Effect of weaning age and commingling after the nursery phase of pigs in a

wean-to-finish facility on growth, and humoral and behavioral indicators of well-being

M. E. Davis, S. C. Sears, J. K. Apple, C. V. Maxwell and Z. B. Johnson
J ANIM SCI 2006, 84:743-756.

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Effect of weaning age and commingling after the nursery phase of pigs
in a wean-to-finish facility on growth, and humoral and behavioral
indicators of well-being1,2
M. E. Davis,3 S. C. Sears, J. K. Apple,4 C. V. Maxwell, and Z. B. Johnson
Department of Animal Science, University of Arkansas, Fayetteville 72701

ABSTRACT: Pigs from one farrowing group in which

gilts were bred to farrow pigs that would be either 14
or 21 d of age at weaning, were divided into older and
younger age groups (108 pigs per group) and penned
12 pigs per pen in a wean-to-finish facility. At the end
of the nursery phase, half the pigs in each age group
were removed, rerandomized, and commingled for the
finishing phase. The other half remained in their original pens. Pigs were fed common Phase 1 (d 0 to 14) and
Phase 2 (d 14 to 35) nursery diets, and a common 4phase program diet during growing/finishing, with
transitions at 45, 68, and 90 kg of BW. The study ended
when the lightest weight block averaged 107 kg. Blood
was obtained on d 0, 2, 10, 27, 37, 44, and 65 after
weaning to determine leukocyte concentrations. In addition, behavior was monitored during the nursery period at weaning (d 0), on d 7, 14, and 27 after weaning,
and during the growing/finishing phase on d 35 (after
commingling following the nursery phase), 38, 44, and
65 after weaning. Older pigs were heavier (P < 0.001)
throughout the nursery period, and the BW difference
between younger and older pigs increased from 2 to 6.5

kg at the start and end of the nursery period, respectively. Older pigs had a greater concentration of white
blood cells (P < 0.05) and lymphocytes (P < 0.10) on d
0, 2, and 10 after weaning than younger pigs. Younger
pigs spent less (P < 0.05) time resting on the day of
weaning, and more (P < 0.05) time active during the
overall nursery phase. During Phase 3 and in the overall finishing phase, younger pigs had greater (P < 0.01)
ADG and G:F than older pigs. Moreover, during Phase
3, ADFI (as fed) decreased (P < 0.05) when older pigs
were commingled compared with older pigs that were
not commingled. There was no difference in ADFI of
younger pigs, regardless of commingling (interaction;
P < 0.10). Results of this study indicate that weaning
age affects growth performance in a wean-to-finish facility, as well as behavioral and immunological responses to weaning and commingling after the nursery
phase. Management strategies should be further explored to optimize these benefits without the detrimental effects on health observed during the nursery period
in this study.

Key words: behavior, growth, immune response, mixing, swine, weaning

2006 American Society of Animal Science. All rights reserved.

INTRODUCTION

J. Anim. Sci. 2006. 84:743756

early weaning systems (Maxwell and Carter, 2000). Although there are no legislative restrictions on weaning
age in the United States, the practice of weaning before
28 d of age is prohibited in Europe (ECCD, 2001). Early
weaning and removal of pigs to a second isolated site for
rearing can decrease the potential for disease transfer
from the dam (Harris, 1988). Pigs reared in isolation
after weaning have decreased immunological stress
(Johnson, 1997), resulting in substantial improvements
in growth and efficiency of feed use compared with those
reared in conventional farrow-to-finish systems (Williams et al., 1997). Similarly, commingling pigs following the nursery phase is a common management practice in the swine industry, and it imposes an additional
stress on the young pig. The advent of wean-to-finish
facilities has potentially alleviated commingling after
the nursery phase; however, the wasted space of placing

Pigs born and raised in conventional swine production systems are routinely weaned as early as 17 d of
age and as early as 12 to 14 d of age in off-site segregated

Research funded by a grant from the National Pork Board, Des

Moines, IA.
2
The authors acknowledge the following Univ. Arkansas staff and
students for their assistance with this project: T. Wistuba, A. Hays,
B. Hinson, C. Bradley, C. Scarborough, B. Scarborough, M. Curran,
R. Harmon, and M. Dirain.
3
Current address: Agtech Products, Inc. W227 N752 Westmound
Dr., Waukesha, WI 53186 (phone: 262-521-1717; fax: 262-521-2442;
e-mail: edavis@agtechproducts.com).
4
Corresponding author: japple@uark.edu
Received September 27, 2004.
Accepted August 10, 2005.

743
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744

Davis et al.

weanling pigs in pens with space allowances for market

hogs has led to the practice of double-stocking pens at
weaning and later moving half the pigs to another pen,
which introduces a commingling stress at an older age
(Wolter et al., 2002). Results supporting wean-to-finish
production systems with regard to piglet well-being are
lacking, and the effect of these systems on pig health
has not been evaluated. Therefore, the objective of this
study was to determine the effect of age at weaning
and of rearing pigs in the same pen in an all-inall-out
wean-to-finish facility vs. commingling pigs after the
nursery phase on growth performance and on humoral
and behavioral indicators of well-being.

MATERIALS AND METHODS

Allocation of Animals
Pigs from one farrowing of 30 litters (DeKalb Line
348 gilts mated to EB sires; Monsanto Choice Genetics,
St. Louis, MO) over a 10-d period were weaned when
the average age was approximately 17 0.3 d. Gilts
were bred to farrow pigs that would average 14 and 21
d of age when weaned on the same day. Gilts were
farrowed in crates in an environmentally controlled farrowing room. Immediately after farrowing, pigs were
ear-notched for individual identification, administered
iron dextran, and tails were docked. Pigs were crossfostered if necessary to equalize litters within their respective age groups. Pigs were divided into 2 age groups
of equal number representing the older (21 d of age)
and younger (14 d of age) groups of pigs (108 pigs in
each group). Pigs within each age group were sorted
into 3 blocks based on BW and gender (36 pigs per
block). Pigs from each gender and BW category within
age group were then allocated randomly to pens within
block in a double-stocked wean-to-finish facility (12 pigs
per pen). This provided a total of 9 replications of each
age group for the nursery study. At the completion of
the nursery phase, half the pigs in each age category
were removed from the double-stocked pens, rerandomized based on gender and BW, and commingled for the
growing-finishing component of the study. Half the pigs
remained in the same wean-to-finish pens. This arrangement of treatments permitted evaluation of the
effects of weaning age in pigs double-stocked in a weanto-finish facility, as well as effects of postnursery commingling on well-being and performance.

Housing and Environment

Pigs were housed in a wean-to-finish facility in totally
slatted pens (1.52 m 3.05 m; 0.39 m2/pig) equipped
with radiant heaters, a 2-hole nursery feeder, and a
wean-to-finish cup waterer. Ambient room temperature
was maintained at 25.6C. In addition, a radiant heater
oriented above rubber mats provided supplemental
heat to a 1.8-m diameter area covering 2 pens per heater
during the nursery phase. Wean-to-finish barns were

Table 1. Composition of Phase 1 (d 0 to 14) and Phase 2

(d 10 to 35) diets fed to pigs during the nursery phase
(%, as-fed basis)
Item
Yellow corn
Steam rolled oats
Dried whey
Lactose
Soy protein concentrate
Soybean meal, 48% CP
Spray-dried plasma protein
Select menhaden fish meal
Soybean oil
Fat
Ethoxyquin
Lysine HCl
Threonine
Methionine
Isoleucine
Antibiotic premix1
Mineral premix2
Vitamin premix3
Zinc oxide
Copper sulfate
Dicalcium phosphate
Calcium carbonate
Salt

Phase 1

Phase 2

27.82
12.00
20.50

8.00
11.00
6.00
8.00
4.00

46.51
5.00
8.00
5.00

20.00
1.75
7.00

4.00
0.03
0.17
0.07
0.07

0.25
0.15
0.25
0.25
0.07
0.65
0.38
0.40

0.03
0.15
0.08
0.10
0.02
0.10
0.15
0.25

1.12
0.38
0.30

1
Provided 0.15 g of neomycin as neomycin sulfate and 0.11 g of
oxytetracycline/kg of feed during Phase 1, and 0.05 g of carbadox/kg
of feed during Phase 2.
2
Supplied 0.30 mg of Se as sodium selenite, 40 mg of Mn as manganous oxide, 165 mg of Zn as zinc oxide, 165 mg of Fe as ferrous sulfate,
17 mg of Cu as copper sulfate, and 0.30 mg of I as calcium iodate
per kilogram of feed.
3
Supplied 11,023 IU of vitamin A, 1,653 IU of vitamin D3 as Dactivated animal sterol, 44 IU of vitamin E, 4.4 mg of vitamin K as
menadione sodium bisulfite complex, 33 mg of pantothenic acid as
D-calcium pantothenate, 55 mg niacin, 10 mg of riboflavin, and 44
g of vitamin B12 per kilogram of feed.

curtain-sided and naturally ventilated, and contained

a pit fan. Temperature was decreased gradually over
the course of the nursery phase according to established
procedures (no more than 1.5C/wk). The temperature
was maintained at approximately 21.1C throughout
the starter and grower I phases, and decreased to 20C
during the grower II and finisher phases. Water and
feed were available ad libitum throughout the study.

Experimental Management
On the day of weaning, pigs were moved from the
farrowing room and distributed to their assigned pen.
Pigs were exposed to natural lighting, with supplemental electrical lighting provided only when animals were
being observed. Pigs were offered ad libitum access to
a Phase I nursery diet for the period of 0 to 14 d and
a Phase II diet for the period of 14 to 35 d (Table 1).
On the day of completion of the nursery phase of the
study, pigs were started on the growing-finishing study.
Pigs were fed a 4-phase diet with transition from starter
to grower I, grower I to grower II, and grower II to
finisher occurring when the mean BW of each block

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Weaning age and commingling in pigs

Table 2. Composition of Phase 1, 2, 3, and 4 diets fed to pigs during the growing/finishing
period (%, as-fed basis)
Item

Phase 1

Phase 2

Phase 3

Phase 4

Yellow corn
Soybean meal, 48% CP
Fat
Ethoxyquin
Lysine
Threonine
Dicalcium phosphate
Calcium carbonate
Mineral premix1
Vitamin premix2
Antibiotic premix3
Salt

71.865
23.00
2.30
0.03
0.15
0.02
0.94
0.77
0.15
0.15
0.125
0.50

75.10
18.10
4.00
0.03
0.15
0.02
0.95
0.80
0.15
0.15
0.05
0.50

80.00
13.50
4.00
0.03
0.15
0.02
0.75
0.70
0.15
0.15
0.05
0.50

83.19
10.50
4.00
0.03
0.15

0.755
0.60
0.125
0.125
0.025
0.50

1
Supplied 0.30 mg of Se as sodium selenite, 40 mg of Mn as manganous oxide, 165 mg of Zn as zinc oxide,
165 mg of Fe as ferrous sulfate, 17 mg of Cu as copper sulfate, and 0.30 mg of I as calcium iodate per
kilogram of feed during Phase 1, 2, and 3; and 0.25 mg of Se as sodium selenite, 33 mg of Mn as manganous
oxide, 138 mg of Zn as zinc oxide, 138 mg of Fe as ferrous sulfate, 14 mg of Cu as copper sulfate, and 0.25
mg of I as calcium iodate per kilogram of feed during Phase 4.
2
Supplied 6,614 IU of vitamin A, 992 IU of vitamin D3, 26 IU of vitamin E, 2.6 mg of vitamin K, 20 mg
of pantothenic acid, 33 mg niacin, 6.0 mg of riboflavin, and 26 g of vitamin B12 per kilogram of feed during
Phase 1, 2, and 3; and 5,512 IU of vitamin A, 827 IU of vitamin D3, 22 IU of vitamin E, 2.2 mg of vitamin
K, 17 mg of pantothenic acid, 28 mg niacin, 5 mg of riboflavin, and 22 g of vitamin B12 per kilogram of
feed during Phase 4.
3
Provided 0.11 g of tylosin/kg of feed during Phase 1, 0.04 g of tylosin/kg of feed during Phases 2 and 3,
and 0.02 g of tylosin/kg of feed during Phase 4.

reached approximately 45, 68, and 90 kg, respectively

(Table 2). All diets met or exceeded NRC (1998) requirements for all nutrients, and were formulated to simulate diets typical of those used in the swine industry.
The study ended when the lightest block reached an
average BW of 107 kg. The experiment was carried out
in accordance with the Animal Care Protocol No. 01015
for swine experiments issued by the University of Arkansas Institutional Animal Care and Use Committee.

Behavioral Measures
Behavioral and postural measures were recorded on
the day of weaning (0) and on d 7, 14, 27, and 35 (after
commingling at the end of nursery phase), as well as
d 38, 44, and 65 of the growing-finishing period. Before
viewing, pigs were numbered (with a permanent
marker) 1 through 12 for behavioral observations during the nursery phase of the study (d 0, 7, 14, and 27
after weaning) and 1 through 6 after commingling (d
35, 38, 44, and 65 after weaning). Three pigs from each
of the observed pens were selected randomly for viewing
during the nursery phase, and another 3 were randomly
selected for viewing following rerandomization after
commingling. The following behaviors were recorded for
each selected pig: 1) acts of aggression (biting, pushing,
head-thrusting, and chasing directed toward penmates); 2) feeding; 3) drinking; 4) resting; 5) active
(standing or locomotion); and 6) belly-nosing (Table 3).
Four pens per treatment were monitored with
mounted camera (installed 2.3 m above floor level) surveillance equipment (Everplex 8CDX; Everfocus Electronics Corp., Pasadena, CA) for 24 h to observe initial

behaviors following weaning. Lenses of the eight cameras were 0.5 cm in diameter. Time-lapse videos (RCA
standard grade, Socorro, TX) were recorded with a timelapse videocassette recorder (Sanyo Electric Co., Ltd,
Moriguchi City, Osaka, Japan) in 12-h recording speed
mode (7/60-s recording interval), and were viewed later
in two 1-h increments (0900 to 1000 and 1400 to 1500).
Duration of time spent by each pig engaged in these
behaviors was recorded from 0900 to 1000 during the
morning and 1400 to 1500 during the afternoon, and
percentages of time were calculated for each pig based
on the total 2-h observation. Percentages of time for
each behavior for each pig were averaged for a mean
percentage of time engaged in each behavior for the
total pen. Behavioral data were analyzed using these
pen means.

Table 3. Observed behaviors and their definitions

Behavior
Resting
Active

Feeding1
Drinking1
Aggression
Belly-nosing

Definition
Lateral and sternal recumbancy
Standing or moving activity that is not defined by
other behavioral categories evaluated
Head positioned in feeder
Snout in contact with nipple waterer
Biting, pushing, head-thrusting, and chasing
directed toward penmates (McGlone, 1986)
Repetitive rooting movement with the snout on
another piglet (Fraser, 1978)

1
Definitions for observed behaviors excerpted from Gardner et al.
(2001).

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746

Davis et al.

Physiological Measures
A 5-mL blood sample was obtained from each pig via
venipuncture on d 0, 2, 7, 13, 27, 37, 42, and 56. Blood
samples were kept on ice for transport approximately
24 km from the University of Arkansas Swine Research
Unit to laboratory facilities at the University of Arkansas Animal Science Department. Blood measurements
of interest (differential leukocyte proportions and neutrophil:lymphocyte ratio) were obtained by analysis on
an automated hematology analyzer (Cell-Dyn 3500SL
System; Abbott, Abbott Park, IL) calibrated for porcine
blood. An additional 15-mL blood sample was obtained
on d 8 of the nursery period from 4 pigs randomly selected from 3 pens from each age group, and peripheral
blood mononuclear cells were isolated to determine lymphocyte blastogenic response, interferon-gamma (IFN) production, and macrophage phagocytosis. This procedure was repeated on d 10 and 12 for the remaining
6 pens in each age group. Data from the 3 sampling
days were pooled and designated as d 10 after weaning.
The same process was repeated postnursery for all pens,
with 2 randomly chosen pigs per pen sampled, and
pooled data were designated as d 44 after weaning.
Lymphocyte Blastogenesis. In vitro cellular proliferative response was measured using a lymphocyte
blastogenesis assay adapted from the methods of
Blecha et al. (1983). Briefly, peripheral blood mononuclear cells were isolated by gradient centrifugation using a Ficoll gradient (Histopaque 1077, density = 1.077
g/mL; Sigma Chemical Co., St. Louis, MO). Remaining
erythrocytes were lysed by adding 1 mL of sterile water
to the isolated cell pellet for 20 s, and isotonicity was
restored by the addition of Roswell Park Memorial Institute medium (RPMI; Sigma). Cells were resuspended in RPMI at 2 106 cells/mL, and plated in
triplicate in 96-well round bottom plates in 100-L aliquots. Phytohemagglutinin (PHA; Sigma), pokeweed
mitogen (PWM; Sigma), and concanavalin A (ConA;
Sigma) were administered in 50-L aliquots to each
well at a concentration of 40, 15, and 25 g/mL, respectively, to stimulate lymphocyte proliferation, whereas
wells containing cells unstimulated with mitogen were
administered 50 L of medium. Concentrations chosen
for each respective mitogen elicited the maximum proliferative response during preliminary trials. Cells were
incubated with the mitogens for 48 h at 39.2C and 5%
CO2. The incubation temperature for all assays conducted in this study was set at 39.2C rather than 37C,
to facilitate monocyte/macrophage activation under
conditions simulating pig body temperature (Natale
and McCullough, 1998). Following the 48-h incubation,
1 Ci of tritiated thymidine in 50 L of RPMI was
added to each well, and the cells were incubated for an
additional 18 h. Cells were harvested on glass fiber
mats and the radioactivity was measured on a liquid
scintillation analyzer (TRI-CARB 2200CA; Packard Instrument Co., Downers Grove, IL). Incubation, labeling
with tritiated thymidine, and cell harvesting followed
procedures outlined by van Heugten et al. (1994).

Interferon- Production. Peripheral blood mononuclear cells were isolated as described for the lymphocyte
blastogenesis assay. Cells were dispensed in duplicate
at a concentration of 2 106 cells/mL in 500-L aliquots
into a 24-well plate. An additional 500 L of RPMI was
added to one of the duplicated wells, and 500 L of
RPMI containing 10 g/mL of ConA was added to the
second well, for unstimulated and stimulated cultures,
respectively. Cell cultures were incubated for 48 h at
39.2C and 5% CO2. Following the 48-h incubation, supernatant fractions were collected from each well and
stored at 20C until analysis for IFN- could be conducted. Interferon- production was determined using
a swine IFN- ELISA kit following the instructions provided by the manufacturer (Biosource International
Inc., Camarillo, CA). The intraassay CV was 17.5%.
Monocyte/Macrophage Phagocytosis. The method
used to measure monocyte/macrophage phagocytic ability of peripheral blood monocytes was adapted from the
methods of Nibbering et al. (1987), Heggen et al. (1998),
and Monteleone et al. (1999). Peripheral blood mononuclear cells from blood were isolated as described for
the lymphocyte blastogenesis assay. Cell suspensions
isolated from blood were diluted to approximately 5
106 cells/mL in LM Hahn (Leibovitzs L-15/McCoys
Hahn media; Atlanta Biologicals, Lawrenceville, GA)
medium. A glass coverslip was added to each well of a
6-well plate, and 2 mL of cell suspension containing
monocytes/macrophages was added to each well in duplicate for each sample. Each coverslip was completely
covered by cell suspension. Cells were incubated for 5
h at 39.2C and 5% CO2. Following the 5-h incubation,
medium from each well was removed and replaced by
2 mL of fresh LM Hahn medium warmed to 39.2C.
Cell cultures were incubated for an additional 19 h.
Following the 24-h total incubation, plates were removed from the incubator, excess medium was removed
from each well, and 2 mL of a 4% sheep red blood cell
(SRBC) suspension were added to each well. Plates
were incubated with SRBC for 2 h, after which coverslips were removed, and nonadherent cells and excess
SRBC were washed from the coverslip by rinsing with
LM Hahn medium warmed to 39.2C. Coverslips were
then stained using the Hema 3 cell staining kit (Fisher
Scientific, Pittsburgh, PA), and the percentage of
phagocytic monocytes/macrophages and the average
number of SRBC consumed by each phagocytic monocyte/macrophage were determined.
Performance Measures
Pigs were weighed as each block reached the projected BW for each phase. Pig BW was averaged by pen
and ADG was calculated. Feed disappearance from each
pen self-feeder was measured and ADFI (as fed) was
calculated for each pen as the difference between feed
added and feed weighed back for each period, and G:F
ratios for each period were determined. Adjusted days
to a common BW (104 kg) were calculated for each pig

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Weaning age and commingling in pigs

Table 4. Behavioral data (presented as percentage of time engaged in each respective

behavior) collected on each of four observation days during the nursery phase from pigs
weaned at either 14 or 21 d of age1
Observation days
Weaning (d 0)
Age at weaning, d:

d 7 after weaning

d 14 after weaning

d 27 after weaning

14

21

14

21

14

21

14

21

SE

26.2d
32.8
1.9
1.8
3.1

45.5c
32.0
3.2
0.0
3.3

32.5
35.4
1.3
9.1
3.7

36.6
29.6
1.5
8.9
1.8

34.2
25.8
1.5d
12.6
1.8

37.9
21.4
4.5c
11.5
1.0

48.3
27.7
1.8
13.3
1.9

63.2
13.5
1.1
7.6
2.0

6.4
5.2
0.80
2.9
1.6

Observation, %
Resting
Active
Drinking
Feeding
Aggression

c,d
Within each observation day and row, least squares means that do not have a common superscript
letter differ, P < 0.05.
1
Data are expressed as the least squares means of 4 pens per age group on each observation day, with
each pen value encompassing the mean of 3 pigs per pen. Because percentages of time for each behavior
were calculated for individual pigs and averaged for each pen, percentages of time for each age group do
not total to 100%.
2
Refer to Table 3 for descriptions of behavioral observations.

using the following formula: (age of pig at the time final

BW was obtained + [desired final BW actual BW]{[age
38]/actual BW}; NSIF, 2002).

Experimental Design
Performance and behavioral data were analyzed as
a randomized complete block design in a 2 2 factorial
arrangement of treatments, with 2 weaning ages and
2 postnursery management systems, pen as the experimental unit, and day of observation as a repeated measure using PROC MIXED of SAS (SAS Inst., Inc., Cary,
NC). When significant day treatment interactions
were observed, ANOVA for each day was performed
using PROC GLM of SAS. Mortality rate was analyzed
by 2 analysis in SAS.

RESULTS
Behavioral Measures
Pigs weaned at 21 d of age spent a greater (P < 0.05)
percentage of time resting on the day of weaning (d 0

postweaning) than pigs weaned at 14 d of age (Table

4). Although the percentage of time spent active did
not differ between pigs of different weaning ages at any
observation day during the nursery period, pigs weaned
at 14 d of age spent a greater (P = 0.05) proportion of
time active during the overall nursery phase than did
pigs weaned at 21 d of age (Table 5). There was no
difference in the percentage of time that pigs weaned
at either age were engaged in aggressive behavior on
any of the sampling days, but the frequency of times
aggressive behaviors were observed was greater (P <
0.05) at weaning (d 0) than on any other observation
day during the nursery period (Figure 1). Very few incidents of belly-nosing by pigs were observed in this
study, and number and duration of belly-nosing incidents did not differ between older and younger pigs.
During the growing/finishing period, the effect of
weaning age and postnursery commingling on feeding
behavior depended on the day of observation (weaning
age commingling date interaction, P < 0.05; Figure
2). During the growing/finishing period, pigs that were
weaned at 14 d of age and remained unmixed after the

Table 5. Summary of behavioral data (presented as percentage of time engaged in each

respective behavior) collected during the overall nursery phase from pigs weaned at either
14 or 21 d of age1
Age at weaning, d
2

Observation, %
Resting
Active
Drinking
Feeding
Aggression

14

21

SE

P-value

36.1
30.1
1.7
11.2
2.9

46.1
22.6
2.5
9.1
2.1

4.00
2.23
0.47
1.69
0.61

0.130
0.054
0.255
0.399
0.387

1
Data are expressed as the least squares means of four pens per age group, with each pen value encompassing the mean of 3 pigs per pen, and averaged across the 4 observation days. Because percentages of
time for each behavior were calculated for individual pigs and averaged for each pen, percentages of time
for each age group do not total to 100%.
2
Refer to Table 3 for description of behavioral observations.

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748

Davis et al.

Figure 1. Frequency of aggressive behaviors observed

on each day of the nursery phase. Bars with an asterisk
differ from those without asterisks, P < 0.05.
nursery period spent a greater (P < 0.05) percentage of
time engaged in feeding activity on d 35 after weaning
than 21-d-old pigs that were unmixed or mixed, regardless of weaning age. The percentage of time spent feeding did not differ among pigs of either weaning age
or postnursery mixing treatment on d 38 or 44 after
weaning; however, on d 65 after weaning, pigs that
were weaned at 21 d of age and mixed and pigs weaned
at 14 d of age and unmixed spent a greater (P < 0.05)
proportion of time engaged in feeding behavior than
pigs in the other two treatments.

Physiological Measures
On the day of weaning (d 0), pigs weaned at 21 d of
age had a greater concentration of white blood cells

Figure 2. Percentage of time spent engaged in feeding

behavior during the growing/finishing phase by pigs
weaned at either 14 or 21 d of age and either commingled
(mixed) following the nursery or remained in original
pens (weaning age mixing date interaction; P < 0.05).
Within a specific day after weaning, bars with an asterisk
differ from those without asterisks, P < 0.05.

(WBC) than pigs weaned at 14 d of age (Table 6). Specifically, pigs weaned at 21 d of age had greater (P <
0.05) concentrations of neutrophils and eosinophils, and
eosinophils occupied a greater (P < 0.01) percentage
of the total WBC concentration in older pigs than in
younger pigs. In addition, pigs weaned at 21 d of age
tended to have a greater (P = 0.09) concentration of
lymphocytes than did pigs weaned at 14 d of age. On
d 2 after weaning, pigs weaned at 21 d of age still had
a greater (P < 0.01) concentration of WBC than pigs
weaned at 14 d of age (Table 7). Specifically, older pigs
had greater (P < 0.05) concentrations of neutrophils and
lymphocytes than younger pigs, although there were
no differences in these cell populations as a percentage
of total WBC. Ten days after weaning, pigs weaned at
21 d of age had a greater (P = 0.05) concentration of
WBC than pigs weaned at 14 d of age (Table 8). This
was a result of older pigs having a greater (P = 0.01)
concentration of lymphocytes than younger pigs. On d
27 after weaning, total WBC concentrations did not
differ between older and younger pigs; however, eosinophil concentrations and eosinophils as a percentage of
total WBC were greater (P < 0.01) in older pigs than
in younger pigs (Table 9). Conversely, the concentration
(P = 0.06) and percentage (P = 0.07) of monocytes tended
to be greater in pigs weaned at 14 d of age compared
with those weaned at 21 d of age.
During the growing/finishing period, WBC concentrations on d 37 after weaning were increased (P < 0.05)
when pigs weaned at 21 d of age were mixed and resorted compared with those remaining in their original
pens, whereas there was no difference (P = 0.90) in
WBC concentrations as a result of mixing when pigs
were weaned at 14 d of age (interaction, P = 0.06; Table
10). The increase in WBC concentrations of older pigs
that were mixed was a result of an increase in lymphocyte concentrations (interaction, P = 0.07). Regardless
of whether the pigs were mixed and re-sorted or remained in their original pens, eosinophil concentration
(0.14 0.01 vs. 0.09 0.01) and eosinophils as a percentage of total WBC (2.10 0.17 vs. 1.23 0.17) were
greater (P < 0.01) in older pigs than in pigs weaned at
14 d of age. Moreover, on d 44 after weaning (Table
11), eosinophil concentration (0.08 0.01 vs. 0.15 0.01)
and eosinophils as a percentage of total WBC (1.77
0.11 vs. 0.97 0.11) were greater (P < 0.01) in older
pigs than in pigs weaned at 14 d of age. At 65 d after
weaning (Table 12), however, WBC concentration (5.81
0.51 vs. 4.43 0.51) tended to be greater (P < 0.07)
when pigs were weaned at 14 d of age compared with
21 d of age. Moreover, neutrophils as a percentage of
total WBC (41.57 4.23 vs. 29.13 4.23) were greater
(P < 0.05) in younger than in older pigs, although older
pigs had a greater (P < 0.05) percentage of lymphocytes
(61.84 3.67 vs. 51.34 3.67) than younger pigs. There
was no effect of weaning age or commingling after the
nursery phase on the neutrophil:lymphocyte ratio on
any of the sampling days (Tables 6 through 12).

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749

Weaning age and commingling in pigs

Table 6. Differential leukocyte concentrations and leukocytes as a percentage of total

white blood cells (WBC) on the day of weaning (d 0) from pigs weaned at 14 or 21 d of age1
Weaning age, d
Item

14

21

SE

P-value

WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils

2.83
0.94
1.56
0.07
0.03

3.51
1.40
1.82
0.10
0.06

0.17
0.11
0.11
0.03
0.01

0.012
0.012
0.097
0.457
0.002

33.54
55.60
1.95
1.08
0.63

35.94
54.37
3.19
1.59
0.76

2.36
1.16
0.81
0.11
0.06

0.480
0.466
0.290
0.004
0.150

WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

1
Twelve pigs per pen were sampled, and pen means were determined. Data are the least squares means
of 9 pens per treatment (age group).

Table 7. Differential leukocyte concentrations and leukocytes as a percentage of total

white blood cells (WBC) on d 2 after weaning from pigs weaned at 14 or 21 d of age1
Weaning age, d
Item

14

21

SE

P-value

WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils

2.80
1.15
1.44
0.06
0.03

3.92
1.68
2.03
0.08
0.04

0.19
0.16
0.14
0.03
0.01

0.001
0.036
0.011
0.691
0.596

40.34
52.13
2.07
1.13
0.86

42.13
51.47
1.94
1.00
1.11

2.94
2.03
0.74
0.13
0.11

0.672
0.820
0.896
0.526
0.115

WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

1
Twelve pigs per pen were sampled, and pen means were determined. Data are the least squares means
of 9 pens per treatment (age group).

Table 8. Differential leukocyte concentrations and leukocytes as a percentage of total

white blood cells (WBC) on d 10 after weaning from pigs weaned at 14 or 21 d of age1
Weaning age, d
Item

14

21

SE

P-value

WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils

5.56
1.90
3.03
0.04
0.03

6.53
1.47
4.24
0.02
0.03

0.33
0.40
0.32
0.02
0.01

0.057
0.455
0.016
0.335
0.661

32.85
54.86
0.83
0.58
1.13

21.94
64.29
0.33
0.56
0.67

5.82
4.03
0.26
0.11
0.21

0.204
0.117
0.194
0.917
0.137

WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

1
Twelve pigs per pen were sampled, and pen means were determined. Data are the least squares means
of 9 pens per treatment (age group).

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Davis et al.

Table 9. Differential leukocyte concentrations and leukocytes as a percentage of total

white blood cells (WBC) on d 27 after weaning from pigs weaned at 14 or 21 d of age1
Weaning age, d
Item

14

21

SE

P-value

WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils

8.92
4.06
4.46
0.08
0.10

8.82
3.76
4.71
0.04
0.20

0.29
0.23
0.30
0.02
0.02

0.821
0.356
0.560
0.063
0.001

45.37
50.14
0.93
1.18
0.98

42.59
53.16
0.47
2.26
0.93

2.61
2.07
0.17
0.16
0.08

0.463
0.318
0.077
0.001
0.628

WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

1
Six randomly selected pigs from each pen of 12 pigs were sampled, and pen means were determined.
Data are the least squares means of 9 pens per treatment (age group).

Table 10. Effect of mixing and age at weaning on peripheral blood leukocyte concentrations
and proportions of pigs on d 37 after weaning1

Item
WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils
WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

Weaning age: 14 d

Weaning age: 21 d

Mixed

Mixed

7.81
2.94
4.37bc
0.35
0.10
37.97
57.13
2.52
1.44
0.78

Unmixed
bc

7.70
2.39
4.80b
0.05
0.07
32.94
60.91
0.64
1.02
0.62

8.56
3.35
4.55b
0.36
0.15
37.64
55.09
2.41
1.94
0.81

Unmixed

P-value
SE

Age

Mixed

Interaction

5.95
2.42
3.13c
0.03
0.14

0.65
0.40
0.49
0.23
0.02

0.45
0.59
0.14
0.99
<0.01

0.04
0.07
0.32
0.17
0.19

0.06
0.63
0.07
0.94
0.68

38.82
53.36
0.41
2.26
0.86

4.31
3.39
1.43
0.24
0.11

0.52
0.17
0.90
<0.01
0.21

0.66
0.76
0.19
0.83
0.61

0.48
0.42
0.97
0.14
0.34

Within a row, least squares means that do not have a common superscript letter differ, P < 0.05.
Three randomly selected pigs were sampled from each of 36 pens, and the values from each pig were
averaged to determine pen means. Data are the least squares means of 9 pens per treatment.
b,c
1

Table 11. Effect of mixing and age at weaning on peripheral blood leukocyte concentrations
and proportions of pigs on d 44 after weaning1

Item
WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils
WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

Weaning age: 14 d

Weaning age: 21 d

P-value

Mixed

Unmixed

Mixed

Unmixed

SE

Age

Mixed

Interaction

9.25
4.13
4.67
0.14
0.08

8.42
3.72
4.30
0.03
0.08

9.35
4.03
4.84
0.03
0.13

8.38
3.83
3.92
0.08
0.17

0.61
0.46
0.40
0.07
0.01

0.96
0.99
0.79
0.68
0.01

0.15
0.51
0.12
0.65
0.23

0.92
0.82
0.49
0.23
0.16

45.14
49.45
1.39
0.92
1.09

43.15
51.82
0.36
1.02
1.08

42.00
51.86
0.36
1.52
1.02

43.19
48.76
1.06
2.02
1.05

3.79
2.87
0.68
0.16
0.14

0.69
0.91
0.81
0.01
0.70

0.92
0.90
0.81
0.08
0.91

0.68
0.35
0.21
0.23
0.90

1
Three randomly selected pigs were sampled from each of 36 pens, and the values from each pig were
averaged to determine pen means. Data are the least squares means of 9 pens per treatment.

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751

Weaning age and commingling in pigs

Table 12. Effect of mixing and age at weaning on peripheral blood leukocyte concentrations
and proportions of pigs on d 65 after weaning1
Weaning age: 14 d

Weaning age: 21 d

Mixed

Unmixed

Mixed

Unmixed

SE

Age

Mixed

Interaction

6.23
2.76
3.04
0.02
0.15

5.40
2.34
2.83
0.02
0.15

5.11
2.27
2.53
0.03
0.18

3.75
1.08
2.40
0.02
0.12

0.73
0.63
0.32
0.01
0.03

0.07
0.18
0.16
0.88
0.99

0.14
0.21
0.60
0.28
0.33

0.71
0.55
0.90
0.51
0.38

40.97
51.23
0.39
2.44
0.93

42.17
51.45
0.36
2.97
0.91

31.33
61.88
0.45
3.30
1.07

26.93
65.79
0.43
3.10
0.72

5.99
5.18
0.07
0.58
0.21

0.05
0.02
0.38
0.40
0.90

0.79
0.69
0.78
0.78
0.40

0.64
0.72
0.95
0.53
0.43

Item
WBC, thousands/mL
Neutrophils
Lymphocytes
Monocytes
Eosinophils
WBC, %
Neutrophils
Lymphocytes
Monocytes
Eosinophils
Neutrophil:Lymphocyte ratio

P-value

1
Three randomly selected pigs were sampled from each of 36 pens, and the values from each pig were
averaged to determine pen means. Data are the least squares means of 9 pens per treatment.

There was no effect of age at weaning or commingling

on macrophage phagocytic capacity or IFN- production
during the nursery or growing/finishing periods (Table
13 and 14, respectively). During the nursery period,
pigs weaned at 14 d of age tended to have a greater
(P = 0.10) lymphocyte proliferation response when peripheral blood mononuclear cells were stimulated with
PWM or ConA (Table 13). Lymphocytes isolated during
the growing/finishing period from pigs weaned at 14 d
of age had greater (P < 0.05) proliferation (1,576 185
vs. 933 185) in unstimulated cultures compared with
older pigs (Table 14), and proliferation in response to
PWM increased (P < 0.05) in lymphocytes isolated dur-

ing the growing/finishing period when pigs weaned at

14 d of age were mixed compared with pigs weaned
at 14 d of age that remained in their original pens.
Conversely, proliferation decreased (P < 0.05) when
older pigs were mixed compared with older pigs that
remained in their original pens (interaction; P < 0.01).

Performance Measures
During the nursery phase of the experiment, pigs
weaned at 21 d of age had greater (P < 0.01) ADG
and ADFI during Phase 1, Phase 2, and in the overall
nursery period than pigs weaned at 14 d of age (Table

Table 13. Effect of age at weaning on macrophage phagocytic capacity, lymphocyte proliferation response, and interferon- production of pigs during the nursery phase1
Weaning age, d
Item

14

Macrophage phagocytosis2
Phagocytic macrophages, %
SRBC consumed
Lymphocyte proliferation,3 cpm
Unstimulated
PHA, 40 g/mL
PWM, 15 g/mL
ConA, 25 g/mL
Interferon-4

7.16
1.40

21
10.71
1.51

SE
1.76
0.11

P-value
0.173
0.491

2,812
130,135
40,413
130,578

518
105,796
28,915
81,635

1,421
13,557
4,628
19,834

0.271
0.222
0.100
0.100

896

828

178

0.792

1
Four randomly selected pigs from each of 18 total pens were sampled to measure macrophage phagocytic
capacity, lymphocyte proliferation response, and interferon- production, and pen means were determined.
Data are the least squares means of 9 pens per treatment (age group).
2
Macrophage phagocytic capacity was determined by evaluating the percentage of macrophages that were
phagocytic, meaning the macrophage consumed at least one sheep red blood cell (SRBC), and the average
number of SRBC consumed by each phagocytic macrophage.
3
Lymphocyte proliferation was measured as counts per minute (cpm) in unstimulated cell cultures and
cultures stimulated with phytohemagglutinin (PHA), pokeweed mitogen (PWM), and concanavalin A (ConA)
to induce proliferation of lymphocytes.
4
Interferon- values are reported as a stimulation index (SI), calculated by subtracting the concentration
of interferon- produced by cell cultures stimulated with concanavalin A from the concentration of interferon produced by unstimulated cell cultures divided by the concentration of the unstimulated cultures.

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Davis et al.

Table 14. Effect of mixing and age at weaning on macrophage phagocytic capacity and lymphocyte proliferation
response of pigs during the growing/finishing phase1
Weaning age: 14 d
Trait

Mixed

Weaning age: 21 d

Unmixed

Mixed

Unmixed

P-value
SE

Age

Mixed

Interaction

0.381
0.938

0.853
0.636

0.985
0.800

262
11,830
3,445
11,115

0.020
0.713
0.864
0.581

0.402
0.567
0.993
0.330

0.231
0.208
0.006
0.121

255

0.980

0.858

0.735

Macrophage phagocytosis
Phagocytic macrophages, %
SRBC consumed
Lymphocyte proliferation,3 cpm
Unstimulated
PHA, 40 g/mL
PWM, 15 g/mL
ConA, 25 g/mL

9.24
1.51
1,847
118,352
40,588e
122,602

Interferon-4

8.87
1.44

11.27
1.49

1,305
96,318
30,322f
93,908

884
98,761
29,760f
98,689

1,249

1,289

1,382

10.81
1.47
981
107,120
39,962e
105,409
1,330

2.24
0.09

Within a row, least squares means that do not have a common superscript letter differ, P < 0.05.
Three pens from each of the four treatments during the growing/finishing phase were selected randomly for sampling, and a blood sample
was obtained from two pigs in each of the selected pens to determine macrophage phagocytic capacity, lymphocyte proliferation response,
and interferon- production. Pen means were determined, and data are the least squares means of 9 pens per treatment.
2
Macrophage phagocytic capacity was determined by evaluating the percentage of macrophages that were phagocytic, meaning the macrophage consumed at least one sheep red blood cell (SRBC), and the average number of SRBC consumed by each phagocytic macrophage.
3
Lymphocyte proliferation was measured as counts per minute (cpm) in unstimulated cell cultures and cultures stimulated with phytohemagglutinin (PHA), pokeweed mitogen (PWM), and concanavalin A (ConA) to induce proliferation of lymphocytes.
4
Interferon- values are reported as a stimulation index (SI), calculated by subtracting the concentration of interferon- produced by cell
cultures stimulated with ConA from the concentration of interferon- produced by unstimulated cell cultures divided by the concentration of
the unstimulated cultures.
e,f
1

15). Pigs weaned at 14 d of age were more efficient (P

< 0.01) than pigs weaned at 21 d of age during Phase
2 of the nursery and throughout the overall nursery
period (Phase 1 and 2), and tended (P = 0.095) to be
more efficient during Phase 1. At the start of the experiment, pig BW was greater (P < 0.01) when pigs were
weaned at 21 d of age compared with pigs weaned at
14 d of age, and as expected, the older pigs continued
to be heavier (P < 0.01) throughout the nursery period;
however, the difference in BW increased from 2.15 kg

Table 15. Average daily gain, ADFI (as fed), and G:F of
pigs in response to weaning age during the nursery
period
Weaning age, d
Trait

14

21

SE

P-value

ADG, g
Phase 1
Phase 2
Phase 1 to 2

262
487
397

383
612
521

15.3
14.7
13.9

<0.001
<0.001
<0.001

ADFI, g
Phase 1
Phase 2
Phase 1 to 2

273
617
474

410
876
689

15.9
29.1
24.1

<0.001
<0.001
<0.001

G:F
Phase 1
Phase 2
Phase 1 to 2
BW, kg
Initial
Phase 1
Phase 2

0.92
0.77
0.81

0.94
0.70
0.76

0.01
0.01
0.01

0.095
0.001
0.001

4.52
8.17
18.39

6.67
12.06
24.92

0.26
0.45
0.72

<0.001
<0.001
<0.001

at the start of the experiment, to approximately 6.5 kg

at the end of the nursery period. In addition, nursery
mortality was greater when pigs were weaned at 14 d
of age compared with pigs weaned at 21 d of age (12
vs. 1%; P < 0.01). Although diagnostic results to determine the causative disease agent were inconclusive,
morbid pigs exhibited clinical signs of Streptococcus
suis, such as wasting, dyspnea, cyanosis, meningitis,
and sudden death (Higgins and Gottschalk, 1999).
During the growing/finishing phase of the experiment, ADG, ADFI, and G:F did not differ between pigs
weaned at 21 or 14 d of age during Phase 1, Phase 2,
or Phase 4 (Table 16). During Phase 3, however, pigs
weaned at 14 d of age had greater (P < 0.05) ADG,
ADFI, and G:F, and greater (P < 0.01) ADG and G:F
across the entire growing/finishing period, than pigs
weaned at 21 d of age. Removal and commingling of
half the pigs from each pen at the end of the nursery
period and re-sorting for the growing/finishing phase
of the experiment did not affect ADG, ADFI, G:F, or pig
BW (Table 17). The ADFI decreased, however, during
Phase 3 (P < 0.05) when pigs weaned at 21 d of age
were mixed and re-sorted compared with those that
remained in original pens, whereas there was no difference in ADFI of pigs weaned at 14 d of age, regardless
of whether they were mixed and re-sorted or were left
in their original pens (interaction; P = 0.08; Figure 3).
Mixing and re-sorting pigs following the nursery phase
of the study had no effect on the calculated number of
days for pigs to reach a common market weight of 104
kg; however, age of pigs at weaning did affect days to
market, such that pigs weaned at 14 d of age reached
a common BW of 104 kg 4 days earlier than pigs weaned
at 21 d of age (143.9 vs. 148.4 1.3 d; P < 0.05).

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Weaning age and commingling in pigs

753

Table 16. Average daily gain, ADFI (as fed), and G:F of
pigs in response to weaning age during the growing/
finishing period
Weaning age, d
Trait

14

21

SE

P-value

ADG, kg
Phase 1
Phase 2
Phase 3
Phase 4
Phase 1 to 4

0.76
1.01
1.04
0.88
0.91

0.77
0.98
0.86
0.86
0.87

0.02
0.02
0.02
0.04
0.01

0.593
0.258
<0.001
0.719
0.005

ADFI, kg
Phase 1
Phase 2
Phase 3
Phase 4
Phase 1 to 4

1.56
2.43
2.76
3.03
2.26

1.61
2.30
2.59
2.88
2.30

0.03
0.05
0.06
0.08
0.04

0.199
0.075
0.040
0.190
0.509

G:F
Phase
Phase
Phase
Phase
Phase

0.49
0.42
0.38
0.29
0.40

0.48
0.43
0.33
0.30
0.38

0.01
0.01
0.01
0.01
0.01

0.396
0.360
0.003
0.757
0.003

1
2
3
4
1 to 4

DISCUSSION
Behavioral Measures
Behavioral observations in this study indicate that
younger pigs spent less time resting on the day of weaning and more time active during the overall nursery
phase, suggesting that younger pigs were slower to ha-

Table 17. Average daily gain, ADFI (as fed), and G:F of
pigs that were mixed and resorted at the initiation of the
growing/finishing period or remained unmixed during
the growing/finishing period
Growing/finishing treatment
Trait

Mixed

Unmixed

SE

P-value

ADG, kg
Phase 1
Phase 2
Phase 3
Phase 4
Phase 1 to 4

0.76
1.02
0.92
0.86
0.88

0.77
0.98
0.97
0.88
0.89

0.02
0.02
0.02
0.04
0.01

0.598
0.242
0.145
0.671
0.607

ADFI, kg
Phase 1
Phase 2
Phase 3
Phase 4
Phase 1 to 4

1.59
2.41
2.64
2.91
2.28

1.58
2.31
2.71
3.01
2.29

0.03
0.05
0.06
0.08
0.04

0.691
0.167
0.345
0.357
0.831

G:F
Phase
Phase
Phase
Phase
Phase

0.48
0.42
0.35
0.30
0.39

0.49
0.42
0.36
0.30
0.39

0.01
0.01
0.01
0.01
0.01

0.198
0.864
0.568
0.887
0.924

1
2
3
4
1 to 4

Figure 3. Effect of mixing and age at weaning on ADFI

(as fed) of pigs during Phase 3 of the growing/finishing
period (interaction; P = 0.08). Bars that do not have a
common letter differ, P < 0.05.

bituate to their new environment than were older pigs.

Worobec et al. (1999) reported similar findings in pigs
weaned at 7 and 14 d of age compared with pigs weaned
at 28 d of age, in which younger pigs exhibited more
escape behavior, less interaction with neighboring pigs,
and less time feeding than pigs weaned at an older age.
This nervousness and unrest may have contributed to
the lower BW gains observed in young pigs as their
unease likely resulted in more energy expended compared with older pigs in the nursery phase of the study,
although the Streptococcus suis disease challenge that
seemed to be more detrimental to younger pigs, also
likely contributed to the decrease in gain.
The incidence of belly-nosing was not affected by age
of pigs at weaning in this study, although many studies
have reported a greater incidence of belly-nosing when
pigs were weaned at 14 d of age (Metz and Gonyou,
1990; Gonyou et al., 1998, Worobec et al., 1999). Nonetheless, the observation times in this study may have
overlooked aberrant behaviors that occurred in either
age group. Moreover, there was no evidence of lesions
present on pigs because of repetitive belly-nosing behavior, indicating that belly-nosing activity was
likely minimal.
During the growing-finishing period, differences in
the percentage of time spent engaged in feeding behavior during the growing/finishing period depended on the
age of pigs at weaning, whether pigs were commingled
after the nursery period, and the time after commingling occurred. Commingling decreased the percentage
of time pigs engaged in feeding behavior directly after
the commingling event for both age groups. This finding
suggests that reestablishment of the social structure
within pens was necessary when pigs were regrouped.
This response was transient, with no observed differences in duration of feeding behavior in the days following the initial postnursery commingling, except on d 65

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Davis et al.

after weaning. Similarly, other research has reported

that altered or agnostic behaviors were present only
during a short time after the commingling stress
(McGlone et al., 1987; Heetkamp et al., 1995). We have
no explanation, however, for the observations on d 65
after weaning, in which pigs weaned at 21 d of age and
mixed, and pigs weaned at 14 d of age and remained
unmixed spent a greater percentage of time engaged in
feeding behavior. However, the observations coincided
with the beginning of the phase 2 period in which feed
intake and gain by pigs weaned at 14 d of age began
to increase during the growing/finishing period.

Physiological Measures
Total WBC concentrations of pigs in this experiment
were lower than values reported previously in the literature (Schmidt and Tumbleson, 1986; Kleinbeck and
McGlone, 1999); however, proportions of individual leukocyte populations were within expected ranges (Duncan and Prasse, 1986; Schmidt and Tumbleson, 1986).
The low concentration of WBC was surprising with the
increased mortality observed due to Streptococcus suis
infection, particularly in the young age group, as a disease challenge would be expected to raise leukocyte
concentrations in the blood. Concentrations of total
WBC vary depending on genetics, rearing environment,
and health status within a herd (Schmidt and Tumbleson, 1986), and the discrepancy between total WBC
values reported in this study and those reported previously is likely a reflection of these differences. Although differences in the concentrations of neutrophils
and lymphocytes were detected between the two age
groups, the percentage of these cells and the neutrophil:lymphocyte ratio did not differ, suggesting that the
observed differences were likely age-related and not
altered due to differing responses to the stress of weaning. Whereas the emphasis in this study was on differences in the neutrophil:lymphocyte ratio as a measure
of stress, the differences in concentrations and percentages of eosinophils on the various sampling days between treatment groups may provide insight into the
pigs immunological response to the stress of weaning
and commingling. Eosinophils decrease in response to
stress (Dvorak, 1968; Schmidt and Tumbleson, 1986),
and the changes in this cell population in response to
stress warrant further investigation.
White blood cell concentrations in this study were
similar between younger and older pigs on d 27 after
weaning, indicating the younger pigs may be advancing
in their immunological development; however, younger
pigs were less immunologically responsive to the stress
of mixing and resorting after the nursery phase than
older pigs, as indicated by the interaction of weaning
age and mixing treatment observed 3 d after commingling. White blood cell concentrations of older pigs that
were commingled were greater compared with older
pigs that remained in their original pens, whereas WBC
concentrations of younger pigs did not differ in response

to the commingling treatment. Although young pigs

did not respond immunologically to commingling, the
difference in the percentage of time engaged in feeding
behavior was very pronounced between mixed and unmixed pigs weaned at 14 d of age, indicating that young
commingled pigs greatly reduced feeding behavior in
response to commingling compared with young, unmixed pigs.
Physiological responses to stress and behavior involve complex interactions of the central nervous system, the endocrine system, and the immune system
(Yang and Glaser, 2000), and these systems all influence health and well-being in response to environmental and management conditions. In an experiment evaluating the effects of stress at birth on cortisol concentrations, pigs responded to stress associated with a
caesarean-section birth with increased serum cortisol,
although this increase was not associated with a resulting increase in ACTH, suggesting a disregulation
of ACTH and cortisol in caesarean-derived pigs (Daniel
et al., 1999). Hohenshell et al. (2000) reported a similar
disregulation of the hypothalamic-pituitary-adrenal
axis between the pituitary and adrenal glands in earlyweaned pigs that was not present in late-weaned pigs.
Although not directly correlated with the sensitivity
of the immune system in response to stressors, this
disregulation of endocrine function could be a cause of
the inability of early-weaned pigs to respond immunologically to the mixing and resorting stress during the
postnursery period in the current study.

Performance Measures
The lower growth rate during the nursery period of
pigs weaned at 14 d of age compared with older pigs
is in contrast to other studies that reported either an
improvement in ADG of early-weaned (10 d) pigs compared with late-weaned (30 d) pigs (Hohenshell et al.,
2000) or no effect of weaning age on rate of the growth in
the overall nursery period (Dritz et al., 1996). Although
Hohenshell et al. (2000) reported greater ADG by earlyweaned pigs at 42, 65, and 102 d of age compared with
late-weaned pigs, early-weaned pigs were fed a nutrient-rich starter diet until the late-weaned pigs were
moved into the nursery, which may have altered their
subsequent performance compared with older pigs. Because gilts in the current study were bred in the same
breeding group to farrow pigs that would average either
14 or 21 d at weaning, pigs were weaned on the same
day to the same wean-to-finish facility and fed the same
diet. Thus, the effect of weaning age should not be confounded by health status of the sow herd, environmental conditions, or management practices. To our knowledge, this is the first experiment to report that earlyweaned pigs overcame a deficit in BW at the end of the
nursery period to reach a common BW in fewer days
than pigs weaned at 21 d of age. Others have reported
no effect of weaning age when evaluating overall gain
from birth to market weight (Hohenshell et al., 2000),

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Weaning age and commingling in pigs

no difference in BW when comparing early- and lateweaned pigs at a common age (Dritz et al., 1996), or
increases in ADG during the nursery and growing/finishing phases with increasing weaning age from 12 to
21.5 d of age (Main et al., 2004).
The difference in mortality between the young and
old pigs during the nursery period is interesting to note,
and it supports the immune suppression reported previously in pigs weaned at 2 to 3 wk of age (Blecha et
al., 1983). Of the pigs that died during the nursery
phase, all exhibited signs of Streptococcus suis, and the
majority was pigs weaned at 14 d of age. Pigs weaned
at a younger age, likely had a less developed, more
naive immune system that was unable to respond to
the disease challenge. This idea is supported by the
greater concentration of WBC observed in older pigs
than in younger pigs initially at weaning, as well as on
d 2 and 10 after weaning.

General Discussion
Whereas the results of this study show that management conditions perceived as stressors, in this case,
weaning and commingling pigs after the nursery period, elicit changes in WBC populations and influence
behavior, there is no indication that any one measure
of the immune response or behavior can quantify wellbeing in pigs. Although there was a tendency for commingled pigs to have greater concentrations of neutrophils, older pigs that were commingled also responded
with elevated concentrations of lymphocytes; however,
this immune response could not be associated with
growth performance or any aggressive behavior observations. A stress-associated increase in cortisol can result in an alteration of blood leukocyte concentrations,
specifically an increase in neutrophils and a concomitant decrease in lymphocytes in the peripheral blood
that would result in an increase in the neutrophil-tolymphocyte ratio in the stressed animal (Gross and
Siegel, 1983; McGlone et al., 1993). In the current study,
the neutrophil-to-lymphocyte ratio was similar among
pigs regardless of weaning age or mixing treatment;
however, the nonsignificant increase in the concentration and percentage of monocytes in response to mixing
on d 37 (2 d following the commingling event) is interesting. To further compound the difficulty in measuring
the immune response or behavior as indications of welfare, there were very few differences in behavioral responses between pigs weaned at different ages or between pigs commingled after the nursery phase and
those that remained unmixed, and immune responses
were inconsistent depending on the age of the pig at
weaning. Although the results of this study raise interesting questions about the effect of management environments and stressors on pigs weaned at varying ages
on subsequent performance and health, no single measure of growth performance, immunity, or behavior was
found to conclusively measure a pigs welfare in response to wean-to-finish management schemes.

In conclusion, results of this study indicate that

weaning age affects growth performance in a wean-tofinish facility, as well as behavioral and immunological
responses to weaning and commingling after the nursery phase. This study did not allow, however, for determination of the physiological changes in response to
weaning at different ages that altered these behavioral
and immunological measurements. Therefore, physiological responses and management strategies should
be further explored to better understand the benefits
and detriments of early weaning in wean-to-finish
facilities.

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