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The law of diminishing returns: consequences for feed enzyme strategy

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The law of diminishing returns:
consequences for feed enzyme strategy
by Dr A J Cowieson and Dr M R Bedford, AB Vista Feed Ingredients, United Kingdom
T
he diets of pigs and poultry are
a complex matrix of nutrients
variably contaminated with
anti-nutrients and diluents. The digesti-
bility of nutrients, such as fat, starch
and protein, is generally high (that is,
over 80%) but this can and does vary
depending on a host of diet, animal
and environmental factors.
Though feed enzymes are purported to
improve the digestibility of various dietary
nutrients, this response will vary depending
largely on the inherent digestibility of the diet
prior to enzyme intervention.
Thus, in diets with a relatively poor starch,
protein, fat or mineral digestibility there is a
greater potential for feed enzymes to elicit a
beneficial response.
A good example of this relationship is the
effect of xylanases on ileal amino acid digest-
ibility. Amino acid digestibility averages 80-85
percent but this is highly dependent on the
amino acid in question with methionine,
arginine and glutamic acid being considerably
more readily digested than cysteine, threo-
nine, serine or proline.
The consequence of this is that feed
enzymes will improve the digestibility of
cysteine and threonine to a much greater
extent than they will for methionine.
So, as with many of life’s principles, the
value of feed enzymes follows a distinct
law of diminishing return (see Figure 1) as
diet nutritional value improves. Thus, when
considering mixtures of enzymes, only the
first can carry its full matrix and all subse-
quent additives must have their contribu-
tions truncated by the contributions of the
incumbents.
By definition this suggests that the effect
of one feed enzyme is highly unlikely to be
16 | September - october 2010
Enzyme Feature Feature Enzyme
general principle that is true for xylanase The difficulty with
cannot be transferred to phytase. simply adding various
Thus, when the effect of phytase on ileal supplements to a ration
amino acid digestibility is plotted as a propor- and ascribing them the
tion of the undigested fraction the conclu- suppliers recommen-
sions are quite different to that of xylanase dations is that in most
(see Figure 4). Indeed, those amino acids instances the magnitude
which benefit most from phytase addition of effect on for exam-
are those amino acids which are present at ple, energetic efficiency
high concentrations in endogenous proteins has been assessed inde-
such as mucins and pepsin. pendent from other
The net effect of this is that when phytase additives. The matrix Figure 2: The effect of xylanase on ileal amino
acid digestibility (IAAD) coefficients from 19 peer-
is present in a diet it improves the digest- values for phytases and
reviewed papers published between 1998 and 2009
ibility of some nutrients more than others. xylanases (see Table 1),
As the response to xylanase is largely driven for example, are gener-
by the digestibility of the diet to which it ally established by feeding diets containing these Total lysine = 1.23%
is added the efficacy of xylanase will be enzymes independent of other feed additives Digestible lysine = 1.15%
fully additive with that of another unless the ibility of starch and fat where between reduced in the presence of phytase, but not and measuring some response criteria such as Undigested lysine = 0.08%
substrates and, importantly, the correspond- 15-20 percent of the undigested fraction is to the same extent for all nutrients. weight gain, conversion, bone ash, or digestibility Phytase: 0.017%
ing nutrients which are released, do not rendered digestible with xylanase. (total tract or ileal). Xylanase: 0.024%
overlap at all. Thus, the magnitude of response is largely The law of diminishing return One practical illustration of this is in the Protease: 0.048%
It is the purpose of the current article to dictated by the inherent digestibility prior to Due to upwards pressure on the price of application of amino acid matrices for feed Using phytase, xylanase and protease
summarise these principles and suggest how xylanase addition but a similar proportion of many commonly used feed ingredients such enzymes used in combination. in combination = 0.089% improvement in
they be applied to the strategic selection of the undigested fraction is captured regardless as corn, fat and protein sources, the use of Taking lysine as an example and applying digestible lysine (i.e. from 1.15% to 1.24%).
feed enzymes, with particular emphasis on of the nutrient in question. several feed additives can be, at least at face real published nutrient release values for vari- In the above example the use of sup-
admixtures of xylanase and phytase. Since xylanase improves the digestibility value, attractive for poultry producers. ous commercially available products: plier’s recommendations for lysine results in
of nutrients based largely on the quantity of
Non-starch polysaccharide- the undigested fraction, any additive which
degrading enzymes improves digestibility (reducing the undigest-
The beneficial effect of carbohydrases ed fraction) will by definition mute xylanase
on performance and nutrient digestibility is efficacy.
thought to be due to reduced intestinal vis-
cosity, reduced integrity of cell wall architec- Phytase
ture, improved microbial balance via removal The mechanism of action of exogenous
of fermentable starch and protein and provi- phytases are quite different to xylanases.
sion of oligomers from fibre digestion which Phytate represents a significant antinutri-
prove pre-biotic in their
nature.
Interestingly, recent
evidence suggests that
a reduced intestinal vis-
cosity may be the most
immediately relevant, even
in diets based on corn.
According to almost 20
peer-reviewed papers
(see Figures 2 and 3)
which report the effect of
xylanase on ileal protein
Figure 1: Illustration of the relationship between
digestibility, a consistent inherent ileal nutrient digestibility and the
15 percent improvement magnitude of response to feed enzymes
in the undigested fraction
is apparent.
This consistency may only be explained ent in the diets of poultry and this antinutri-
by a mass average mechanism such as tive effect is expressed through a physi-
improved diffusion of nutrients in the lumen ological cascade involving changes in pepsin,
as no preference is evident for nutrients of mucin and NaHCO3 production.
endogenous or exogenous origin. These As the effect of phytate and phytase is
principles also hold true for the ileal digest- quite specific (rather than mass average) the
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 Enzyme Feature

a new digestible lysine which is in excess ofTotal lysine = 1.23%

total lysine. Digestible lysine = 1.15%
Clearly this cannot possibly be the caseUndigested lysine = 0.08%
Phytase: 0.017%
and the error is in assumed additivity of matrix
New undigested lysine = 0.063%
values across all additives. In truth only the
New xylanase effect: (16%
first additive will carry its full matrix and sub-
sequent products will have their effect mutedof 0.063%) = 0.010
as a consequence of the improvements con- New undigested lysine = 0.053%
ferred by the current incumbent. New protease effect: (max 30% of
Thus, it is more appropriate to assign remaining undigested) = 0.016%
Using phytase, xylanase and protease
nutrient matrices as a proportion of the undi- Figure 3: The effect of exogenous xylanase
in combination = 0.043% improvement
gested fraction as this automatically accounts on ileal amino acid digestibility expressed as
a proportion of the undigested fraction. The
in digestible lysine (i.e. from 1.15% to
for any reduction in the undigested fraction
undigested fraction ranges from 12% for
1.19%).
associated with the use of other feed addi- methionine to 28% for cysteine but xylanase
tives. For example, in the above example: In this second example some delivers around 15-16% of this fraction regardless
acknowledgement of the
Table 1: Example* phytase1, xylanase2 and improved digestibility of
phytase+xylanase nutrient release values the diet is made when
demonstrating sub-additivity in response considering the likely
Econase Quantum effect of the next addi-
Quantum tive.
Nutrient (16,000 +
(500FTU)
U/Kg) Econase The author submits
that this process is more
logical than simply adding
AvP, % 0.130 0.000 0.130
supplier’s recommenda- Figure 4: The effect of exogenous phytase
Ca, % 0.130 0.000 0.130 on ileal amino acid digestibility expressed
tions together and takes
ME Kcal/Kg 45.000 100.000 116.000 into account that the as a proportion of the undigested fraction.
The undigested fraction ranges from 12%
Protein, % 0.365 0.374 0.592 opportunity for further for methionine to 28% for cysteine. Unlike
Cys, % 0.027 0.027 0.043 improvement in digest- xylanase, phytase delivers between 7 and
ibility declines as each 17% improvement in this fraction, depending
Met, % 0.006 0.011 0.014
new product is added. on the amino acid
Thr, % 0.029 0.025 0.043
The order in which prod-
Lys, % 0.015 0.019 0.027 ucts are added to the that realistic assumptions are made about the
Ile, % 0.022 0.019 0.033 diet will to an extent dictate extent of improved digestibility of this fraction.
Val, % 0.020 0.018 0.030 their value (the first product Achieving ileal starch, protein and fat digest-
Gly, % 0.023 0.027 0.041
being more valuable than ibility of 100 percent is extremely unlikely and
subsequent). the literature would suggest that only up to
Asp, % 0.024 0.024 0.038
However, this approach around 30 percent of the undigested frac-
Ser, % 0.026 0.018 0.036 reduces the likelihood of tion may be rendered digestible by enzyme
Ala, % 0.017 0.020 0.030 overvaluing combinations intervention. With these principals in mind it
Pro, % 0.016 0.023 0.032 of additives whose indi- is possible to assign enzyme matrices that are
His, % 0.021 0.024 0.036 vidual contributions have dynamic and consider both the sub-additive
been assessed independent incremental advantage of each new additive
Tyr, % 0.013 0.017 0.024
of one another but whose as well as the quantity of substrate remaining.
Trp, % 0.017 0.015 0.026 combined effect is less than Further research is warranted to explore
Phe, % 0.018 0.018 0.028 the sum of the various parts. the less tangible effects of the ingestion
Leu, % 0.018 0.015 0.026 of enzymes such as physiological changes,
Glu, % 0.015 0.012 0.022 Conclusion secretory and absorptive function differences
Substantial confusion per- and ultimately the effect of enzymes on
Arg, % 0.011 0.013 0.020
sists regarding the additivity nutrient requirement and the net value of
Na, % 0.030 0.000 0.030
of matrix values for feed addi- energy and other nutrients.
*
Note that phytase and xylanase matrices are not constant tives.
and will vary depending on the nature of the diet which is
As there is a law of dimin- More inforMation:
fed including e.g. dietary phytate-P concentration and the
quality of the corn. Thus these matrices are for illustrative ishing return for the incre-
Dr A J Cowieson and Dr M R Bedford
purposes only and should not be considered to be absolute mental addition of each new AB Vista Feed Ingredients
for all diets. additive it is prudent to cal- 3 Woodstock Court, Blenheim Road
1 Quantum phytase (500 FTU/kg) culate the undigested fraction Marlborough Business Park
of the diet at the level of the Marlborough, Wiltshire, SN8 4AN
2 Econase XT xylanase (16,000 BXU/kg)
terminal ileum and ensure United Kingdom

18 | September - october 2010 Grain &feed millinG technoloGy

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