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Cattle temperament: Persistence of assessments and associations with productivity,
                      efficiency, carcass and meat quality traits
  L. M. Cafe, D. L. Robinson, D. M. Ferguson, B. L. McIntyre, G. H. Geesink and P. L.
                                      Greenwood


                            J ANIM SCI 2011, 89:1452-1465.
        doi: 10.2527/jas.2010-3304 originally published online December 17, 2010




The online version of this article, along with updated information and services, is located on
                                    the World Wide Web at:
                             http://jas.fass.org/content/89/5/1452




                                             www.asas.org




                           Downloaded from jas.fass.org at UNESP on July 20, 2011
Cattle temperament: Persistence of assessments and associations
       with productivity, efficiency, carcass and meat quality traits1
L. M. Cafe,*†2 D. L. Robinson,*† D. M. Ferguson,*‡ B. L. McIntyre,*§ G. H. Geesink,*#
                                and P. L. Greenwood*†

 *Australian Cooperative Research Centre for Beef Genetic Technologies, University of New England, Armidale,
 New South Wales 2351, Australia; †Industry & Investment New South Wales, Beef Industry Centre, Armidale,
    New South Wales 2351, Australia; ‡CSIRO Livestock Industries, FD McMaster Laboratories, Armidale,
          New South Wales 2350, Australia; §Department of Agriculture and Food, Western Australia,
              South Perth, Western Australia 6151, Australia; and #Department of Meat Science,
                   University of New England, Armidale, New South Wales 2351, Australia



ABSTRACT: Relationships between temperament                             calpain-system markers for beef tenderness. Tempera-
and a range of performance, carcass, and meat quality                   ment was not related (most P > 0.05) to tenderness
traits in young cattle were studied in 2 experiments                    gene marker status in Brahman cattle and was not (all
conducted in New South Wales (NSW) and Western                          P > 0.26) modified by the growth promotant treatment
Australia (WA), Australia. In both experiments, growth                  in either breed. The Brahman cattle had greater indi-
rates of cattle were assessed during backgrounding on                   vidual variation in, and greater correlations within and
pasture and grain finishing in a feedlot. Carcass and                   between, repeated assessments of FS and CS than did
objective meat quality characteristics were measured                    the Angus cattle. Correlations for repeated measures of
after slaughter. Feed intake and efficiency during grain                FS were greater than for repeated assessments of CS,
finishing were also determined in NSW. Brahman (n =                     and the strength of correlations for both declined over
82 steers and 82 heifers) and Angus (n = 25 steers and                  time. Average FS or CS for each experiment and loca-
24 heifers) cattle were used in the NSW experiment. In                  tion (NSW or WA × backgrounding or finishing) were
NSW, temperament was assessed by measuring flight                       more highly correlated than individual measurements,
speed [FS, m/s on exit from the chute (crush)] on 14                    indicating that the average values were a more reliable
occasions, and by assessing agitation score during con-                 assessment of cattle temperament than any single mea-
finement in the crush (CS; 1 = calm to 5 = highly                       sure. In Brahman cattle, increased average FS and CS
agitated) on 17 occasions over the course of the ex-                    were associated with significant (P < 0.05) reductions
periment. Brahman (n = 173) and Angus (n = 20)                          in backgrounding and feedlot growth rates, feed intake
steers were used in the WA experiment. In WA, tem-                      and time spent eating, carcass weight, and objective
perament was assessed by measuring FS on 2 occasions                    measures of meat quality. In Angus cattle, the associa-
during backgrounding and on 2 occasions during grain                    tions between temperament and growth rates, feed in-
feeding. At both sites, a hormonal growth promotant                     take, and carcass traits were weaker than in Brahmans,
(Revalor-H, Virbac, Milperra, New South Wales, Aus-                     although the strength of relationships with meat qual-
tralia) was applied to one-half of the cattle at feedlot                ity were similar.
entry, and the Brahman cattle were polymorphic for 2
                  Key words: carcass, cattle, flight speed, meat quality, productivity, temperament

©2011 American Society of Animal Science. All rights reserved.	 J. Anim. Sci. 2011. 89:1452–1465
	doi:10.2527/jas.2010-3304


   1	
    This work was possible because of the financial and in-kind         stock Australia (North Sydney, New South Wales, Australia); South
support and efforts of many individuals from the Cooperative Re-        Australian Research and Development Institute (Urrbrae, South
search Centre for Beef Genetic Technologies (Armidale, New South        Australia, Australia); Victorian Department of Primary Industries
Wales, Australia); Industry & Investment NSW (Armidale, New             (Melbourne, Victoria, Australia); the Australian Brahman Breeders’
South Wales, Australia); Queensland Department of Employment,           Association (Rockhampton, Queensland, Australia); John Dee Ab-
Economic Development and Innovation (City East, Queensland,             attoir (Warwick, Queensland, Australia); and Harvey Beef (Harvey,
Australia); CSIRO Livestock Industries (St. Lucia, Queensland,          Western Australia, Australia).
                                                                           2	
Australia); the University of New England (Armidale, New South              Corresponding author: linda.cafe@industry.nsw.gov.au
Wales, Australia); Western Australia Department of Agriculture and         Received July 8, 2010.
Food (South Perth, Western Australia, Australia); Meat and Live-           Accepted December 8, 2010.

                                                                   1452

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Cattle temperament and productivity                                          1453
                 INTRODUCTION                                       ing practices. The Brahman cattle, treatments, data,
                                                                    and sample collection and their management through-
   Cattle vary in their behavioral response to stressful            out the experiments are described in detail by Cafe et
events, and this trait is defined as temperament. Ex-               al. (2010a,b). Results are also reported here for Angus
treme or reactive responses can be detrimental to cattle            cattle, which were treated identically to the Brahman
welfare and the safety of human handlers. Evidence is               cattle in both experiments, with the 2 breeds managed
emerging that cattle with calmer temperaments have                  together in combined replicates throughout the experi-
improved productivity; however, the effects of tempera-             ments. The numbers of animals in the experiments are
ment on economically important traits can be variable,              presented in Table 1.
and the biological basis for the effects is not well under-            Briefly, the experiments were conducted at Industry
stood (Ferguson et al., 2006).                                      & Investment New South Wales, Agricultural Research
   Several tests have been developed to measure tem-                and Advisory Station [Glen Innes, New South Wales
perament by using the escape and avoidance behaviors                (NSW); 29°44′ S, 151°42′ E, altitude 1,057 m, n =
that cattle display when responding to stressful events,            213 cattle] and at the Western Australian Department
such as handling by humans (reviewed by Burrow,                     of Agriculture and Food’s Vasse Research Station near
1997). Two tests, which are simple and safe to mea-                 Busselton, Western Australia (WA; 33°45′ S, 115°21′ E,
sure and are being used by the Australian beef cattle               altitude 25 m, n = 193). Brahman cattle were selected
industry to select for calmer temperament, are flight               for experimental groups based on their genotype for the
speed (FS; Burrow et al., 1988) and crush score (CS;                calpastatin (Barendse, 2002) and calpain 3 (Barendse
Grandin, 1993). It is likely that these tests measure dif-          et al., 2008) tenderness gene markers. At both sites,
ferent combinations of aspects of cattle temperament,               a small group of Angus cattle with only the favorable
including general agitation and fear of humans, but this            alleles for the calpastatin and calpain 3 gene markers
remains a topic of discussion (Burrow, 1997; Kilgour et             were included as positive controls for biological studies
al., 2006; Petherick et al., 2002, 2009a). Faster FS have           on the calpain system. Equal numbers of heifers and
been shown to lead to slower growth rates, particularly             steers were used in the NSW experiment, whereas only
under more intensive management conditions (Burrow                  steers were used in the WA experiment. One-half the
and Dillon, 1997; Petherick et al., 2009b); reduced feed            cattle in each experiment were treated with a HGP
conversion efficiency (Petherick et al., 2002); and re-             (Revalor-H, Virbac, Milperra, New South Wales, Aus-
duced yield of poorer quality meat (King et al., 2006).             tralia) at feedlot entry.
Similar effects have been shown with greater CS (Voi-                  The Brahman weaner cattle were sourced from re-
sinet et al., 1997a,b).                                             search and commercial herds (n = 15 herds) in NSW,
   The present study was conducted to investigate the               WA, and Queensland. Angus weaner cattle were sourced
persistency over time of cattle temperament, as assessed            from research herds (n = 3) in NSW and WA. All cattle
by FS and CS, and to assess relationships between tem-              were weaned at approximately 6 to 8 mo of age, but be-
perament and a comprehensive range of performance                   cause of differing production systems in their regions of
traits in young Brahman and Angus cattle. The experi-               origin, the Angus cattle were approximately 2 mo older
mental design also allowed potential interactions be-               than the Brahman cattle at both sites. The cattle were
tween temperament and tenderness gene marker status,                grown (backgrounded) on pasture for approximately 6
sex, hormonal growth promotant (HGP) treatment,                     mo, then grain finished in a feedlot for 80 d in WA
and cattle management and meat processing practices                 and 117 d in NSW. In NSW, the cattle were trans-
to be studied.                                                      ported approximately 160 km to the Australian Coop-
                                                                    erative Research Centre for Beef Genetic Technologies
        MATERIALS AND METHODS                                       “Tullimba” research feedlot near Kingstown (30°20′ S,
                                                                    151°10′ E, altitude 560 m) for grain finishing. In WA,
   Use of animals and the procedures performed in this              the cattle were transferred to the feedlot facility at the
study were approved by the Orange Agricultural Insti-               Vasse Research Station for grain finishing. Feed intake
tute Animal Ethics Committee of Industry & Invest-                  and feeding behavior were measured in the NSW feed-
ment New South Wales, the Rockhampton Animal Ex-                    lot by using an automatic individual feeding system, as
perimentation Ethics Committee of the Commonwealth                  described by Bindon (2001), and feed efficiency traits
Scientific and Industrial Research Organisation, and                were calculated as described in detail by Cafe et al.
the Animal Ethics Committee of the Western Austra-                  (2010a).
lian Department of Agriculture and Food.                               Cattle from each experiment were transported to
                                                                    their respective commercial abattoirs the day before
Animals and Experimental Designs                                    slaughter, with no mixing of pens during transport or
                                                                    lairage. For each experiment, one-half of the replicates
  The present study was conducted as a part of 2                    were slaughtered on each of 2 slaughter dates, with the
concurrent experiments designed to study the effects                remaining replicates slaughtered 2 d later. Slaughter
and mechanisms of action of tenderness gene markers,                was conducted through captive bolt stunning and ex-
and their interaction with management and process-                  sanguination. Electrical stimulation of the carcasses

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1454                                                                        Cafe et al.

Table 1. Descriptive statistics for the major traits assessed in Brahman and Angus cattle in the New South Wales
(NSW) and Western Australia (WA) experiments
                                            NSW Brahman                           WA Brahman                             NSW Angus                          WA Angus

Variable                                n         Mean          SD          n         Mean          SD          n        Mean          SD          n        Mean          SD

Growth, kg                                                                                                                                                            
  Background start BW                   164       218          36.0         173       208          59.2         49       295          28.3         20       293           9.4
  Feedlot start BW                      164       321          38.1         173       343          35.6         49       419          41.6         20       403          25.0
  Feedlot end BW                        164       435          55.8         173       449          51.1         49       578          59.1         20       520          28.5
  Background ADG                        164         0.72        0.119       173         0.64        0.169       49         0.71        0.134       20         0.52        0.097
  Feedlot ADG                           164         1.01        0.294       173         1.28        0.345       49         1.43        0.282       20         1.42        0.242
Carcass                                                                                                                                                                     
  Carcass wt, kg                        164       244          32.3         173       242          25.7         49       321          36.6         20       270          15.1
 Dentition1                             164         0.05        0.310       173         0.72        1.032       49         0.61        0.931       20         1.90        0.447
  LLM area,2 cm2                        164        59.9         8.57        143        63.6         6.25        49        67.0         8.93        17        66.1         4.57
  Rump fat, mm                          164        12.0         2.61        173         8.0         2.56        49        18.3         5.44        20         8.9         2.67
  Rib fat,2 mm                          164         6.2         2.08        143         5.3         2.38        49         9.4         2.81        17         8.2         3.26
  Marble score2                         164       261          66.2         143       293          61.9         49       424          72.8         17       321          41.2
  Meat color score2                     164         2.8         1.07        143         2.7         1.05        49         3.2         0.74        17         2.8         0.83
  Ultimate pH2                          164         5.49        0.051       143         5.57        0.085       49         5.49        0.054       17         5.57        0.047
Shear force,3 N                                                                                                                                                             
  AT 1-d aged LLM                       161        78.2        18.53        140        52.2        11.46        49        61.7        11.78        16        44.1         7.44
  AT 7-d aged LLM                       161        68.1        17.63        133        49.5        10.25        46        51.7        10.79        15        44.4         7.80
  TS 1-d aged LLM                       164        47.2         5.61        141        51.6        11.81        49        37.0         4.43        17        41.2         6.39
  TS 7-d aged LLM                       163        45.6         5.56        128        46.0         9.63        49        36.5         4.01        14        40.4         5.29
Feed intake and efficiency                                                                                                                                                  
  Feedlot DMI, kg of DM/d               160         8.0         1.36         —          —            —          49        11.0         1.34        —          —            —
  FCR, kg of DM/kg of BW gain           160         7.5         2.39         —          —            —          49         7.5         2.22        —          —            —
 NFI,4 kg of DM/d                       160       −0.07         0.830        —          —            —          49         0.23        0.97        —          —            —
  Feedlot ADG, kg                       160         1.13        0.314        —          —            —          49         1.53        0.322       —          —            —
  Feeding time, min/d                   160        73.4        20.53         —          —            —          49       106.5        23.63        —          —            —
  Feeding sessions, n/d                 160        11.7         6.14         —          —            —          49         8.8         3.90        —          —            —
  1
   Dentition = number of erupted permanent incisors.
  2
   Meat Standards Australia (2009) grading data, where marble score is from 100 to 1,100 in increments of 10; meat color score is from 1 (lightest)
to 9 (darkest), and ultimate pH is the pH at grading.
  3
   AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum.
  4
   NFI = net feed intake.


was limited to that necessary for the hide removal pro-                                 cattle were being handled through the yards for other
cess at both sites, plus immobilization during exsan-                                   management or data collection purposes. Cattle were
guination in WA. Standard AUS-MEAT carcasses were                                       confined for a period of at least 5 s in a single-animal
prepared (AUS-MEAT, 2007) and split into 2 sides,                                       weighing crate before being released. Crush score was
and the right sides were resuspended by the pelvis [ten-                                assessed visually during the period in the weighing
derstretch (TS) suspension method; Thompson, 2002].                                     crate, using a 5-point scale of agitation based on the
Sides were graded according to Meat Standards Aus-                                      behavioral scoring system described by Grandin (1993),
tralia (2009) procedures, and at bone-out, the musculus                                 which was applied to cattle restrained in a squeeze
longissimus lumborum (LLM) and musculus semiten-                                        chute (crush) and head bail. Minor modifications were
dinosus (STN) were taken from the Achilles-suspended                                    made so that it was more suitable for loosely restrained
(AT) sides, and the LLM were removed from the TS                                        cattle. The scale used was as follows: 1= calm, stand-
sides. These muscles were divided and aged at 1°C for                                   ing still, head mostly still, slow movements; 2 = slightly
either 1 or 7 d before freezing at −20°C. Sample prep-                                  restless, looking around more quickly, moving feet; 3 =
aration and measurement of texture (shear force and                                     restless, moving backward and forward, shaking crate;
compression), cooking loss, CIELAB meat color, and                                      4 = nervous, continuous vigorous movement backward
intramuscular fat percentage (determined by near-in-                                    and forward, snorting; 5 = very nervous, continuous
frared spectrophotometry) were performed as described                                   violent movement, attempting to jump out. All CS as-
by Perry et al. (2001a).                                                                sessments throughout the experiment were made by the
                                                                                        same person.
Temperament Assessments                                                                   When the cattle were released from the weighing
                                                                                        crate, flight time was measured over a distance of 1.7 m
  Temperament was assessed by FS (Burrow et al.,                                        at both sites, and converted to FS (m/s) for analyses.
1988) in both NSW and WA and also by CS (Grandin,                                       Flight speeds of 1 to 1.5 m/s equated to cattle leav-
1993) in NSW. The measurements were taken when the                                      ing the crush at a walk, FS of 2 to 2.5 m/s equated to

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Cattle temperament and productivity                                          1455
cattle leaving the crush at a trot, and FS of 3 to 3.5            methodology (Robinson, 1987), with animal fitted as a
m/s equated to cattle leaving the crush at a run. Dur-            random term.
ing backgrounding in NSW, the yard design required                   The average temperaments during backgrounding
the cattle to turn right at 90° into a side yard upon re-         and finishing (FS and CS for NSW, and FS alone for
lease from the crate. In this case, the FS measurement            WA) were used in the analyses of temperament effects
began after the animals had made the turn and were                on other traits because of the differences in the way
traveling in a straight line. Portable yard panels were           that FS and CS characterized the temperament of cat-
used in the side yard to narrow the exit sufficiently to          tle; the changes in FS and CS between backgrounding
keep the cattle moving in a direct route over the 1.7-            and grain finishing; and the greater reliability of av-
m flight distance. At the feedlot in NSW and during               erages compared with individual assessments. The ef-
both phases in WA, FS measurements were taken in                  fects of temperament on production, carcass traits, and
a straight line directly ahead of the point of release            meat quality traits were assessed using linear mixed
from the weigh crate. In WA, the same set of yards was            models in Genstat. Separate analyses were carried out
used to handle the cattle during the backgrounding and            for each breed (Brahman and Angus) and experimental
feedlot phases.                                                   site (WA and NSW) combination because of the differ-
   In NSW, FS was measured on 5 occasions during                  ences in experimental designs and residual variances.
backgrounding and on 9 occasions during grain finish-             To ensure all aspects of the experimental design were
ing (FS 1 to 14); CS was assessed on 6 occasions during           accounted for, the full models included the fixed effects
backgrounding and on 11 occasions during feedlot fin-             of the tenderness marker genotypes, HGP treatment,
ishing (CS 1 to 17). The timing of the temperament as-            and, for the NSW herd, sex. Random effects included in
sessments and the timing of the more invasive handling            the models were property of origin, backgrounding rep-
events are shown for the NSW experiment in Figure                 licate, feedlot replicate, slaughter group within slaugh-
1. Blood sampling from the tail vein was conducted in             ter day, and the first-order interactions. The effect of
the race before weighing, and then the temperament                temperament was fitted as a covariate in the full model
measures were made as described above. Ultrasound                 for each site × breed combination, with the average
scanning was conducted on 3 occasions with the cattle             temperament variables (FS or CS during background-
caught in the head bail; CS was assessed on each occa-            ing or finishing) fitted as single covariates in separate
sion during the final 30 s of the scanning process, which         analyses, and both the linear and quadratic fits were
took approximately 2 min; and FS was measured after               tested. Main effects and interactions were considered
release from the crush on 1 of these occasions. Muscle            significant at P < 0.05 and were considered a tendency
biopsy was performed on the LLM, STN, and musculus                toward significance at P < 0.10.
semimembranosus under local anesthetic using a drill
biopsy technique (Gardner et al., 2001) on 2 occasions
with the cattle caught in the head bail. Temperament                                          RESULTS
assessments were not made when biopsies were per-
                                                                    The primary purpose of this paper is to report the
formed.
                                                                  assessments of temperament in Brahman and Angus
   In WA, FS was measured on 2 occasions during back-
                                                                  cattle in NSW and WA and their relationships with
grounding and on 2 occasions during grain finishing
                                                                  productivity, carcass traits, and meat quality traits, for
(FS 1 to 4). Flight speed 1 was measured 12 wk after
                                                                  which descriptive statistics are presented in Table 1.
the commencement of backgrounding, when the cattle
                                                                  Results for the effects of HGP, tenderness gene marker
were in the yard for weighing; FS 2 was measured after
                                                                  status, and sex on the measured traits in the Brahman
ultrasound scanning 10 wk later; FS 3 was measured at
                                                                  cattle are presented by Cafe et al. (2010a,b).
feedlot entry a further 8 wk later; and FS 4 was mea-
sured after ultrasound scanning a further 10 wk later.
                                                                  Relationships Between Temperament, HGP
                                                                  Treatment, Tenderness Gene Marker
Statistical Analyses                                              Status, and Sex
  The consistency of FS and CS in ranking animals                    No interactions were observed between HGP treat-
throughout the experiment was assessed using Pearson              ment and temperament assessments (all P > 0.26) in
correlations in Genstat (VSN International Ltd., Hemel            Brahman or Angus cattle at either site. No association
Hempstead, UK). The consistency of FS was analyzed                was observed between CS and tenderness gene marker
for both sites (NSW, 14 measures; WA, 4 measures),                status (all P ≥ 0.12) in Brahman cattle in NSW, and
and the consistency of CS was analyzed using the 17               no consistent association was observed between FS and
assessments made in NSW. The significance of the day              tenderness gene marker status (Cafe et al., 2010a) in
of measurement effects for the repeated measures of               Brahman cattle at either site. Where there were indica-
FS at both sites and of CS in NSW were conducted                  tions of sex differences in NSW, heifers always had nu-
in Genstat using linear mixed models and the REML                 merically greater temperament scores than did steers,


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1456                                                            Cafe et al.

but the differences were small and rarely significant.                   FS and CS over Time
The effect of sex on FS was not significant for either
breed (all P ≥ 0.09). Brahman heifers had greater CS                       FS in NSW. Means for all 14 FS measurements on
than steers during backgrounding (2.15 vs. 1.98, SED                     NSW cattle are shown graphically in Figure 1a, with
= 0.079, P = 0.037) and in the feedlot (1.59 vs. 1.45,                   means and SD of a representative 8 (selected to provide
SED = 0.066, P = 0.045); no significant effect of sex on                 an even spread over time) presented in Table 2. A sig-
CS was observed in the Angus cattle (all P ≥ 0.18). Be-                  nificant effect of day of measurement on FS (P < 0.001)
cause of the lack of interactions between temperament                    was observed in both breeds.
and these effects, further discussion on temperament is                    In the Brahmans, FS decreased during background-
made without reference to them.                                          ing (FS 1 = 2.1 to FS 5 = 1.6 m/s, SED = 0.05 m/s, P




   Figure 1. Mean (±SEM) a) flight speed (FS) and b) crush score (CS) for Angus (●, ○) and Brahman (■, □) cattle in the New South Wales
experiment during backgrounding (solid symbols) and finishing in a feedlot (open symbols). Time of management, ultrasound scan (Scan), and
tissue (Biopsy) and blood (Bleed) sampling events and are also shown.


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Cattle temperament and productivity                                                  1457
                                                                                              1
Table 2. Mean (±SD) and timing of a spread of individual flight speed (FS, m/s) measurements taken during
backgrounding or at the feedlot, and correlations between FS measurements for the Brahman (below diagonal, n
= 164) and Angus (above diagonal, n = 49) cattle in the New South Wales experiment
Item              FS 1            FS 2            FS 3             FS 4             FS 6              FS 9        FS 12           FS 14

Brahman        2.1 ± 0.99      2.0 ± 0.74      1.8 ± 0.75       1.5 ± 0.74       2.4 ± 0.78       2.1 ± 0.92   1.9 ± 0.92      2.1 ± 0.77
Angus          1.3 ± 0.44      1.3 ± 0.53      1.5 ± 0.53       1.0 ± 0.42       2.0 ± 0.51       1.9 ± 0.62   2.0 ± 0.62      2.0 ± 0.49
Day2              0             30              91              126              182              203          231             252
FS 1                              0.26†           0.43*            0.50*            0.33*            0.28†        0.26†           0.29†
FS 2              0.65*                           0.35*            0.54*            0.33*            0.15       −0.06             0.15
FS 3              0.55*           0.67*                            0.70*            0.19             0.27†        0.12            0.25
FS 4              0.62*           0.66*           0.70*                             0.40*            0.29†        0.14            0.25
FS 6              0.48*           0.52*           0.54*            0.53*                             0.51*        0.41*           0.30†
FS 9              0.44*           0.45*           0.51*            0.52*            0.66*                         0.50*           0.41*
FS 12             0.37*           0.47*           0.54*            0.48*            0.61*            0.75*                        0.49*
FS 14             0.36*           0.45*           0.48*            0.50*            0.57*            0.63*        0.71*               
  1
   FS 1 to 5 conducted during backgrounding, and FS 6 to 14 conducted at the feedlot. A subset of 8 FS were chosen to illustrate the range of
correlations in the entire set of 14 measurements.
  2
   Days from first FS measurement.
  †P < 0.10; *P < 0.05.


< 0.001), with no pattern for the SD except that it was                   < 0.001) and Angus (P = 0.008) cattle. Little change
greater at FS1. In the feedlot (where FS was measured                     was observed in SD over time in either breed. In Brah-
as the cattle exited the crush in a straight line, unlike                 man cattle, correlations ranged from 0.41 to 0.52 (all P
backgrounding, where cattle had to turn right at 90°                      < 0.001). In the Angus cattle, correlations ranged from
before measurement), the FS was slightly faster, but                      −0.02 to 0.55 (P < 0.001 to P < 0.9), with the weakest
no consistent change over time was observed in means                      being those involving the first measurement.
or SD. The first feedlot (FS 6) and FS 13 measure-                           CS in NSW. Means for all CS assessments for the
ments (measured after the animals had been scanned                        NSW cattle are shown graphically in Figure 1b, with
and biopsied the previous week) were the greatest (P                      means and SD for a representative 8 of 17 CS assess-
< 0.001). Angus cattle had slower FS than Brahmans.                       ments (selected to provide an even spread over time)
Flight speed in the Angus decreased during back-                          shown in Table 3.
grounding (FS 1 = 1.3 to FS 5 = 1.1 m/s, SED = 0.07,                         A significant effect of day of assessment was ob-
P < 0.001), with no change in the SD over time. Flight                    served. In the Brahman cattle, CS decreased during
speed of the Angus cattle was also faster in the feedlot,                 backgrounding (CS 1 = 2.5 to CS 6 = 1.7, SED = 0.07,
again with no pattern of change over time in means or                     P < 0.001), but the SD were variable. The second as-
SD. The fastest FS was FS 13 (P < 0.001), measured                        sessment (CS 2), when the animals were ultrasound
after the animals had been scanned and biopsied the                       scanned for the first time, was the greatest (P < 0.001).
previous week.                                                            At the feedlot, CS were less in the second half of the
   Correlations for 8 of 14 FS measurements (selected                     feeding period, except for CS 15, which was assessed
to provide an even spread over time) are presented in                     during ultrasound scanning (P < 0.001). In Angus cat-
Table 2. The moderate to high correlations were all                       tle, CS decreased slightly during backgrounding (CS 1
significant (all P < 0.001) for the Brahman cattle, and                   = 1.6 to CS 5 = 1.4, SED = 0.10, P < 0.001), except
were greatest between measurements from the same lo-                      for CS 2, taken during ultrasound scanning, which was
cation (i.e., backgrounding or finishing). For the Angus                  the greatest. The SD was greater for CS 1 and CS 2.
cattle, correlations were not as strong; 90% of the cor-                  Crush scores were less at the feedlot, and again, they
relations of FS at the same location were significant                     were slightly less in the second half of the feeding pe-
(P < 0.001 to P < 0.21), but only approximately 30%                       riod. No pattern of change in SD in the feedlot was
at different locations were significant (P < 0.001 to P                   observed for either breed.
< 0.95). For both breeds, correlations decreased with                        Correlations for 8 of the 17 CS assessments are pre-
increasing time between measures.                                         sented in Table 3. Overall, correlations for CS were less
   FS in WA. Flight speed was measured 4 times in                         than for FS, and were less in the Angus cattle. In the
the WA herd: 86, 155, 210, and 282 d after the begin-                     Brahmans, correlations were generally greater between
ning of backgrounding. Means and SD were 1.7 ± 0.45,                      CS assessments at the same location, but most correla-
1.5 ± 0.52, 1.5 ± 0.52, and 1.5 ± 0.52 m/s for Brah-                      tions between backgrounding and feedlot assessments
mans and were 1.7 ± 0.30, 1.4 ± 0.38, 1.5 ± 0.41, and                     of CS were also significant. Correlations were not as
1.4 ± 0.45 m/s for Angus on these respective days. The                    strong for the Angus cattle, with 30% of those for the
effect of day of measurement was significant, with the                    same location being significant and 25% for different
second measurement being slowest in both Brahman (P                       locations being significant.


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1458                                                                  Cafe et al.

Table 3. Mean (±SD) and timing of a spread of individual crush score1 (CS, score 1 to 5) assessments taken dur-
ing backgrounding or at the feedlot, and correlations between CS assessments for the Brahman (below diagonal, n
= 164) and Angus (above diagonal, n = 49) cattle in the New South Wales experiment
Item              CS 1               CS 2            CS 3              CS 4              CS 7                CS 10          CS 14             CS 17

Brahman        2.5 ± 0.93      2.8 ± 0.73         1.5 ± 0.63       2.0 ± 0.84        1.5 ± 0.59      1.6 ± 0.59           1.4 ± 0.57        1.4 ± 0.60
Angus          1.6 ± 0.64      2.1 ± 0.69         1.3 ± 0.48       1.3 ± 0.48        1.2 ± 0.37      1.2 ± 0.43           1.1 ± 0.33        1.1 ± 0.35
Day2            30              71                 91              126               182             203                  231               252
CS 1                              0.24†              0.31*            0.24†             0.27†         −0.10                  0.13              0.25†
CS 2              0.20*                              0.44*            0.25†             0.04            0.09                 0.06              0.22
CS 3              0.40*           0.19*                               0.28†             0.37*           0.08                 0.25†             0.32*
CS 4              0.37*           0.21*              0.37*                              0.03            0.08                 0.25†             0.19
CS 7              0.20*           0.08               0.26*            0.15                              0.26†                0.17              0.29*
CS 10             0.32*           0.23*              0.36*            0.27*             0.35*                                0.37*             0.04
CS 14             0.26*           0.18*              0.33*            0.22*             0.35*           0.47*                                  0.20
CS 17             0.19*           0.21*              0.24*            0.32*             0.34*           0.48*                0.38*                 
  1
   CS 1 to 6 conducted during backgrounding, and CS 7 to 17 conducted at the feedlot. A subset of 8 CS were chosen to illustrate the range of
correlations in the entire set of 17 measurements.
  2
   Days from first flight speed measurement.
  †P < 0.10; *P < 0.05.


Average FS and CS                                                               ships were weaker for Angus cattle; only the correlation
                                                                                between average FS and CS during backgrounding was
   Averages of FS and CS assessed during background-                            significant (r = 0.39, P = 0.006).
ing and at the feedlot in NSW, and average FS assessed
during backgrounding and at the feedlot in WA are                               FS and Productivity Traits
presented in Table 4. In line with the changes over time
described above, average feedlot FS was faster than av-                            The relationships between average FS and produc-
erage backgrounding FS in NSW (P < 0.001), whereas                              tion, efficiency, carcass traits, and objective meat qual-
CS was less at the feedlot (P < 0.001). In WA Brah-                             ity traits are indicated by estimates of the linear covari-
mans, FS was slightly slower at the feedlot than during                         ates in Tables 6, 7, and 8; quadratic relationships are
backgrounding (P = 0.024). In WA Angus cattle, no                               reported only when significant.
difference was observed in FS measured during back-                                Growth, Intake, and Efficiency. Effects of av-
grounding or at the feedlot (P = 0.35).                                         erage backgrounding and average feedlot FS on produc-
   Correlations between backgrounding and feedlot aver-                         tion and efficiency traits are presented in Table 6. For
ages within assessment type (e.g., 0.69 for FS and 0.41                         NSW Brahmans, cattle with faster backgrounding FS
for CS in the NSW Brahmans; Table 5) were greater                               had reduced BW at all times during the experiment (all
than for any individual pair of measurements from dif-                          P ≤ 0.046) and reduced ADG during backgrounding
ferent locations. This indicates that the averages gave a                       (P = 0.043) and finishing (P = 0.001). The quadratic
more accurate assessment of both FS and CS than did                             relationship between background FS and background
any single measure. Similarly, in WA the correlation                            ADG was stronger (P = 0.009) than the linear relation-
between the average backgrounding and feedlot FS was                            ship, with most of the decline in ADG occurring for
0.66 (P < 0.001) for Brahmans and 0.51 (P = 0.02) for                           background FS of >2.5 m/s. Increasing background FS
Angus.                                                                          was also related to reduced DMI (P = 0.012) and less
   In the NSW Brahmans, correlations between average                            time spent eating (P = 0.046). Increasing feedlot FS
FS and CS, ranging from 0.41 to 0.49 (all P < 0.001),                           was related to reduced BW at the midpoint (P = 0.040)
provide an indication that the 2 different temperament                          and at the end of the feedlot period (P = 0.030) and to
measurements ranked the cattle similarly. The relation-                         decreased feedlot ADG (P = 0.007). Increasing feedlot


Table 4. Average flight speed (m/s) and crush score (1 to 5) during backgrounding and feedlot finishing in Brah-
man and Angus cattle in the New South Wales (NSW) and Western Australia (WA) experiments
                                                       Flight speed                                                          Crush score

Item                   NSW Brahman              NSW Angus          WA Brahman            WA Angus                    NSW Brahman           NSW Angus

Location                                                                                                                                
 Background                  1.84                  1.24                 1.61               1.49                          2.12                 1.56
 Feedlot                     2.09                  1.97                 1.54               1.42                          1.58                 1.21
SED                          0.042                 0.063                0.028              0.080                         0.034                0.04
P-value                     <0.001                <0.001                0.024              0.35                         <0.001               <0.001


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Cattle temperament and productivity                                                             1459
                 Table 5. Correlations between average flight speed (FS, m/s) and crush score (CS,
                 score 1 to 5) determined during backgrounding and feedlot finishing in Brahman (be-
                 low diagonal, n = 164) and Angus (above diagonal, n = 49) cattle in the New South
                 Wales experiment
                 Item                    Background FS           Feedlot FS             Background CS               Feedlot CS

                 Background FS                                       0.42*                     0.39*                  0.23
                 Feedlot FS                   0.69*                                            0.08                   0.24
                 Background CS                0.49*                  0.41*                                            0.62*
                 Feedlot CS                   0.42*                  0.41*                     0.58*                     
                    *P < 0.05.


FS was also associated with reduced time spent eating                 dency toward reduced BW at the end of the feedlot
(P = 0.040) and tended to reduce DMI (P = 0.07). In                   period (P = 0.06).
the WA Brahman cattle, increasing background FS was                      Carcass Characteristics. Effects of average
associated with reduced BW at the beginning and end                   backgrounding and feedlot FS on carcass traits are pre-
of the feedlot period (both P = 0.008), reduced back-                 sented in Table 7. In the NSW Brahman cattle, in-
ground ADG (P = 0.025), and a tendency for reduced                    creasing background FS was associated with significant
feedlot ADG (P = 0.07). In addition, increasing feedlot               reductions in carcass weight (P = 0.001), rib fat (P =
FS was associated with reduced BW at the beginning                    0.016), and ultimate pH (P = 0.014), and an increase
(P = 0.023) and end (P = 0.015) of the feedlot period                 in the temperature at which the carcass reached pH 6
and with a tendency toward reduced feedlot ADG (P                     (P < 0.001). Carcass weight also tended to be reduced
= 0.07).                                                              with increasing feedlot FS (P = 0.09). In the WA Brah-
  In the NSW Angus cattle, increasing background FS                   man cattle, increasing background FS was associated
was related to lighter BW at the beginning of back-                   with reduced carcass weight (P = 0.013), reduced LLM
grounding (P = 0.045), and increasing feedlot FS was                  area (P = 0.035), and darker meat color (P = 0.028).
related to reduced time spent eating (P = 0.038) and                  It also tended to influence carcass pH decline, with the
a tendency toward a decreased feed conversion ratio                   carcasses tending to take longer to reach pH 6 (P =
(FCR; P = 0.07). In WA Angus cattle, no significant                   0.07) and at reduced carcass temperatures (P = 0.09).
relationships were observed between background FS                     Increased feedlot FS also tended to be associated with
and any growth trait, but increased feedlot FS was as-                reduced carcass weight (P = 0.09). For the WA Brah-
sociated with reduced background ADG (P = 0.003),                     man cattle, the quadratic relationship between FS and
reduced BW at feedlot entry (P = 0.018), and a ten-                   carcass weight was slightly stronger than the linear re-


Table 6. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot
finishing on live traits in Brahman and Angus cattle in the New South Wales (NSW) and Western Australia (WA)
experiments
                                                                                     Background FS                            Feedlot FS

Breed        Site                Trait                                       Slope        SE       P-value            Slope       SE       P-value

Brahman      NSW (n = 164)       Beginning background BW, kg                  −6.9        3.45          0.046                                  
                                 Beginning feedlot BW, kg                    −11.5        3.61          0.002                                  
                                 Mid feedlot BW, kg                          −19.1        4.48         <0.001         −9.3        4.50      0.040
                                 End feedlot BW, kg                          −21.0        5.047        <0.001        −11.1        5.07      0.030
                                 Background ADG, kg                           −0.02       0.011         0.043                                  
                                 Feedlot ADG, kg                              −0.08       0.025         0.001         −0.07       0.025     0.007
                                 Feedlot DMI, kg of DM/d                      −0.37       0.146         0.012         −0.26       0.143     0.07
                                 Feeding time, min/d                          −4.7        2.33          0.046         −4.68       2.259     0.040
             WA (n = 173)        Beginning feedlot BW, kg                    −14.3        5.26          0.008        −11.6        5.06      0.023
                                 End feedlot BW, kg                          −20.9        7.78          0.008        −18.3        7.45      0.015
                                 Background ADG, kg                           −0.05       0.020         0.025                                  
                                 Feedlot ADG, kg                              −0.10       0.056         0.07          −0.08       0.046     0.07
Angus        NSW (n = 49)        Beginning background BW, kg                 −23.0       11.08          0.045                                  
                                 Feedlot FCR,1 kg of DM/kg of gain                                                    −1.5        0.81      0.07
                                 Feeding time, min/d                                                                 −17.6        8.19      0.038
             WA (n = 20)         Beginning feedlot BW, kg                                                            −34.8       12.70      0.018
                                 End feedlot BW, kg                                                                  −27.6       13.13      0.06
                                 Background ADG, kg                                                                   −0.16       0.043     0.003
 1
  FCR = feed conversion ratio.


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1460                                                               Cafe et al.

Table 7. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot
finishing on carcass traits in Brahman and Angus cattle in the New South Wales (NSW) and Western Australia
(WA) experiments
                                                                                             Background FS                              Feedlot FS

Breed            Site                     Trait                                  Slope            SE         P-value            Slope      SE        P-value

Brahman          NSW (n = 163)            Carcass wt, kg                         −9.9             2.92        0.001             −5.0       2.93       0.09
                                          Rib fat,1 mm                           −5.7             0.23        0.016                                      
                                          Ultimate pH1                           −0.01            0.006       0.014                                      
                                          Temperature at pH 6, °C                  0.81           0.068      <0.001                                      
                 WA (n = 143)             Carcass wt, kg                         −9.7             3.85        0.013             −6.4       3.73       0.09
                                          LLM area,1 cm2                         −2.67            1.20        0.035                                      
                                          Meat color score1                        0.48           0.213       0.028                                      
                                          Temperature at pH 6, °C                −1.1             0.64        0.09                                       
                                          Time to pH 6, h                          0.22           0.120       0.07                                       
Angus            NSW (n = 49)             Carcass wt, kg                        −27.0            14.47        0.07                                       
                                          Rump fat, mm                           −4.0             2.03        0.06                                       
                                          Time to pH 6, h                          0.56           0.093      <0.001                                      
                 WA (n = 17)              LLM area,1 cm2                                                                        −5.9       1.61       0.005
                                          Marble score1                         −79.4            28.27        0.020                                      
  1
   Meat Standards Australia (2009) grading data, where marble score is from 100 to 1,100 in increments of 10; meat color score is from 1 (lightest)
to 9 (darkest), and ultimate pH is the pH at grading. LLM = musculus longissimus lumborum.


lationship, with most of the decline in carcass weight                        Objective Meat Quality. Effects of average back-
occurring for cattle with a background or feedlot FS of                     grounding and feedlot FS on objective meat quality
>2 m/s (P = 0.012 and 0.06, respectively).                                  traits are presented in Table 8. In the NSW Brahmans,
  In the NSW Angus cattle, increasing background FS                         increasing background FS was related to increased cook-
tended to reduce carcass weight (P = 0.07) and rump                         ing loss in TS 1-d aged LLM (P = 0.036), and increas-
fat (P = 0.06) and significantly increased the time to                      ing feedlot FS tended to be related to increased shear
reach pH 6 (P < 0.001), but no significant effects of                       force (SF) of AT 1-d aged LLM (P = 0.05). In the WA
feedlot FS on carcass traits were observed. In the WA                       Brahmans, increasing background FS was related to in-
Angus cattle, increasing background FS was associated                       creased SF of TS 7-d aged LLM (P = 0.044), increased
with reduced marbling score (P = 0.020), and increas-                       compression in AT 1-d aged STN (P = 0.043), and in-
ing feedlot FS was associated with reduced LLM area                         creased cooking loss (P = 0.023) and darker meat color
(P = 0.005).                                                                (P = 0.006) of AT 7-d aged LLM. Increasing feedlot


Table 8. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot
finishing on objective meat quality traits in Brahman and Angus cattle in the New South Wales (NSW) and West-
ern Australia (WA) experiments
                                                                                        Background FS                                   Feedlot FS

Breed            Site                    Trait1                                 Slope           SE         P-value             Slope       SE        P-value

Brahman          NSW (n = 161)           AT 1-d aged LLM SF, N                                                                  4.2       2.10        0.050
                                         TS 1-d aged LLM cook, %                 0.84          0.396        0.036                                        
                 WA (n = 137)            TS 7-d aged LLM SF, N                   4.3           2.11         0.044               4.2       1.86        0.027
                                         AT 1-d aged STN comp, N                 1.4           0.70         0.043                                        
                                         AT 1-d aged LLM cook, %                                                                0.65      0.292       0.028
                                         AT 7-d aged LLM cook, %                 0.65          0.284        0.023                                        
                                         TS 7-d aged LLM cook, %                                                                0.70      0.286       0.016
                                         AT 1-d aged LLM color L*                                                              −0.84      0.461       0.07
                                         AT 7-d aged LLM color L*               −1.6           0.56         0.006              −1.1       0.53        0.034
Angus            NSW (n = 48)            AT 1-d aged LLM SF, N                  11.2           4.32         0.013                                        
                                         AT 7-d aged LLM SF, N                   7.3           4.04         0.08                                         
                 WA (n = 16)             AT 7-d aged LLM SF, N                  15.8           7.70         0.07                                         
                                         TS 1-d aged LLM SF, N                  13.1           5.89         0.050               6.6       2.65        0.032
                                         AT 7-d aged LLM comp, N                 4.6           2.18         0.07                                         
                                         AT 1-d aged STN comp, N                 7.1           3.06         0.045                                        
                                         TS 1-d aged LLM cook, %                                                                2.4       0.92        0.027
                                         AT 7-d aged LLM pH                      0.10          0.042        0.049                                        
  1
   AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum; STN = musculus semitendinosus; SF =
shear force; comp = compression; cook = cooking loss; color L* = CIELAB color scale, where 0 = dark and 100 = light.

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Cattle temperament and productivity                                                   1461
Table 9. Significant effects of average crush score (CS, score 1 to 5) determined during either backgrounding or
feedlot finishing on live traits in Brahman and Angus cattle in the New South Wales experiment
                                                                           Background CS                          Feedlot CS

Breed                 Trait                                        Slope        SE        P-value       Slope        SE        P-value

Brahman (n = 164)     Beginning background BW, kg                                                       −11.2       5.25        0.034
                      Mid feedlot BW, kg                          −13.8        6.17        0.027        −30.5       6.80       <0.001
                      End feedlot BW, kg                          −11.9        6.94        0.09         −30.0       7.83       <0.001
                      Background ADG, kg                           −0.04       0.015       0.016                                  
                      Feedlot ADG, kg                                                                    −0.12      0.040       0.003
                      Feedlot DMI, kg of DM/d                                                            −0.75      0.226       0.001
Angus (n = 49)        Feed intake per session, kg of DM            −0.74       0.302       0.020         −0.98      0.484       0.05
                      Feeding sessions, No./d                         3.7      1.46        0.016            4.9     2.32        0.043



FS was also related to increasing SF of TS 7-d aged                 sions increasing with both increasing background (P
LLM (P = 0.027) and to increased cooking loss in AT                 = 0.016) and feedlot (P = 0.043) CS. Intake per ses-
1-d aged LLM (P = 0.028) and TS 7-d aged LLM (P =                   sion decreased (P = 0.020) with increasing background
0.016). Increasing feedlot FS was also related to darker            CS and tended to decrease (P = 0.05) with increasing
meat color in AT 7-d aged LLM (P = 0.034) and tend-                 feedlot CS.
ed to be related to darker meat color in AT 1-d aged                   Carcass Characteristics. Effects of average
LLM (P = 0.07).                                                     backgrounding and feedlot CS on carcass traits in the
  In the NSW Angus cattle, increasing background FS                 NSW experiment are presented in Table 10. In Brah-
was associated with increased SF in AT 1-d aged LLM                 man cattle, increasing feedlot CS was associated with a
(P = 0.013) and a tendency toward increased SF in AT                reduction in carcass weight (P < 0.001), and increasing
7-d aged LLM (P = 0.08). No significant relationships               background and feedlot CS were associated with a re-
were observed between feedlot FS and objective meat                 duction in rib fat (P = 0.012 and 0.017). No significant
quality traits. In WA Angus cattle, increasing FS tend-             effects of CS were observed on carcass traits in the
ed to be associated with increased SF in AT 7-d aged                Angus cattle.
LLM (P = 0.07), increased SF in TS 1-d aged LLM                        Objective Meat Quality. Effects of average back-
(P = 0.05), and increased compression in AT 7-d aged                grounding and feedlot CS on objective meat quality
LLM (P = 0.07). It was also related to increased com-               traits in the NSW experiment are presented in Table
pression in AT 1-d aged STN (P = 0.045) and increased               10. In Brahman cattle, as background CS increased,
pH in the laboratory sample of the AT 7-d aged LLM                  so did SF in TS 7-d aged LLM (P = 0.048) and com-
(P = 0.049). Increasing feedlot FS was also associated              pression in AT 1-d aged LLM (P = 0.019). As feedlot
with increased SF (P = 0.032) and cooking loss (P =                 CS increased, SF in AT 1-d aged LLM increased (P =
0.027) in TS 1-d aged LLM.                                          0.024) and tendencies were observed for increased SF
                                                                    with increasing feedlot CS in AT and TS 7-d aged LLM
CS and Productivity Traits                                          (P = 0.08 and 0.09). Cooking loss in AT 1-d aged LLM
                                                                    also increased (P = 0.001) with increasing feedlot CS in
  Growth, Intake, and Efficiency. Effects of                        the Brahman cattle.
average backgrounding and feedlot CS on production                     In Angus cattle, greater background CS was associ-
and feed efficiency traits in the NSW experiment are                ated with increased compression in AT 7-d aged LLM
presented in Table 9. In Brahman cattle, increasing                 (P = 0.04) and with a tendency toward increased SF
background CS was associated with reduced mid feed-                 and compression in AT 1-d aged LLM (P = 0.05 and P
lot BW (P = 0.027), reduced background ADG (P =                     = 0.06). Increasing feedlot CS led to increased SF (P
0.016), and a tendency toward reduced BW (P = 0.09)                 = 0.047) and compression (P = 0.045) in AT 1-d aged
at the end of the feedlot period. The relationship be-              LLM and to a tendency toward increased SF in AT 7-d
tween background CS and feedlot ADG and DMI was                     aged LLM (P = 0.09).
quadratic, with most of the decline in carcass ADG
and intake occurring in cattle with a background CS                                             DISCUSSION
>3 (P = 0.006 and 0.031, respectively). Increased feed-
lot CS was related to reduced BW at the beginning of                   This study shows that the temperament of cattle, as
backgrounding (P = 0.034), mid feedlot (P < 0.001),                 assessed by FS and CS, was persistent over time, and
and at the end of the feedlot period (P < 0.001), and               that cattle with faster FS or greater CS (flightier tem-
to reduced feedlot ADG (P = 0.003) and DMI (P =                     peraments) had inferior performance across a compre-
0.001).                                                             hensive range of beef production traits. Flight speed (or
  In Angus cattle, increasing CS was related only to                its inverse, flight time) and CS are simple to conduct
feeding behavior, with the number of daily feeding ses-             on farm, and their use is encouraged by various Aus-

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1462                                                       Cafe et al.

Table 10. Significant effects of average crush score (CS, score 1 to 5) during either backgrounding or feedlot finish-
ing on carcass and meat quality traits in Brahman and Angus cattle in the New South Wales experiment
                                                                           Background CS                           Feedlot CS

Breed                      Trait1                                  Slope        SE        P-value       Slope         SE        P-value

Brahman (n = 161)          Carcass wt, kg                                                               −16.6        4.50       <0.001
                           Rib fat, mm                             −0.77       0.302        0.012        −0.90       0.370       0.017
                           AT 1-d aged LLM SF, N                                                           7.6       3.34        0.024
                           AT 7-d aged LLM SF, N                                                           5.5       3.10        0.08
                           TS 7-d aged LLM SF, N                    1.5        0.74         0.048          1.6       0.97        0.09
                           AT 1-d aged LLM comp, N                  0.8        0.35         0.019                                  
                           AT 1-d aged LLM cook, %                                                          0.98     0.293       0.001
Angus (n = 48)             AT 1-d aged LLM SF, N                    9.4        4.64         0.05         16.0        7.78        0.047
                           AT 7-d aged LLM SF, N                                                         12.1        6.98        0.09
                           AT 1-d aged LLM comp, N                  1.4        0.72         0.06            2.4      1.16        0.045
                           AT 7-d aged LLM comp, N                  1.1        0.53         0.04                                   
  1
   AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum; SF = shear force; comp = compression,
cook = cooking loss.


tralian beef cattle breed societies [for example, Limou-                handling events, the cattle showed calmer behavioral
sin (CS and pen score), Angus (CS and flight time),                     responses, presumably as they habituated to handling.
and Brahman (flight time); http://breedplan.une.edu.                    This is consistent with results for repeated tempera-
au] to allow selection for a calmer temperament or do-                  ment assessments reported by other authors (Burrow
cility. Despite this, still relatively few published stud-              and Dillon, 1997; Curley et al., 2006; Kilgour et al.,
ies have described relationships between temperament                    2006; Petherick et al., 2009a). The day of measurement
and commercially important traits, and the biological                   effect, significant for both FS and CS, can be attrib-
mechanisms that underpin these associations are not                     uted, at least in part, to the fact that on some days,
well understood (Ferguson et al., 2006).                                the data collection procedures involved closer and more
   Regular FS and CS measurements were taken                            prolonged handling.
throughout the NSW experiment to study the consis-                         Correlations between repeated measures for FS and
tency of the measures over time with changes in loca-                   CS in Brahman cattle were usually significant and were
tion and during various husbandry and sample collec-                    greater than for Angus cattle. The variation was also
tion procedures. Management was more intensive than                     consistently greater for both FS and CS at each as-
for commercial herds because of the data and sample                     sessment in the Brahman than in the Angus cattle,
collection required for other aspects of the experiment,                indicating that the Brahman cattle had greater indi-
but all handling of the animals was conducted as calm-                  vidual variation in temperament than the Angus cattle
ly as possible.                                                         in the present study. This finding would account for
                                                                        the poorer correlations in Angus cattle among individ-
Temperament over Time                                                   ual measures of FS and CS, and between averages for
                                                                        FS and CS. For both breeds, the strength of correla-
   The decreased average feedlot vs. backgrounding CS                   tions declined over time, indicating small, consistent
in NSW, and slower feedlot vs. backgrounding FS in                      changes over time. Because the behavioral response is
WA indicate that the general response of the cattle to                  a combination of genetic and environmental influences
handling declined over the duration of the experiment.                  (Boissy et al., 2005), small changes over time would be
In contrast, FS in NSW was faster at the feedlot than                   expected. In this regard, it was also notable that the
during backgrounding, where FS was measured after                       largest decline in the strength of correlations occurred
the cattle had turned 90° after exiting the chute. The                  with the change in location between backgrounding and
differences in the way in which backgrounding FS was                    feedlot finishing. It is also important to note that the
measured resulted in slower speeds, but nonetheless                     use of average values for FS and CS resulted in greater
provided a meaningful measure of FS.                                    correlations, indicating that average measures provided
   Much of the decline in the mean and variation within                 a more reliable assessment of cattle temperament than
FS was seen after the first 3 measurements, suggest-                    did any single measure, as suggested by Grandin (1993)
ing that the variation between animals stabilized after                 and Burrow and Dillon (1997).
some initial familiarization with handling and the fa-
cilities. The mean and variation for CS were also great-                Relationships Between Temperament
er during initial measurements, after which both were                   and Other Traits
again very consistent for both breeds. Hence, the pat-
tern of change in mean and variation for both FS and                      Relationships between temperament and other pro-
CS indicated that after a small number of consistent                    duction traits were assessed using average FS and CS

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Cattle temperament and productivity                                           1463
during backgrounding or finishing. Where effects of               of time spent eating. A similar but slightly smaller ef-
temperament were significant, cattle with greater FS or           fect was obtained using feedlot FS as the measure of
CS grew more slowly, produced smaller carcasses with              temperament. Increasing CS was also associated with a
less fat cover, and had darker meat that was greater              reduced feed intake (of 750 g of DM/d with each unit
in SF and compression. It is important to note that               increase in feedlot CS) in the NSW Brahman cattle.
all significant effects of a more reactive temperament            These effects were not accounted for entirely by the
(increasing FS or CS) on economically significant traits          BW differences at the beginning of the feedlot period,
were detrimental.                                                 and they remained or tended to remain evident when
   Faster FS was associated with reduced BW and                   feedlot entry BW was fitted as a covariate. However,
growth rates throughout the experiment in the Brah-               there was no evidence of FS or CS being related to dif-
man cattle in NSW and WA. Estimates of the reduc-                 ferences in FCR or NFI in the Brahman cattle. Togeth-
tion in feedlot exit BW were 20.0 and 20.9 kg with each           er, this suggests that temperament plays a significant
1 m/s increase in background FS in the NSW and WA                 role in controlling feed intake and time spent eating,
Brahman cattle, respectively. Increasing CS had similar           but that it has lesser effects on efficiency of utilization
effects in the NSW Brahmans, with a 1-score increase in           of feed; hence, poor temperament reduces DMI and
background CS leading to an 11.9-kg decrease in feedlot           ADG through behavioral rather than metabolic mecha-
exit BW. Carcass weights were reduced with increasing             nisms. This conclusion is in agreement with recent work
FS in the Brahmans, by 9.9 kg in NSW and 9.7 kg in                by Nkrumah et al. (2007) and Elzo et al. (2009), who
WA for each 1 m/s increase in FS. The NSW Brahmans                found that young cattle of mixed breeds with faster
also had a 16.6-kg reduction in carcass weight per unit           feedlot FS had less feedlot DMI but showed no differ-
increase in feedlot CS. There was also some indication            ence in FCR or NFI.
of reduced carcass fatness and LLM area with increas-                In the NSW herd, temperament had less effect on
ing FS, and of reduced carcass fatness with increasing            feed intake in Angus than in Brahman cattle. However,
CS, in the NSW Brahmans. The inclusion of carcass                 in the Angus cattle, each meter per second increase in
weight as a covariate in the statistical models showed            feedlot FS was associated with a 17.6 min/d reduction
that these differences in composition were mostly ex-             in feeding time and a tendency toward reduced FCR by
plained by the differences in carcass weight. In the An-          1.5 kg of DM/kg of BW gain. This reduction in FCR is
gus cattle, increasing FS also tended to reduce BW and            the only result relating to poorer temperament within
growth rate, but the relationships were much weaker               the present study that might be considered beneficial.
than in Brahmans. In line with the weak trends toward             Although caution is required, because of the P-value
lighter BW with increasing FS in the Angus cattle, ten-           of 0.07, it is possible that among the Angus cattle, a
dencies for reductions in BW-related carcass traits were          slightly decreased intake with increasing FS allowed for
observed. However, no significant relationships were ob-          more efficient digestion (Herd et al., 2004).
served between CS and BW-related traits in the Angus                 The determinants of eating quality of the meat are
cattle in the NSW herd.                                           complex and multifactorial, and pre- and postmortem
   Previously reported results for relationships between          events can have major effects on beef tenderness (Mal-
cattle temperament and growth have been variable.                 tin et al., 2003; Ferguson and Warner, 2008). Cattle
Slower growth rates have been reported in cattle with             temperament is assumed to be related to stress respon-
faster FS, greater CS, or both in studies conducted un-           siveness, and it is likely that the stress response to han-
der more intensive management systems (Burrow and                 dling and transport is greater in temperamental cattle,
Dillon, 1997; Voisinet et al., 1997b; Müller and von              resulting in depletion of muscle glycogen before slaugh-
Keyserlingk, 2006; Behrends et al., 2009). Others have            ter and hence reduced meat quality because of greater
found little relationship between temperament and                 carcass pH and the associated darker color (Ferguson et
growth rates in herds in which the ranges in FS and CS            al., 2006). In the present study, increasing FS or CS was
were small and cattle were generally docile (Graham et            related to darker LLM meat color and increased muscle
al., 2001; Elzo et al., 2009). Our results are consistent         pH, shear force, compression, and cooking loss, effects
with the above studies in that the Angus cattle were              all considered detrimental to the eating quality of beef
generally more docile than the Brahmans, and greater              (Perry et al., 2001b).
effects of temperament on growth were observed in the                The relationships between temperament and meat
Brahmans.                                                         quality were strongest for the WA cattle. Differences
   It has previously been postulated that cattle with             between experimental sites could, at least in part, be
a more reactive temperament may grow more slowly                  due to processing differences resulting in differences in
because of the greater energy expenditure associated              postmortem pH or temperature declines, as discussed
with, for example, more vigilant behavior, resulting in           by Cafe et al. (2010b). Greater effects of temperament
poorer FCR or net feed intake (NFI; Burrow and Dil-               on meat quality traits were evident in both breeds in
lon, 1997; Petherick et al., 2002). In the NSW Brah-              WA, where the carcasses had much faster pH declines
mans, each meter per second increase in background FS             and slower cooling rates than in NSW. Relationships
was associated with a reduction in feed intake of 370 g           between temperament and shear force tended to be
of DM/d, and a reduction of 4.7 min/d in the amount               less significant with 7-d aging. This may indicate that

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1464                                                     Cafe et al.

temperament was related to variation in tenderness be-                              LITERATURE CITED
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result was that the Angus cattle, with smaller numbers            AUS-MEAT. 2007. AUS-MEAT National Accreditation Standards.
of animals and less variation in temperament, showed                   2007 Edition. AUS-MEAT Ltd., Brisbane, Queensland, Aus-
stronger effects of temperament on SF than did the                     tralia.
                                                                  Barendse, W. J. 2002. DNA markers for meat tenderness. The Com-
Brahman cattle in both herds.                                          monwealth Scientific and Industrial Research Organization,
   Overall, most relationships with temperament were                   The State of Queensland through its Department of Primary
linear, indicating that selection of cattle with calmer                Industries, The University of New England, The State of New
temperaments, and not only culling of cattle with the                  South Wales through its Department of Agriculture, and Meat
most reactive temperaments, can improve productivity                   and Livestock Australia Limited, assignees. Int. Patent No. W0
                                                                       02/064820.
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safety for human handlers. Other studies have shown                    2008. Variation at the calpain 3 gene is associated with meat
both linear (Burrow and Dillon, 1997) and quadratic                    tenderness in zebu and composite breeds of cattle. BMC Gen-
(Müller and von Keyserlingk, 2006) relationships be-                   et. 9:41.
tween temperament and productivity traits. However,               Behrends, S. M., R. K. Miller, F. M. Rouquette Jr., R. D. Randel,
                                                                       B. G. Warrington, T. D. A. Forbes, T. H. Welsh, H. Lippke, J.
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exist within the herd, with the presence of extremely                  tenderness in beef steers. Meat Sci. 81:433–438.
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                                                                       quality research (1998–2000). Aust. J. Exp. Agric. 41:843–
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management at the time of measurement. The earlier                     2005. Genetics of fear in ruminant livestock. Livest. Prod. Sci.
(backgrounding) assessments had stronger and more                      93:23–32.
frequent relationships with productivity and carcass              Burrow, H. M. 1997. Measurements of temperament and their rela-
                                                                       tionships with performance traits of beef cattle. Anim. Breed.
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assessments showed significant relationships with meat            Burrow, H. M., and R. D. Dillon. 1997. Relationships between tem-
quality. Other authors have reported weak to no re-                    perament and growth in a feedlot and commercial carcass traits
lationships between temperament measurements taken                     of Bos indicus crossbreds. Aust. J. Exp. Agric. 37:407–411.
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                                                                       nique for measuring temperament in cattle. Proc. Aust. Soc.
al., 2009). In the present study, the uniform measure-                 Anim. Prod. 17:154–157.
ment procedures and frequent handling of the cattle               Cafe, L. M., B. L. McIntyre, D. L. Robinson, G. H. Geesink, W.
throughout the experiment for intensive data and sam-                  Barendse, and P. L. Greenwood. 2010a. Production and pro-
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have allowed variation in temperament to continue to                   in Brahman cattle: 1. Growth, efficiency, temperament and car-
                                                                       cass characteristics. J. Anim. Sci. 88:3047–3058.
be detected at a later stage of production.                       Cafe, L. M., B. M. McIntyre, D. L. Robinson, G. H. Geesink, W.
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                                                                  Curley, K. O., Jr., J. C. Paschal, T. H. Welsh Jr., and R. D. Randel.
measured by reduced FS and CS, had superior perfor-                    2006. Technical note: Exit velocity as a measure of cattle tem-
mance across a comprehensive range of beef produc-                     perament is repeatable and associated with serum concentra-
tion traits. Repeated assessments of temperament using                 tion of cortisol in Brahman bulls. J. Anim. Sci. 84:3100–3103.
FS and CS were correlated, with the strength of the               Elzo, M. A., D. G. Riley, G. R. Hansen, D. D. Johnson, R. O. Myer,
correlations declining over time. Correlations between                 S. W. Coleman, C. C. Chase, J. G. Wasdin, and J. D. Driver.
                                                                       2009. Effect of breed composition on phenotypic residual feed
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tween repeated assessments of CS. A stronger relation-                 crossbred cattle. J. Anim. Sci. 87:3877–3886.
ship was observed between temperament and growth                  Ferguson, D. M., D. Johnston, H. M. Burrow, and A. Reverter. 2006.
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                                                                       2001. The impact of nutrition on bovine muscle glycogen me-
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Herd, R. M., V. H. Oddy, and E. C. Richardson. 2004. Biological ba-              and taste panel assessment of beef quality. Aust. J. Exp. Agric.
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                                               Downloaded from jas.fass.org at UNESP on July 20, 2011
References       This article cites 30 articles, 6 of which you can access for free at:
                 http://jas.fass.org/content/89/5/1452#BIBL




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Seminário temp bov

  • 1. Cattle temperament: Persistence of assessments and associations with productivity, efficiency, carcass and meat quality traits L. M. Cafe, D. L. Robinson, D. M. Ferguson, B. L. McIntyre, G. H. Geesink and P. L. Greenwood J ANIM SCI 2011, 89:1452-1465. doi: 10.2527/jas.2010-3304 originally published online December 17, 2010 The online version of this article, along with updated information and services, is located on the World Wide Web at: http://jas.fass.org/content/89/5/1452 www.asas.org Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 2. Cattle temperament: Persistence of assessments and associations with productivity, efficiency, carcass and meat quality traits1 L. M. Cafe,*†2 D. L. Robinson,*† D. M. Ferguson,*‡ B. L. McIntyre,*§ G. H. Geesink,*# and P. L. Greenwood*† *Australian Cooperative Research Centre for Beef Genetic Technologies, University of New England, Armidale, New South Wales 2351, Australia; †Industry & Investment New South Wales, Beef Industry Centre, Armidale, New South Wales 2351, Australia; ‡CSIRO Livestock Industries, FD McMaster Laboratories, Armidale, New South Wales 2350, Australia; §Department of Agriculture and Food, Western Australia, South Perth, Western Australia 6151, Australia; and #Department of Meat Science, University of New England, Armidale, New South Wales 2351, Australia ABSTRACT: Relationships between temperament calpain-system markers for beef tenderness. Tempera- and a range of performance, carcass, and meat quality ment was not related (most P > 0.05) to tenderness traits in young cattle were studied in 2 experiments gene marker status in Brahman cattle and was not (all conducted in New South Wales (NSW) and Western P > 0.26) modified by the growth promotant treatment Australia (WA), Australia. In both experiments, growth in either breed. The Brahman cattle had greater indi- rates of cattle were assessed during backgrounding on vidual variation in, and greater correlations within and pasture and grain finishing in a feedlot. Carcass and between, repeated assessments of FS and CS than did objective meat quality characteristics were measured the Angus cattle. Correlations for repeated measures of after slaughter. Feed intake and efficiency during grain FS were greater than for repeated assessments of CS, finishing were also determined in NSW. Brahman (n = and the strength of correlations for both declined over 82 steers and 82 heifers) and Angus (n = 25 steers and time. Average FS or CS for each experiment and loca- 24 heifers) cattle were used in the NSW experiment. In tion (NSW or WA × backgrounding or finishing) were NSW, temperament was assessed by measuring flight more highly correlated than individual measurements, speed [FS, m/s on exit from the chute (crush)] on 14 indicating that the average values were a more reliable occasions, and by assessing agitation score during con- assessment of cattle temperament than any single mea- finement in the crush (CS; 1 = calm to 5 = highly sure. In Brahman cattle, increased average FS and CS agitated) on 17 occasions over the course of the ex- were associated with significant (P < 0.05) reductions periment. Brahman (n = 173) and Angus (n = 20) in backgrounding and feedlot growth rates, feed intake steers were used in the WA experiment. In WA, tem- and time spent eating, carcass weight, and objective perament was assessed by measuring FS on 2 occasions measures of meat quality. In Angus cattle, the associa- during backgrounding and on 2 occasions during grain tions between temperament and growth rates, feed in- feeding. At both sites, a hormonal growth promotant take, and carcass traits were weaker than in Brahmans, (Revalor-H, Virbac, Milperra, New South Wales, Aus- although the strength of relationships with meat qual- tralia) was applied to one-half of the cattle at feedlot ity were similar. entry, and the Brahman cattle were polymorphic for 2 Key words: carcass, cattle, flight speed, meat quality, productivity, temperament ©2011 American Society of Animal Science. All rights reserved. J. Anim. Sci. 2011. 89:1452–1465 doi:10.2527/jas.2010-3304 1 This work was possible because of the financial and in-kind stock Australia (North Sydney, New South Wales, Australia); South support and efforts of many individuals from the Cooperative Re- Australian Research and Development Institute (Urrbrae, South search Centre for Beef Genetic Technologies (Armidale, New South Australia, Australia); Victorian Department of Primary Industries Wales, Australia); Industry & Investment NSW (Armidale, New (Melbourne, Victoria, Australia); the Australian Brahman Breeders’ South Wales, Australia); Queensland Department of Employment, Association (Rockhampton, Queensland, Australia); John Dee Ab- Economic Development and Innovation (City East, Queensland, attoir (Warwick, Queensland, Australia); and Harvey Beef (Harvey, Australia); CSIRO Livestock Industries (St. Lucia, Queensland, Western Australia, Australia). 2 Australia); the University of New England (Armidale, New South Corresponding author: linda.cafe@industry.nsw.gov.au Wales, Australia); Western Australia Department of Agriculture and Received July 8, 2010. Food (South Perth, Western Australia, Australia); Meat and Live- Accepted December 8, 2010. 1452 Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 3. Cattle temperament and productivity 1453 INTRODUCTION ing practices. The Brahman cattle, treatments, data, and sample collection and their management through- Cattle vary in their behavioral response to stressful out the experiments are described in detail by Cafe et events, and this trait is defined as temperament. Ex- al. (2010a,b). Results are also reported here for Angus treme or reactive responses can be detrimental to cattle cattle, which were treated identically to the Brahman welfare and the safety of human handlers. Evidence is cattle in both experiments, with the 2 breeds managed emerging that cattle with calmer temperaments have together in combined replicates throughout the experi- improved productivity; however, the effects of tempera- ments. The numbers of animals in the experiments are ment on economically important traits can be variable, presented in Table 1. and the biological basis for the effects is not well under- Briefly, the experiments were conducted at Industry stood (Ferguson et al., 2006). & Investment New South Wales, Agricultural Research Several tests have been developed to measure tem- and Advisory Station [Glen Innes, New South Wales perament by using the escape and avoidance behaviors (NSW); 29°44′ S, 151°42′ E, altitude 1,057 m, n = that cattle display when responding to stressful events, 213 cattle] and at the Western Australian Department such as handling by humans (reviewed by Burrow, of Agriculture and Food’s Vasse Research Station near 1997). Two tests, which are simple and safe to mea- Busselton, Western Australia (WA; 33°45′ S, 115°21′ E, sure and are being used by the Australian beef cattle altitude 25 m, n = 193). Brahman cattle were selected industry to select for calmer temperament, are flight for experimental groups based on their genotype for the speed (FS; Burrow et al., 1988) and crush score (CS; calpastatin (Barendse, 2002) and calpain 3 (Barendse Grandin, 1993). It is likely that these tests measure dif- et al., 2008) tenderness gene markers. At both sites, ferent combinations of aspects of cattle temperament, a small group of Angus cattle with only the favorable including general agitation and fear of humans, but this alleles for the calpastatin and calpain 3 gene markers remains a topic of discussion (Burrow, 1997; Kilgour et were included as positive controls for biological studies al., 2006; Petherick et al., 2002, 2009a). Faster FS have on the calpain system. Equal numbers of heifers and been shown to lead to slower growth rates, particularly steers were used in the NSW experiment, whereas only under more intensive management conditions (Burrow steers were used in the WA experiment. One-half the and Dillon, 1997; Petherick et al., 2009b); reduced feed cattle in each experiment were treated with a HGP conversion efficiency (Petherick et al., 2002); and re- (Revalor-H, Virbac, Milperra, New South Wales, Aus- duced yield of poorer quality meat (King et al., 2006). tralia) at feedlot entry. Similar effects have been shown with greater CS (Voi- The Brahman weaner cattle were sourced from re- sinet et al., 1997a,b). search and commercial herds (n = 15 herds) in NSW, The present study was conducted to investigate the WA, and Queensland. Angus weaner cattle were sourced persistency over time of cattle temperament, as assessed from research herds (n = 3) in NSW and WA. All cattle by FS and CS, and to assess relationships between tem- were weaned at approximately 6 to 8 mo of age, but be- perament and a comprehensive range of performance cause of differing production systems in their regions of traits in young Brahman and Angus cattle. The experi- origin, the Angus cattle were approximately 2 mo older mental design also allowed potential interactions be- than the Brahman cattle at both sites. The cattle were tween temperament and tenderness gene marker status, grown (backgrounded) on pasture for approximately 6 sex, hormonal growth promotant (HGP) treatment, mo, then grain finished in a feedlot for 80 d in WA and cattle management and meat processing practices and 117 d in NSW. In NSW, the cattle were trans- to be studied. ported approximately 160 km to the Australian Coop- erative Research Centre for Beef Genetic Technologies MATERIALS AND METHODS “Tullimba” research feedlot near Kingstown (30°20′ S, 151°10′ E, altitude 560 m) for grain finishing. In WA, Use of animals and the procedures performed in this the cattle were transferred to the feedlot facility at the study were approved by the Orange Agricultural Insti- Vasse Research Station for grain finishing. Feed intake tute Animal Ethics Committee of Industry & Invest- and feeding behavior were measured in the NSW feed- ment New South Wales, the Rockhampton Animal Ex- lot by using an automatic individual feeding system, as perimentation Ethics Committee of the Commonwealth described by Bindon (2001), and feed efficiency traits Scientific and Industrial Research Organisation, and were calculated as described in detail by Cafe et al. the Animal Ethics Committee of the Western Austra- (2010a). lian Department of Agriculture and Food. Cattle from each experiment were transported to their respective commercial abattoirs the day before Animals and Experimental Designs slaughter, with no mixing of pens during transport or lairage. For each experiment, one-half of the replicates The present study was conducted as a part of 2 were slaughtered on each of 2 slaughter dates, with the concurrent experiments designed to study the effects remaining replicates slaughtered 2 d later. Slaughter and mechanisms of action of tenderness gene markers, was conducted through captive bolt stunning and ex- and their interaction with management and process- sanguination. Electrical stimulation of the carcasses Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 4. 1454 Cafe et al. Table 1. Descriptive statistics for the major traits assessed in Brahman and Angus cattle in the New South Wales (NSW) and Western Australia (WA) experiments NSW Brahman WA Brahman NSW Angus WA Angus Variable n Mean SD n Mean SD n Mean SD n Mean SD Growth, kg                           Background start BW 164 218 36.0 173 208 59.2 49 295 28.3 20 293 9.4   Feedlot start BW 164 321 38.1 173 343 35.6 49 419 41.6 20 403 25.0   Feedlot end BW 164 435 55.8 173 449 51.1 49 578 59.1 20 520 28.5   Background ADG 164 0.72 0.119 173 0.64 0.169 49 0.71 0.134 20 0.52 0.097   Feedlot ADG 164 1.01 0.294 173 1.28 0.345 49 1.43 0.282 20 1.42 0.242 Carcass                           Carcass wt, kg 164 244 32.3 173 242 25.7 49 321 36.6 20 270 15.1  Dentition1 164 0.05 0.310 173 0.72 1.032 49 0.61 0.931 20 1.90 0.447   LLM area,2 cm2 164 59.9 8.57 143 63.6 6.25 49 67.0 8.93 17 66.1 4.57   Rump fat, mm 164 12.0 2.61 173 8.0 2.56 49 18.3 5.44 20 8.9 2.67   Rib fat,2 mm 164 6.2 2.08 143 5.3 2.38 49 9.4 2.81 17 8.2 3.26   Marble score2 164 261 66.2 143 293 61.9 49 424 72.8 17 321 41.2   Meat color score2 164 2.8 1.07 143 2.7 1.05 49 3.2 0.74 17 2.8 0.83   Ultimate pH2 164 5.49 0.051 143 5.57 0.085 49 5.49 0.054 17 5.57 0.047 Shear force,3 N                           AT 1-d aged LLM 161 78.2 18.53 140 52.2 11.46 49 61.7 11.78 16 44.1 7.44   AT 7-d aged LLM 161 68.1 17.63 133 49.5 10.25 46 51.7 10.79 15 44.4 7.80   TS 1-d aged LLM 164 47.2 5.61 141 51.6 11.81 49 37.0 4.43 17 41.2 6.39   TS 7-d aged LLM 163 45.6 5.56 128 46.0 9.63 49 36.5 4.01 14 40.4 5.29 Feed intake and efficiency                           Feedlot DMI, kg of DM/d 160 8.0 1.36 — — — 49 11.0 1.34 — — —   FCR, kg of DM/kg of BW gain 160 7.5 2.39 — — — 49 7.5 2.22 — — —  NFI,4 kg of DM/d 160 −0.07 0.830 — — — 49 0.23 0.97 — — —   Feedlot ADG, kg 160 1.13 0.314 — — — 49 1.53 0.322 — — —   Feeding time, min/d 160 73.4 20.53 — — — 49 106.5 23.63 — — —   Feeding sessions, n/d 160 11.7 6.14 — — — 49 8.8 3.90 — — — 1 Dentition = number of erupted permanent incisors. 2 Meat Standards Australia (2009) grading data, where marble score is from 100 to 1,100 in increments of 10; meat color score is from 1 (lightest) to 9 (darkest), and ultimate pH is the pH at grading. 3 AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum. 4 NFI = net feed intake. was limited to that necessary for the hide removal pro- cattle were being handled through the yards for other cess at both sites, plus immobilization during exsan- management or data collection purposes. Cattle were guination in WA. Standard AUS-MEAT carcasses were confined for a period of at least 5 s in a single-animal prepared (AUS-MEAT, 2007) and split into 2 sides, weighing crate before being released. Crush score was and the right sides were resuspended by the pelvis [ten- assessed visually during the period in the weighing derstretch (TS) suspension method; Thompson, 2002]. crate, using a 5-point scale of agitation based on the Sides were graded according to Meat Standards Aus- behavioral scoring system described by Grandin (1993), tralia (2009) procedures, and at bone-out, the musculus which was applied to cattle restrained in a squeeze longissimus lumborum (LLM) and musculus semiten- chute (crush) and head bail. Minor modifications were dinosus (STN) were taken from the Achilles-suspended made so that it was more suitable for loosely restrained (AT) sides, and the LLM were removed from the TS cattle. The scale used was as follows: 1= calm, stand- sides. These muscles were divided and aged at 1°C for ing still, head mostly still, slow movements; 2 = slightly either 1 or 7 d before freezing at −20°C. Sample prep- restless, looking around more quickly, moving feet; 3 = aration and measurement of texture (shear force and restless, moving backward and forward, shaking crate; compression), cooking loss, CIELAB meat color, and 4 = nervous, continuous vigorous movement backward intramuscular fat percentage (determined by near-in- and forward, snorting; 5 = very nervous, continuous frared spectrophotometry) were performed as described violent movement, attempting to jump out. All CS as- by Perry et al. (2001a). sessments throughout the experiment were made by the same person. Temperament Assessments When the cattle were released from the weighing crate, flight time was measured over a distance of 1.7 m Temperament was assessed by FS (Burrow et al., at both sites, and converted to FS (m/s) for analyses. 1988) in both NSW and WA and also by CS (Grandin, Flight speeds of 1 to 1.5 m/s equated to cattle leav- 1993) in NSW. The measurements were taken when the ing the crush at a walk, FS of 2 to 2.5 m/s equated to Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 5. Cattle temperament and productivity 1455 cattle leaving the crush at a trot, and FS of 3 to 3.5 methodology (Robinson, 1987), with animal fitted as a m/s equated to cattle leaving the crush at a run. Dur- random term. ing backgrounding in NSW, the yard design required The average temperaments during backgrounding the cattle to turn right at 90° into a side yard upon re- and finishing (FS and CS for NSW, and FS alone for lease from the crate. In this case, the FS measurement WA) were used in the analyses of temperament effects began after the animals had made the turn and were on other traits because of the differences in the way traveling in a straight line. Portable yard panels were that FS and CS characterized the temperament of cat- used in the side yard to narrow the exit sufficiently to tle; the changes in FS and CS between backgrounding keep the cattle moving in a direct route over the 1.7- and grain finishing; and the greater reliability of av- m flight distance. At the feedlot in NSW and during erages compared with individual assessments. The ef- both phases in WA, FS measurements were taken in fects of temperament on production, carcass traits, and a straight line directly ahead of the point of release meat quality traits were assessed using linear mixed from the weigh crate. In WA, the same set of yards was models in Genstat. Separate analyses were carried out used to handle the cattle during the backgrounding and for each breed (Brahman and Angus) and experimental feedlot phases. site (WA and NSW) combination because of the differ- In NSW, FS was measured on 5 occasions during ences in experimental designs and residual variances. backgrounding and on 9 occasions during grain finish- To ensure all aspects of the experimental design were ing (FS 1 to 14); CS was assessed on 6 occasions during accounted for, the full models included the fixed effects backgrounding and on 11 occasions during feedlot fin- of the tenderness marker genotypes, HGP treatment, ishing (CS 1 to 17). The timing of the temperament as- and, for the NSW herd, sex. Random effects included in sessments and the timing of the more invasive handling the models were property of origin, backgrounding rep- events are shown for the NSW experiment in Figure licate, feedlot replicate, slaughter group within slaugh- 1. Blood sampling from the tail vein was conducted in ter day, and the first-order interactions. The effect of the race before weighing, and then the temperament temperament was fitted as a covariate in the full model measures were made as described above. Ultrasound for each site × breed combination, with the average scanning was conducted on 3 occasions with the cattle temperament variables (FS or CS during background- caught in the head bail; CS was assessed on each occa- ing or finishing) fitted as single covariates in separate sion during the final 30 s of the scanning process, which analyses, and both the linear and quadratic fits were took approximately 2 min; and FS was measured after tested. Main effects and interactions were considered release from the crush on 1 of these occasions. Muscle significant at P < 0.05 and were considered a tendency biopsy was performed on the LLM, STN, and musculus toward significance at P < 0.10. semimembranosus under local anesthetic using a drill biopsy technique (Gardner et al., 2001) on 2 occasions with the cattle caught in the head bail. Temperament RESULTS assessments were not made when biopsies were per- The primary purpose of this paper is to report the formed. assessments of temperament in Brahman and Angus In WA, FS was measured on 2 occasions during back- cattle in NSW and WA and their relationships with grounding and on 2 occasions during grain finishing productivity, carcass traits, and meat quality traits, for (FS 1 to 4). Flight speed 1 was measured 12 wk after which descriptive statistics are presented in Table 1. the commencement of backgrounding, when the cattle Results for the effects of HGP, tenderness gene marker were in the yard for weighing; FS 2 was measured after status, and sex on the measured traits in the Brahman ultrasound scanning 10 wk later; FS 3 was measured at cattle are presented by Cafe et al. (2010a,b). feedlot entry a further 8 wk later; and FS 4 was mea- sured after ultrasound scanning a further 10 wk later. Relationships Between Temperament, HGP Treatment, Tenderness Gene Marker Statistical Analyses Status, and Sex The consistency of FS and CS in ranking animals No interactions were observed between HGP treat- throughout the experiment was assessed using Pearson ment and temperament assessments (all P > 0.26) in correlations in Genstat (VSN International Ltd., Hemel Brahman or Angus cattle at either site. No association Hempstead, UK). The consistency of FS was analyzed was observed between CS and tenderness gene marker for both sites (NSW, 14 measures; WA, 4 measures), status (all P ≥ 0.12) in Brahman cattle in NSW, and and the consistency of CS was analyzed using the 17 no consistent association was observed between FS and assessments made in NSW. The significance of the day tenderness gene marker status (Cafe et al., 2010a) in of measurement effects for the repeated measures of Brahman cattle at either site. Where there were indica- FS at both sites and of CS in NSW were conducted tions of sex differences in NSW, heifers always had nu- in Genstat using linear mixed models and the REML merically greater temperament scores than did steers, Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 6. 1456 Cafe et al. but the differences were small and rarely significant. FS and CS over Time The effect of sex on FS was not significant for either breed (all P ≥ 0.09). Brahman heifers had greater CS FS in NSW. Means for all 14 FS measurements on than steers during backgrounding (2.15 vs. 1.98, SED NSW cattle are shown graphically in Figure 1a, with = 0.079, P = 0.037) and in the feedlot (1.59 vs. 1.45, means and SD of a representative 8 (selected to provide SED = 0.066, P = 0.045); no significant effect of sex on an even spread over time) presented in Table 2. A sig- CS was observed in the Angus cattle (all P ≥ 0.18). Be- nificant effect of day of measurement on FS (P < 0.001) cause of the lack of interactions between temperament was observed in both breeds. and these effects, further discussion on temperament is In the Brahmans, FS decreased during background- made without reference to them. ing (FS 1 = 2.1 to FS 5 = 1.6 m/s, SED = 0.05 m/s, P Figure 1. Mean (±SEM) a) flight speed (FS) and b) crush score (CS) for Angus (●, ○) and Brahman (■, □) cattle in the New South Wales experiment during backgrounding (solid symbols) and finishing in a feedlot (open symbols). Time of management, ultrasound scan (Scan), and tissue (Biopsy) and blood (Bleed) sampling events and are also shown. Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 7. Cattle temperament and productivity 1457 1 Table 2. Mean (±SD) and timing of a spread of individual flight speed (FS, m/s) measurements taken during backgrounding or at the feedlot, and correlations between FS measurements for the Brahman (below diagonal, n = 164) and Angus (above diagonal, n = 49) cattle in the New South Wales experiment Item FS 1 FS 2 FS 3 FS 4 FS 6 FS 9 FS 12 FS 14 Brahman 2.1 ± 0.99 2.0 ± 0.74 1.8 ± 0.75 1.5 ± 0.74 2.4 ± 0.78 2.1 ± 0.92 1.9 ± 0.92 2.1 ± 0.77 Angus 1.3 ± 0.44 1.3 ± 0.53 1.5 ± 0.53 1.0 ± 0.42 2.0 ± 0.51 1.9 ± 0.62 2.0 ± 0.62 2.0 ± 0.49 Day2 0 30 91 126 182 203 231 252 FS 1   0.26† 0.43* 0.50* 0.33* 0.28† 0.26† 0.29† FS 2 0.65*   0.35* 0.54* 0.33* 0.15 −0.06 0.15 FS 3 0.55* 0.67*   0.70* 0.19 0.27† 0.12 0.25 FS 4 0.62* 0.66* 0.70*   0.40* 0.29† 0.14 0.25 FS 6 0.48* 0.52* 0.54* 0.53*   0.51* 0.41* 0.30† FS 9 0.44* 0.45* 0.51* 0.52* 0.66*   0.50* 0.41* FS 12 0.37* 0.47* 0.54* 0.48* 0.61* 0.75*   0.49* FS 14 0.36* 0.45* 0.48* 0.50* 0.57* 0.63* 0.71*   1 FS 1 to 5 conducted during backgrounding, and FS 6 to 14 conducted at the feedlot. A subset of 8 FS were chosen to illustrate the range of correlations in the entire set of 14 measurements. 2 Days from first FS measurement. †P < 0.10; *P < 0.05. < 0.001), with no pattern for the SD except that it was < 0.001) and Angus (P = 0.008) cattle. Little change greater at FS1. In the feedlot (where FS was measured was observed in SD over time in either breed. In Brah- as the cattle exited the crush in a straight line, unlike man cattle, correlations ranged from 0.41 to 0.52 (all P backgrounding, where cattle had to turn right at 90° < 0.001). In the Angus cattle, correlations ranged from before measurement), the FS was slightly faster, but −0.02 to 0.55 (P < 0.001 to P < 0.9), with the weakest no consistent change over time was observed in means being those involving the first measurement. or SD. The first feedlot (FS 6) and FS 13 measure- CS in NSW. Means for all CS assessments for the ments (measured after the animals had been scanned NSW cattle are shown graphically in Figure 1b, with and biopsied the previous week) were the greatest (P means and SD for a representative 8 of 17 CS assess- < 0.001). Angus cattle had slower FS than Brahmans. ments (selected to provide an even spread over time) Flight speed in the Angus decreased during back- shown in Table 3. grounding (FS 1 = 1.3 to FS 5 = 1.1 m/s, SED = 0.07, A significant effect of day of assessment was ob- P < 0.001), with no change in the SD over time. Flight served. In the Brahman cattle, CS decreased during speed of the Angus cattle was also faster in the feedlot, backgrounding (CS 1 = 2.5 to CS 6 = 1.7, SED = 0.07, again with no pattern of change over time in means or P < 0.001), but the SD were variable. The second as- SD. The fastest FS was FS 13 (P < 0.001), measured sessment (CS 2), when the animals were ultrasound after the animals had been scanned and biopsied the scanned for the first time, was the greatest (P < 0.001). previous week. At the feedlot, CS were less in the second half of the Correlations for 8 of 14 FS measurements (selected feeding period, except for CS 15, which was assessed to provide an even spread over time) are presented in during ultrasound scanning (P < 0.001). In Angus cat- Table 2. The moderate to high correlations were all tle, CS decreased slightly during backgrounding (CS 1 significant (all P < 0.001) for the Brahman cattle, and = 1.6 to CS 5 = 1.4, SED = 0.10, P < 0.001), except were greatest between measurements from the same lo- for CS 2, taken during ultrasound scanning, which was cation (i.e., backgrounding or finishing). For the Angus the greatest. The SD was greater for CS 1 and CS 2. cattle, correlations were not as strong; 90% of the cor- Crush scores were less at the feedlot, and again, they relations of FS at the same location were significant were slightly less in the second half of the feeding pe- (P < 0.001 to P < 0.21), but only approximately 30% riod. No pattern of change in SD in the feedlot was at different locations were significant (P < 0.001 to P observed for either breed. < 0.95). For both breeds, correlations decreased with Correlations for 8 of the 17 CS assessments are pre- increasing time between measures. sented in Table 3. Overall, correlations for CS were less FS in WA. Flight speed was measured 4 times in than for FS, and were less in the Angus cattle. In the the WA herd: 86, 155, 210, and 282 d after the begin- Brahmans, correlations were generally greater between ning of backgrounding. Means and SD were 1.7 ± 0.45, CS assessments at the same location, but most correla- 1.5 ± 0.52, 1.5 ± 0.52, and 1.5 ± 0.52 m/s for Brah- tions between backgrounding and feedlot assessments mans and were 1.7 ± 0.30, 1.4 ± 0.38, 1.5 ± 0.41, and of CS were also significant. Correlations were not as 1.4 ± 0.45 m/s for Angus on these respective days. The strong for the Angus cattle, with 30% of those for the effect of day of measurement was significant, with the same location being significant and 25% for different second measurement being slowest in both Brahman (P locations being significant. Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 8. 1458 Cafe et al. Table 3. Mean (±SD) and timing of a spread of individual crush score1 (CS, score 1 to 5) assessments taken dur- ing backgrounding or at the feedlot, and correlations between CS assessments for the Brahman (below diagonal, n = 164) and Angus (above diagonal, n = 49) cattle in the New South Wales experiment Item CS 1 CS 2 CS 3 CS 4 CS 7 CS 10 CS 14 CS 17 Brahman 2.5 ± 0.93 2.8 ± 0.73 1.5 ± 0.63 2.0 ± 0.84 1.5 ± 0.59 1.6 ± 0.59 1.4 ± 0.57 1.4 ± 0.60 Angus 1.6 ± 0.64 2.1 ± 0.69 1.3 ± 0.48 1.3 ± 0.48 1.2 ± 0.37 1.2 ± 0.43 1.1 ± 0.33 1.1 ± 0.35 Day2 30 71 91 126 182 203 231 252 CS 1   0.24† 0.31* 0.24† 0.27† −0.10 0.13 0.25† CS 2 0.20*   0.44* 0.25† 0.04 0.09 0.06 0.22 CS 3 0.40* 0.19*   0.28† 0.37* 0.08 0.25† 0.32* CS 4 0.37* 0.21* 0.37*   0.03 0.08 0.25† 0.19 CS 7 0.20* 0.08 0.26* 0.15   0.26† 0.17 0.29* CS 10 0.32* 0.23* 0.36* 0.27* 0.35*   0.37* 0.04 CS 14 0.26* 0.18* 0.33* 0.22* 0.35* 0.47*   0.20 CS 17 0.19* 0.21* 0.24* 0.32* 0.34* 0.48* 0.38*   1 CS 1 to 6 conducted during backgrounding, and CS 7 to 17 conducted at the feedlot. A subset of 8 CS were chosen to illustrate the range of correlations in the entire set of 17 measurements. 2 Days from first flight speed measurement. †P < 0.10; *P < 0.05. Average FS and CS ships were weaker for Angus cattle; only the correlation between average FS and CS during backgrounding was Averages of FS and CS assessed during background- significant (r = 0.39, P = 0.006). ing and at the feedlot in NSW, and average FS assessed during backgrounding and at the feedlot in WA are FS and Productivity Traits presented in Table 4. In line with the changes over time described above, average feedlot FS was faster than av- The relationships between average FS and produc- erage backgrounding FS in NSW (P < 0.001), whereas tion, efficiency, carcass traits, and objective meat qual- CS was less at the feedlot (P < 0.001). In WA Brah- ity traits are indicated by estimates of the linear covari- mans, FS was slightly slower at the feedlot than during ates in Tables 6, 7, and 8; quadratic relationships are backgrounding (P = 0.024). In WA Angus cattle, no reported only when significant. difference was observed in FS measured during back- Growth, Intake, and Efficiency. Effects of av- grounding or at the feedlot (P = 0.35). erage backgrounding and average feedlot FS on produc- Correlations between backgrounding and feedlot aver- tion and efficiency traits are presented in Table 6. For ages within assessment type (e.g., 0.69 for FS and 0.41 NSW Brahmans, cattle with faster backgrounding FS for CS in the NSW Brahmans; Table 5) were greater had reduced BW at all times during the experiment (all than for any individual pair of measurements from dif- P ≤ 0.046) and reduced ADG during backgrounding ferent locations. This indicates that the averages gave a (P = 0.043) and finishing (P = 0.001). The quadratic more accurate assessment of both FS and CS than did relationship between background FS and background any single measure. Similarly, in WA the correlation ADG was stronger (P = 0.009) than the linear relation- between the average backgrounding and feedlot FS was ship, with most of the decline in ADG occurring for 0.66 (P < 0.001) for Brahmans and 0.51 (P = 0.02) for background FS of >2.5 m/s. Increasing background FS Angus. was also related to reduced DMI (P = 0.012) and less In the NSW Brahmans, correlations between average time spent eating (P = 0.046). Increasing feedlot FS FS and CS, ranging from 0.41 to 0.49 (all P < 0.001), was related to reduced BW at the midpoint (P = 0.040) provide an indication that the 2 different temperament and at the end of the feedlot period (P = 0.030) and to measurements ranked the cattle similarly. The relation- decreased feedlot ADG (P = 0.007). Increasing feedlot Table 4. Average flight speed (m/s) and crush score (1 to 5) during backgrounding and feedlot finishing in Brah- man and Angus cattle in the New South Wales (NSW) and Western Australia (WA) experiments Flight speed Crush score Item NSW Brahman NSW Angus WA Brahman WA Angus NSW Brahman NSW Angus Location                Background 1.84 1.24 1.61 1.49   2.12 1.56  Feedlot 2.09 1.97 1.54 1.42   1.58 1.21 SED 0.042 0.063 0.028 0.080   0.034 0.04 P-value <0.001 <0.001 0.024 0.35   <0.001 <0.001 Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 9. Cattle temperament and productivity 1459 Table 5. Correlations between average flight speed (FS, m/s) and crush score (CS, score 1 to 5) determined during backgrounding and feedlot finishing in Brahman (be- low diagonal, n = 164) and Angus (above diagonal, n = 49) cattle in the New South Wales experiment Item Background FS Feedlot FS Background CS Feedlot CS Background FS   0.42* 0.39* 0.23 Feedlot FS 0.69*   0.08 0.24 Background CS 0.49* 0.41*   0.62* Feedlot CS 0.42* 0.41* 0.58*   *P < 0.05. FS was also associated with reduced time spent eating dency toward reduced BW at the end of the feedlot (P = 0.040) and tended to reduce DMI (P = 0.07). In period (P = 0.06). the WA Brahman cattle, increasing background FS was Carcass Characteristics. Effects of average associated with reduced BW at the beginning and end backgrounding and feedlot FS on carcass traits are pre- of the feedlot period (both P = 0.008), reduced back- sented in Table 7. In the NSW Brahman cattle, in- ground ADG (P = 0.025), and a tendency for reduced creasing background FS was associated with significant feedlot ADG (P = 0.07). In addition, increasing feedlot reductions in carcass weight (P = 0.001), rib fat (P = FS was associated with reduced BW at the beginning 0.016), and ultimate pH (P = 0.014), and an increase (P = 0.023) and end (P = 0.015) of the feedlot period in the temperature at which the carcass reached pH 6 and with a tendency toward reduced feedlot ADG (P (P < 0.001). Carcass weight also tended to be reduced = 0.07). with increasing feedlot FS (P = 0.09). In the WA Brah- In the NSW Angus cattle, increasing background FS man cattle, increasing background FS was associated was related to lighter BW at the beginning of back- with reduced carcass weight (P = 0.013), reduced LLM grounding (P = 0.045), and increasing feedlot FS was area (P = 0.035), and darker meat color (P = 0.028). related to reduced time spent eating (P = 0.038) and It also tended to influence carcass pH decline, with the a tendency toward a decreased feed conversion ratio carcasses tending to take longer to reach pH 6 (P = (FCR; P = 0.07). In WA Angus cattle, no significant 0.07) and at reduced carcass temperatures (P = 0.09). relationships were observed between background FS Increased feedlot FS also tended to be associated with and any growth trait, but increased feedlot FS was as- reduced carcass weight (P = 0.09). For the WA Brah- sociated with reduced background ADG (P = 0.003), man cattle, the quadratic relationship between FS and reduced BW at feedlot entry (P = 0.018), and a ten- carcass weight was slightly stronger than the linear re- Table 6. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot finishing on live traits in Brahman and Angus cattle in the New South Wales (NSW) and Western Australia (WA) experiments Background FS Feedlot FS Breed Site Trait Slope SE P-value Slope SE P-value Brahman NSW (n = 164) Beginning background BW, kg −6.9 3.45 0.046           Beginning feedlot BW, kg −11.5 3.61 0.002             Mid feedlot BW, kg −19.1 4.48 <0.001   −9.3 4.50 0.040     End feedlot BW, kg −21.0 5.047 <0.001   −11.1 5.07 0.030     Background ADG, kg −0.02 0.011 0.043             Feedlot ADG, kg −0.08 0.025 0.001   −0.07 0.025 0.007     Feedlot DMI, kg of DM/d −0.37 0.146 0.012   −0.26 0.143 0.07     Feeding time, min/d −4.7 2.33 0.046   −4.68 2.259 0.040   WA (n = 173) Beginning feedlot BW, kg −14.3 5.26 0.008   −11.6 5.06 0.023   End feedlot BW, kg −20.9 7.78 0.008   −18.3 7.45 0.015     Background ADG, kg −0.05 0.020 0.025             Feedlot ADG, kg −0.10 0.056 0.07   −0.08 0.046 0.07 Angus NSW (n = 49) Beginning background BW, kg −23.0 11.08 0.045           Feedlot FCR,1 kg of DM/kg of gain         −1.5 0.81 0.07     Feeding time, min/d         −17.6 8.19 0.038   WA (n = 20) Beginning feedlot BW, kg         −34.8 12.70 0.018   End feedlot BW, kg         −27.6 13.13 0.06     Background ADG, kg         −0.16 0.043 0.003 1 FCR = feed conversion ratio. Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 10. 1460 Cafe et al. Table 7. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot finishing on carcass traits in Brahman and Angus cattle in the New South Wales (NSW) and Western Australia (WA) experiments Background FS Feedlot FS Breed Site Trait Slope SE P-value Slope SE P-value Brahman NSW (n = 163) Carcass wt, kg −9.9 2.92 0.001   −5.0 2.93 0.09   Rib fat,1 mm −5.7 0.23 0.016             Ultimate pH1 −0.01 0.006 0.014             Temperature at pH 6, °C 0.81 0.068 <0.001           WA (n = 143) Carcass wt, kg −9.7 3.85 0.013   −6.4 3.73 0.09   LLM area,1 cm2 −2.67 1.20 0.035             Meat color score1 0.48 0.213 0.028             Temperature at pH 6, °C −1.1 0.64 0.09             Time to pH 6, h 0.22 0.120 0.07         Angus NSW (n = 49) Carcass wt, kg −27.0 14.47 0.07           Rump fat, mm −4.0 2.03 0.06             Time to pH 6, h 0.56 0.093 <0.001           WA (n = 17) LLM area,1 cm2         −5.9 1.61 0.005   Marble score1 −79.4 28.27 0.020         1 Meat Standards Australia (2009) grading data, where marble score is from 100 to 1,100 in increments of 10; meat color score is from 1 (lightest) to 9 (darkest), and ultimate pH is the pH at grading. LLM = musculus longissimus lumborum. lationship, with most of the decline in carcass weight Objective Meat Quality. Effects of average back- occurring for cattle with a background or feedlot FS of grounding and feedlot FS on objective meat quality >2 m/s (P = 0.012 and 0.06, respectively). traits are presented in Table 8. In the NSW Brahmans, In the NSW Angus cattle, increasing background FS increasing background FS was related to increased cook- tended to reduce carcass weight (P = 0.07) and rump ing loss in TS 1-d aged LLM (P = 0.036), and increas- fat (P = 0.06) and significantly increased the time to ing feedlot FS tended to be related to increased shear reach pH 6 (P < 0.001), but no significant effects of force (SF) of AT 1-d aged LLM (P = 0.05). In the WA feedlot FS on carcass traits were observed. In the WA Brahmans, increasing background FS was related to in- Angus cattle, increasing background FS was associated creased SF of TS 7-d aged LLM (P = 0.044), increased with reduced marbling score (P = 0.020), and increas- compression in AT 1-d aged STN (P = 0.043), and in- ing feedlot FS was associated with reduced LLM area creased cooking loss (P = 0.023) and darker meat color (P = 0.005). (P = 0.006) of AT 7-d aged LLM. Increasing feedlot Table 8. Significant effects of average flight speed (FS, m/s) determined during either backgrounding or feedlot finishing on objective meat quality traits in Brahman and Angus cattle in the New South Wales (NSW) and West- ern Australia (WA) experiments Background FS Feedlot FS Breed Site Trait1 Slope SE P-value Slope SE P-value Brahman NSW (n = 161) AT 1-d aged LLM SF, N         4.2 2.10 0.050   TS 1-d aged LLM cook, % 0.84 0.396 0.036           WA (n = 137) TS 7-d aged LLM SF, N 4.3 2.11 0.044   4.2 1.86 0.027   AT 1-d aged STN comp, N 1.4 0.70 0.043             AT 1-d aged LLM cook, %         0.65 0.292 0.028     AT 7-d aged LLM cook, % 0.65 0.284 0.023             TS 7-d aged LLM cook, %         0.70 0.286 0.016     AT 1-d aged LLM color L*         −0.84 0.461 0.07     AT 7-d aged LLM color L* −1.6 0.56 0.006   −1.1 0.53 0.034 Angus NSW (n = 48) AT 1-d aged LLM SF, N 11.2 4.32 0.013           AT 7-d aged LLM SF, N 7.3 4.04 0.08           WA (n = 16) AT 7-d aged LLM SF, N 15.8 7.70 0.07           TS 1-d aged LLM SF, N 13.1 5.89 0.050   6.6 2.65 0.032     AT 7-d aged LLM comp, N 4.6 2.18 0.07             AT 1-d aged STN comp, N 7.1 3.06 0.045             TS 1-d aged LLM cook, %         2.4 0.92 0.027     AT 7-d aged LLM pH 0.10 0.042 0.049         1 AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum; STN = musculus semitendinosus; SF = shear force; comp = compression; cook = cooking loss; color L* = CIELAB color scale, where 0 = dark and 100 = light. Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 11. Cattle temperament and productivity 1461 Table 9. Significant effects of average crush score (CS, score 1 to 5) determined during either backgrounding or feedlot finishing on live traits in Brahman and Angus cattle in the New South Wales experiment Background CS Feedlot CS Breed Trait Slope SE P-value Slope SE P-value Brahman (n = 164) Beginning background BW, kg         −11.2 5.25 0.034 Mid feedlot BW, kg −13.8 6.17 0.027   −30.5 6.80 <0.001   End feedlot BW, kg −11.9 6.94 0.09   −30.0 7.83 <0.001   Background ADG, kg −0.04 0.015 0.016           Feedlot ADG, kg         −0.12 0.040 0.003   Feedlot DMI, kg of DM/d         −0.75 0.226 0.001 Angus (n = 49) Feed intake per session, kg of DM −0.74 0.302 0.020   −0.98 0.484 0.05 Feeding sessions, No./d 3.7 1.46 0.016   4.9 2.32 0.043 FS was also related to increasing SF of TS 7-d aged sions increasing with both increasing background (P LLM (P = 0.027) and to increased cooking loss in AT = 0.016) and feedlot (P = 0.043) CS. Intake per ses- 1-d aged LLM (P = 0.028) and TS 7-d aged LLM (P = sion decreased (P = 0.020) with increasing background 0.016). Increasing feedlot FS was also related to darker CS and tended to decrease (P = 0.05) with increasing meat color in AT 7-d aged LLM (P = 0.034) and tend- feedlot CS. ed to be related to darker meat color in AT 1-d aged Carcass Characteristics. Effects of average LLM (P = 0.07). backgrounding and feedlot CS on carcass traits in the In the NSW Angus cattle, increasing background FS NSW experiment are presented in Table 10. In Brah- was associated with increased SF in AT 1-d aged LLM man cattle, increasing feedlot CS was associated with a (P = 0.013) and a tendency toward increased SF in AT reduction in carcass weight (P < 0.001), and increasing 7-d aged LLM (P = 0.08). No significant relationships background and feedlot CS were associated with a re- were observed between feedlot FS and objective meat duction in rib fat (P = 0.012 and 0.017). No significant quality traits. In WA Angus cattle, increasing FS tend- effects of CS were observed on carcass traits in the ed to be associated with increased SF in AT 7-d aged Angus cattle. LLM (P = 0.07), increased SF in TS 1-d aged LLM Objective Meat Quality. Effects of average back- (P = 0.05), and increased compression in AT 7-d aged grounding and feedlot CS on objective meat quality LLM (P = 0.07). It was also related to increased com- traits in the NSW experiment are presented in Table pression in AT 1-d aged STN (P = 0.045) and increased 10. In Brahman cattle, as background CS increased, pH in the laboratory sample of the AT 7-d aged LLM so did SF in TS 7-d aged LLM (P = 0.048) and com- (P = 0.049). Increasing feedlot FS was also associated pression in AT 1-d aged LLM (P = 0.019). As feedlot with increased SF (P = 0.032) and cooking loss (P = CS increased, SF in AT 1-d aged LLM increased (P = 0.027) in TS 1-d aged LLM. 0.024) and tendencies were observed for increased SF with increasing feedlot CS in AT and TS 7-d aged LLM CS and Productivity Traits (P = 0.08 and 0.09). Cooking loss in AT 1-d aged LLM also increased (P = 0.001) with increasing feedlot CS in Growth, Intake, and Efficiency. Effects of the Brahman cattle. average backgrounding and feedlot CS on production In Angus cattle, greater background CS was associ- and feed efficiency traits in the NSW experiment are ated with increased compression in AT 7-d aged LLM presented in Table 9. In Brahman cattle, increasing (P = 0.04) and with a tendency toward increased SF background CS was associated with reduced mid feed- and compression in AT 1-d aged LLM (P = 0.05 and P lot BW (P = 0.027), reduced background ADG (P = = 0.06). Increasing feedlot CS led to increased SF (P 0.016), and a tendency toward reduced BW (P = 0.09) = 0.047) and compression (P = 0.045) in AT 1-d aged at the end of the feedlot period. The relationship be- LLM and to a tendency toward increased SF in AT 7-d tween background CS and feedlot ADG and DMI was aged LLM (P = 0.09). quadratic, with most of the decline in carcass ADG and intake occurring in cattle with a background CS DISCUSSION >3 (P = 0.006 and 0.031, respectively). Increased feed- lot CS was related to reduced BW at the beginning of This study shows that the temperament of cattle, as backgrounding (P = 0.034), mid feedlot (P < 0.001), assessed by FS and CS, was persistent over time, and and at the end of the feedlot period (P < 0.001), and that cattle with faster FS or greater CS (flightier tem- to reduced feedlot ADG (P = 0.003) and DMI (P = peraments) had inferior performance across a compre- 0.001). hensive range of beef production traits. Flight speed (or In Angus cattle, increasing CS was related only to its inverse, flight time) and CS are simple to conduct feeding behavior, with the number of daily feeding ses- on farm, and their use is encouraged by various Aus- Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 12. 1462 Cafe et al. Table 10. Significant effects of average crush score (CS, score 1 to 5) during either backgrounding or feedlot finish- ing on carcass and meat quality traits in Brahman and Angus cattle in the New South Wales experiment Background CS Feedlot CS Breed Trait1 Slope SE P-value Slope SE P-value Brahman (n = 161) Carcass wt, kg         −16.6 4.50 <0.001 Rib fat, mm −0.77 0.302 0.012   −0.90 0.370 0.017   AT 1-d aged LLM SF, N         7.6 3.34 0.024   AT 7-d aged LLM SF, N         5.5 3.10 0.08   TS 7-d aged LLM SF, N 1.5 0.74 0.048   1.6 0.97 0.09   AT 1-d aged LLM comp, N 0.8 0.35 0.019           AT 1-d aged LLM cook, %         0.98 0.293 0.001 Angus (n = 48) AT 1-d aged LLM SF, N 9.4 4.64 0.05   16.0 7.78 0.047 AT 7-d aged LLM SF, N         12.1 6.98 0.09   AT 1-d aged LLM comp, N 1.4 0.72 0.06   2.4 1.16 0.045   AT 7-d aged LLM comp, N 1.1 0.53 0.04         1 AT = Achilles-suspended side; TS = tenderstretched side; LLM = musculus longissimus lumborum; SF = shear force; comp = compression, cook = cooking loss. tralian beef cattle breed societies [for example, Limou- handling events, the cattle showed calmer behavioral sin (CS and pen score), Angus (CS and flight time), responses, presumably as they habituated to handling. and Brahman (flight time); http://breedplan.une.edu. This is consistent with results for repeated tempera- au] to allow selection for a calmer temperament or do- ment assessments reported by other authors (Burrow cility. Despite this, still relatively few published stud- and Dillon, 1997; Curley et al., 2006; Kilgour et al., ies have described relationships between temperament 2006; Petherick et al., 2009a). The day of measurement and commercially important traits, and the biological effect, significant for both FS and CS, can be attrib- mechanisms that underpin these associations are not uted, at least in part, to the fact that on some days, well understood (Ferguson et al., 2006). the data collection procedures involved closer and more Regular FS and CS measurements were taken prolonged handling. throughout the NSW experiment to study the consis- Correlations between repeated measures for FS and tency of the measures over time with changes in loca- CS in Brahman cattle were usually significant and were tion and during various husbandry and sample collec- greater than for Angus cattle. The variation was also tion procedures. Management was more intensive than consistently greater for both FS and CS at each as- for commercial herds because of the data and sample sessment in the Brahman than in the Angus cattle, collection required for other aspects of the experiment, indicating that the Brahman cattle had greater indi- but all handling of the animals was conducted as calm- vidual variation in temperament than the Angus cattle ly as possible. in the present study. This finding would account for the poorer correlations in Angus cattle among individ- Temperament over Time ual measures of FS and CS, and between averages for FS and CS. For both breeds, the strength of correla- The decreased average feedlot vs. backgrounding CS tions declined over time, indicating small, consistent in NSW, and slower feedlot vs. backgrounding FS in changes over time. Because the behavioral response is WA indicate that the general response of the cattle to a combination of genetic and environmental influences handling declined over the duration of the experiment. (Boissy et al., 2005), small changes over time would be In contrast, FS in NSW was faster at the feedlot than expected. In this regard, it was also notable that the during backgrounding, where FS was measured after largest decline in the strength of correlations occurred the cattle had turned 90° after exiting the chute. The with the change in location between backgrounding and differences in the way in which backgrounding FS was feedlot finishing. It is also important to note that the measured resulted in slower speeds, but nonetheless use of average values for FS and CS resulted in greater provided a meaningful measure of FS. correlations, indicating that average measures provided Much of the decline in the mean and variation within a more reliable assessment of cattle temperament than FS was seen after the first 3 measurements, suggest- did any single measure, as suggested by Grandin (1993) ing that the variation between animals stabilized after and Burrow and Dillon (1997). some initial familiarization with handling and the fa- cilities. The mean and variation for CS were also great- Relationships Between Temperament er during initial measurements, after which both were and Other Traits again very consistent for both breeds. Hence, the pat- tern of change in mean and variation for both FS and Relationships between temperament and other pro- CS indicated that after a small number of consistent duction traits were assessed using average FS and CS Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 13. Cattle temperament and productivity 1463 during backgrounding or finishing. Where effects of of time spent eating. A similar but slightly smaller ef- temperament were significant, cattle with greater FS or fect was obtained using feedlot FS as the measure of CS grew more slowly, produced smaller carcasses with temperament. Increasing CS was also associated with a less fat cover, and had darker meat that was greater reduced feed intake (of 750 g of DM/d with each unit in SF and compression. It is important to note that increase in feedlot CS) in the NSW Brahman cattle. all significant effects of a more reactive temperament These effects were not accounted for entirely by the (increasing FS or CS) on economically significant traits BW differences at the beginning of the feedlot period, were detrimental. and they remained or tended to remain evident when Faster FS was associated with reduced BW and feedlot entry BW was fitted as a covariate. However, growth rates throughout the experiment in the Brah- there was no evidence of FS or CS being related to dif- man cattle in NSW and WA. Estimates of the reduc- ferences in FCR or NFI in the Brahman cattle. Togeth- tion in feedlot exit BW were 20.0 and 20.9 kg with each er, this suggests that temperament plays a significant 1 m/s increase in background FS in the NSW and WA role in controlling feed intake and time spent eating, Brahman cattle, respectively. Increasing CS had similar but that it has lesser effects on efficiency of utilization effects in the NSW Brahmans, with a 1-score increase in of feed; hence, poor temperament reduces DMI and background CS leading to an 11.9-kg decrease in feedlot ADG through behavioral rather than metabolic mecha- exit BW. Carcass weights were reduced with increasing nisms. This conclusion is in agreement with recent work FS in the Brahmans, by 9.9 kg in NSW and 9.7 kg in by Nkrumah et al. (2007) and Elzo et al. (2009), who WA for each 1 m/s increase in FS. The NSW Brahmans found that young cattle of mixed breeds with faster also had a 16.6-kg reduction in carcass weight per unit feedlot FS had less feedlot DMI but showed no differ- increase in feedlot CS. There was also some indication ence in FCR or NFI. of reduced carcass fatness and LLM area with increas- In the NSW herd, temperament had less effect on ing FS, and of reduced carcass fatness with increasing feed intake in Angus than in Brahman cattle. However, CS, in the NSW Brahmans. The inclusion of carcass in the Angus cattle, each meter per second increase in weight as a covariate in the statistical models showed feedlot FS was associated with a 17.6 min/d reduction that these differences in composition were mostly ex- in feeding time and a tendency toward reduced FCR by plained by the differences in carcass weight. In the An- 1.5 kg of DM/kg of BW gain. This reduction in FCR is gus cattle, increasing FS also tended to reduce BW and the only result relating to poorer temperament within growth rate, but the relationships were much weaker the present study that might be considered beneficial. than in Brahmans. In line with the weak trends toward Although caution is required, because of the P-value lighter BW with increasing FS in the Angus cattle, ten- of 0.07, it is possible that among the Angus cattle, a dencies for reductions in BW-related carcass traits were slightly decreased intake with increasing FS allowed for observed. However, no significant relationships were ob- more efficient digestion (Herd et al., 2004). served between CS and BW-related traits in the Angus The determinants of eating quality of the meat are cattle in the NSW herd. complex and multifactorial, and pre- and postmortem Previously reported results for relationships between events can have major effects on beef tenderness (Mal- cattle temperament and growth have been variable. tin et al., 2003; Ferguson and Warner, 2008). Cattle Slower growth rates have been reported in cattle with temperament is assumed to be related to stress respon- faster FS, greater CS, or both in studies conducted un- siveness, and it is likely that the stress response to han- der more intensive management systems (Burrow and dling and transport is greater in temperamental cattle, Dillon, 1997; Voisinet et al., 1997b; Müller and von resulting in depletion of muscle glycogen before slaugh- Keyserlingk, 2006; Behrends et al., 2009). Others have ter and hence reduced meat quality because of greater found little relationship between temperament and carcass pH and the associated darker color (Ferguson et growth rates in herds in which the ranges in FS and CS al., 2006). In the present study, increasing FS or CS was were small and cattle were generally docile (Graham et related to darker LLM meat color and increased muscle al., 2001; Elzo et al., 2009). Our results are consistent pH, shear force, compression, and cooking loss, effects with the above studies in that the Angus cattle were all considered detrimental to the eating quality of beef generally more docile than the Brahmans, and greater (Perry et al., 2001b). effects of temperament on growth were observed in the The relationships between temperament and meat Brahmans. quality were strongest for the WA cattle. Differences It has previously been postulated that cattle with between experimental sites could, at least in part, be a more reactive temperament may grow more slowly due to processing differences resulting in differences in because of the greater energy expenditure associated postmortem pH or temperature declines, as discussed with, for example, more vigilant behavior, resulting in by Cafe et al. (2010b). Greater effects of temperament poorer FCR or net feed intake (NFI; Burrow and Dil- on meat quality traits were evident in both breeds in lon, 1997; Petherick et al., 2002). In the NSW Brah- WA, where the carcasses had much faster pH declines mans, each meter per second increase in background FS and slower cooling rates than in NSW. Relationships was associated with a reduction in feed intake of 370 g between temperament and shear force tended to be of DM/d, and a reduction of 4.7 min/d in the amount less significant with 7-d aging. This may indicate that Downloaded from jas.fass.org at UNESP on July 20, 2011
  • 14. 1464 Cafe et al. temperament was related to variation in tenderness be- LITERATURE CITED cause of factors other than proteolysis. An unexpected result was that the Angus cattle, with smaller numbers AUS-MEAT. 2007. AUS-MEAT National Accreditation Standards. of animals and less variation in temperament, showed 2007 Edition. AUS-MEAT Ltd., Brisbane, Queensland, Aus- stronger effects of temperament on SF than did the tralia. Barendse, W. J. 2002. DNA markers for meat tenderness. The Com- Brahman cattle in both herds. monwealth Scientific and Industrial Research Organization, Overall, most relationships with temperament were The State of Queensland through its Department of Primary linear, indicating that selection of cattle with calmer Industries, The University of New England, The State of New temperaments, and not only culling of cattle with the South Wales through its Department of Agriculture, and Meat most reactive temperaments, can improve productivity and Livestock Australia Limited, assignees. Int. Patent No. W0 02/064820. and the safety and welfare of cattle and can improve Barendse, W., B. E. Harrison, R. J. Bunch, and M. B. Thomas. safety for human handlers. Other studies have shown 2008. Variation at the calpain 3 gene is associated with meat both linear (Burrow and Dillon, 1997) and quadratic tenderness in zebu and composite breeds of cattle. BMC Gen- (Müller and von Keyserlingk, 2006) relationships be- et. 9:41. tween temperament and productivity traits. However, Behrends, S. M., R. K. Miller, F. M. Rouquette Jr., R. D. Randel, B. G. Warrington, T. D. A. Forbes, T. H. Welsh, H. Lippke, J. differences in results between studies may simply de- M. Behrends, G. E. Carstens, and J. W. Holloway. 2009. Re- pend on the extent to which extremes of temperament lationship of temperament, growth, carcass characteristics and exist within the herd, with the presence of extremely tenderness in beef steers. Meat Sci. 81:433–438. reactive animals likely to result in nonlinear relation- Bindon, B. M. 2001. Genesis of the Cooperative Research Centre for ships. the Cattle and Beef Industry: Integration of resources for beef quality research (1998–2000). Aust. J. Exp. Agric. 41:843– Flight speed and CS were consistent measures of 853. temperament during the experiment, even with varying Boissy, A., A. D. Fisher, J. Bouix, G. N. Hinch, and P. Le Neindre. management at the time of measurement. The earlier 2005. Genetics of fear in ruminant livestock. Livest. Prod. Sci. (backgrounding) assessments had stronger and more 93:23–32. frequent relationships with productivity and carcass Burrow, H. M. 1997. Measurements of temperament and their rela- tionships with performance traits of beef cattle. Anim. Breed. traits, but both the backgrounding and the finishing Abstr. 65:477–495. assessments showed significant relationships with meat Burrow, H. M., and R. D. Dillon. 1997. Relationships between tem- quality. Other authors have reported weak to no re- perament and growth in a feedlot and commercial carcass traits lationships between temperament measurements taken of Bos indicus crossbreds. Aust. J. Exp. Agric. 37:407–411. later in life and growth and meat quality (Behrends et Burrow, H. M., G. W. Seifert, and N. J. Corbet. 1988. A new tech- nique for measuring temperament in cattle. Proc. Aust. Soc. al., 2009). In the present study, the uniform measure- Anim. Prod. 17:154–157. ment procedures and frequent handling of the cattle Cafe, L. M., B. L. McIntyre, D. L. Robinson, G. H. Geesink, W. throughout the experiment for intensive data and sam- Barendse, and P. L. Greenwood. 2010a. Production and pro- ple collection, and the young age of the cattle, may cessing studies on calpain-system gene markers for tenderness have allowed variation in temperament to continue to in Brahman cattle: 1. Growth, efficiency, temperament and car- cass characteristics. J. Anim. Sci. 88:3047–3058. be detected at a later stage of production. Cafe, L. M., B. M. McIntyre, D. L. Robinson, G. H. Geesink, W. Barendse, D. W. Pethick, J. M. Thompson, and P. L. Green- Conclusions wood. 2010b. Production and processing studies on calpain-sys- tem gene markers for tenderness in Brahman cattle 2. Objective In this study, cattle with a calmer temperament, as meat quality. J. Anim. Sci. 88:3059–3069. Curley, K. O., Jr., J. C. Paschal, T. H. Welsh Jr., and R. D. Randel. measured by reduced FS and CS, had superior perfor- 2006. Technical note: Exit velocity as a measure of cattle tem- mance across a comprehensive range of beef produc- perament is repeatable and associated with serum concentra- tion traits. Repeated assessments of temperament using tion of cortisol in Brahman bulls. J. Anim. Sci. 84:3100–3103. FS and CS were correlated, with the strength of the Elzo, M. A., D. G. Riley, G. R. Hansen, D. D. Johnson, R. O. Myer, correlations declining over time. Correlations between S. W. Coleman, C. C. Chase, J. G. Wasdin, and J. D. Driver. 2009. Effect of breed composition on phenotypic residual feed repeated measurements of FS were greater than be- intake and growth in Angus, Brahman, and Angus × Brahman tween repeated assessments of CS. A stronger relation- crossbred cattle. J. Anim. Sci. 87:3877–3886. ship was observed between temperament and growth Ferguson, D. M., D. Johnston, H. M. Burrow, and A. 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