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Introduction
The 1990s saw a steady rise in the use of certain biotechnological products as feed
additives, and in research on new possibilities. Bonneau and Laarveld (1999)
reviewed this whole area and its potential impact on farm animal nutrition, physiol-
ogy and health and readers are referred to that paper for a comprehensive review of
the subject.
Amongst the biotechnological additives, feed enzymes are seen perhaps as hav-
ing made most progress and impact in the past decade, following the extensive and
increasing use of crystalline amino acids, many of which are also produced from
industrial fermentation. Bonneau and Laarveld (1999) identified the use of enzymes
in a number of applications for improving feed characteristics, such as: removal of
specific anti-nutritional factors; improving digestibility of nutrients; aiding the
digestibility of non-starch polysaccharides; supplementing endogenous enzymes for
improved digestion; reducing effluent losses.
The weaner pig has always provided a unique set of nutritional challenges that
need to be overcome in order to maximize the young animal’s growth potential.
Some of the principal factors that limit post-weaning growth rate have been out-
lined by others (Partridge and Gill, 1993; Thacker, 1999a) and include: inadequate
levels of endogenous enzyme secretion; reduced absorptive capacity in the gut; inad-
equate acid secretion; and inadequate food and water intake. Many of these factors
are clearly interrelated and all, potentially, can be influenced to varying degrees by
the application of feed enzyme technology.
Young pig nutrition is also being increasingly influenced by current trends in
agricultural production, specifically to find potential alternatives to therapeutic and
sub-therapeutic antibiotics at all stages of the production cycle. There is increasing
research interest towards a better understanding of the role that certain feed addi-
tives, or combinations of them, can play in manipulating the gut microflora in a
positive way against a background of reduced use of antibiotics.
124 G. Partridge
This chapter reviews the role for feed enzymes in the nutrition of the weaner
pig. It focuses on the main enzyme activities for which there is a growing body of
published data: carbohydrases (with emphasis on xylanases and -glucanases), phy-
tases and proteases. It also examines current views on the modes of action of these
enzymes and how they can offer particular benefits in the young, newly weaned ani-
mal. The interaction between feed enzymes and the gut microflora is receiving
increasing attention in current research, using novel microbiological techniques,
and this topic is considered in the light of the above constraints on antibiotic usage.
Carbohydrases
Carbohydrase enzymes for application in diets for the pig after weaning are targeted
primarily against the non-starch polysaccharide (NSP) and starch components.
Maize, wheat and barley, the main grain components of post-weaning pig feeds
globally, all contain significant cell wall material, but they differ in both their level
and composition (Table 7.1).
NSP levels in maize and sorghum (where used) are similar (9–10%; Dierick and
Decuypere, 1994), with insoluble arabinoxylans comprising over 40% of this total.
Standard milling procedures (e.g. roller or hammer milling through screens of vary-
ing diameter) are known to result in intact packages of cell wall material, with
enclosed nutrients, entering the stomach and small intestine. As the cell wall struc-
ture of these particular grains is predominantly insoluble fibre and unresponsive to
endogenous enzymes, varying quantities of ‘packaged’ material have been observed
at the end of the small intestine, entering the hindgut (M.R. Bedford, Finnfeeds,
2000, personal communication). The observation that the post-weaned pig, in par-
ticular, appears to respond positively in growth rate and feed utilization to progres-
sively fine grinding of maize and sorghum (Healy et al., 1994) is, most likely, a
reflection of reductions in nutrient ‘packaging’ by this cell wall material. Although
Table 7.1. The starch and fibre composition of the key grains used in weaner feeds
globally (g kg1) (from various sources).
The effect of exogenous enzymes on diets for weaned pigs based on barley, wheat or
mixtures of these grains has received far greater attention than maize or sorghum as
Weaner Pig Chap 07 26/9/01 2:02 PM Page 126
126 G. Partridge
58.0
39.1
8.4
3.9
3.5 5.5
Time after feeding (h)
Maize starch
Fig. 7.1. Concentration of starch (mg g1 dry matter) in caecal digesta of pigs fed
diets containing maize starch or high amylose starch (mean of two pigs per
treatment) (Topping et al., 1997).
substrates in the scientific literature. This is unsurprising, given the history of feed
enzyme application, which has been dominated by the use of, predominantly,
xylanase and -glucanase activities into diets for poultry and pigs containing these
‘viscous’ grains in Europe, Canada and Australia over the period 1990–2000. Hulled
barley contains around 180 g NSPs kg1, with -glucans and arabinoxylans constitut-
ing the majority of this cell wall material (Table 7.1). The concentration of soluble -
glucans is highest in the endosperm of the grain. Insoluble fibre predominates in the
outer (aleurone) layer, with the surface of the aleurone cell walls containing high levels
of xylan, surrounding a core of mixed-linked -glucans (Autio et al., 1996). For this
reason optimal responses in weaner pig diets based on, or containing, barley are only
likely to be seen when both xylanase and -glucanase activities are supplied in suffi-
cient quantities (van Lunen and Schulze, 1996; Ramaswamy et al., 1996).
The last few years has seen increasing interest from certain markets, particularly
Canada, in the opportunities for the use of hull-less barley as a feed ingredient in
diets for weaned pigs. Hull-less barley differs from conventional barley grain in that
the hull is less firmly attached to the kernel and therefore becomes detached during
threshing. Despite, consequently, having lower insoluble fibre levels and higher levels
of protein, its feeding value for piglets and poultry has often failed to live up to its
theoretical potential (Baidoo et al., 1998b; Thacker, 1999b). The reason appears to
be that removal of the hull effectively concentrates the soluble fibre fraction in the
grain endosperm, typically giving -glucan levels of 4.5–7% vs. 3.5–4.5% in hull-
less and hulled barley, respectively. These soluble, high molecular weight fibres are a
problem for the pig in the small intestine: they can interfere with effective nutrient
digestion and absorption, slow digesta transit rate and encourage microbial prolifera-
tion in the foregut. Feed processing (e.g. conditioning/pelleting/expansion) can exac-
Weaner Pig Chap 07 5/10/01 8:57 AM Page 127
erbate this situation by further solubilizing more fibre (Fig. 7.2). Feed processing can
therefore be seen as a double-edged sword in these situations – positive in terms of its
effects on feed hygiene and starch gelatinization (depending on moisture content
during processing and the temperatures achieved) and negative in terms of some of
these effects on dietary fibre, and their potential consequences.
Not surprisingly, exogenous enzymes have been found to be particularly benefi-
cial in barley-based diets for young pigs (Table 7.2). Several studies used -glu-
canase activities alone; although positive effects were almost invariably seen, it is
questionable whether they were maximized based on the structure of the barley
aleurone layer, as described above.
The predominant NSPs present in the endosperm and aleurone layers of wheat are
arabinoxylans, comprising 50–60% of the total NSP content (Table 7.1). In contrast to
barley and oats, -glucan is a minor component. The arabinoxylans found in the thin
cell walls within the endosperm are water-soluble to varying degrees, depending on vari-
ety, growing conditions, etc. In contrast, those in the thick cell walls of the aleurone
layer are predominantly insoluble where they encircle, and complex with, potentially
useful nutrients. Xylanases are therefore the main type of exogenous enzyme used for
wheat-based diets and a number of trials in recent years have looked at their effects in
diets for young pigs (Table 7.3). Overall, the consistency of response in published stud-
ies appears to be more variable than that of the barley-based trials (Table 7.2) but,
equally, few studies have attempted to superimpose some predetermined aspect of wheat
quality before appraising enzyme response. In the few recent studies where this has been
attempted (Choct et al., 1999; Partridge et al., 1999) the ability of xylanase to raise sig-
nificantly the productive performance of a ‘poor’ wheat to that of a ‘good’ wheat was
clearly demonstrated, irrespective of the age of pigs studied (Table 7.4).
The relative importance of soluble arabinoxylans to the response to xylanase in
the young pig is thrown into question by a number of studies where rye was the
predominant grain (Thacker et al., 1991, 1992a; Bedford et al., 1992; Thacker and
Baas, 1996). Dusel et al. (1997b) showed that the use of a mixed grain diet based
Cont’d on p. 137
3.91 cP
3.9
Meal Pellets
Fig. 7.2. The effect of conditioning and pelleting on solubilization of fibre and
resultant viscosity value. Measured by ‘Avicheck’TM (Finnfeeds), an estimation of
viscosity in the water-soluble phase, following a simulated predigestion step in
vitro. Eleven paired pig feed samples, wheat- and barley-based mixtures (cP =
centipoise).
Weaner Pig Chap 07
128
Table 7.2. Some published studies (1990–2000) on the effects of enzyme preparations on barley-based diets for weaner pigs (note that where digestibility
measurements were made, data are presented preferentially at the ileal level).
26/9/01
(P) or Pig
Basal diet, meal-mash weight Inclusion Area of Plus
References main ingredients (M) (kg) Enzyme rate (t1) response Control enzymes
2:02 PM
wheat, -glucanase, performance
SBM -amylase, (g day1) 381 440
glucoamylase Feed:gain 1.88 1.70*
Bedford et al. Hull-less M 12–18 -Glucanase, 2 kg Weight gain (kg) 5.24 6.14*
(1992) barley, (1086 U g1) Feed:gain 1.75 1.55
Page 128
SBM (P = 0.10)
G. Partridge
Thacker et al. Hull-less P ( < 60°C) 08–18 -Glucanase 2.5 kg Growth
(1992b) barley, (750 U g1) performance
SBM (g day1)
(C1 = no C1 254 274
acid; C2 275 275
C2 = + Feed:gain
fumaric C1 2.20 2.03
acid) C2 1.99 1.91
Faecal
digestibility of
protein (%)
C1 78.4 77.8
C2 77.6 81.7
Faecal
digestibility of
energy (%)
C1 82.6 81.9
C2 81.0 82.3
Weaner Pig Chap 07
Inborr (1994) Hulled or M 9.5–14 -Glucanase 2.5 kg Growth
hull-less 89–95 mU g–1 performance
barley, SBM feed (g day–1)
Hull-less 223 229
Feed:gain 1.60 1.52
26/9/01
Hulled 200 208
Feed:gain 1.71 1.65
Effect of enzyme
on growth
(P = 0.07)
2:02 PM
Effect of enzyme
on feed:gain
(P = 0.06)
Power et al. (1996) Hulled barley, wheat, P (~70°C) 6–15 -Glucanase Growth
Page 129
2000 ppm) and digestibility
at optimum
inclusion level
(2000 ppm)
(g day–1) 201 227*
Feed:gain 2.11 1.95*
Faecal
digestibility
of protein (%) 77.4 80.3
Li et al. (1996b) Hull-less barley, M 6.6–11 -Glucanase 2.5 kg (this Faecal
SBM (1000 U g–1) highest digestibility
inclusion of protein (%) 81.6 88.5*
gave max. (significant linear
response) effects of dose
rate were seen)
Faecal
digestibility
of energy (%) 85.2 89.5*
(significant linear
effects of dose
129
rate were seen)
Continued
Weaner Pig Chap 07
130
Table 7.2. Continued
26/9/01
References main ingredients (M) (kg) Enzyme response Control enzymes
2:02 PM
Ileal
digestibility
of energy (%) 64.9 71.1*
Baidoo et al. Hull-less barley M 14 (ileal Cellulase; 100 g Ileal
(1998b) (varieties ‘Condor’ cannulated) -glucanase; digestibility
Page 130
and ‘Buck’), xylanase of protein (%) 57.6 61.8*
G. Partridge
canola meal (11 U g–1; Effects were
27 U g–1; seen on both
43 U g–1) varieties, hence
these are overall
values
Ileal
digestibility
of energy (%) 57.1 63.3*
Effects were
seen on both
varieties, hence
these are overall
values
Baidoo et al. Hull-less barley, M or P (80°C) 9–60 Cellulase; 100 g Growth
(1998b) canola meal, -glucanase; performance
SBM xylanase (g day–1) 721 782*
(11 U g–1; Feed:gain 2.01 1.84*
27 U g–1; Positive effects
43 U g–1) to enzyme addition
Weaner Pig Chap 07
were seen in both
mash and pelleted
feed, hence overall
values are shown
Baidoo, et al. Hull-less barley M 10–14 E1: Xylanase Ileal
26/9/01
(1998a) (Falcon) (ileal E2: -glucanase digestibility
cannulated) E3: Xylanase; of protein (%) 59.6
-glucanase; E1 60.1
protease E2 66.7*
E3 67.2*
2:02 PM
Ileal
digestibility
of energy (%) 62.1
E1 64.6
Page 131
E3 71.2*
Baidoo et al. Hull-less barley M 14 (ileal Xylanase; Ileal
(1998a) (Buck, Condor cannulated) -glucanase; digestibility
or Falcon), canola protease of protein (%)
meal Buck 60.3 64.5
Condor 54.8 60.7
Falcon 67.6 67.2
Ileal
digestibility
of energy (%)
Buck 57.7 67.0
Condor 56.2 66.3
Falcon 60.6 61.3
Gill et al. (2000) Hulled barley, P (75°C) 8–18 -glucanase; 1 kg Growth
SBM xylanase; performance
amylase (g day–1) 329 350
Feed:gain 1.60 1.53
DE (MJ kg ) –1 14.86 15.08
Faecal
digestibility
131
of protein (%) 76.8 75.1
SBM = soybean meal.
* P < 0.05.
Weaner Pig Chap 07
132
Table 7.3. Some published studies (1990–2000) on the effects of enzyme preparations on wheat-, rye- or triticale-based diets for weaner pigs (where digestibility
measurements were made, data are presented preferentially at the ileal level).
26/9/01
Basal diet, meal-mash weight Inclusion Area of Plus
References main ingredients (M) (kg) Enzyme rate (t1) response Control enzymes
2:02 PM
SBM -amylase; (g day–1) 411 458
glucoamylase Feed:gain 1.81 1.56*
Bedford et al. Rye, SBM M 11–15 Xylanase 2 kg Weight gain (kg) 4.24 4.24
(1992) (850 U g–1) Feed:gain 1.64 1.57
McLean et al. Wheat, SBM Weaner E1: Amylase; 700 g Ileal
Page 132
(1992) pigs xylanase; digestibility
G. Partridge
cellulase; of protein (%) 74.0
pectinase E1 76.6
E2 73.1
E2: -glucanase; 1 kg Ileal
pectinase; digestibility
cellulase; of energy (%) 70.8
hemicellulase E1 73.5*
E2 72.9
Inborr et al. Wheat:barley M ~6–10 E1: -glucanase # 1; 1 kg Growth
(1993) (50:50), SBM xylanase; amylase performance
(35; 590; 3300 U g–1) (g day–1) 204
E2: -glucanase # 2; 950 g Feed:gain 1.38
xylanase; amylase Feed:gain 1.38
(41; 740; 3300 U g–1) E1 223
E2 205
1.39
Ileal
digestibility
of protein (%) 76.8
E1 78.4
Weaner Pig Chap 07
E2 77.9
Ileal
digestibility
of starch (%) 94.9
E1 97.7*
26/9/01
E2 96.8
Officer (1995) Wheat, fishmeal, M ~6.5– 14 E1: protease; 2 kg Growth
meat meal, SBM lipase; performance
-glucanase; (after weaning)
amylase; cellulase (g day–1)
2:02 PM
E2: protease; 2 kg E1 309 319
lipase; Feed:gain 1.54 1.40
-glucanase; (P < 0.10)
amylase; E2 299 282
Page 133
hemicellulase; E3 361 332
pentosanase Feed:gain 1.56 1.49
E3: amylase; 2 kg (P < 0.10)
xylanase;
-glucanase;
pectinase
Campbell et al. Wheat, lupin (W/L) P (70°C) 6–13 Xylanases; 400 g Growth
(1995) or wheat, SBM (W/S) pentosanases; performance
-glucanases W/L 355 388
Feed:gain 1.21 1.15
W/S 291 353
Feed:gain 1.29 1.23
Overall effects
of enzyme on
growth rate
(P < 0.05)
and feed:gain
(P = 0.08)
Continued
133
Weaner Pig Chap 07
Table 7.3. Continued
134
Pelleted Enzymes added Responses
(P) or Pig
Basal diet, meal-mash weight Inclusion Area of Plus
References main ingredients (M) (kg) Enzyme rate (t1) response Control enzymes
26/9/01
Schulze et al. Wheat, SBM P 8–22 E1: Xylanase # 1 1 kg Growth
(1996) (5000 U g–1) performance
E2: Xylanase # 1/ 1 kg (g day–1) 304
protease Feed:gain 1.69
2:02 PM
(5000/500 U g–1) E1 394*
E3: Xylanase # 2 1 kg Feed:gain 1.49
(5000 U g–1) E2 385*
Feed:gain 1.49*
E3 402*
Page 134
G. Partridge
Feed:gain 1.45*
Li et al. (1996a) Wheat, SBM M 7–11 -glucanase 2 kg Ileal
(1000 U g–1) digestibility
of protein (%) 68.8 75.9
Ileal
digestibility
of energy (%) 66.9 71.2
Li et al. (1996b) Wheat, SBM (W) or M 6–11 -glucanase, 2 kg (W) Faecal
rye, SBM (R) 1000 U g–1 1 kg (R) digestibility
(A dose–response of protein (%)
study – the W 85.1 89.1
numerically max. R 87.0 89.3
responses are Faecal
shown) digestibility
of energy (%)
W 86.8 88.4
R 87.2 88.1
Dusel et al. Wheat (high extract P 10–25 E1: Xylanase 1 kg Growth
(1997b) and viscosity), rye (5000 U g–1) performance
Jeroch et al. barley, SBM or (g day–1) 354
(1999) E2: Xylanase 1 kg Feed:gain 1.92
Weaner Pig Chap 07
(5000 U g–1) E* 462*
Protease Feed:gain 1.66*
(500 U g–1) E* 480*
Feed:gain 1.65*
DE (measured
26/9/01
MJ kg–1 dry
matter) 13.87 15.08*
Partridge et al. Wheat:barley P 8–30 Xylanase; 1 kg Growth
(1998) (50:50), SBM – C1 -glucanase; performance
Wheat:corn P amylase; (g day–1)
2:02 PM
(50:50), wheat pectinase C1 575 581
pollard, SBM – C2 (4000; 150; Feed:gain 1.67 1.59*
1000; 25 U g–1) C2 512 557*
Feed:gain 1.69 1.62*
Page 135
(1999) diet), low (L) or (5000 U g–1) performance
high (H) extract (g day–1)
viscosity L 376 383
Feed:gain 1.96 1.93
H 398 400
Feed:gain 1.90 1.80
DE (measured,
MJ kg–1 dry
matter)
L 16.23 16.29
H 15.91 16.07
Choct et al. High (H), 7–16 Xylanase; 120 g Growth
(1999) medium (M) -glucanase; performance
and low (L) cellulase (g day–1)
feed intake L 230 466*
wheats – pre- Feed:gain 1.38 1.23
determined M 425 445
Feed:gain 1.27 1.20
H 460 479
Feed:gain 1.14 1.20
135
Continued
Weaner Pig Chap 07
136
Table 7.3. Continued
5/10/01
References main ingredients (M) (kg) Enzyme response Control enzymes
Gill et al. (2000) Wheat/SBM (W) P (75°C) 8–18 Xylanase; amylase; 1 kg Growth
Wheat/beetpulp/ pectinase (W) performance
SBM (WSBP) -glucanase; (g day–1)
9:05 AM
amylase; (W) 338 360
pectinase (WSBP) Feed:gain 1.53 1.48
(WSBP) 345 363
Feed:gain 1.56 1.48
Faecal
Page 136
digestibility
G. Partridge
of protein (%)
(WSBP) 72.1 73.9
DE (MJ kg1)
(WSBP) 14.10 14.33
Table 7.4. Effect of xylanase addition to wheats pre-selected for productive performance
(determined in prior growth studies with weaner pigs, 28–60 kg) (Partridge et al., 1999).
*P < 0.05.
NS, non-significant.
ab Means with different superscripts in the same column are significantly different (P <
0.05).
Table 7.5. Effect of enzyme addition to a wheat-based diet with a high extract viscosity,
piglets 10–25 kg (Dusel et al.,1997b, and personal communication).
Viscosity in
Wheat extract Daily gain Daily feed the small
viscosity* (g) (g) Feed:gain intestine (cP)
on a combination of wheat (high extract viscosity), rye and barley gave responses to
enzyme addition that went beyond the apparent effects on viscosity reduction
(Table 7.5). As in the studies of Choct et al. (1999) and Partridge et al. (1999),
addition of the xylanase or xylanase/protease combination in these diets released an
apparent limitation on voluntary food intake in the animal such that the growth
response seen was a cumulative effect of both increased nutrient availability and
intake. Other, more subtle changes in the microbial flora may also have influenced
the response to the enzyme, a factor that will be examined later in this chapter. Pigs
in the xylanase/protease-supplemented group appeared to show decreased bacterial
proliferation in the small intestine (Fig. 7.3) when compared with both control
diets, suggesting that some degree of nutrient ‘sparing’ may also have contributed to
the productive response seen. These studies illustrate the difficulty in interpreting
responses, or apparent lack of them, to exogenous enzymes in the pig unless many
parameters are examined synchronously. The ‘microbial environment’ in which the
enzyme is used will almost certainly influence the response seen, which has implica-
tions for trials based on ‘low-challenge’ research vs. more commercial situations.
Weaner Pig Chap 07 26/9/01 2:02 PM Page 138
138 G. Partridge
0
Stomach Duodenum Jejunum IIeum Caecum
Fig. 7.3. Effect on microbial populations in digesta when piglets are fed diets based
on high viscous wheat + xylanase/protease, or low viscous wheat. Microscopical
analysis of digesta (VTT Biotechnology and Food Research, Finland); samples from
the studies of Dusel et al. (1997a). Y-axis is microbial loading in digesta; 0 = few
microbes present; 4 = high levels of microbes present.
Phytases
Of the total phosphorus found in common ingredients used in pig and poultry
feeds, around 60–80% is in the form of phytates, which are salts of phytic acid. The
bioavailability of phosphorus in cereal grains, their by-products and legumes is very
low (e.g. 10–30% in maize/soybean diets) because both pigs and poultry have a very
limited capability to utilize phytate phosphorus (Kornegay, 1996). Phytic acid also
has chelating potential in the gut, forming a wide variety of insoluble salts with di-
and trivalent cations at neutral pH. Manganese, iron and magnesium can all be com-
plexed, but zinc and copper have the strongest binding affinity. This binding poten-
tially renders these minerals less available for intestinal absorption. Under neutral
conditions, the carboxyl groups of some amino acids may also bind to phytate
through a divalent or trivalent mineral. Phytate–protein or phytate–mineral–protein
complexes may therefore reduce, to some degree, the utilization of dietary protein.
The phosphate groups of phytate must be hydrolysed before the phosphorus
can be utilized by pigs or poultry and the supplementation of feeds with microbially
synthesized phytase offers this opportunity. The last decade has seen a big invest-
ment in research on phytase, and latterly in its application in diets for different
classes of stock when it is economically interesting relative to inorganic phosphorus,
or desirable for environmental reasons.
Benefits of phytase supplementation have been found in a number of studies in
the weaned pig (e.g. Beers and Jongbloed, 1992; Mroz et al., 1994; Yi et al., 1996;
Qian et al., 1996; Cadogan et al., 1999). In the latter study superior feed:gain ratios
were seen in phytase-supplemented feeds, where calcium and available phosphorus
levels were reduced by 0.12% relative to the positive control diet (Table 7.6). The
opportunity that phytase supplementation offers for reductions in acid-binding
capacity, due to reduced calcium levels, is also considered a bonus in formulations
for young pigs with limited acid-secreting capability.
Weaner Pig Chap 07 26/9/01 2:02 PM Page 139
Table 7.6. The effect of phytase addition and lysine level on the performance of male pigs
over 21 days after weaning (Cadogan et al., 1999).
Available
Phytase level lysine Daily feed
(FTU kg–1) (g MJ1 DE) Daily gain (g) intake (g) Feed:gain
Proteases
Maximizing the digestibility of raw materials is a primary concern in the formula-
tion of diets for young pigs immediately after weaning, especially in segregated
early-weaning systems, where animals will have a particularly immature digestive
system. Protein sources in the most effective diets for weaned pigs have historically
been highly processed and relatively expensive raw materials, such as low tempera-
ture-processed fishmeal, milk products and plasma products, coupled with limited
amounts of certain isolated vegetable proteins. Although these diets undoubtedly
work successfully, there comes a stage in most production systems at which the
young animal has to adapt to simpler diets heavily based on vegetable protein, usu-
ally soybeans, in extracted or full fat form.
Concerns about the extensive use of soya in young pig diets are mainly fuelled
by the residual anti-nutrients it can contain (e.g. trypsin inhibitors, lectins), which
are known to stimulate excessive endogenous nitrogen losses (Jansman et al., 1994),
as well as its tendency to induce damaging, transient hypersensitivity reactions in
the immature gut (Li et al., 1991; Friesen et al., 1993). The main storage proteins
in the bean (glycinin and -conglycinin) have been particularly implicated in this
respect. Tokach et al. (1994) outlined the practical dilemma, under USA pro-
duction conditions, whereby feeding all-milk protein diets after weaning merely
seemed to postpone this damaging hypersensitivity response giving no net growth
advantage and a clear economic disadvantage over a 5-week period after weaning
(Table 7.7). There seems, therefore, to be a physiological benefit in introducing the
young animal to soybean protein as soon as possible but without inducing these
problems, which seem to be further influenced by the degree of immunological
‘stress’ that the young pigs are under, such as health status and presence or absence
of an antibiotic in the diet (Yen and Pond, 1987; Friesen et al., 1993).
Weaner Pig Chap 07 26/9/01 2:02 PM Page 140
140 G. Partridge
Table 7.7. The effect of post-weaning exposure to a soybean-based diet on starter pig
performance over a 5-week period (Tokach et al., 1994). All pigs received the same phase
II diet from days 14 to 35.
Days 0–14
Daily gain (g) 299 218 < 0.01
Daily feed intake (g) 295 277 < 0.05
Food conversion ratio 0.99 1.27 < 0.01
Days 14–35
Daily gain (g) 404 503 < 0.01
Daily feed intake (g) 739 807 < 0.01
Food conversion ratio 1.83 1.60 < 0.01
Days 0–35
Daily gain (g) 363 390 < 0.05
Daily feed intake (g) 562 594 < 0.05
Food conversion ratio 1.54 1.51 NS
Milk-based diet: maize 44%; skimmed milk 20%; whey powder 20%; casein 7%; soya oil
6%.
Soybean-based diet: maize 25%; soybean meal 40%; lactose 24%; soya oil 6%.
Phase II diet (days 14–35): maize 56%; soybean meal 23%; whey powder 10%; fishmeal
4%; soya oil 4%.
Table 7.8. Effect of protease addition on total soluble matter, soluble crude protein and
soybean trypsin inhibitor level (SBTI) under the optimum temperature (50°C),
concentration 1 mg g1 soybean meal and pH conditions (pH 4.5) defined by Caine et al.
(1998) (in vitro studies with subtilisin protease).
Control
(no enzyme addition) + Protease
211 210
198
191
1.84
175 1.74
155**
1.48
141** 1.40 1.35
129 121
95
Growth rate (g day–1) Dry matter feed intake (g day–1) Dry matter feed:gain
Fig. 7.4. The effect of pre-treatment of soybean meal with acid + protease on
piglet performance in the first 7 days after weaning (Rooke et al., 1996). Soybean
meal treated for 3 h at 50°C, pH 4.5 and at 20% dry matter; product neutralized
and then dried (65°C) before inclusion into a maize diet + amino acids +
vitamin/mineral, lysine 1.2%, digestible energy 14.5 MJ kg1 (3465 kcal). **P <
0.01.
Weaner Pig Chap 07 26/9/01 2:02 PM Page 142
142 G. Partridge
1479b
1442b
1373b
1250a
2.32a 2.22a
1.87b
1.71b
845c
792c
620b
540a
Daily dry matter intake (g) Daily gain (g) Dry matter feed:gain
Fig. 7.5. The effect of protease pretreatment of raw or autoclaved full-fat soybeans
on grower pig performance (32 kg start weight) (Beal et al., 1998b).
a–c within parameters, means with different superscripts are significantly different (P
< 0.05).
8.7 36.3
5.7
Fig. 7.6. Effect of wheat variety on intestinal viscosity (mPa) in broilers, turkeys and
piglets (adapted from Danicke et al., 1999). Ileum = small intestine (lower two-
thirds for the pig); wheat var. 1 = ‘Ibis’ (11 g soluble arabinoxylan kg1 dry matter);
wheat var. 2 = ‘Alidos’ (17 g soluble arabinoxylan kg1 dry matter).
6.4
4.2
3.3
3.1*
2.2 1.8(*)
2.2*
1.7 1.5* 1.7*
Fig. 7.7. Effects of enzymes on viscosity in the small intestine of piglets. Ref. 1,
Inborr (1994); Ref. 2, Sudendey and Kamphues (1995) wheat/piglet and
barley/grower; Ref. 3, Dusel et al. (1997a); Ref. 4, Partridge, et al. (1998).
Other physiological effects have been found following the addition of exoge-
nous enzymes to diets for young pigs. Sudendey and Kamphues (1995) (Fig. 7.8)
described studies in which viscosity reduction following enzyme addition in both
wheat- and barley-based diets was associated with significant increases in digesta dry
matter flow rate from the stomach down into the small intestine. Ellis et al. (1996)
noted that an increase in gastric viscosity in the pig can lead to a failure in the ‘siev-
ing mechanism’ whereby larger digesta particles normally fall to the bottom of the
stomach – a feature seemingly unique to the pig. In a viscous environment, these
large particles stay suspended and are thereby more likely to exit the pylorus before
Weaner Pig Chap 07 26/9/01 2:02 PM Page 144
144 G. Partridge
(*) 3.36 *
3.22
2.62
2.27 2.26
1.70
Control
Fig. 7.8. Effect of enzyme addition to wheat- or barley-based diets on dry matter
outflow rate from the stomach (Sudendey and Kamphues, 1995). Values are dry
matter (g) leaving the stomach per hour per kilogram body weight.
being exposed to gastric secretions or particle size reduction, which may have nega-
tive implications for subsequent digestion. Further down the gastrointestinal tract,
viscosity is also known to disturb peristalsis, creating laminar rather than disturbed
flow, and also leads to disrupted pancreatic secretion via its effects on gastric
inhibitory polypeptide. Viscous NSPs may also physically coat starch granules, fur-
ther reducing the rate of digestion (Ellis et al., 1996).
In successful pig trials using enzyme supplementation, a stimulation of voluntary
food intake is frequently an important contributor to the observed improvements in
daily liveweight gain (Haberer et al., 1998). The fact that gastric emptying rate and dis-
tension of the stomach are two of a number of factors involved in satiety signals in the
pig (Forbes, 1995) is unlikely to be coincidental. It is proposed that some of the posi-
tive responses to exogenous enzymes on voluntary food intake could be due to various
influences on digesta flow rate as well as to improvements in nutrient availability and
digestibility in the small intestine that appear to have an influence on food intake. For
example, Whittemore (1993) proposed the following relationship between food intake
and diet digestibility in the pig:
Limit to voluntary food intake (kg day1) ~
– [0.013 liveweight (kg)]/
(1 digestibility coefficient).
Feedback loops involving gut hormones could also be contributors to carbohydrase
effects, as outlined by Bedford and Schulze (1998). In this case it was proposed that
caecal fermentation, enhanced by breakdown products from xylanase addition (fer-
mentable oligomers), could influence enteroglucagon concentrations, which in turn
influence stomach motility by depressing gastrin concentrations. These many possibili-
ties suggest that exogenous enzymes could be potent influencers of gastrointestinal
physiology of the young pig through a variety of different mechanisms, but these
aspects have been little studied to date.
The water-holding capacity (WHC) of feed raw materials has also been shown
to influence voluntary food intake in young pigs (Kyriazakis and Emmans, 1995),
with feed intake falling as WHC in the diet rises. As carbohydrase enzymes have
Weaner Pig Chap 07 26/9/01 2:02 PM Page 145
–12% –29% –31% –19% –12% –17% –52% –8% –14% –5% –75%
10
s
l
ur
rm
er
l
y
d
s
M
ze
ea
ea
ke
rle
at
ea
SB
flo
w
ai
ge
m
O
fla
lo
Ba
Br
M
at
nf
ss
ze
to
ee
he
Su
ra
ai
ta
es
W
G
M
Po
ap
No Enzyme R
+ Enzyme complex (xylanase, β-glucanase; protease)
been found to reduce the WHC of feedstuffs in vitro (Fig. 7.9), it follows that some
of the benefits of enzyme addition on voluntary food intake in piglets may arise
from influences on these physical factors in the gut. These benefits would be in
addition to the potential benefits of enzyme use arising from both water and water-
soluble nutrient release from dietary fibre residues in the gut.
146 G. Partridge
Other studies have reported positive interactions between the use of feed
enzymes and the incidence of diet-induced diarrhoea, particularly in the young pig
after weaning (Inborr and Ogle, 1988; Flourou-Paneri et al., 1998; Kantas et al.,
1998). These, together with reports of potential synergies between enzymes and thera-
peutic and sub-therapeutic addition of antibiotics (Flourou-Paneri et al., 1998; Kantas
et al., 1998; Gollnisch et al., 1999) offer interesting opportunities for future produc-
tion methods where strategic rather than routine antibiotic use will become the norm.
Recent advances in methodologies to appraise the gut microflora have enabled
quantification of the changes that exogenous enzymes can illicit. These methods
overcome the limitations of conventional bacterial plating techniques, allowing
changes and disturbances in bacterial populations to be studied in the presence or
absence of various feed additives. Apajalahti and Bedford (1998), for example,
described the use of DNA-based techniques to identify changes in the populations
of bacterial genera in the guts of broilers fed enzymes. The addition of xylanase to
wheat-based diets altered the relative abundance of bacterial DNA collected from
the caecal flora (Table 7.9), with an increase in abundance of Peptostreptococcus,
Bacteroides, Proprionibacterium, Eubacterium and Bifidobacterium and decreased
abundance of Clostridium, Enterobacteriacaea and Campylobacter. The production
of volatile fatty acids (VFAs) in the caecum was also changed, with a relative rise in
proprionate production and stimulation of total VFAs produced. Huge shifts in
bacterial groups were also reported, with changes in wheat:rye ratio in the broiler
(Table 7.9), with clear implications for bird performance and the likelihood of dis-
ease incidence. Total microbial numbers in the small intestine of broilers fed wheat-
based diets were also reduced by addition of xylanase compared with control
animals, though to a lesser extent than the addition of an antimicrobial (avoparcin).
There are clear parallels in this work with the increasing interest in non-
digestible oligosaccharides (NDOs) as potential ‘prebiotics’ (functional foods) for
use in human foods and animal feeds (Mul and Perry, 1994; Grizard and
Barthomeuf, 1999; van Loo et al., 1999; Mosenthin et al., 1999). Xylo-oligosaccha-
rides are one such group of potentially beneficial NDOs and it is of interest in the
studies of Apajalahti and Bedford (1998) that addition of xylanase resulted in
increased concentrations of short-chain xylo-oligomers in the ileum of broilers fed
wheat-based diets (Table 7.9). Whether these breakdown products of xylanase addi-
tion influence the microbial population in a beneficial way remains an active area of
current research and hints at potential opportunities for in situ prebiosis, if it can be
adequately characterized and manipulated.
Future Perspectives
Some of the nutritional problems faced by the young pig in the post-weaning
period have proved to be positively influenced by the use of an increasingly effective
range of specific exogenous enzymes over the past decade. The most cost-effective
use of feed enzymes will always demand a good understanding of the anti-nutri-
tional factors that influence raw material variability and (of equal importance
Weaner Pig Chap 07 26/9/01 2:02 PM Page 147
Table 7.9. The effect of xylanase addition to wheat- (and rye)-based broiler feeds on
microflora changes in the caecum and xylo-oligomer production in the ileum. Data from
Apajalahti and Bedford (1998).
(a) Relative abundance (%) of bacterial DNA collected from the caeca of broiler
chickens fed wheat-based diets with or without 2500 U xylanase kg1 feed.
(b) Influence of the proportion of wheat:rye in broiler diets on the relative abundance of
bacterial DNA (%) in the caeca of broiler chickens.
(c) Effect of the addition of xylanase to wheat-based broiler diets on the concentration (g
kg1 digesta) of xylo-oligomers of varying degrees of polymerization in the ileum.
Degree of polymerization
commercially) their rapid estimation at realistic cost. This remains an area of active
research, with considerable scope for improvements in animal performance.
Changes in feeding systems for young pigs, specifically the potential use of wet-
feeding (where applicable), also offer a good opportunity for enzyme solutions and,
ultimately, more cost-effective production.
Finally, the changing nature of animal production with an increasing emphasis
on prophylactic nutrition focuses even more attention on understanding the
Weaner Pig Chap 07 26/9/01 2:02 PM Page 148
148 G. Partridge
complex interrelationships between the animal, its microflora and the feed it
receives. New methodologies in this area offer much scope for improvements in
nutritional welfare, particularly for the young newly weaned animal.
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