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Contents lists available at ScienceDirect

Animal Feed Science and Technology

journal homepage: www.elsevier.com/locate/anifeedsci

The effects of lactose inclusion and seaweed extract derived from

Laminaria spp. on performance, digestibility of diet components
and microbial populations in newly weaned pigs
J.V. O’Doherty ∗ , S. Dillon, S. Figat, J.J. Callan, T. Sweeney
School of Agriculture, Food Science & Veterinary Medicine, College of Life Sciences, Lyons Research Farm,
Newcastle, Co. Dublin, Ireland

a r t i c l e i n f o a b s t r a c t

Article history: A 2 × 2 factorial experiment was conducted to investigate the interactions between two
Received 21 January 2009 different lactose (L) levels (150 g/kg vs. 250 g/kg) and seaweed extract (SWE): (0 g/kg vs.
Received in revised form 1 March 2010
2.8 g/kg; containing laminarin and fucoidan) derived from Laminaria spp. on growth per-
Accepted 4 March 2010
formance, coefficient of total tract apparent digestibility (CTTAD) and faecal microbial
Available online xxx
populations in the weanling pig. Two hundred and forty pigs (120 male and 120 female)
were selected after weaning (24 days of age, 7.6 ± 0.9 kg live weight) and blocked on the
Keywords:
basis of live weight and within each block assigned to one of the four dietary treatments. The
Lactose
Laminarin pigs were offered the following diets on an ad libitum basis for 25 days: (T1) 150 g L/kg; (T2)
Fucoidan 150 g L/kg + SWE; (T3) 250 g L/kg; (T4) 250 g L/kg + SWE. Pigs offered diets supplemented
Seaweed with SWE had a higher average daily gain (ADG): (322 g vs. 281 g, s.e. ± 9.0; P<0.01) and gain
Pig to feed (G:F) ratio (669 g/kg vs. 611 g/kg, s.e. ± 19.0; P<0.05) between days (d) 0–25 com-
pared with pigs offered non-SWE diets. Pigs offered high L diets had a higher ADG (319 g
vs. 283 g, s.e. ± 9.0; P<0.05) and average daily feed intake (ADFI) between d 0–25 (480 g
vs. 447 g, s.e. ± 11.0; P<0.05) compared with pigs offered the low L diets. The inclusion of
SWE increased (P<0.001) the CTTAD of nitrogen (N) and gross energy (GE) and reduced the
counts of Escherichia coli in the faeces compared with non-SWE diets. Pigs fed the high L
diets had increased CTTAD of GE (P<0.001) and N (P<0.05) and decreased the counts of E. coli
in the faeces compared with pigs offered low L diets. Summarising, the inclusion of either
a high dietary concentration of L or a laminarin–fucoidan extract increased the CTTAD of
diet components, decreased the counts of E. coli in the faeces and improved performance
of pigs after weaning in the current study.
© 2010 Elsevier B.V. All rights reserved.

1. Introduction

At weaning, the pig is subjected to an array of stressors of nutritional, environmental and social origin (Pluske et al.,
1996). These stressors have been shown to be associated with reduced growth rates (Pluske et al., 1997), changes in the gut
morphology (Li et al., 1990; Pluske et al., 1996) and a change in the microbial populations in the gut, leading to increased

Abbreviations: ADG, average daily gain; ADFI, average daily feed intake; AIA, acid insoluble ash; AGP, antibiotic growth promoter; CS, cleanliness scores;
CTTAD, coefficient of total tract apparent digestibility; d, days; DM, dry matter; FS, faecal scores; GE, gross energy; G:F, gain to feed; HS, health scores; L,
lactose; N, nitrogen; NDF, neutral detergent fibre; OM, organic matter; SWE, seaweed extract; VFA, volatile fatty acid.
∗ Corresponding author. Tel.: +353 17167128; fax: +353 17161103.
E-mail address: john.vodoherty@ucd.ie (J.V. O’Doherty).

0377-8401/$ – see front matter © 2010 Elsevier B.V. All rights reserved.
doi:10.1016/j.anifeedsci.2010.03.004

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
Laminaria spp. on performance, digestibility of diet components and microbial populations in newly weaned pigs. Anim.
Feed Sci. Technol. (2010), doi:10.1016/j.anifeedsci.2010.03.004
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susceptibility to scouring and disease (Hampson, 1986). In the past, these problems have been managed in a controlled
fashion by the use of prophylactic antibiotics (Williams et al., 2001). Also high concentrations of dietary minerals, e.g. zinc
in the form of zinc oxide, were shown to decrease the incidence of nonspecific post-weaning scours (Poulsen, 1995) and
to improve growth performance of newly weaned pigs (O’Doherty et al., 2005). However, the use of high concentrations of
inorganic minerals has raised some environmental concerns.
Much research exists regarding the benefits of lactose (L) supplementation in terms of improving pig performance
(O’Doherty et al., 2005; Pierce et al., 2006). The inclusion of high levels of L in weaning pig diets resulted in a reduction
in intestinal pH and increases in Lactobacilli spp. and short-chain fatty acids and reductions in coliforms (Pierce et al., 2006).
Recently seaweeds and sea weed extracts (SWE) have been researched as an alternative to in-feed antibiotics in pig diets
(Gardiner et al., 2008; Reilly et al., 2008; Gahan et al., 2009). Seaweeds are rich in undigestible polysaccharides and are a
potential source of soluble dietary fibres. The most abundant polysaccharides in brown seaweeds are laminarin, fucoidan
and alginic acid (Zvyagintseva et al., 2003). Laminarin has a chemical structure consisting mainly of a linear ␤-(1 → 3)-linked
glucose backbone with some random ␤-(1 → 6)-linked glucose side-chains depending on the variety of seaweed (Brown
and Gordon, 2005). Recently, Lynch et al. (2010) showed that laminarin had antimicrobial properties. Deville et al. (2007)
noted that laminarin can influence the adherence and the translocation of bacteria across the epithelial wall and seems to
be a modulator of the intestinal metabolism by its effects on mucus composition, intestinal pH and short-chain fatty acid
production, especially butyrate. Fucoidans are sulfated polysaccharides extracted from the cell wall of various species of
brown seaweeds (Berteau and Mulloy, 2003). Fucoidans have been shown to have antitumor (Zhuang et al., 1995), antiviral
(Lee et al., 2004) and antibacterial (McClure et al., 1992) properties. Thus, fucoidan and laminarin have a wide spectrum of
biological actions which may alleviate the need for high concentrations of L (O’Doherty et al., 2004) in antibiotic growth
promoter (AGP)-free diets.
The hypothesis of this study is that the inclusion of a seaweed extract containing laminarin and fucoidan from Lami-
naria spp. will reduce the counts of Escherichia coli, allowing levels of L to be reduced while maintaining post-weaning pig
performance in AGP-free diets.

2. Materials and methods

All procedures described in this experiment were conducted under experimental licence from the Irish Department of
Health in accordance with the Cruelty to Animals Act 1876 and the European Communities (Amendments of the Cruelty to
Animals Act 1976) Regulations, 1994.

2.1. Experimental design and dietary treatments

This experiment was designed as a 2 × 2 factorial arrangement of dietary treatments with the respective factors being
two different levels of L: low (150 g/kg) and high (250 g/kg); and the presence (2.8 g/kg) or absence of SWE (AGP-free diets).
This experiment was carried out over two periods of 25 days. Two hundred and forty pigs (120 male and 120 female) were
selected after weaning at 24 days of age and blocked on the basis of weaning weight and within each block assigned to one
of four dietary treatments (equal numbers of males and females were allocated to each pen). The pigs in periods 1 and 2
had an initial live weight of 7.2 and 7.8 kg (s.d. 0.9 kg) respectively. During the experiment (d 0–25) the pigs were offered
the following diets: (T1) 150 g L/kg; (T2) 150 g L/kg + SWE; (T3) 250 g L/kg; (T4) 250 g L/kg + SWE. The SWE was included
at 2.8 g/kg and derived from Laminaria digitata. The extract contained both laminarin (112 g/kg), fucoidan (89 g/kg) and
ash (799 g/kg) and was sourced from Bioatlantis Ltd. (Tralee, County Kerry, Ireland). The concentration of SWE used was
deemed as the optimum dietary inclusion level based on results of work from Gahan et al. (2009) and Reilly et al. (2008). The
composition and chemical analysis of the experimental diets are presented in Table 1. Diets were formulated to have similar
concentrations of digestible energy (16 MJ/kg) and standardised ileal digestible lysine (14 g/kg). All amino acid requirements
were met relative to lysine (Close, 1994).

2.2. Animals and management

Pigs were housed in groups of four (15 replicates per treatment, 9 replicates in period 1 and 6 replicates in period 2)
on fully slatted pens (1.68 m × 1.22 m). House temperature was maintained at 30 ◦ C in the first week and then reduced by
2 ◦ C per week. Pigs were weighed at the beginning of the experiment (day of weaning = day 0), day 7, day 14 and day 25.
The pigs were fed on an ad libitum basis from a four space feeder with precautions taken to avoid wastage of feed. Water
was available on an ad libitum basis from nipple drinkers. Food remained in the feeders until the time pigs were weighed,
and then was weighed back for the purpose of calculating G:F ratio. During the experiment, feed samples were collected at
time of feeding and were retained for chemical analysis. Multiple fresh faecal samples were collected daily from all pens on
days 10–15 and mixed with sodium benzoate and phenylmethylsulfonyl fluoride, in order to stop any bacterial activity and
minimize the effects of post-thawing fermentation on resulting volatile fatty acid (VFA) concentrations. The samples were
then stored at −20 ◦ C until analysis for VFA and acid insoluble ash (AIA) contents. Chromic oxide (150 mg/kg) was added to the
feed at manufacture. Multiple fresh faecal samples were collected from all pens on day 10 and stored in sterile containers

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
Laminaria spp. on performance, digestibility of diet components and microbial populations in newly weaned pigs. Anim.
Feed Sci. Technol. (2010), doi:10.1016/j.anifeedsci.2010.03.004
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Table 1
Composition and chemical analysis of experimental starter diets (g/kg as fed).

Treatment

T1 T2 T3 T4
Lactose 150 (g/kg) 150 (g/kg) 250 (g/kg) 250 (g/kg)
Seaweed extract − + − +

Ingredients (g/kg)
Whey permeate 107 107 215 215
Wheat 445 445 349 349
Soya bean meal 160 160 160 160
Whey protein isolate 130 130 130 130
Full fat soya bean 80 80 80 80
Soya oil 65 65 55 55
Lysine HCL 4.5 4.5 3.7 3.7
dl-Methionine 1.6 1.6 1.6 1.6
l-Threonine 2.2 2.2 1.8 1.8
Vitamin and mineral premixa 4 4 4 4

Analysed composition (g/kg)

Dry matter 902 899 913 903
Lactose 148 153 247 249
Crude protein (N*6.25) 204 201 198 196
Neutral detergent fibre 85.9 96.2 85.6 78.5
Ash 43 45 50 50
Gross energy (MJ/kg) 17.7 17.8 17.5 17.3
Lysineb 16 16 16 16
Methionine and cysteineb 8.8 8.8 8.8 8.8
Threonineb 10.4 10.4 10.4 10.4
Laminarin (mg/kg) 0 300 0 300
Fucoidan (mg/kg) 0 236 0 236

Lactofeed 70: Volac International Ltd., Orwel, Royston, SG8 5QX, UK. The chemical analysis is as follows (g/kg unless otherwise stated): DM 955, crude
protein 125, oil 50, ash 90, fibre 10, GE content of 15.5 MJ/kg and a pH of 6.5–7.
a
Provided (mg/kg complete diet): Cu 100, Fe 140, Mn 47, Zn 120, I .6, Se 0.3, retinol 1.8, cholecalciferol 0.025, alpha-tocopherol 67, phytylmenaquinone 4,
cyanocobalamin 0.01, riboflavin 2, nicotinic acid 12, pantothenic acid 10, choline chloride 250, thiamine 2, pyridoxine 0.015. Chromium Ш oxide included
at 150 mg/kg complete diet.
b
Calculated from total amino acid analysis (Sauvant et al., 2004).

(Sarstedt, Wexford, Ireland) on dry ice and transported to the laboratory within 3 h, for the enumeration of E. coli and
Lactobacilli.

2.3. Faeces scoring and morbidity

Faecal scores (FS), cleanliness scores (CS) and health scores (HS) were recorded daily (09.00 h) in the individual pens,
by the same operator throughout the experiment using a subjective score on a one-point scale ranging from 1 to 4. FS:
1 = firm, 2 = soft, spreads slightly, 3 = very soft, spreads readily, 4 = watery liquid consistency. CS: 1 = clean, free of faecal
contamination, 2 = some indication of faecal contamination, 3 = contaminated with faecal material, 4 = heavily contaminated
with faecal material. Scores were given daily for individual pens and the average faecal score value per pen was given. HS:
1 = no indication of impaired health, lameness or scours, 2 = some indication of impaired health (e.g. slow to respond to
human presence), lameness or scours, 3 = clear indications of impaired health (e.g. shivering), lameness or scours, 4 = serious
signs of impaired health (e.g. showing pain), lameness or scours.

2.4. Laboratory analysis

Both concentrates and faeces were analysed for N, dry matter (DM), ash, GE, neutral detergent fibre (NDF) and
AIA. Following collection, faeces were dried at 100 ◦ C for 48 h. The concentrates and dried faeces samples were milled
through a 1 mm screen (Christy and Norris hammer mill). The DM of faeces and feed were determined after drying
over night at 103 ◦ C. Ash was determined after ignition of a known weight of concentrates or faeces in a muffle fur-
nace (Nabertherm, Bremen, Germany) at 500 ◦ C. The N content of both feed and faeces was determined using the LECO
FP 528 instrument (Leco Instruments, U.K Ltd., Newby Road, Hazel Grove, Stockport, SK7 5DA, Cheshire, UK). The NDF
was determined using a Fibertec extraction unit (Tecator, Hoganans, Sweden). The GE of the feed and faeces was deter-
mined using a Parr 1201 oxygen bomb calorimeter (Parr, Moline, IL, USA). The AIA analysis was determined according
to the method of McCarthy et al. (1974). The total laminarin content in the diets was determined using a Megazyme
kit (Megazyme, Bray, Ireland). The fucoidan level in the diets was determined according to the method described
by Usov et al. (2001). Thawed faecal samples were analysed for VFA concentration and profile using the method of

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
Laminaria spp. on performance, digestibility of diet components and microbial populations in newly weaned pigs. Anim.
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O’Connell et al. (2006). The L concentration of the diets was determined using the method of Birch and Mwangelwa
(1974).

2.5. Microbiology

A 1.0 g sample was removed from the faecal sample, serially diluted (1:10) in 9.0 ml aliquots of maximum recovery diluent
(MRD, Oxoid, Basingstoke, UK), and spread plated (0.1 ml aliquots) onto selective agars, as follows. Lactobacillus spp. were
isolated on de Man, Rogosa and Sharp (MRS) agar (Oxoid) with overnight (18–24 h) incubation at 37 ◦ C in an atmosphere
enriched with CO2 5%, as recommended by the manufacturers (Oxoid). The API 50 CHL (BioMerieux, France) kit was used to
confirm suspect Lactobacilli spp. The E. coli species were isolated on MacConkey agar (Oxoid), following aerobic incubation
at 37 ◦ C for 18–24 h. Suspect colonies were confirmed with API 20E (BioMerieux, France). This API system identifies the
suspect colonies by measuring their ability of produce cytochrome oxidase. Typical colonies of each bacteria on each agar
were counted and the numbers of bacteria expressed per gram of faeces after being serially diluted.

2.6. Statistical analysis

The experimental data was analysed as a 2 × 2 factorial using the General Linear Model procedure of Statistical Analysis
Systems Institute (1985). The models for both performance and digestibility analysis included the main effects of L level, SWE
inclusion, period and the interaction between L and SWE. Both the performance and CTTAD data were adjusted for initial
live weight by covariance analysis. All the data were checked initially for outliers and normality using the PROC Univariate
procedure of Statistical Analysis Systems Institute (1985). The microbial counts were log transformed prior to statistical
analysis. The least significant difference test was used to separate main effect means. The probability level that denotes
significance is P<0.05. The data in the tables are presented as least square interaction means ± standard error of the mean.

3. Results

3.1. Dietary analysis

The concentration of laminarin and fucoidan in T2 and T4 diets was 300 and 236 mg/kg respectively. The AIA content of
the diets ranged from 1.7 to 1.8 g/kg and 989 g/kg of AIA present in the diet was recovered in the faeces.

3.2. Performance

The effect of L concentration and SWE inclusion on ADG, ADFI and G:F ratios are presented in Table 2. Pigs offered diets
containing SWE gained more weight than pigs offered non-SWE diets on d 0–7 (165 g vs. 123 g, s.e. ± 12.0; P<0.05) and d
0–25 (322 g vs. 281 g, s.e. ± 9.0; P<0.01). Pigs offered high L diets had a higher (P<0.05) ADG between d 0–7 (164 g vs. 123 g,

Table 2
The effect of lactose (L) level and seaweed extract (SWE) on pig performance (average daily gain (ADG), average daily feed intake (ADFI) and gain to feed
(G:F) ratio) (least square interaction means ± standard error of mean).

Treatment SEM Significance

T1 T2 T3 T4 L SWE L × SWE
150 (g/kg)a 150 (g/kg)a 250 (g/kg)a 250 (g/kg)a
−b +b −b +b

ADG (g)
d 0–7 100 148 146 183 18.0 * * ns
d 7–14 302 303 325 387 21.0 * ns ns
d 14–25 438 427 388 455 20.0 ns ns *
d 0–25 275 293 287 350 13.0 * ** ns

ADFI (g)
d 0–7 242 257 239 271 15.0 ns ns ns
d 7–14 415 426 450 472 19.0 * ns ns
d 14–25 682 663 683 737 25.0 ns ns ns
d 0–25 446 449 458 502 14.0 * ns ns

(G:F) ratio (g/kg)

d 0–7 413 558 589 659 62.0 * ns ns
d 7–14 747 705 729 832 55.0 ns ns ns
d 14–25 633 636 569 622 27.0 * ns ns
d 0–25 603 633 619 691 62.0 ns * ns

Probability of significance: *P<0.05; **P<0.01; ns: P>0.05.

a
L.
b
SWE.

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
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Table 3
The effect of dietary treatment on the coefficient of total tract apparent digestibility (CTTAD) of organic matter (OM), nitrogen (N), gross energy (GE) and
neutral detergent fibre (NDF) (least square interaction means ± standard error of mean).

Treatment SEM Significance

T1 T2 T3 T4 L SWE L × SWE
150 (g/kg)a 150 (g/kg)a 250 (g/kg)a 250 (g/kg)a
−b +b −b +b

Digestibility coefficientsc
OM 0.862 0.901 0.898 0.937 0.005 *** *** ns
N 0.821 0.880 0.853 0.907 0.011 * *** ns
GE 0.839 0.889 0.882 0.925 0.007 *** *** ns
NDF 0.375 0.650 0.619 0.746 0.027 *** *** *

Probability of significance: *P<0.05; ***P<0.001; ns: P>0.05.

a
L.
b
SWE.
c
Samples collected once daily from all pens on days 10–15.

s.e. ± 12.0), d 7–14 (355 g vs. 302 g, s.e. ± 14.0) and d 0–25 (319 g vs. 283 g, s.e. ± 9.0) than pigs offered low L diets. There was
a L × SWE interaction on ADG between d 14–25 (P<0.05). The addition of the SWE to the high L diet increased ADG compared
with the high L diet only. However the addition of the SWE had no effect at the low L inclusion.
Pigs offered the high L diets had a higher (P<0.05) ADFI between d 7–14 (461 g vs. 420 g, s.e. ± 13.0) and d 0–25 (480 g vs.
447 g, s.e. ± 11.0) compared to pigs offered the low L diets.
Dietary inclusion of SWE increased the G:F ratio between d 0–25 (669 g/kg vs. 611 g/kg, s.e. ± 19.0; P<0.05). The inclusion
of high L increased the G:F ratio between d 0–7 (623 g/kg vs. 485 g/kg, s.e. ± 44.0; P<0.05).

3.3. Coefficient of total tract apparent digestibility (CTTAD)

The effects of L level and SWE on the CTTAD are presented in Table 3. The supplementation of SWE increased (P<0.001)
the CTTAD of organic matter (OM) (0.919 vs. 0.880, s.e. ± 0.004), N (0.893 vs. 0.827, s.e. ±0.007) and GE (0.907 vs. 0.860,
s.e. ± 0.005) compared with the non-SWE diets. The inclusion of high L in the diet increased (P<0.001) the CTTAD of OM
(0.917 vs. 0.881, s.e. ± 0.004) and GE (0.903 vs. 0.864, s.e. ± 0.005), and also increased (P<0.05) CTTAD of N (0.880 vs. 0.850,
s.e. ± 0.007) compared with pigs offered low L diets. There was an L × SWE interaction on the CTTAD of NDF (P<0.05). Pigs
offered the high lactose diet had higher CTTAD of NDF compared with pigs offered the low L diets. However, when the SWE
was supplemented there was no difference in the CTTAD of NDF between both L levels.

3.4. Microbiology and volatile fatty acids

The effect of L level and SWE on selected microbial populations in the faeces of post-weaned pigs is presented in
Table 4. The inclusion of SWE decreased (P<0.05) the counts of E. coli in the faeces compared with non-SWE diets (5.1 vs.
6.0 log10 c.f.u./g, s.e. ± 0.25). Also high L diets decreased the counts of E. coli in the faeces (5.1 vs. 6.0 log10 c.f.u./g, s.e. ± 0.25;
P<0.05) compared with low L diets. There was a significant L × SWE interaction on the numbers of Lactobacilli spp. (P<0.05).
Dietary inclusion of SWE increased the Lactobacilli spp. numbers in pigs fed the high L diets, however there was no effect of
SWE inclusion at the low L concentration. There was no effect of treatment on VFA concentrations or profile.

3.5. Faeces scoring

There was no effect of treatment on FS during the experiment. Pigs offered the high L diets were more contaminated in
faecal material (d 0–25) compared with pigs offered the low L diets (1.18 vs. 1.31, s.e. ± 0.03; P<0.05). There was an interaction

Table 4
The effect of dietary treatment on counts of Lactobacilli and Escherichia coli (least square interaction means ± standard error of mean).

Treatment SEM Significance

T1 T2 T3 T4 L SWE L × SWE
150 (g/kg)a 150 (g/kg)a 250 (g/kg)a 250 (g/kg)a
−b +b −b +b

Bacterial counts (log10 c.f.u./g faeces)

Lactobacilli spp. 8.46 8.63 8.19 8.84 0.12 ns *** *
E. coli 6.30 5.80 5.70 4.50 0.42 * * ns

Probability of significance: *P<0.05; ***P<0.001; ns: P>0.05.

a
L.
b
SWE.

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
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between L level and SWE on pig health between d 7–14 (P<0.05) and CS between d 14–25 (P<0.05). Pigs offered the high L
diets showed more signs of impaired health (d 7–14) and were more contaminated in faecal material (d 14–25) compared
with pigs offered the low L diets. However when SWE was supplemented there were no differences in HS or CS between
both L levels.

4. Discussion

The objective of the experiment was to investigate the effect of inclusion of laminarin and fucoidan, as sources of algal
polysaccharides in the diet of the newly weaned pig on performance, faecal populations of selected microbiota, the CTTAD
of diet components and VFA concentrations at two levels of L inclusion. The diets used in the current experiment were
formulated to create the greatest nutritional challenge possible to the post-weaned pig in the absence of in-feed antibiotics.
These diets contained a high crude protein level, high level of soya bean meal and low level of L. A high level of dietary
crude protein has been suggested to predispose the pig to post-weaning colibacillosis (Prohaszka and Baron, 1980). Soya
bean protein is not included in large quantities in weanling pig diets. It has been proposed that many of the morphological
changes that take place in the young pig’s gut after weaning are due to a transient hypersensitivity to antigenic components
present in the diet (Miller et al., 1984). Pierce et al. (2006) found that the inclusion of high levels of L in weaned pig diets
resulted in improved intestinal health mainly through a reduction in intestinal pH and increases in lactobacilli and VFA
concentrations, and reduction in coliforms.
The inclusion of the laminarin and fucoidan extract in the current experiment increased both ADG and G:F ratio of the
pigs throughout the duration of the experiment. This positive response to SWE may be due to a number of reasons. First,
the improved performance may be attributable to a reduction in E. coli numbers in the gut of the laminarin and fucoidan
extract-fed pigs. The inclusion of the SWE decreased the counts of E. coli in the faeces compared with non-SWE diets.
Similarly, McDonnell et al. (2010) demonstrated that the inclusion of a laminarin extract had an inhibitory effect on the
counts of E. coli in weaner pigs.
The decrease in E. coli numbers as a result of laminarin and fucoidan inclusion was particularly evident in pig health
between days 7–14. Pigs offered the high L diets showed more signs of impaired health (d 7–14) and were more contaminated
in faecal material (d 14–25) compared with pigs offered the low L diets. However when the SWE was supplemented there
were no differences in HS or CS between both L levels. The addition of the SWE to the high L diet also increased ADG from d
7to 14 compared with the high L diet only. It is not unreasonable to suggest that the high levels of L (250 g/kg) may act as a
substrate for E. coli (Huber and Hurlburt, 1984) to multiply in the gastrointestinal tract, and possible contribute to diarrhea
and impaired health. However, the inclusion of the SWE would have reduced E. coli numbers in this situation and improved
pig health.
Second, the improved performance may be attributable to the improved CTTAD of diet components. The supplementation
of SWE increased the CTTAD of OM, N and GE compared with the non-SWE diets. Little research has been conducted regarding
the effects of SWE on digestibility or gut physiology in pigs. The increased digestibility may be due to the increase in the
Lactobacilli spp. numbers observed with the SWE fed pigs. This increase in faecal Lactobacilli spp. and CTTAD was particularly
evident with high L inclusion. The SWE increased the numbers of lactic acid bacteria in the high L diets. These bacteria have an
ability to produce a wide range of cell-associated polysaccharide depolymerases and glycosidases which may aid in nutrient
digestion, and in particular will aid in the degradation of plant cell wall structural polysaccharides (Macfarlane et al., 1990). It
has also been reported that SWE may influence nutrient digestibility by maintaining the function and structure of the small
intestine, leading to an increased digestive capacity of the gut (Reilly et al., 2008). However, the digestibility measurements
were made over the entire gastrointestinal tract, something that may not be accurate because of microbial fermentation in
the large intestine. It has been shown that there is significant metabolism of dietary nutrients in the large intestine (Sauer
and Ozimek, 1986) and subsequently the apparent faecal digestibility method has been labeled as an inaccurate method for
the measurement, especially of protein and amino acid digestibility.
The inclusion of both SWE and high L diets increased the CTTAD of OM, N and GE compared with non-SWE and low L
diets. In general, the CTTAD values measured in the current study are high. However, the CTTAD values of N and GE are
0.821 and 0.839 respectively for the control diet (T1). These values are very similar to previous published work using sim-
ilar diets and chromium oxide as an indigestible marker (Pierce et al., 2005). In the current study, no additional material
(e.g. celite) was added to the feed to increase the level of AIA. However, chromium will make up part of the AIA con-
tent. McCarthy et al. (1974) examined the addition of celite (5 g/kg) to increase AIA levels in feed. They concluded that
there was no benefit to the addition of celite where AIA occurred in sufficient quantities. However, they also suggested
that the method becomes more reliable with an increase in AIA levels in the feed. In the present study, AIA level in the
diet was low (1.7–1.8 g/kg), at less than 2 g/kg, suggesting that a possible benefit could have been derived from celite
inclusion.
The results of this experiment indicate the beneficial effects of using high L levels in post-weaning pig diets. This may
be attributed to a number of reasons. First, this improved performance may be attributable to an observed increase in ADFI.
Second, L increased the counts of Lactobacilli spp. and decreased the E. coli numbers in the current experiment. Third, L can
be efficiently utilised by the newly weaned pig by maintaining the function and structure of the small intestine, leading to
an increased digestive capacity of the gut (Pierce et al., 2006). However, there is an economic implication in using high levels
of L in pig diets. The results of the current experiment would indicate that the inclusion of a SWE containing laminarin and

Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
Laminaria spp. on performance, digestibility of diet components and microbial populations in newly weaned pigs. Anim.
Feed Sci. Technol. (2010), doi:10.1016/j.anifeedsci.2010.03.004
G Model
ANIFEE-12232; No. of Pages 8 ARTICLE IN PRESS
J.V. O’Doherty et al. / Animal Feed Science and Technology xxx (2010) xxx–xxx 7

fucoidan could substitute for L in the absence of in-feed antibiotics. However, there was still a synergistic response to the
SWE at high L inclusion levels.
High L levels were found to improve the CTTAD of diet components in the current study. The improved N digestibility
observed indicates that the high L level determines conditions in the gut conducive to proteolytic activity. The microbial
fermentation of L to lactic acid creates an environment ideal for the growth of lactic acid bacteria (Pollmann et al., 1980). The
fermentation of the feed may affect the digestibility of the feed components. The increased NDF disappearance observed in
the current study may be the result of increased bacterial fermentation due to the increased L inclusion.
Despite the increase in the Lactobacilli population with high L diets, there was no dietary effect on VFA concentration or
profiles. Faecal VFA concentrations are the outcome of production, absorption and degradation within the gut. Faecal VFA
concentrations may not be a totally accurate way to demonstrate fermentation intensity in the large intestine.

5. Conclusion

The inclusion of either a high dietary concentration of L or a laminarin-fucoidan extract improved performance parameters
of weaner pigs, increased the CTTAD of diet components and reduced the counts of E. coli in the faeces. Further studies will
explore the potential of these diets to prevent the negative impact of pathogen challenges.

Acknowledgements

Funding for this research was provided under the National Development Plan through the Research Stimulus Fund
administered by the Irish Department of Agriculture, Fisheries and Food.

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Please cite this article in press as: O’Doherty, J.V., et al., The effects of lactose inclusion and seaweed extract derived from
Laminaria spp. on performance, digestibility of diet components and microbial populations in newly weaned pigs. Anim.
Feed Sci. Technol. (2010), doi:10.1016/j.anifeedsci.2010.03.004