Genetic Variation in Pure Lines and Crosses of Large-Bodied Turkey Lines.

1. Body Weight, Walking Ability, and Body Measurements of Live Birds

1
K. E. Nestor,
2
J. W. Anderson, and S. G. Velleman
Department of Animal Sciences, Ohio Agricultural Research and Development Center,
The Ohio State University, Wooster, Ohio 44691
ABSTRACT An experimental line of turkeys selected
for increased BW (F) was reciprocally crossed with sire
lines (designated A and B) from each of two major com-
mercial breeders in order to study the inheritance of
growth traits. All genetic groups were grown intermin-
gled in connement with the sexes reared in different
houses. Traits measured included BW at 8, 16, and 20 wk
of age; shank length, width, and depth; breast width; and
walking ability scores at 16 wk of age.
The F line had a different growth pattern than the two
commercial sire lines. The F line had higher (males) or
similar (females) BW in comparison to Line A at 8 wk of
age, but by 16 wk of age, Line A was heavier than Line
F in both sexes. The difference in BW between the F and
A lines increased from 16 to 20 wk of age. Lines F and B
did not differ in BW at 8 wk of age, but at 16 and 20 wk
of age, Line B birds were heavier than those of F line.
(Key words: turkey, body weight, breast width, inheritance, walking ability score)
2001 Poultry Science 80:10871092
INTRODUCTION
Commercial turkeys are the result of a cross of a com-
mercial sire line (or sire line cross) and a commercial dam
line (or dam line cross). In general, early research has
indicated that nonadditive genetic variation is not a major
contributor to total genetic variation for growth traits. In
earlier studies, heterosis for BW has been observed in
some crosses (Asmundson, 1942, 1948; Jerome et al., 1960;
Friars et al., 1963) but not in other crosses (Knox and
Marsden, 1944; Kondra and Shoffner, 1955; Jerome et al.,
1960; Clark, 1961; Nestor, 1971). In some cases, heterosis
was observed for BW only at certain ages (Asmundson
and Pun, 1954; Friars et al., 1963). Using diallel crosses,
McCartney and Chamberlin (1961) concluded that addi-
tive genetic variance was much more important than non-
2001 Poultry Science Association, Inc.
Received for publication September 11, 2000.
Accepted for publication April 9, 2001.
1
Salaries and research support were provided by state and federal
funds appropriated to the Ohio Agricultural Research and Development
Center, The Ohio State University.
2
To whom correspondence should be addressed: Nestor.1@osu.edu.
1087
Reciprocal effects, a measure of sex linkage and mater-
nal effects, were noted only for breast width of females
in crosses of Lines A and F and for shank width and
depth of males and 8-wk BW and shank depth of females
in crosses of Lines B and F. Heterosis was an important
source of variation in BW for males from both crosses.
The percentage heterosis at the various ages ranged from
3.1 to 7.5. For females, heterosis (range = 2.6 to 4.9%) was
only signicant at younger ages (8 wk for the crosses of
the A and F lines and 8 and 16 wk for the crosses of the
B and F lines). No heterosis for breast width was observed
in either cross. Heterosis in walking ability scores was
signicant only for males from the crosses of Lines B
and F. The presence of heterosis was inconsistent for
shank measurements.
additive genetic variance for BW and body conforma-
tion measurements.
More recently, Emmerson et al. (1991) observed sig-
nicant heterosis for BW, particularly at older ages, and
shank length in F
1
reciprocal crosses of an experimental
line (F) selected long-term for increased 16-wk BW and
a commercial sire line (N) no longer in commercial use.
No heterosis for BW was observed in a cross of an experi-
mental egg line and the F line at 8, 16, and 20 wk of age,
and heterosis was observed only at 20 wk of age in a
cross of the egg and Nlines (Nestor et al., 1997). In crosses
of an experimental line (FL) selected for increased shank
width and an unimproved commercial sire line, Ye et al.
(1997) observed signicant heterosis for BW and shank
length. When the FL line was crossed with two improved
commercial sire lines (A and B), signicant heterosis in
BW was observed for females of the cross involving Line
A and for males of the cross involving Line B (Nestor and
Anderson, 1998). The results of the most recent studies
Abbreviation Key: A, B, and N = commercial sire lines; F = experi-
mental line selected long-term for increased 16-wk BW; FL = subline
of F selected for increased shank width.
NESTOR ET AL. 1088
suggest that nonadditive genetic variation may be evolv-
ing into a more important contributor to variation in BW
than was observed in earlier studies as would be the case
if additive genetic variation in BW is decreasing. In the
F line, Nestor et al. (2000) foundthat the realized heritabil-
ity, a measure of additive genetic variation, of 16-wk BW
declined slightly over 30 generations of selection. The
purpose of the present study was to study the genetic
variation of growth traits in the F, A, and B lines and in
reciprocal crosses among them.
MATERIALS AND METHODS
Genetic Stocks
The F line was started from a randombred control pop-
ulation by mass selection only for increased 16-wk BW.
Details of the maintenance of the F line and response to
selection have been previously reported (Nestor, 1977,
1984; Nestor et al., 1996, 2000). The F line had been se-
lected for 30 generations at the time of the present study.
A sample of a sire line from each of two major interna-
tional turkey breeders was obtained as unpedigreed eggs.
The commercial sire lines, designated A and B, were re-
produced in the rst generation by selecting 15 females
and 10 males from each sire line with minimal selection
pressure for increased 16-wk BW and increased 16-wk
breast width to try to maintain the lines at current perfor-
mance levels. Approximately 50% of the males with the
largest BW and breast width were selected, whereas no
selection was practiced in females. The pure sire lines
were reproduced by articially inseminating 15 females
with pooled semen from the 10 males. The A and B lines
were then reciprocally crossed with the F line.
Offspring from the pure lines and reciprocal crosses
were produced in two hatches, each of which represented
a 2-wk collection of eggs. The F line was produced by
weekly articial mating of 36 sires to 72 dams with each
sire being mated to two dams. The number of offspring
produced was 91 males and 122 females. The pure A and
B lines were produced from six dams. The number of
sires used in the production of the A and B lines was 11
and 9, respectively. Each week, the sire used for articial
insemination of each hen was changed so that as large a
genetic base as possible was obtained. The number of
offspring obtained for the A line was 25 males and 35
females and for the B line was 23 males and 24 females.
To produce crosses involving F-line sires and A- and B-
line dams, the same sires were used to mate dams from
both lines and were sires that were used to reproduce
the F line. A different F-line sire was used to inseminate
each dam each week, making a total of 28 F-line sires
being used in each cross. The number of offspring from
each cross and sex subgroup ranged from 18 to 38 with
an average of 29 per group. For the reciprocal cross, nine
F-line dams were used in each cross. The same sires used
in the production of the A and B lines were also used in
the production of the crosses, and the sire assigned to
each damwas changed weekly to ensure as wide a genetic
base as possible. The number of offspring for the crosses
and sex subgroups involving F line dams ranged from
27 to 40 with an average of 33.
Management of Birds
and Measurements Made
The different genetic groups were grown intermingled
in connement with the sexes reared in separate houses.
All birds were provided a declining protein ve-ration
system(Naber andTouchburn, 1970) basedonthe feeding
schedule for males. Continuous lighting was provided
from hatching to 6 wk of age, at which time the photope-
riod was reduced to 12 h per day. At 16 wk of age, the
amount of light per day was reducedto 10 handremained
at this level until 20 wk of age.
Body weight was recorded at 8, 16, and 20 wk of age.
At 16 wk of age, measurements of shank length, shank
width (laterally at the dewclaw), shank depth (perpendic-
ular at the dewclaw), and breast width were made. Breast
width was measured at 6.35 cm of body depth at a point
approximately 3.18 cmfromthe anterior point of the keel.
Walking ability at 16 wk of age was estimated by the
method of Nestor et al. (1985) in which each bird was
given a score of 1 to 5, with 1 representing birds with no
leg defects and no difculty walking and 5 indicating
birds with exhibited extreme lateral deviations of the legs
and great difculty walking. Ratings of 2, 3, and 4 repre-
sented intermediate values.
Statistical Analysis
The data were analyzed within sexes for the crosses of
Line F with Lines Aand B using the general linear models
procedure of SAS software (SAS Institute, 1988) with
genetic group (pure lines and crosses) and hatch as
sources of variation. Orthogonal contrasts (SAS Institute,
1988) were used to estimate additive genetic effects (con-
trast of A vs. B, F vs. A, and F vs. B), heterotic effect
(contrast of average of the parental lines with the average
of the reciprocal crosses), and sex-linked and maternal
effects (contrast of reciprocal crosses). The data were ana-
lyzed in one analysis, but for simplicity, the results of the
F and A line comparisons and F and B line comparisons
will be presented separately.
RESULTS
Hatch effects were signicant for BW at 8, 16, and 20
wk of age; breast width; and walking ability scores of
males (data not shown). For females, hatches differed
signicantly for 8-wk BW, shank measurements (length,
depth, and width), and breast width (data not shown).
Additive Genetic Effects
Comparison of Lines A and B. For males, Line B was
larger at 8, 16, and 20 wk of age; had wider breasts; and
had poorer walking ability (higher walking ability score)
than Line A (Tables 1 and 3). There was no difference
GENETIC VARIATION OF GROWTH TRAITS IN TURKEYS 1089
TABLE 1. Effect of reciprocally crossing commercial sire Line A (A) and a line (F) selected long-term
for increased 16-wk BW on performance of males
Parental lines Reciprocal crosses Additive
genetic Reciprocal Percentage
Variable A F x A F F A x effect
1
effect
2
heterosis
3
SEM
Body weight, kg
8 wk 3.90 4.20 4.05 4.35 4.30 4.32 ** NS 6.7** 0.074
16 wk 14.2 13.7 13.9 14.6 14.9 14.8 * NS 6.2*** 0.151
20 wk 18.2 17.2 17.7 19.1 19.1 19.1 ** NS 7.5*** 0.236
Walking ability score
4,5
2.40 2.93 2.66 2.54 2.71 2.62 ** NS 1.3 0.134
Shank
Length,
5
cm 21.6 22.3 21.9 21.9 22.1 22.0 *** NS 0.4 0.101
Width,
5
mm 16.2 16.7 16.4 16.8 17.2 17.0 NS NS 3.6** 0.183
Depth,
5
mm 24.7 25.1 24.9 25.2 25.6 25.4 NS NS 1.9* 0.209
Breast width,
5
cm 15.7 14.4 15.1 15.1 15.9 15.5 *** * 2.8 0.235
1
Measured by contrast of parental lines.
2
Measured by contrast of reciprocal crosses.
3
Percentage heterosis = [(average of reciprocal crosses average of parental lines)/average of parental lines] 100.
4
Birds were subjectively rated from 1 to 5, with 1 representing birds whose legs did not have any defects and had no difculty walking and 5
indicating birds whose legs exhibited extreme lateral deviations or had great difculty walking. Ratings of 2, 3, and 4 represented intermediate
values.
5
Measurements made at 16 wk of age.
*P 0.05; **P 0.01; ***P 0.001.
between lines for any shank measurements of males. For
females, Line B had wider breasts and wider shanks than
Line A, but there were no signicant line differences in
BW at any age, shank length and depth, and walking
ability scores (Tables 2 and 4).
Comparison of Lines A and F. The F line was heavier
than the A line at 8 wk of age with the difference being
signicant (P 0.01) only for males (Tables 1 and 2). By
16 wk of age, males and females of the Aline were heavier
than those of the F line, and the absolute line difference
continued to increase at 20 wk of age. For males, the
F line had longer shanks, narrower breasts, and higher
walking scores than the A line (Table 1). There was no
TABLE 2. Effect of reciprocally crossing commercial sire Line A (A) and a line (F) selected long-term
for increased 16-wk BW on performance of females
Parental lines Reciprocal crosses Additive
genetic Reciprocal Percentage
Variable A F x A F F A x effect
1
effect
2
heterosis
3
SEM
Body weight, kg
8 wk 3.63 3.70 3.66 3.82 3.83 3.83 NS NS 4.4** 0.051
16 wk 11.0 10.2 10.6 10.7 10.7 10.7 *** NS 1.4 0.103
20 wk 13.5 12.2 12.9 12.7 12.9 12.8 *** NS 0.6 0.134
Walking ability score
4,5
2.31 2.08 2.20 1.98 2.16 2.07 NS NS 5.9 0.108
Shank
Length,
5
cm 17.4 17.8 17.6 17.7 17.8 17.7 *** NS 0.9 0.075
Width,
5
mm 15.3 14.9 15.1 15.0 15.5 15.2 * NS 1.0 0.175
Depth,
5
mm 21.9 21.4 21.6 21.7 21.6 21.6 * NS 0.1 0.124
Breast width,
5
cm 15.0 13.4 14.2 14.2 14.3 14.2 *** NS 0.1 0.155
1
Measured by contrast of parental lines.
2
Measured by contrast of reciprocal crosses.
3
Percentage heterosis = [(average of reciprocal crosses average of parental lines)/average of parental lines] 100.
4
Birds were subjectively rated from 1 to 5, with 1 representing birds whose legs did not have any defects and had no difculty walking and 5
indicating birds whose legs exhibited extreme lateral deviations or had great difculty walking. Ratings of 2, 3, and 4 represented intermediate
values.
5
Measurements made at 16 wk of age.
*P 0.05; **P 0.01; ***P 0.001.
line difference in shank width and shank depth of males.
Females of the F line had longer and narrower shanks
than females of the A line, but walking ability score did
not differ between lines (Table 2). The breasts of females
were narrower in the F line than in the A line.
Comparison of Lines B and F. The B and F lines were
similar in BW at 8 wk of age, but the B line was larger
than the F line at 16 and 20 wk of age (Tables 3 and 4).
At 16 wk of age, the B line had wider breasts in both
sexes. The shank was longer for both sexes in the F line
than in the B line. There was no signicant line difference
in walking ability scores of either sex. Shank width of the
B line was greater than in the F line only for females.
NESTOR ET AL. 1090
TABLE 3. Effect of reciprocally crossing commercial sire Line B (B) and a line (F) selected long-term
for increased 16-wk BW on performance of males
Parental lines Reciprocal crosses Additive
genetic Reciprocal Percentage
Variable B F x B F F B x effect
1
effect
2
heterosis
3
SEM
Body weight, kg
8 wk 4.21 4.20 4.21 4.45 4.45 4.45 NS NS 5.8** 0.074
16 wk 15.2 13.7 14.4 14.9 14.9 14.9 *** NS 3.1* 0.151
20 wk 19.0 17.2 18.1 19.0 18.6 18.8 *** NS 3.5* 0.236
Walking ability score
4,5
3.21 2.93 3.07 2.62 2.73 2.67 NS NS 12.9** 0.134
Shank
Length,
5
cm 21.3 22.3 21.8 21.9 22.2 22.1 *** NS 1.2* 0.101
Width,
5
mm 16.5 16.7 16.6 16.8 17.7 17.3 NS ** 4.1*** 0.183
Depth,
5
mm 25.0 25.1 25.1 25.4 26.3 25.9 NS * 3.2*** 0.209
Breast width,
5
cm 16.7 14.4 15.5 15.4 15.4 15.4 *** NS 1.1 0.235
1
Measured by contrast of parental lines.
2
Measured by contrast of reciprocal crosses.
3
Percentage heterosis = [(average of reciprocal crosses average of parental lines)/average of parental lines] 100.
4
Birds were subjectively rated from 1 to 5, with 1 representing birds whose legs did not have any defects and had no difculty walking and 5
indicating birds whose legs exhibited extreme lateral deviations or had great difculty walking. Ratings of 2, 3, and 4 represented intermediate
values.
5
Measurements made at 16 wk of age.
*P 0.05; **P 0.01; ***P 0.001.
Reciprocal Effects
No difference between reciprocal crosses was noted in
the crosses of the A and F lines for any trait, except for
breast width of males (Tables 1 and 2). For the crosses of
the B and F lines, reciprocal effects were signicant for
shank width and depth of males (Table 3) and for 8-wk
BW and shank depth of females (Table 4).
Heterotic Effects
Heterosis was an important source of variation in BW
of males from the crosses of the A and F lines (Table 1)
TABLE 4. Effect of reciprocally crossing commercial sire Line B (B) and a line (F) selected long-term
for increased 16-wk BW on performance of females
Parental lines Reciprocal crosses Additive
genetic Reciprocal Percentage
Variable B F x B F F B x effect
1
effect
2
heterosis
3
SEM
Body weight, kg
8 wk 3.83 3.70 3.76 4.07 3.83 3.95 NS ** 4.9** 0.051
16 wk 11.1 10.2 10.6 10.9 10.9 10.9 *** NS 2.6* 0.103
20 wk 13.3 12.2 12.8 12.9 13.0 13.0 *** NS 1.6 0.134
Walking ability score
4,5
2.13 2.08 2.10 2.04 2.30 2.17 NS NS 3.3 0.108
Shank
Length,
5
cm 17.1 17.8 17.4 17.6 17.7 17.6 *** NS 1.0* 0.075
Width,
5
mm 16.3 14.9 15.6 15.8 15.8 15.8 *** NS 1.6 0.175
Depth,
5
mm 21.5 21.4 21.5 21.4 22.0 21.7 NS * 1.0 0.124
Breast width,
5
cm 15.5 13.4 14.5 14.3 14.6 14.4 *** NS 0.1 0.155
1
Measured by contrast of parental lines.
2
Measured by contrast of reciprocal crosses.
3
Percentage heterosis = [(average of reciprocal crosses average of parental lines)/average of parental lines] 100.
4
Birds were subjectively rated from 1 to 5, with 1 representing birds whose legs did not have any defects and had no difculty walking and 5
indicating birds whose legs exhibited extreme lateral deviations or had great difculty walking. Ratings of 2, 3, and 4 represented intermediate
values.
5
Measurements made at 16 wk of age.
*P 0.05; **P 0.01; ***P 0.001.
andof the BandF lines (Table 3). The percentage heterosis
at the various ages ranged from 6.2 to 7.5 for the A F
crosses and 3.1 to 5.8 for the B F crosses. Overdominance
(i.e., the weight of the crosses was greater than either
parental line) for male BW occurred at all ages in the A
F crosses and at 8 wk of age in the B F crosses.
For females of the A F crosses, heterosis in BW was
signicant with overdominance only for 8-wk BW (4.4%,
Table 2), whereas in the B F crosses, heterosis in BW
was signicant at 8 (4.9%) and 16 (2.6 %) wk of age
(Table 4).
In the crosses of the A and F lines, heterosis was not
signicant for walking ability scores, shank length, and
GENETIC VARIATION OF GROWTH TRAITS IN TURKEYS 1091
breast width of both sexes (Tables 1 and 2). Heterosis for
shank width and shank depth was signicant for males
but not females in the Aand F crosses. Negative heterosis
for walking ability score was observed in males from the
B F crosses (Table 3), but no signicant heterosis was
observed in females of this cross (Table 4). Heterosis was
signicant for all shank measurements in males from the
B F crosses (Table 3) but only for shank length of females
(Table 4). No signicant heterosis was observed for breast
width in either sex of the B F crosses (Tables 3 and 4).
DISCUSSION
The F line apparently had a different growth curve
than the two commercial sire lines. At 8 wk of age, the
F line had similar BW or was heavier than the sire lines
but was smaller than both commercial lines at older ages.
The two commercial sire lines might have also differed
in their growth curves. The difference in BW between the
F and A lines increased to a greater extent from 16 to 20
wk of age than the difference between the F and B lines.
Based on commercial birds of the same breeders from
which the sire lines were obtained in the present study,
Barbour and Lilburn (1996) observed a difference in
growth pattern of males through 82 d of age. However,
in an earlier study using commercial females of the same
breeders, Barbour and Lilburn (1995) did not observe an
interaction of the two commercial strains and age for BW
from 14 to 145 d, suggesting no difference in growth
pattern between the commercial lines. In comparisons of
commercial birds from two major turkey breeders, one
of which was not involved in the current study, differ-
ences in growth curves were noted for males (Lilburn
and Emmerson, 1993) and females (Lilburn et al., 1992).
Additive genetic variation, as indicated by differences
among lines, was an important source of variation, as
expected, for BW at 16 and 20 wk of age, shank length,
shank width (females only), and breast width. It has been
shown that large gains in BW (McCartney et al., 1968;
Nestor, 1977, 1984; Nestor et al., 1996), breast width (Nes-
tor et al., 1969), and shank width (Nestor et al., 1985) can
be made by genetic selection.
Nonadditive genetic variation was evident for BW of
males at 8, 16, and 20 wk of age in the crosses of the
F line and two commercial sire lines. In females, such
variation was important only at 8 or 16 wk of age. Age-
specic heterosis for BW has been previously observed
(Asmundson, 1942; Asmundson and Pun, 1954). Signi-
cant heterosis was also observed in shank length, shank
width and depth in some comparisons, and walking abil-
ity score for males from the crosses of the F and B lines.
The F line is not closely related to the commercial sire
Lines A and B even though the F line was started from
a randombred control that was developed fromreciprocal
crosses of two commercial strains of turkeys (Nestor et
al., 1969). The F line differed greatly from Lines A and B
in the frequency of MHC haplotypes (Zhu et al., 1995,
1996b) and had a level of band sharing of DNA nger-
prints with Lines A or B that was less than the average
of all lines compared (Ye et al., 1998). The frequency of
shared bands for DNA ngerprints of Lines A and B was
greater than the average of all lines in the study of Ye et
al. (1998).
Inbreeding, as measured by band sharing of DNA n-
gerprints (Kuhnlein et al., 1990; Zhu et al., 1996a), was
greater in the commercial sire lines than in the F line (Ye
et al., 1998). Accumulated inbreeding in the F line, as
calculated by variation in family size, was 27.6% (unpub-
lished data) when the crosses were made with the com-
mercial sire lines. Therefore, because the lines involved
in the crosses were moderately inbred, the heterosis ob-
served in the present study may have been due to the
elimination of inbreeding effects by crossing relatively
unrelated lines. Based on a simple dominance model,
a linear relationship is expected between the degree of
heterosis and the level of inbreeding (Hill, 1982). The
magnitude of heterosis is inversely related to the degree
of genetic resemblance between parental populations
(Wilhelm and Pollak, 1985).
Nonadditive genetic variation in BW of turkeys is usu-
ally not an important source of genetic variation (Kondra
andShoffner, 1955; Clark, 1961; McCartney andChamber-
lin, 1961; Nestor, 1971), but heterosis has been observed
in some crosses in which the parents differed greatly in
body conformation (Asmundson, 1945, 1948; Emmerson
et al., 1991; Ye et al., 1997; Nestor and Anderson, 1998).
There were large differences in breast width between the
F and A lines and F and B lines and the lines differed in
body shape (unpublished data). The differences in confor-
mation and body shape between the experimental F line
andthe commercial sire lines might have beenresponsible
for the heterosis in BW observed. It is unknown why
heterosis was expressed to a greater extent in males than
in females.
No heterosis in breast width was observed in the cur-
rent study. Similarly, Asmundson (1948), Ye et al. (1997),
andNestor andAnderson (1998) didnot observe heterosis
in crosses of turkey lines differing greatly in breast width.
By using diallel crosses among six strains, McCartney
andChamberlin(1961) foundthat the nonadditive genetic
variation accounted for 8% of the total genetic variation
in breast width of males, but there was no nonadditive
genetic variation in breast width in females.
Heterosis in walking ability scores (12.9%) was sig-
nicant only for males from the reciprocal crosses of the
Band F lines. Negative heterosis for walking ability scores
has been observed previously in reciprocal crosses of the
N and F lines (Emmerson et al., 1991) and in a cross of
an unimproved commercial sire line and the FL line (Ye
et al., 1997). When the FL line was reciprocally crossed
with the two commercial sire lines used in the present
study, heterosis was negative and signicant only for
males in one of the crosses. The results in the present
study and those reported in the literature indicate that
nonadditive genetic variation in walking ability scores
may be an important source of variation in certain crosses.
In summary, nonadditive genetic variation was an im-
portant source of variation in BWof males fromreciprocal
NESTOR ET AL. 1092
crosses of an experimental growth line and two commer-
cial sire lines. Heterosis values ranged from 3.1 to 7.5%
at 8, 16, and 20 wk of age. For females, heterosis was
present in the crosses only at 8 or 16 wk of age. Signicant
heterosis was observed for shank length and, in some
cases, for shank width and shank depth. No signicant
heterosis was observed for breast width.
ACKNOWLEDGMENTS
The authors thank Hybrid Turkeys Inc. (Kitchner, On-
tario, Canada N2K) and Nicholas Turkey Breeding Farms
(Sonoma, CA 95476) for graciously providing the hatch-
ing eggs from their sire lines. Financial support for this
project was also provided by British United Turkeys of
America (Lewisburg, WV 14901) and Hybrid Turkeys.
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