ABSTRACT

The biggest single expense in any system of poultry production is feed

accounting for up to 70% of total production cost per bird. Poultry naturally
synthesizeenzymes to aid the digestion of feed nutrients. However, they do not
have enzyme to break down fibre completely and need exogenous enzymes in
feed to aid digestion. This review covers the information on enzymes and their
uses in poultry nutrition. Enzymes are biological catalyst composed of amino
acids with vitamins and minerals. They bring about biochemical reactions
without themselves undergoing any change. The benefits of using enzymes in
poultry diets include not only enhanced bird performance and feed conversion
but also less environmental problems due to reduced output of excreta. In
addition, this review demonstrates that enzyme isa very useful tool in the study
of physiological and metabolic mechanisms. Such reviewwill enhance our
understanding regarding the role of dietary enzymes in poultry nutrition. The
information presented here demonstrates the significant potential of enzymes in
poultry feed.
INTRODUCTION
The use ofenzymes in poultrynutrition is one of the major nutritional advances
in the last fifty years (Paloheimo et al., 2010). It is the culmination of something
that nutritionists realized for a long time but until 1980's, it remained beyond
their reach. Indeed, the theory of feed enzymes is simple. Plants contain some
compounds that either the animal cannot digest or which hinder its digestive
system, often because the animal cannot produce the necessary enzyme to
degrade them. Nutritionists can help the animal by identifying these indigestible
compounds and feeding a suitable enzyme. These enzymes come from
microorganisms that are carefully selected for the task and grown under
controlled conditions (Wallis, 1996).
Such change is driven by least costfeed formulation processes, and endeavours
to providemaximum nutrient density for minimum cost. Inevitably, this results
in diets reliant on the mostavailable cereal and vegetable protein, which
translates into maize, wheat or sometimes barleyas the dominant cereal, and
soyabean meal or rapeseed meal as the dominant protein source.
Each of these ingredients contains varying quantities of several anti-nutritional
factors (ANF),the consequence of which is minimal if fed at low concentrations
but considerable if theirthreshold is exceeded (Jansman etal.,1994).
Modem feed formulation practices tend to create such a problem, whereby the
dietary concentrationof a given anti-nutritional factor (ANF)often approaches
its threshold due to the economically attractivenature of the cereal or protein
meal for achieving the goal of nutrient provision. Because ofnatural variation in
the content of ANF (Matthaus, 1997) these thresholds are sometimesunwittingly
exceeded.
The activity of such ANF depends upon the anti-nutrient concerned and the
concentration inthe final ration (Tamminga etal.,1995).
Many ANF are anti-nutritional because they are not degraded sufficiently or
indeed at all by theconditions and digestive enzyme array in the gastrointestinal
tract(Reference). In such cases, any processwhich will reduce ANF
concentrations will enhance performance and relax some of the constraintson
feed formulation. It is under such circumstances that exogenous enzymes can be
ofbenefit. They may simply supplement those enzymes that are present in the
digestive tract to alevel that is effective, or provide hydrolytic capacity that is
totally absent. Addition of exogenous amylases and proteinases has been shown
to be ofbenefit in both the pig and chicken (Ritz etal.,1995; McNab etal.,1996)
and suggests that thisis an area requiring more attention.
The poultry industry readily accepts enzymes as a standard dietary component,
especially in wheat and barley-based rations. But still many questions are
partially answered. For example, how do enzymes work? Do growth rates
reflect differences in the potency of different enzyme preparations? What is the
link between gut viscosity, enzyme action and growth rates? And are enzymes
necessary in all poultry rations? This review article aims to supply some
background information about enzyme and its usage in poultry nutrition and
also help to answer some of commonly asked questions regarding enzymes.
 OVERVIEW OF ENZYME
Enzymes
Enzymes are one of the many types of protein in biological systems. Their
essential characteristic is to catalyze the rate of a reaction but is not themselves
altered by it. They are involved in all anabolic and catabolic pathways of
digestion and metabolism. Enzymes tend to be very specific catalysts that act on
oneor, at most, a limited group of compounds known as substrates. Enzymes are
not living organisms and are not concerned about viability or cross
infection(Reference). They are stable at 80-85 degree centigrade for short time.
Another important feature of enzymes is that the rate of an enzyme catalyzed
reaction increases with increasing substrate concentration, to the point where
there is no further response and the enzyme is said to be saturated. Therefore,
we need to match the amount of enzyme with the quantity of substrate
(Acamovic and McCleary, 1996).
It is common practice to name enzymes by adding the suffix ase to the name of
the principal substrate. For example, β-glucanase is an enzyme that splits β-
glucans, and proteases break protein. We may also broadly categorize the
digestive enzymes as endogenous or exogenous - referring to those produced by
the animal and those administered from outside, respectively. For example,
pancreatic lipase, which splits fat or lipid into glycerol and fatty acids, is an
endogenous enzyme. Those enzymes added to feed as a supplement are
exogenous (Classen, 1996; Classen and Bedford, 1991).

Sources of Enzymes
Enzymes were used in the preparation of foods long before there was any
awareness of enzymes as such, possibly as long ago as 10,000 years. The
industrial exploitation of microbial enzymes in the Western world started 100
years ago with the patenting of a process for the production of alpha-amylase
(Taka mine) from the fungus Aspergillus oryzae. Enzymes are produced in
every living organism from the highest developed animals and plants to the
simplest unicellular forms of life, as they are essential for metabolic
process(Reference). Most of the enzymes currently used in the food
andbeverage industry are from Aspergillus, but hemicellulases and cellulases
are derived from Trichoderma. Recently, genes encoding for different enzymes,
including phytases, ß-glucanases, and xylanases, have been cloned and
expressed in different commercial systems (microorganisms and plants). It is
possible to produce large amounts of cheap enzyme by continually selecting
favorable microbes, growing them in advanced fermentation systems and by
streamlining the extraction and purification of the enzyme (Wallis, 1996).
Microorganisms that generally involved in production of enzymes are; Bacteria
(Bacillus subtilis, Bacillus lentus, Bacillus amyloliquifaciens and Bacillus
stearothermophils), Fungus (Triochoderma longibrachiatum, Asperigillus
oryzae and Asperigillus niger) and Yeast (S. cerevisiae)

Exogenous Enzymes in Poultry Nutrition

The use of enzymes in animal feed is of great importance. Consistent increase in
the price of feed ingredients has been a major constraint in most of the
developing countries. As a consequence cheaper and non-conventional feed
ingredients have to be used which contain higher percentage of Non-Starch
Polysaccharides (soluble and insoluble/crude fibre) along with starch. Non
Starch Polysaccharides (NSPs) are polymeric carbohydrates which differ in
composition and structure from starch (Morgan et al., 1995) and possess
chemical cross linking among them therefore, are not well digested by poultry
(Adams and Pough, 1993; Annison, 1993). A part of these NSPs is water-
soluble which is notorious for forming a gel like viscous consistency in the
intestinal tract (Ward, 1995) thus by reducing gut performance. Predominantly
water solubleand viscous arabinoxylans, which belong to pentosan group, are
assumed to be the factor responsible. These pentosans also greatly increase the
water intake by the birds, which lead to unmanageable litter problems caused by
wet and sticky droppings. This deteriorates the hygienic conditions and carcass
quality (Dunn, 1996). On the other hand, ß-glucans adversely affect all
nutrients, especially protein and starch utilization and are known to give rise
highly viscous conditions in the small intestine of the chicks (Hasselman and
Aman, 1986).
Poultry do not produce enzymes for the hydrolysis of Non-Starch
Polysaccharide present in the cell wall of the grains and they remain un-
hydrolyzed. This results in low feed efficiency. Research work has suggested
that the negative effects of NSPs can be overcome by dietary modifications
including supplementation of diets with suitable exogenous enzyme
preparations (Creswell, 1994). Enzymes break down the NSPs, decreases
intestinal viscosity and eventually improve the digestibility of nutrients by
improving gut performance.

Types of Exogenous Enzyme Available for Poultry Nutrition

Some of the enzymes that have been used over the past several years or have
potential for use in the feed industry include cellulase (ß-glucanases), xylanases
and associated enzymes, phytases, proteases, lipases, and galactosidases (Table
1). Enzymes in the feed industry have mostly been used for poultry to neutralize
the effects of the viscous, non-starch polysaccharides in cereals such as barley,
wheat, rye, and triticale. These anti-nutritive carbohydrates are undesirable, as
they reduce digestion and absorption of all nutrients in the diet, especially fat
and protein. Recently, considerable interest has been shown in the use of
phytase as a feed additive, as it not only increases the availability of phosphate
in plants but also reduces environmental pollution. Several other enzyme
products are currently being evaluated in the feed industry, including protease to
enhance protein digestion, lipases to enhance lipid digestion, ß-galactosidases to
neutralize certain anti-nutritive factors in non-cereal feedstuffs, and amylase to
assist in the digestion of starch in early-weaned animals.

Table 1. ExogenousEnzymes Used in Poultry Feeds

Enzymes Substrate
ß-glucanases Barley Oats
Xylanases Wheat, Rye,
Triticale Rice
bran
ß-galactosidases Grain legumes
Lupins
Phytases Plant feedstuffs
Proteases Proteins
Lipases Lipids
Amylases Starch

Benefits ofExogenous Enzymes in Poultry Nutrition

Benefits of using feed enzymes to poultry diets include; reduction in digesta
viscosity, enhanced digestion and absorption of nutrients especially fat and
protein, improved Apparent Metabolizable Energy (AME) value of the diet,
increased feed intake, weight gain, and feed–gain ratio, reduced beak impaction
and vent plugging, decreased size of gastrointestinal tract,altered population of
microorganisms in gastrointestinal tract, reduced water intake, reduced water
content of excreta, reduced production of ammonia from excreta, reduced
output of excreta, including reduced N and P (Ouhida et al, 2000; Gill, 2001;
Odetallah, 2002; Gracia, et al., 2003; Saleh, et al., 2003; Odetallah, et. al., 2005
and Wang et al., 2005).

Reduction in Digesta Viscosity

Enzymes added to poultry diets; especially diets containing cereals rich in NSP
such as wheat, barley, and rye, reduced viscosity in the diet and digesta. Morgan
et al., (1995) and Muramatsu et al., (1992) found that that enzyme
supplementation of wheat based diets significantly reduced foregut digesta
viscosity of birds. The reduction in foregut digesta viscosity was achieved
primarily by reducing the molecular weight through hydrolysis of xylan
backbone by endo-xylanase into smaller compounds and thus reduction in
viscous effects of the feed because foregut digesta viscosity is directly
proportional to the molecular weight of wheat arabinoxylans (Bedford and
Classen, 1993). As a result of endo-xylanase and ß-glucanase supplementation,
the long backbones of the arabinoxylans and ß-glucans are cleaved into shorter
fragments, thereby reducing their viscosity (Gruppen et al. 1993).
Similar findings on digesta viscosity were also reported by Bedford and Classen
(1993); Bhatt et al., (1991) and Dunn, (1996) who inferred that the high
viscosity in the gut contents caused by the pentosans led to increased water
intake of the birds, which resulted in the wet and sticky droppings.
Increase in Available Energy
One of the main reasons for supplementing wheat- and barley-based poultry
diets with enzymes is to increase the available energy content of the diet.
Increased availability of carbohydrates for energy utilization is associated with
increased energy digestibility (Partridge and Wyatt 1995; Van der Klis et al.
1995). The AME of wheat has been extensively studied and found to have a
considerable range i.e 9 500–16 640 kJ/kg (Ward 1995). Enzyme
supplementation improves this range by enhancing carbohydrate digestibility,
reducing gut viscosity, and improving fat utilization (Almirall et al. 1995). The
improvements in AME resulting from enzyme supplementation are variable
because of the variability in the NSP content of wheat. Classen et al. (1995),
Schutte et al. (1995), and Van der Klis et al. (1995) reported improvements of
5–16, 3.1–4.5, and 4.5–12.4%, respectively. The increase in AME with the use
of enzymes is difficult to predict, as nutrient ratios, such as energy–protein, and
other factors also play an important part in poultry-feed formulations. The AME
value of wheat has been correlated with its content of water-soluble NSPs
(Annison 1991), which in turn affects gut viscosity (Bedford et al. 1991).
Unfortunately, NSP analyses are relatively lengthy processes, and in a
commercial situation rapid testing of incoming grains is required. No chemical
test or detectable physical characteristic can be used to rapidly predict the AME
value of wheat or to estimate the improvements to be expected from the use of
enzymes. This is part of the difficulty in trying to accurately estimate the energy
content of wheat or barley in poultry feeds and compensate for the deficiency
by adding enzymes.
Adding adequate activity levels of α-amylase, β-glucanase, and xylanase to
broiler starter and grower corn-soybean diets with a 3% reduction in dietary ME
allowed full restoration of growth performance of broilers comparable to those
fed the adequate energy (Yu and Chung, 2004).
Improvement in Nutrient Digestibilities
Enzymes have been shown to improve performance and nutrient digestibility
when added to poultry diets containing cereals, such as barley (Marquardt et al.
1994), maize (Saleh et al., 2003), oats (Friesen et al. 1992), rye (Friesen et al.
1992; Bedford and Classen 1992; Marquardt et al. 1994), and wheat (Marquardt
et al. 1994), and to those containing pulses, such as lupins (Brenes et al. 1993).
The effect of enzyme supplementation on Dry Matter Digestibilities (DMD) in
pigs and poultry depends on the type of diet and the type of animal: increases in
DMD range from 0.9 (Schutte et al. 1995) to 17% (Annison and Choct 1993) in
poultry.
The enzymes currently used in monogastric diets are predominantly glycanases,
which cleave NSPs into smaller polymers, thereby removing their ability to
form viscous digesta and enhancing nutrient digestibilities. The effects of
glycanases are generally nonspecific, except for their effect on fat (greater effect
on saturated fat than on unsaturated fat). Another enzyme used in feed is
phytase, which increases the utilization of phytate phosphorus. The ability of
phytase to improve the digestion of phytate phosphorus and subsequently to
reduce the output of organic phosphorus to the environment has attracted a great
deal of scientific and commercial interest. In poultry use of phytase was
reported to reduce phosphorus excretion by as much as 40% for broilers. When
phytase was added to layer diets, increased egg production and positive effects
on egg weight and tibia ash were also noted (Simons and Versteegh, 1991).

Reduction in Excreta Moisture:

A reduction in the moisture content of poultry excreta is often noted when
glycanases are included in the diet. Supplementing the NSP-enriched diet with
three different commercial glycanase products improved performance, but their
effectiveness in reducing the moisture levels of the excreta differed from 10 to
29%. This supports the view that different glycanases have similar
performance-enhancement effects in monogastric animals but the site of the
breakdown of the NSPs in the gut and the molecular sizes of the released
products differ (Choct and Annison, 1990). Graham, (1996) also reported an
increased water uptake and excretion in broilers given diets containing higher
levels of viscous cereal grains.

Health improvement
Morgan and Bedford (1995) reported that coccidiosis problems could be
prevented by using enzymes. Birds fed a wheat-based diet with and without
glycanase supplementation showed vastly different responses to coccidiosis
challenge. Growth was depressed by 52.5% in the control group but by only
30.5% in the enzyme group, which also had a much better lesion score. An
increase in digesta passage rate and a reduction in excreta moisture are often
noted when glycanases are added to poultry diets, which may be detrimental to
the life cycle of the organism.

Precision and Flexibility in Least Cost Feed Formulation:

Enzymes provide greater flexibility in feed formulation and allow the use of a
wide range of ingredients without compromising bird performance and hence
provide great flexibility in least-cost feed formulation. The nutritive value of
cereal grains for poultry varies greatly, and no suitable assays are currently
available for rapid in-mill testing. For instance, the variability in the AME of
wheat for poultry can be as great as 4 MJ/kg DM (Rogel et al. 1987). This
problem can be largely overcome by using glycanases to bring the AME of
different wheats to comparable levels (Choct et al. 1995).
Impact on Environment
Enzymes have been approved for use in poultry feed because they are natural
products of fermentation and therefore pose no threat to the animal or the
consumer.
Enzymes not only will enable livestock and poultry producers to economically
use new feedstuffs, but will also prove to be environmentally friendly, as they
reduce the pollution associated with animal production. As well as contributing
to improved poultry production, feed enzymes can have a positive impact on the
environment. In areas with intensive poultry production, the phosphorus output
is often very high, resulting in environmental problems such as eutrophication.
This happens because most of the phosphorus contained in typical feedstuffs
exists as the plant storage form phytate, which is indigestible for poultry. The
phytase enzyme frees the phosphorus in feedstuffs and also achieves the release
of other minerals (e.g. Ca, Mg), as well as proteins and amino acids bound to
phytate. Thus, by releasing bound phosphorus in feed ingredients, phytase
reduces the quantity of inorganic phosphorus needed in diets, makes more
phosphorus available for the bird, and decreases the amount excreted into the
environment.

Factors Affecting the Benefits of enzyme

The degree of improvement obtained by adding enzymes to the diet depends on
many factors (Bedford, 1996), including the type and amount of cereal in the
diet; the level of antinutritive factor in the cereal, which can vary within a given
cereal (for example, low- versus high-ß-glucan barley); the spectrum and
concentration of enzymes used; the type of animal (poultry tend to be more
responsive to enzyme treatment than pigs); and the age of the animal (young
animals tend to respond better to enzymes than older animals); type of gut
micro flora present and the physiology of the bird. Older birds, because of the
enhanced fermentation capacity of the micro flora in their intestines, have a
greater capacity to deal with negative viscosity effects (Allen et al. 1995; Choct
et al. 1995; Vukic Vranjes and Wenk 1993).
Use of enzymes in layers:
Although the majority of research trials were conducted on broilers. However,
the responses of laying hens to enzyme-supplemented feeds are also well
documented. Typically, enzymes added to layer feed appear to have little effect
on egg mass but improve feed efficiency (Benabdeljelil and Arbaoui 1994;
Vukic Vranjes and Wenk 1993), energy utilization (Wyatt and Goodman 1993).
Wyatt and Goodman (1993) reported that corn-fed layers exhibited better feed
efficiency than those fed enzyme-supplemented barley-based diets.
Nevertheless, enzyme supplementation improved the utilization of barley diets.
Increased energy utilization in laying hens appears to be due to microbial
fermentation of solubilized NSPs and the subsequently higher absorption of
volatile fatty acids (Choct et al., 1995). Wet litter arising from the use of barley
and newly harvested wheat can result in an increased incidence of dirty egg
shells and in ammonia build-up in poultry barns. Adding enzymes to both
wheat- and barley-based diets has been shown to reduce the moisture content of
fecal matter in layers (Marquardt et al., 1994).

Need for addition of Exogenous enzyme on poultry nutrition

Least cost formulation of livestock rations is crucial in minimizing production
costsand maximizing profit margins. Feed accounts for approximately 70% of
the total cost of pigand poultry production (Barletta, 2010). With the rising costs
of dietary energy sources,mainly soybeans, corn, and oil, alternative means of
deriving energy from feed ingredientsare being investigated. Enzymes have
potential to increase overall feeding value, potentiallyimproving or maintaining
poultry performance while allowing the use of lower costingredients. Pectinase,
cellulase, β-mannanase, xylanase, α-amylase, and β-glucanase areamong
enzymes that are often utilized in poultry diets containing soybean meal to
potentiallyalleviate the anti-nutritional NSP and to increase nutrient utilization
in corn-soybean mealbased diets (Bach Knudsen 1997; Kocher et al., 2003).
Researchers examined broiler performance from 1 to 21 d with soybean meal or
asoybean meal sunflower combination as the primary protein source.
Replacement ofapproximately 25% soybean meal with sunflower meal
significantly improved growth rate,feed intake, and feed conversion. When an
enzyme cocktail containing pectinase,galactanase, arabinanase, β-glucanase,
and polygalacturonase (380, 420, 140, 480, 530units/g) were added to these
same soybean meal or sunflower meal combination diets noimprovements in
performance were observed. Enzyme addition did increase theconcentrations of
raffinose and sucrose in the proximal small intestine. Overall, when birdswere
fed soybean meal as the primary protein source, an increase in the concentration
ofoligosaccharides present in the digesta from the distal small intestine
demonstratedsignificant negative impacts on broiler performance (Irish and
Balnave, 1993).
In vitro research utilizing a multi-enzyme preparation, isolated from Aspergillus
aculeatus, determined type 1,4-β-arabinogalactanase led to effective
solubilization of drymatter in soybean meal. Another enzyme, 1,4-β-
galactomannanase, demonstrated release ofsugar from untreated (lacking
previous pepsin degradation) soybean meal. Synergisticimpacts relating to use
of galactanase and mannanase on endo-polygalacturonases were alsonoted
(Vahjen et al., 2005). In vivo work was also conducted with galactanase
originatingform H. insolens solely and in combination with a mannanase
enzyme. From 7 to 35 dbroilers were fed corn and soybean meal (48% CP)
rations supplemented with a combinationof both enzymes. Compared to
galactanase alone, additional supplementation of mannanasedecreased live
weight gain and feed intake. Researchers concluded that 1,4-β-
arabinogalactanase enzymes are promising for use in soybean meal based diets,
andcombination with endo-polygalacturonases should also be considered. In
this experimentpartial hydrolysis of NSP did result in enhanced digestibility;
however, increased hydrolysis,with increased NSP degrading enzyme
(mannanase) addition, may produce oligo- andmonomers with anti-nutritive
impacts (Vahjen et al., 2005).
Exogenous enzyme products increase availability of nutrients already present
infeedstuffs (Zanella et al., 1999; Kocher et al., 2003; Rutherfurd, 2007).
Poultry have limitedendogenous secretion of NSP-degrading enzymes, reducing
utilization of these nutrients inthe small intestine (Angkanaporn et al., 1994).
Exogenous enzyme addition has been showneffective in improving ileal
digestible energy of soybean meal (Douglas et al.,2000).
Apparent ileal digestibility of CP was increased by 2.9% when male Hubbard
broilers werefed an exogenous enzyme cocktail (0.1% inclusion of 800 μ/g
xylanase, 6,000 μ/g protease,and 2,000 μ/g amylase) from 1 to 45 d (Zanella et
al., 1999). Apparent ileal digestibility ofnitrogen was also observed to increase
by 6.6% when 0.5% exogenous enzyme (α-amylase,β-glucanase, and xylanase)
was fed from 21 to 28 d to male Ross 308 broiler chickens(Rutherfurd et al.,
2007). Nutrients are often bound by cell walls rich in NSP limitingendogenous
digestive enzyme access to these nutrients, thereby decreasing digestibility.
Theadvantageous effects of exogenous enzymes on complementing and
increasing endogenousenzyme activity are beneficial for improving overall
feeding value of soy products (Bedford,1996).
Figure 2.5. Substrates and anti-nutrients targeted by some commonly utilized
exogenousenzymes. Source: Bedford and Partridge (2010).
SOME EXOGENOUS ENZYMES EXPERIMENTED IN POULTRY
NUTRITION
Exogenous xylanase amylase protease enzyme
Exogenous supplementation of xylanase, amylase, and protease enzymes are
used to target NSP, starch, and protein consequently improving digestion in
monogastricanimals fed corn-soybean meal diets (Burnett, 1996). Xylanase is a
general descriptor forseveral classes of enzymes that have varying modes of
action but all result in the hydrolysisof the xylan component of NSP present in
dietary ingredients (Paloheimo et al., 2010).
Hydrolysis of soluble NSP, mainly in wheat based diets, results in decreased
digestiveviscosity leading to increased digesta passage rate, which decreases
bacterial proliferation inthe small intestine, endogenous secretions, and enzyme
production. Collectively, this resultsin increased nutrient sparing and
digestibility (Brenes et al., 1993; Jozefiak et al., 2007;Pirgozliev et al., 2010).
Hydrolysis of both soluble and insoluble NSP decreases waterholding capacity
of the gastrointestinal tract resulting in increased digesta flow and
nutrientdiffusion, and hydrolysis of insoluble NSP can also increase cell wall
permeability leading toincreased nutrient diffusion and digestibility (Bedford
1996; Zanella et al., 2004).
Breakdown of plant cell wall components allows increased access by pancreatic
amylases and proteases which are capable of releasing starch and protein.
Release of starchand protein can result in increased animal performance through
increased growth and feedutilization along with decreased gastrointestinal tract
maintenance (Choct and Cadogan,2001). Exogenous protease and amylase
enzyme products are often supplemented as acocktail with xylanase and have
been shown to increase utilization of protein, in the uppergastrointestinal tract
of broiler chickens, as well as digestion of starch encapsulated inendosperm
(Zanella et al., 1999).
The addition of XAP enzymes were evaluated in corn-soybean meal diets
usingdigestibility assays with 15 month old White Leghorn roosters for a 37 d
period (Zanella etal., 1999). In this experiment birds were fed corn and soy
(soybean meal, extruded full fatsoybeans, or roasted full-fat soybeans) and
supplemented with exogenous xylanase amylaseprotease (XAP) enzyme
product (containing 800 μ/g xylanase, 2,000 μ/g amylase, and 6,000μ/g
protease) included at 0.1% of the diet. Ileal contents were sampled and the
addition ofXAP improved overall CP digestibility by 2.9% in comparison to the
control diet, along withspecific improvements in valine and threonine
digestibility of 2.3 and 3.0%, respectively.
The enzyme complex also improved ileal starch digestibility by 1.8% and ileal
fatdigestibility by 1.6%. These same corn and soy diets supplemented with XAP
were fed in aperformance based trials to day old Hubbard male broilers for a 45
d period. Enzymesupplementation reduced feed conversion ratio by 2.2% and
improved body weight by 1.9%.
Silversides and Bedford (1999) hypothesized that supplementation of
exogenous multienzymecocktails in corn-soybean meal diets may increase
starch utilization in the proximalsmall intestine, by increasing the time
endogenous secretions and the microbial flora have toact on nutrients.
A commercial XAP enzyme (300 U xylanase/g diet, 400 U amylase/g diet, and
4,000U protease/g diet) was utilized in a series of experiments at 0.10%
inclusion. Diets containinghigh NSP wheat cereal grain were fed to turkey hens
from 3 to 9 wk of age. Addition of XAPresulted in significant improvement in
feed conversion ratio and AMEn, over diets withoutenzyme supplementation. In
addition to the increased AMEn, starch disappearance from theproximal small
intestine was also increased in these hens (Persia et al., 2002).
Anotherexperiment utilizing XAP enzyme (800 U/g xylanase, 2,000 U/g
amylase, and 6,000 U/gprotease) showed significant improvement in ileal
digestible energy of the corn-soybeanmeal diet when fed to commercial male
broilers from 8 to 21 d. No significant effect ofenzyme supplementation on bird
weight gain was observed and impact of XAP enzymesupplementation varied
depending on soybean meal source in this experiment (Douglas et al.,2000).
Although XAP is a common combination of enzymes for use in commercial
poultryproduction, other combinations have been evaluated. Marsman and
others (1997) evaluatedthe effects of protease and carbohydrase combination,
on growth performance and ilealnutrient digestibility of broiler chickens. Both
enzymes were supplemented alone and incombination to diets containing
soybean meal as the primary protein source. Day old femaleRoss broilers were
fed a control diet with casein as the main protein source from 1 to 7 d.From 7 to
25 d birds were transitioned to experimental diets. Performance parameters,
bodyweight gain, feed intake, and feed conversion were not affected by enzyme
treatment.
Carbohydrase enzyme treatment significantly increased ileal NSP degradability
and overallenzyme treatment improved apparent ileal crude protein digestibility
over the treatmentgroups with no enzyme. Increases observed in CP and NSP
degradation may result inimproved body weight gain and overall performance
parameters if enzyme wassupplemented from day of hatch.
The influence of supplementation of exogenous xylanase (200 mg/kg diet),
derivedfrom Thermomyces lanuginosus, and protease (200 mg/kg diet) from the
fermentation ofBacillus licheniformis and transcribed genes from Nocardiopsis
prasina on performance,nutrient utilization, and intestinal morphology in broiler
chickens fed a wheat-soybean mealbased diet was examined by Kalmendal and
Tauson (2012). Day old straight run Ross 308broiler chicks were fed for a 34 d
period. Results show enzyme supplementation had nosignificant effect on body
weight gain and feed intake of birds at any time point. Xylanaseand protease
supplementation, alone or the combination, significantly improved
feedconversion ratio (1.43 g/g) and feed AMEn (3,389 kcal/kg) compared to the
control FCR andAMEn of 1.47 g/g and 3,271 kcal/kg respectively. Apparent
digestibility of starch and fatwas significantly improved with all enzyme
treatments; however, apparent retention of CPwas not affected.

Another experiment utilized the same specific xylanase (derived from

Thermomyceslanuginosus) provided at 100 g/kg diet, and showed significantly
increased body weight gainin Ross 208 broilers fed wheat based diets from 15
to 42 d (Engberg et al., 2004). Increasedwheat ingredients in the second
experiment combined with xylanase may have resulted inincreased hydrolysis
of soluble NSP and reduced digesta viscosity (Kocher et al., 2002)leading to
increased nutrient diffusion and energy availability for growth (Bedford, 1996).
Breakdown of plant cell wall carbohydrates by NSP enzymes eventually leads
toproduction of short-chain oligosaccharides (di- and tri-saccharides).
Oligosaccharidesprovide substrate for bacterial fermentation, and can also be
beneficial in volatile fatty acidproduction (Choct and Cadogan, 2001;
Paloheimo et al., 2010). When NSP degradingenzymes have been used in
higher concentrations, work suggests that oligosaccharides can befurther
degraded to monosaccharides (Fuller, 2001; Choct.et al., 2010).
Excessmonosaccharides in the digestive tract have the potential to cause
osmotic diarrhea as well aspoor performance (Schutte, 1990). The effects of
feeding graded amounts (25, 50, and 75g/kg) of D-xylose or L-arabinose fed a
diet based on corn, corn starch, and soybean isolate onperformance was
examined. All diets were formulated to be isocaloric and were provided
tobroiler chickens from 6 to 27 d of age. A significant negative effect of both
pentose sugars(xylose and arabinose) was reported for body weight gain. Birds
fed D-xylose diets hadsignificantly decreased feed intake with increasing
pentose sugar concentration. Water intakeincreased linearly as the dietary
concentrations of both pentose sugars was increased, andconsequently dry
matter content of the droppings decreased.
Supplementation of exogenous XAP targets a number of substrates and
complementsendogenous enzymes potentially leading to increased nutrient
digestibility and metabolicefficiency. Variation in bird performance and nutrient
digestibility response has beenobserved when feeding soy ingredients and
supplementing with multi-enzyme cocktails.
Hydrolysis of NSP may lead to production of oligosaccharides that have
potential topositively modulate the gastrointestinal tract and immune function.
However an overabundanceof short-chain oligosaccharides or pentose sugars in
the gastrointestinal tract maylead to digestive inefficiency. All potential impacts
on digestibility and efficiency, bothpositive and negative, should be considered
when supplementing multi-enzyme combinationsto broiler chickens.

Exogenous β-mannanase
Identifying and alleviating anti-nutritional factors that reduce or inhibit
nutrientutilization is a necessity for successful poultry production. Among these
anti nutrients areNSP or fiber which includes β-mannans (Daskiran et al., 2004).
Beta-mannanase hydrolyzesβ-mannan and is the main activity of the
fermentation product of Bacillus lentus. Two of themain β-mannans in soybean
meal are β-galactomannan and β-glucomannan (Zou et al.,2006). It is thought
that β-mannanase has three primary modes of action against β-mannans,these
mechanisms include decreased viscosity, improved energy metabolism, and
reducedinnate immune stimulation (Jackson et al., 2004).
Evaluation of β-mannanase on corn-soybean meal based diets with variable β
mannancontent has been conducted using guar gum to alter β-mannan levels as
its chemical structureis similar to that of β-mannan in soy (Daskiran et al.,
2004). Experiment one focused onevaluating the effects of varying β-mannan
concentration (0, 0.5, 1, and 2%) on performance,MEn, and net energy for gain
of day old Cobb x Cobb male broilers supplemented with a β-mannanase.
Experimental diets were fed for 2 wk and β-mannanase enzyme activities
weredetermined to be 152.7, 127.7, 145.9, and 141.5 million units per ton for
the four treatmentgroups. Data show enzyme supplementation improved feed
efficiency, MEn, and net energygain at all β-mannan inclusions, relative to the
control group. As expected, increasing guargum (β-mannan) inclusion
decreased bird weight gain especially at the 2% inclusion asweight gain was
only 153.8 g in comparison to the 178.6 g of the control fed birds.
Similarbroiler performance results have been reported with 2% guar gum levels
in previous research(Vohra and Kratzer, 1964; Ray et al., 1982).
A second set of experiments, utilizing the same broiler stock type (Cobb x
Cobb)evaluated the effect of incrementally increasing β-mannanase
supplementation (0, 0.05, 0.1,0.15%) with activities of 1.4, 145.9, 225.8, 349.8
million units/ ton respectively. Enzymeswere added to a corn-soybean meal
based diet with 1% guar gum inclusion from 1 to 14 d.Increasing enzyme
supplementation did not influence final body weight, but did
significantlyimprove feed:gain ratio at all inclusions (1.533 g/g compared to
1.495, 1.493, and 1.454 g/g).
As in the first set of experiments MEn improved with increasing β-
mannanasesupplementation (control at 2,971 kcal/kg compared to 0.15%
inclusion at 3,061 kcal/kg)(Daskiran et al., 2004). As seen with previous
exogenous enzyme supplementation research,increasing duration of enzyme
inclusion could produce differences in body weight gainespecially due to
significant differences in MEn values compared to control andsupplemented
diets.
Degradation of β-mannans can produce mannan oligosaccharide (MOS) which
hasbeen shown to positively influence the immune system in poultry. Zou and
others (2006)hypothesized that the inclusion of β-mannanase in corn-soybean
meal based broiler dietswould improve growth performance and immunity.
Results showed inclusion of exogenousβ-mannanase (0, 0.025, 0.05, and
0.075%) increased weight gain and feed conversion ratio inbirds during the 4 to
6 wk and 0 to 6 wk periods. All inclusions of β-mannanase resulted
inimprovement in weight gain compared to the non-supplemented control. Feed
conversionratio was significantly improved (1.637 g/g) at 0.05% inclusion
compared to all othertreatments from 1.684 to 1.720 g/g. Performance
improvements may have been observed dueto the extended experimental trial
duration compared to previously described research. SerumIgA, IgG, and IgM
concentrations were also measured to determine the impact ofsupplementation
on immune factors. A significant increase of 0.286% IgM was noted for0.025%
inclusion during the 3 to 6 wk period; however, no difference in IgG and
IgAconcentrations occurred during the trial period. T lymphocyte production
was significantlyimproved in 6 wk old broilers at 0.05% β-mannanase inclusion
(Zou et al. 2006). Resultsfrom this experiment demonstrate the potential
beneficial impacts, on performanceparameters and immune function, of
including β-mannanase in broiler diets containing cornand soybean meal.
Another manuscript reported supplementing exogenous β-mannanase and
pectinaseenzymes targeted the breakdown of soy cell wall components of the
diet, to a greater extentcompared to corn (Jackson et al., 2004). Four
experimental treatments were utilizedsupplying 0, 50, 80, and 110 MU (1
MU=106 enzyme activity units) of β-mannanase tobroilers fed corn and
soybean meal based rations. Feed intake was significantly increased indiets
containing 80 MU/ton compared to 50 MU/ton β-mannanase. Despite the
reduced feedintake, both 80 MU/ton and 110 MU/ton inclusions resulted in
significantly increased weightgain and feed efficiency in comparison to 50
MU/ton. Regression analysis indicated, forevery 100 MU/ton β-mannanase
supplemented, weight gain was improved by 120 g and feedconversion ratio
improved by 0.067 g of feed/g of gain. Clearly β-mannanasesupplementation
has the ability to impact broiler performance; however, concentration of β-
mannanase supplemented is important in understanding the bird response.
Hydrolysis of mannans and galactans by commercial galactanase and
mannanaseexogenous enzymes in diets containing corn and soybean meal have
resulted in improvedbroiler performance when fed from 1 to 28 d (Centeno et
al., 2006). Day old Ross 308broilers were supplemented with three separate
exogenous enzyme preparations designed totarget arabinoxlyans, β-1,3-1,4-
glucans, and raffinose an oligosaccharides. Supplementeddiets resulted in
shortened ceca. Dietary fiber can cause an increase in muscle developmentof the
intestine in response to indigestible nutrients and microbial activity.
Thereforeshortening of the ceca may indicate improved intestinal hydrolysis of
mannans and galactansdecreasing the amount of fermentation occurring in the
caecum.
Utilization of β-mannanase supplements in broiler rations has the potential
topositively influence bird performance, nutrient utilization, and immune
function. Theseeffects could lead to an increase in broiler efficiency, decreased
endogenous secretions, andreduced cecal fermentation. When β-mannanase is
supplemented into broiler rationscontaining mainly corn and soybean meal at
hatch it has the ability to increase productivityand therefore profitability of the
bird for the remainder of its production cycle. Howeverresearch regarding the
effects of increased soy ingredient inclusion along with combinationsof β-
mannanase and other exogenous enzymes is novel.

Conclusion
The use of enzymes as a feed additive has rapidly expanded. In the last decade,
extensive studies have been conducted to study the effects of feeding exogenous
enzymes on the performance of poultry. By compiling these studies into a single
focused work, this review provides evidence that enzyme is a significant
instrument for the use in poultry feed.
Although the economic and social benefits of enzymes have been well
established and the future of feed enzymes is a bright one. However, further
research is required if enzymes are to reach their full potential inthe industry
and to answer some of the questions that this article raises, particularly those
regarding the mode of action of enzymes, how best to match the levels of
enzyme and substrate and how enzymes counter the variable environments in
the animal's gut. Ultimately, in vitro tests will tell us not only whether we need
exogenous enzymes, but also the correct enzyme cocktail for a specific batch of
a feed ingredient. This will allow the formulation of better rations using a wider
range of ingredients. Any advances in this field must ultimately improve the
welfare of chickens, reduce the production of wastes and conserve resources.

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