Bone ossification and carcass characteristics of wethers given silastic implants

containing estradiol
R. A. Field, G. Maiorano, F. C. Hinds, W. J. Murdoch and M. L. Riley
J Anim Sci 1990. 68:3663-3668.

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BONE OSSIFICATION AND CARCASS CHARACTERISTICS

OF WETHERS GIVEN SILASTIC IMPLANTS
CONTAINING ESTRADIOL
R. A. Field', G. MaioranG, F. C. Hinds',
W. J. Murdoch' and M. L. Riley'
University of Wyoming, Laramie 82071
ABSTRACT

Administration of growth promotants with estrogenic activity via hastening closure of

the growth plate could have an economic impact on lambs because closure results in ovine
carcasses being classified as yearlings. Twelve wether lambs approximately 12 mo old
were given silastic implants filled with estradiol-17P and allotted randomly to be
slaughtered 30, 60,Wor 220 d after implantation to determine time of growth plate closure
in relation to date of implanting. Seven comparable wethers not implanted served as
controls. Four wethers implanted at 2 mo of age and slaughtered 220 d later also were
included. Implanted wethers had serum concentrations of estradiol averaging 13.9 pg/ml
over the 2204 implant period; controls averaged 2.7 p g / d Estradiol implants increased
carcass maturity scores but fat deposition was not altered. Growth plate widths decreased
(P < .05) as wethers grew older and implants were in place for longer periods of time.
Metacarpal growth plates in 12-mo-old lambs were completely ossified 220 d after
implanting, but control wethers and wethers implanted at 2 mo maintained growth plate
widths. Neither metacarpal nor metatarsal bone lengths differed (P > .05) between control
and implanted wethers. Growth plate ossification was not complete until 570 d of age in
implanted lambs, even though bone length had stopped increasing by 408 d. Therefore,
even though estradiol and other growth stimulants with estrogenic activity increase rate of
ossification of the metacarpal growth plate, bone length or mature size is not limited. Even
with estrogen implantation, growth plate closure in wethers does not occur until after 450 d
of age, an age older than that at which lambs normally are marketed.
(Key Words: Sheep, Growth, Bones, Estrogens, Maturity).
J. Anim. Sci. 1990. 68:366>3668

of age, whereas a sizeable percentage of

wethers still have break joints at 21 mo of age.
Ossification of growth plates on distal ends Stout (1974) concluded that after 280 d of age
of the metacarpals results in formation of spool break joint maturity scores were decidedly
joints. Ovine carcasses with spool joints rather biased against ewes. Oberbauer et al. (1988)
than break joints on both metacarpals are indicated that metacarpal closure in ewe lambs
classified as yearling mutton or mutton occurred between 360 and 480 d of age,
(USDA, 1982); this classification lowers car- whereas closure in rams exceeded 500 d
cass value. The USDA (1982) reported that
(Oberbauer, 1985).
some ewes have spool joints as early as 9 mo
Because estrogen inhibits cartilage proliferation, promotes mineralization and speeds
ossification (Silbermann, 1983; Van Sickle,
1985). we hypothesized that estradiol-17P
lDept. of ~ n i m .Sci., univ. of wyoming.
*Univmittt degli Shdi del Molise, Pacolttt di A@%
(estradiol) implants in wethers would increase
Dipartimento delle Scienze e Technologi AgriAlimentari,
the rate of ossification of the metacarpal
Vis S. Antonio Dei Lazzari, 86100 Campobasso, Italy.
growth plates and cause formation of spool
Received January 8, 1990.
Accepted March 30, 1990.
joints at an earlier age. If this happened,
lntroductlon

3663

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3664

FIELD ET AL..

carcasses would be classified as yearlings at an

earlier age. Growth promotants with estrogenic
activity also could result in earlier ossification
of the metacarpal growth plate, in earlier
formation of spool joints, and in sheep with
smaller mature sizes. The observation that
some ewe lambs carcasses have spool joints as
early as 9 mo (USDA, 1982) might be linked
to estrogens.
The purpose of this study was to determine
the influence on ossification of the metacarpal
growth plate of silastic implants that release a
constant amount of estradiol. Maturity and
carcass characteristics of implanted and control
wethers also were studied.
Methods

Western grade white-faced wethers, castrated at 2 wk of age, were fed restricted

amounts of a lowenergy diet until they were
approximately 12 mo old. Nineteen of the
wethers weighing 37 f 3 kg were allotted
randomly to control and 30-, 60-, 90- or
220-d slaughter groups and were given ad
libitum access to suncured alfalfa pellets.
Seven wethers not implanted served as
220-d controls; the remaining 12 wethers were
divided into three per group and given silastic
implants filled with estradiol-17P as described
by F. J. Karsch (personal communication).
Inside diameter of the silastic tubing containing estradiol was 3.35 mm, outside diameter
was 4.64 mm, and the tubing was cut into
3.8-cm segments; the ends were plugged with
medical grade elastomer (Karsch et al., 1973).
When ovariectomized ewes were treated with
similar silastic implants, levels of serum
estradiol increased (Karsch et al., 1977), and
the increased levels were maintained for over 1
yr (Legan et al., 1977). Lambs were implanted
in the unwooled axillary area between the right
front leg and the breast. In addition to the 12
wethers, four 2-mo-old grade white-faced
wethers were implanted on the same date to
determine whether growth plate ossification in
young and older lambs would occur at the
same time after implanting. These wethers
were given ad libitum access to sun-cured
alfalfa pellets and housed in the same pen as

kE2D1, doubk antibody, Diagnostic Products, Los

Angeles, CA.

%eelman Instruments, Palo Alto, CA.

the 19 described previously. Our plan was to

slaughter the younger lambs when ossification
of growth plates in the older lambs occurred.
Jugular blood samples were collected from
each control and implanted wether 30, 60,90
and 220 d after implanting. Individual samples
were allowed to clot and the serum was
harvested and frozen at -20C. Serum concentrations of estradiol were quantified by radioimmunoassay using a commercially available
kit3 modified for use with ovine serum as
described below. Duplicate l-ml serum samples were extracted twice in 5 mt of diethyl
ether. Efficiency of extraction was greater than
92%. Ether extract from 1 ml of charcoaltreated serum collected from wethers was
added to each tube containing the standard and
dried under a stream of air. Assay tubes were
incubated overnight at 4'C. Iodinated (1251)
estradiol tracer then was added and tubes were
incubated for 1 h at 25'C. After the addition of
precipitating antisera, tubes were incubated for
10 min at 25'C and centrifuged (1.500 x g, 20
min, 4'C). The Supernatant fraction was
discarded and bound radioactivity was determined using a Beckman Model 4OOO gamma
count&. The assay was sensitive to .78 pg/
tube. The supplier reported that the antisera
against estradiol had negligible (< 6%) crossreactivity with related compounds. Inhibition
curves obtained with standard and serial
dilutions of sera were parallel. Inter- and intraassay coefficients of variation were 11.6% and
3.3%, respectively. Because no differences (P
> .05) in levels of serum estradiol by time after
implanting were detected, only the mean level
of serum estradiol for implanted and control
wethers over all periods is reported.
Control and treated wethers were given ad
libitum access to pelleted, sun-cured alfalfa
until slaughter at 30, 60, 90 or 220 d after
implanting. AU seven older control wethers
and the four wethers implanted at 2 mo of age
were slaughtered 220 d after implanting.
Carcasses were chilled at 2 to 4'C for 24 h
prior to assigning scores for flank color and
texture, break joint maturity, rib shape and
color and overall maturity (USDA, 1982). Fat
depth was measured over the center of the
longissimus muscle at the 12th rib and
bodywall thickness was measured 5 cm from
the lateral edge of the longissimus muscle.
Kidney and kidney fat were removed from the
chilled carcass and weighed so that percentage
of kidney fat could be calculated. Yield grade

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CHARACTERISTICS OF WETHERS GIVEN ESTRADIOL

was calculated using leg conformation, adjusted fat thickness, and actual kidney fat
percentage (USDA, 1982).
Growth plate width on the distal end of the
right metacarpal was measured after silver
nitrate staining (Hinds,1959) of five longitudinal slices 2 mm thick cut on the sagittal plane.
Three width measurements per bone slice were
made on different anatomical locations 114, 1/2
and 314 the distance across the bone slice.
Therefore, 15 measurements on each metacarpal growth plate were averaged. If no epiphyseal cartilage could be detected upon microscopic examination, the plate was considered
ossified.
The left metacarpal and left metatarsal were
cleaned of all connective tissue, measured for
length and weighed. Dry weight was recorded
after 7 d at 100C in a drying oven.
Statistical analysis of data was completed
using General Linear Model procedures (SAS,
1985). A oneway analysis of variance was
used to determine means for the following
treatments: 2-mo-old lambs slaughtered 220 d
after implanting; 12-mo-old lambs slaughtered
30, 60, 90 or 220 d after implanting; and
12-mo-old controls slaughtered 220 d after
their counterparts were implanted. Differences
between means for the six treatments were
tested using pairwise contrasts.
Results and Discussion

Wethers implanted with silastic implants

containing estradiol had serum estradiol concentrations averaging 13.9 f 4.2 p g / d over
the 220-d implant period. Control wethers had
lower (P < .05) concentrations of serum
estradiol, averaging 2.7 f 1.5 pg/ml. We were
surprised to find detectable estradiol in serum
of control wethers, because the adrenals would
not be expected to secrete much. Nevertheless,
serum estradiol values were five times higher
in sem of implanted wethers than in that of
control wethers.
Age, weight and maturity characteristics of
estradiol-implanted and control wethers are
presented in Table 1. Live weight increased
with age in the 12-mo-old lambs, but the
younger, 281dald wethers were as heavy as
12-mo-old lambs implanted for 90 d. Control
wethers were lighter (P < .05) than wethers of
similar age that had been implanted for 220 d.
Muir et al. (1983) cited several reports
showing that diethylstilbestrol, a synthetic

3665

estrogen, increased growth rate in sheep.

Zeranol, also with estrogenic activity, increased rate of gain in sheep (Wilson et al.,
1972; Larson et al., 1983). However, Cheng et
al. (1953) found estrogenic activity in alfalfa
hay and suggested that the presence of
estrogens in hay can explain why control
lambs fed alfalfa hay grew at the same rate as
lambs treated with diethylstilbestrol.
Subjective carcass maturity scores for flank
color and texture, break joint, rib shape and
color, and overall maturity of implanted
wethers increased with age (Table 1). Maturity
characteristics of control wethers scored younger than those of wethers implanted for 220 d,
but 2-mo-old wethers implanted for 220 d
were much younger than all other groups.
Maturity scores of the younger wethers were
typical of those from lambs of approximately
the same age in the study by Ho et al. (1989).
It is clear that estradiol implants increase
carcass maturity scores in older wether carcasses.
Carcass weight and measures of fatness
increased with age and(or) time after implanting (Table 2). Fat depth, bodywall thickness,
kidney fat percentage and yield grade values
indicate that 2-mo-old wethers and 1-yr-old
wethers both became excessively fat after
being fed suncured alfalfa pellets for 220 d.
No differences (P > .05) in fat measures
existed between estradiol-implanted and control wethers of similar age. The findings
support those of Muir et al. (1983) with
diethylstilbestrol and Wilson et al. (1972) with
zeranol, which showed that substances with
estrogenic activity tend not to alter fat deposition in lambs. However, Preston (1975) concluded that estrogen, particularly diethylstilbestrol, increased protein deposition and
decreased fat deposition in lambs. Failure to
find a decrease in fat deposition of estradiolimplanted lambs in the present study may be a
result of the older age of the lambs. Similar
values for longissimus muscle area, leg conformation and quality grade between control
wethers and wethers implanted with estradiol
for 220 d support previous work (Wilson et al.,
1972; Larsm et al., 1983).
Metacarpal growth plate widths of estradiolimplanted wethers are shown in Table 3.
Growth plate width decreased as wethers 12
mo of age grew older and the implants were in
place for longer periods of time. The growth
plates were completely ossified in 57Od-old

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3666

mELD ET AL..
TABLE 1. AGE, WEIGHT AND MATURITY CHAFWCTEFSSTICS OF WETHERS
AFIER IMPLANTING WITH ESTRADIOL

Item

2-mwId
220 d

NUmber
Age, d
Live wt, kg

4
28 1
61.4d

Day after implanting, 12-mo-old

30
3

408'

60

90

3
413'

448d

Pooled

220

Controls

SE*

3
57oe
82.6f

7
565'
75.8'

3.76
2.22

43.9

58.F

62.6d

1.4'

1.7d
l.Bd

1.9d
1.9d

2.3'
2.3'

3.M
3.28

2.7f
2.6

.09
.06

1.4'
1.4'

1.8d
1.Y

1.8d
1.9d

2.4'
2.3'

32f
3.18

2.F
2.Bf

.w

Maturity
Flank color
and textureb
Break jointb
Rib shape and
colorb
overaumaturitvb

1.4'

.ll

'Standard error of the mean.

b1.5 = A5', 2.3 = BJo, 3.1 = yearling1o.
c*ae*f*gMeansin the same row that do not have a common superscript leaer differ (P < .05).

wethers implanted for 220 d. However, the later-thanexpected age (570 d) for growth
565-d-old control wethers and 2-mo-old im- plate ossification and the lack of ossification in
planted wethers maintained growth plate 2-mo-old wethers implanted for 220 d conwidths comparable to those of other wethers of firmed the observation of Garrigus (1965) that
similar age in previous studies (Ho et al., in order for estrogens to hasten growth plate
1989; Field et al., 1990). According to ossifcation, lambs must be nearing the age at
Garrigus (1965), diethylstilbestrol hastens ossi- which ossification would occur normally. This
fication in lambs nearing the age at which the observation casts doubt on the ability of an
growth plate would normally ossify. Implant- estrogen to limit mature size of lambs by
ing 12-mo-old wether lambs with estradiol causing earlier ossification of growth plates
might be expected to result in growth plate than is normal. Perhaps bone growth in lamb
ossification at 480 d age, as was observed in must stop before elevated levels of estrogens
ewe lambs (Ho et al., 1989). Perhaps differ- can cause complete ossification of the growth
ences in the endocrine milieu to which ewes plate. Conversely, estrogens result in rapid
and estradiol-implanted wethers are exposed is cessation of growth, even though the epiphyses
the reason why this did not occur. However, remain open (Van Wyk, 1984). Four

TABLE 2. CARCASS CL4RACTEIUSTICS OF WE-S

Item
Hot ciucass wt, Q

Fat depth. mm
Loogissimus area, cm2
Bodywall t h i b s s , mm
Kidney fat, %
Leg conformationb
carcass conformationb
mank fat streaLingc

ww

srdeb

Yield grade

AFTER IMPLANTING WITH ESTRADIOL

Day after implanting, 12-mo-old

2-mwld
220 d

30

60

90

220

controls

34.6'
7.4'
14.9&
33.68
2.Y
3.e
3.e
3.v
3.4=
3.F

21.3'
1.8d
13.Sd
9.4d
2.2d
3.2d
3.4d
2Sd
3.od
2.ld

30.8'
3.6&
14.8&
16.e
2.3d
3.F
3.F
2.Sd
3.1d
2.Sd

33.c

49.6
12.2f
18.1C
43.28
4.F
3.F
3.8"
52f
3.F
5.4f

46.3f

7.1'

16.1'
%.Ief
2.4d
3.F
3.F
3.F
3.3&
3.6'

1l.lf

18.1'
38.11
5 .4'
3.8'
3.IF
4.7f
3.4e
5.4f

'Lstandard e m x of he mean.
9 . 1 = Good",
'2.5 = traces"

3.5 = Choice50. 4.4 = Primem.

3.4 = slightm, 4.4 = smallm, 5.6 = modestm.

a'*f*BMeans in the same row that do not have a common superscript letter differ (P

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.05).

Pooled
sEa

1.67
1.21

.so

2.60
.39
.10

.os

.20
.18
.25

3667

CHARACTERISTICS OF WBTH@RS GIVEN ESTRADIOL

IMPLANTZNG 'WITH ESTRADIOL

TABLE 3. BONE CHARACTERISTICS OF \KETHERS

Day after implanting, lZmo-oId

2-mo-old
220 d

30

60

90

220

Controls

Pooled
SE'

54.4b

36.3'

22.4d

19.7&

Closed

18.3'

3.7

Length,
Fresh wt, g
Moisture. %

11.9b
44.9b
20.2b

12.3b
44.lb
24.7b

12.7b
53.F
20.9b

12.6b
53.3'
24.7b

13.1bc
54.7
12.8'

13.3bc
54.8'
14.2'

.33
2.1
1.7

Metatarsal
Lenglh, cm
Fresh wt, g
Moisture, %

12.4b
47.4b
20.2b

12.Sb
46.Sb
25 .9b

13.4bc
56.p
25.2b

12.9b
55.3'
23.3b

13Sk
62.pd
14.9

13.Sbc
63.3d
15.F

.40
2.4
1.8

Item
Growth plate width, mm
Metaca$al

'Standard error of the mean.

b,'*'%feans on the same line that do not have a common superscript letter differ (P < .05).

2-mo-old grade white-faced wethers implanted

with estradiol at the same time the yearliig
wethers were implanted all had cartilaginous
growth plates when slaughtered 220 d after
implanting. Growth plate width as well as
bone length, weight and moisture content for
these four 9-meold wethers were similar to
characteristics of 9-mo-old wethers that we
have studied in the past (Ho et al., 1989).
Metacarpal and metatarsal lengths did not
change significantly between the 30- and
220-d implant periods (Table 3). In addition,
neither length nor fresh weight of the bones
differed (P > .05) between the control and
implanted group that were slaughtered 220 d
after implanting. Ho et al. (1989) previously
reported that bone lengths and weights of
sheep slaughtered between 459 and 652 d of
age were similar. Oberbauer (1985) found that
95% of the mature metacarpal length in
Suffolk and Dorset rams was attained by 226 d
and 165 d, respectively. Growth plates, however, did not begin to ossify until 390 d, and
closure was not complete in all animals until
480 d, indicating that metacarpal growth rate
was dissociated temporally from growth plate
closure. Therefore, even though estradiol, and
presumably other growth stimulants with estrogenic activity, increases ossification of the
metacarpal growth plate, there is no evidence
that bone length or mature s u e is limited.
Because mature frame size plays a significant
role in muscle development and composition
of gain to a given weight (Now et al., 1981;
Nour and Thonney, 1987; Thonney, 1987), the
finding that estrogenic implants do not limit
mature size is impomnt. Percentage moisture
in metacarpal and metatarsal bones of control
wethers and wethers implanted for 220 d was

lower (P < .OS) than that of wethers implanted

30, 60 or 90 d (Table 3). Lower moisture
values probably are a result of less hemopoietic marrow and more fatty marrow in bones
from the older wethers; fatty marrow contains
much less moisture than hemopoietic marrow
(Field et al., 1974).
implications

Implanting wethers with estradiol hastens

ossification of the growth plate; presumably
any substance with estrogenic activity administered to promote growth would have the same
effect. This could have economic implications
for 9- to 16-mo-old ewe lambs because
ossification of metacarpal growth plates at this
age could hasten formation of spool joints and
change classification of ovine carcasses from
lamb to yearling mutton. However, wether
lambs normally do not have ossification of
growth plates until 650 d of age, so reducing
age at ossification by 100 d would not have an
economic impact because wethers are seldom
fed to this age. Bone size of wethers 570 d of
age was not reduced by implanting with
estradiol. Nevertheless, estrogens are believed
to cause rapid cessation of growth. More work
is needed to determine whether implanting or
feeding estrogens limit mature size.
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