min of pressure
toasting, which highly increased the net absorbable protein DVE value 76, 90, 64 and 45%
for lupin seeds, faba beans, peas and soybeans, respectively, as AA source in the intestines.
When the OEB value of a feed is positive, it indicates a potential N loss from the ru-
men. When negative, microbial protein synthesis is predicted to be impaired because of
a shortage of N in the rumen of this feed. The studies (Table 8) showed that raw lupin
seeds, horse beans, peas and soybeans all had high OEB values (142, 101, 88 and 186 g/kg
DM, respectively), which indicated a potential imbalance between feed N degradation and
utilization and indicated a potentially large N loss from rumen. Pressure toasting reduced
the OEB values with increasing time and temperature, but did not cause them to become
negative (142–44; 101–32; 88–25; 186–128 g/kg DM for lupin seeds, horse beans, peas and
soybeans, respectively). High positive OEB values in the most treatments except 136 ◦ C for
15 min for horse beans and peas indicated that there were still large potential losses of N
in the rumen. In other words, the treatments of 100 ◦ C/7, 15 or 30 min, 118 ◦ C/3, 7, 15 or
30 min and 136 ◦ C/3 or 7 min were not sufficient to reduce N loss in the rumen if they are
used as rumen undegradable protein sources for high production ruminants.
The above results indicate that with pressure toasting, temperature and/or duration could
go still higher than 136 ◦ C and/or longer than 15 min with soybeans and lupins (due to their
high OEB values, 128.2 and 44.4 g/kg DM, respectively) to further prevent potential N loss
in the rumen if total tract digestion is not depressed. But pressure toasting up to 136 ◦ C
for 15 min might cover the optimal treatment range for peas and faba beans (under certain
circumstances) in terms of treating to achieve target values for potential high net absorbable
protein in the small intestine while holding any N loss in the rumen to a low level.
Yu et al. (1999) used the same approach by using the DVE/OEB model to predict changes
of potential nutrient supply to ruminants from whole lupin seeds and whole faba beans after
dry roasting at various conditions, which are commonly used for ruminants on farm basis
in Australia (Table 9) and to determine the optimal dry roasting conditions under certain
circumstances for whole lupin seeds and whole faba beans) (in terms of treating to achieve
target values for potential high net absorbable protein in the small intestine while holding
any N loss in the rumen to a low level), and found that dry roasting of whole lupin seeds and
whole faba beans increased BCP, BSt, TPSI, ABCP and DVE values and decreased FOM,
E MP, AMP, N MP and OEB with increasing temperature and time. Though dry roasting
was effective in shifting protein degradation from rumen to intestine, and increased net
absorbable protein DVE value, dry roasting up to 150 ◦ C for 45 min could not fully prevent
potential N loss in the rumen for whole faba beans (due to the high OEB value). But dry
roasting up to 150 ◦ C for 45 min of whole lupin seeds might cover the optimal treatment
range (Table 9) due to its high net absorbable DVE value (increasing from 110 to 197 g/kg
DM) and lowest OEB value (reducing from 157 to 26 g/kg DM).
All the results reported here are output from a model with inputs based on in sacco and
mobile bag techniques measurements. The challenge is to apply the predictions and eval-
uate them in an animal performance experiment. However, the number of such studies in
this area available to challenge the model is extremely limited. Part of reason is that the
information on DVE and OEB values of each feedstuff in the diets, or data from which
these are derived is extremely limited.
8. Conclusions
Legume seeds (lupin seeds, faba beans, peas and soybeans) have high ratio of available
N to available carbohydrates (>25–33 g N/kg DM of optimum ratio). The soluble or rapidly
degradable protein and starch of legume seeds is too high, resulting in unnecessary potential
nutrient losses from the rumen.
The rumen degradation characteristics of protein in raw legume seeds have been studied
extensively, but little information was found on the rumen degradation characteristics of
starch in legume seeds. This suggests the need for further investigation.
Feed processing, involving heat methods, has potential for application as an effective
means to improve ruminal digestion and nutrient conservation of legume seed components.
The site of digestion of starch and protein of legume seeds can also be manipulated by
processing.
Heat treatment (pressure toasting, dry roasting, extrusion, etc.), except of pelleting, can
reduce the rate and extent of rumen degradation of both protein and starch in legume seeds,
thus resulting in a potential increase in the supply of protein (as AA source) and/or starch
(as glucose source) to the small intestines. However, optimal processing conditions have
not yet been established for individual legume seeds in terms of maximizing AA absorption
in the small intestines.
The newly developed DVE/OEB model appears to have characteristics that can provide
relatively accurate information on the quantitative aspects of both ruminal and post-ruminal
feed protein digestion in ruminants for each feedstuff. With the DVE/OEB model, it is pos-
sible to predict the potential nutrient supply to the animal from legume seeds as affected by
heat processing.
To evaluate the accuracy of the model predictions, animal performance experiments are
needed. However, the number of such studies in this area available to challenge the model
are extremely limited. Part of the reason is that the information on DVE and OEB values of
each feedstuff, or data from which these can be obtained, are limited.
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An Analysis of The Nutritive Value of Heat Processed Legume Seeds For Animal Production Using The DVE-OEB Model A Review

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An Analysis of The Nutritive Value of Heat Processed Legume Seeds For Animal Production Using The DVE-OEB Model A Review

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Animal Feed Science and Technology

An analysis of the nutritive value of heat processed

University of Melbourne, Parkville, Vic. 3052, Australia

∗ Corresponding author. Tel.: +1-306-966-4150; fax: +1-306-966-4151.

E-mail address: yupe@sask.usask.ca (P. Yu).

2. Several important aspects of ruminant digestion and nutrient utilization of

2.1. Protein of legume seeds

2.2. Starch of legume seeds

Starch, as a main component in non-structural carbohydrates of legume seeds, plays

2.3. Optimization of rumen fermentation

3. The modern evaluation system for describing digestive behavior of

3.1. Rumen degradation model

R(t) = U + (100 − S − U ) × e−Kd ×(t−T0 )

R(t) = (100 − S) × e−Kd ×t

3.2. A modern protein evaluation system: the DVE/OEB model

4. Heat processing methods for legume seeds

4.1. Dry roasting

4.3. Pressure toasting

4.5. Expander treatment

5. Digestive characteristics and behaviors of legume seeds as ruminant feedstuffs

5.1. Legume seeds and common and alternative names

5.2. Nutrient values of legume seeds

5.3. Rumen degradation characteristics of raw legume seeds

5.3.1. Rumen protein degradation of raw legume seeds

Crude protein 35.0 30.6 26.1 39.4

Table 3 presents the rumen protein degradation characteristics (S, D, U, Kd , T0 , BCP) of

5.3.2. Rumen starch degradation of raw legume seeds

6. Manipulation of digestive behavior and characteristics of protein and/or

6.1. Effect of heat treatments on protein digestive behavior of legume seeds

50 Increase hydration, some loss of crystalline structure

6.2. Effect of heat treatments on starch digestive behavior of legume seeds

6.3. Physicochemical characteristics of starches between legumes and cereals grains

7. The prediction of changes of potential nutrient supply to ruminants from legume

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