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Growth hormone 1 gene (
GH1
) polymorphisms as possible markers
of the production potential of beef cattle using the Brazilian Canchim breed
as a model
Luiz Guilherme G. Silveira1, Luiz R. Furlan1, Rogério A. Curi1, André Luiz J. Ferraz1,
Maurício M. de Alencar2, Luciana C.A. Regitano2, Cyntia L. Martins1, Mário de Beni Arrigoni1,
Liliane Suguisawa1, Antônio C. Silveira1and Henrique N. de Oliveira1
1Departamento de Melhoramento e Nutrição Animal, Faculdade de Medicina Veterinária e Zootecnia,
Universidade Estudual Paulista “Júlio de Mesquita Filho”, Botucatu, SP, Brazil.
2Empresa Brasileira de Pesquisa Agropecuária, Centro de Pesquisa de Pecuária do Sudeste,
Fazenda Canchim, São Carlos, SP, Brazil.
Abstract
The growth hormone 1 gene (
GH1
) is a candidate gene for body weight and weight gain in cattle since it plays a fun-
damental role in growth regulation. We investigated the
GH1
gene
Alu
I and
Dde
I restriction enzyme polymorphisms,
located 149 bp apart in the cattle genome, as possible markers of the production potential of Canchim crossbreed
cattle, a 5/8 Charolais (
Bos taurus
) and 3/8 Nelore (
Bos indicus
) breed developed in Brazil, by evaluating the birth
weight, weaning weight, yearling weight and plasma insulin-like growth factor-1 (IGF-1) concentration of 7 month to
10 months old Canchim calves (n = 204) of known genealogy and which had been genotyped for the
Alu
I and
Dde
I
markers. Our results showed significant effect (p < 0.05) between the homozygous
Dde
I+/
Dde
I+ polymorphism and
the estimated breeding value for weaning weight (ESB-WW), while the
Alu
I leucine homozygous (L/L) and
leucine/valine (L/V) heterozygous polymorphisms showed no significant effect on the traits studied. The restriction
sites of the two enzymes led to the formation of haplotypes which also exerted a significant effect (p < 0.05) on the
ESB-WW, with the largest difference being 8.5 kg in favor of the homozygous L plus
Dde
I+/L plus
Dde
I+ genotype
over the heterozygous L plus
Dde
I-/V plus
Dde
I+ genotype.
Key words: beef cattle, polymorphism, growth hormone, candidate gene.
Received: September 14, 2007; Accepted: March 13, 2008.
Introduction
The growth hormone 1 (GH1) gene is a candidate
gene for body weight and weight gain in cattle since it plays
a fundamental role in growth regulation. The GH protein is
a single-chain polypeptide consisting of 191 amino acids
and is synthesized and secreted by the anterior pituitary
gland under the hypothalamic control of two hormones,
GH-releasing hormone (GHRH), which increases the se-
cretion of GH, and somatotropin release-inhibiting factor
(SRIF, also called somatostatin) which inhibits its secretion
(Nicoll et al., 1986). It is known that GH is the main regula-
tor of postnatal somatic growth, stimulating anabolic pro-
cesses such as cell division, skeletal growth and protein
synthesis and is involved in nutrient partition by way of
regulating the oxidation rate of fats (lipolytic activity), inhi-
bition of glucose transport to peripheral tissues and the reg-
ulation of ribosomal activity involved in translation, which,
in turn, influences protein synthesis (Goodman, 1993).
The effects of some GH1 gene polymorphisms have
been widely studied in beef cattle (Switonski, 2002) and the
proximity between some of these polymorphisms, which
can be characterized using different restriction enzymes,
suggests a strong linkage between them. The presence of
the AluI restriction site corresponds to the presence of the
amino acid leucine (L) at position 127 in the polypeptide
chain of cattle GH, whereas the absence of this site indi-
cates the presence of valine (V) at the same position. The
presence of the DdeI restriction site corresponds to the
presence of an adenine nucleotide in the cattle GH1 gene
sequence (Ddel+), while the absence of this site (DdeI-) in-
dicates the presence of a cytosine nucleotide at the same po-
Genetics and Molecular Biology, 31, 4, 874-879 (2008)
Copyright © 2008, Sociedade Brasileira de Genética. Printed in Brazil
www.sbg.org.br
Send correspondence to Henrique N. de Oliveira. Departamento de
Melhoramento e Nutrição Animal, Faculdade de Medicina Veteriná-
ria e Zootecnia, Universidade Estudual Paulista “Júlio de Mesquita
Filho”, 18618-000 Botucatu, SP, Brazil. E-mail: hnunes@fca.
unesp.br.
Research Article
sition. These two polymorphisms can be used as markers
but, however, no studies are available in the literature re-
garding the combined segregation of these markers or the
effects of the haplotypes formed on cattle production traits.
The objective of the study described in this paper was
to characterize GH1 gene polymorphisms as possible
markers of the production potential of Canchim cattle by
evaluating the individual and combined effects associated
with these markers on body weight traits and the plasma
concentration of the polypeptide hormone insulin-like
growth factor-1 (IGF-1), which has important effects on
growth.
Material and Methods
We used 108 male and 96 female (n = 204) Canchim
calves, ranging in age from 7 months to 10 months, of
known genealogy and belonging to the Brazilian Agricul-
tural Research Corporation – Embrapa (Empresa Brasileira
de Agropecuária -Embrapa) herd at Canchim Farm, Mu-
nicipality of São Carlos, São Paulo, Brazil. The calves were
weaned during the May, June and July (autumn/winter in
the southern hemisphere) of 1999 and they represent the
progeny of 10 bulls with the half-sib group varying be-
tween 8 and 34 animals. Canchim crossbreed cattle, devel-
oped in Brazil, are 5/8 Charolais (Bos taurus) and 3/8
Nelore (Bos indicus).
Blood samples were collected from the calves for the
separation of the leukocyte layer, used for extraction of
DNA, and the blood plasma layer, used for the quantifica-
tion of IGF-1. Genomic DNA was extracted from leuko-
cytes according to the method of Zadworny and Kuhnlein,
modified by Miretti MM (1998, MSc Dissertation, Univer-
sity of São Paulo, Ribeirão Preto, SP, Brazil). Plasma
IGF-1 concentrations were determined by radioimmuno-
assay using the Active IGF1 DSL-5600 kit (Diagnostic
Systems Laboratories, Inc., USA) according to the instruc-
tions of the manufacturer.
The GH1 gene was genotyped using the polymerase
chain reaction and restriction fragment length polymor-
phism (PCR – RFLP) analysis using the forward (5’-TAG
GGG AGG GTG GAA AAT GGA-3’) and reverse
(5’-GAC ACC TAC TCA GAC AAT GCG-3’) primer pair
published by Gordon et al. (1983) to amplify a 404-bp frag-
ment located between positions +1405 and +1808, com-
prising the end of the fourth intron, the fifth exon and the
initial portion of the 3’ UTR region. The Alu I and Dde Ire
-
striction enzyme map for the fragment being shown in Fig-
ure 1. The amplification reactions were carried out in a final
volume of 25 μL containing 100 ng of DNA, 0.5 μMof
each primer, 2.5 μL of 10x PCR buffer (10 mM Tris-HCl,
pH 9.0, 1.5 mM MgCl2and 50 mM KCl), 100 μM of dNTPs
and 0.5 units of Taq DNA polymerase (Invitrogen, USA).
After initial denaturation at 94 °C for 120 s, amplification
was carried out using 40 cycles at 94 °C for 30 s, 59 °C for
80 s and 72 °C for 90 s, followed by a final extension step at
72 °C for 5 min. Aliquots of the amplification products
(15 μL) were digested, separately, with 5 units of AluI
(Invitrogen, USA) and 5 units of DdeI (Invitrogen, USA) at
37 °C for 2.5 h and DNA fragments separated on 1.8%
(w/v) agarose gel in a horizontal electrophoresis system us-
ing a 100-bp molecular weight standard (Invitrogen, USA)
to calculate the size of the fragments, which were visual-
ized by ethidium bromide staining and exposure to ultravi-
olet light. The possible genotypes, characterized in
function of the restriction fragments, for both GH1 gene
polymorphism are presented in Table 1.
Analysis of the data consisted of the calculation of the
allele and genotype frequencies for the loci determined
with each restriction enzyme. Due to the proximity of the
two restriction sites (149 bp), the independent segregation
of the two loci was tested. After the confirmation of the for-
mation of the haplotypes we applied a chi-square test of the
SAS program (SAS, 1999) to the data to verify if the
haplotype segregation conformed to the Hardy-Weinberg
law.
The isolated effects of the genotypes and the com-
bined effects of the two loci on estimated breeding values
for birth weight, weaning weight, yearling weight and
plasma IGF-1 concentration were evaluated using the GLM
procedure of the SAS program (SAS, 1999). These breed-
ing values were estimated based on data obtained from the
Canchim Farm, with the genetic parameters being esti-
mated on the same basis. For the analysis of IGF-1 concen-
Silveira et al. 875
Figure 1 - Restriction map for the AluI and DdeI enzymes between posi-
tions 1405 and 1808 of the cattle growth hormone 1 (GH1) gene (GenBank
Gi: 163091). Three restriction sites are shown for each enzyme (positions
1442, 1493 and 1678 for AluI and 1450, 1642 and 1799 for DdeI), with the
AluI polymorphism being identified when nucleotide change occurs at po-
sition 1493 and the DdeI polymorphism when it occurs at position 1642.
Table 1 - Polymerase chain reaction restriction fragment length polymor-
phism (PCR-RFLP) informative fragments of genotyping of the growth
hormone 1 gene (GH1) polymorphisms of Canchim calves.
Polymorphism and genotype Informative fragments (bp)
AluI
L/L 185, 131, 51 and 37
L/V 236, 185,131, 51 and 37
L/L 236, 131 and 37
DdeI
DdeI+/DdeI+ 192, 157, 45 and 9
DdeI+/DdeI- 349, 192, 157, 45 and 9
DdeI-/DdeI- 349, 45 and 9
tration, genetic parameters estimated for the taurine Angus
breed were used (Davis et al., 2003).
Using the same genetic parameters, a mixed model
analysis was performed to determine the average effects of
the AluIDdeI haplotypes. For these analyses we considered
the presence of three haplotypes in the population, identi-
fied as L plus DdeI+, L plus DdeI- and V plus DdeI+, and
regression of the number of L plus DdeI+ and L plus DdeI-
haplotypes in the genotype of each calve was added to the
statistical model. Regression of the third haplotype was not
included since the number of V plus DdeI+ haplotype in the
genotype is completely determined by the sum of the num-
ber of the other two haplotypes. The average effects of
haplotype substitution were obtained as regression coeffi-
cients. The MTDFREML program (Boldman et al., 1995)
was used for these analyses and the model included the
combined effect of sex and month of birth, haplotype ef-
fects and random effect of the animal.
Results and Discussion
The L and V genetic variants of the AluI site polymor-
phism were observed in the Canchim calves studied but the
V/V genotype was not detected, while for the DdeI site
polymorphism both DdeI+ and DdeI- alleles were observed
and the three possible genotypes (DdeI+/DdeI+,
DdeI+/DdeI-, DdeI-/DdeI-) were detected. The genotype
and allele frequencies (f) at the loci determined with the
two restriction enzymes are shown in Table 2. For the AluI
polymorphism the most frequent genotype was L/L
(f = 0.82), whereas the heterozygous L/V genotype was
present at only a low frequency (f = 0.18). The distribution
of DdeI genotypes in the calves showed the heterozygous
DdeI+/DdeI- genotype being the most frequent (f = 0.50),
followed by the homozygous DdeI-/DdeI- (f = 0.26) and
DdeI+/DdeI+ (f = 0.24) genotypes.
Simple observation of the combined distribution of
the genotypes determined by the two polymorphisms
showed that they were not independent. This finding was
expected because of the proximity of the restriction sites of
the two enzymes and, since the population was derived
from crossings, there was a strong linkage disequilibrium
which was confirmed by the chi-square test. Based on the
combined distribution, only three haplotypes were ob-
served in the population. The absence of calves with the L
plus DdeI-/V plus DdeI- genotype suggests that no gametes
without the restriction sites for the two enzymes were
formed. Thus the haplotypes L plus DdeI+ (f = 0.40), L plus
DdeI- (f = 0.51) and V plus DdeI+ (f = 0.09) were detected
but not V plus DdeI-. The observed and expected genotypes
of haplotypes showed that the population studied was in
equilibrium for this compound locus (Table 3).
The AluI genotypes showed no significant differences
(p > 0.05) regarding any of the estimated breeding values or
the plasma IGF-1 levels, although the homozygous L/L ge-
notype did show higher weaning and yearling weights and
IGF-1 values than the L/V genotype (Table 4). With respect
to the DdeI polymorphism shown in Table 4, significant
differences between genotypes (p < 0.05) only occurred for
weaning weight, with the DdeI+/DdeI+ genotype showing
an approximately 5 kg higher weaning weight than the het-
erozygous DdeI+/DdeI- genotype. It is also interesting to
note that the weaning and yearling weights for the
DdeI-/DdeI- homozygotes were intermediate between the
DdeI+/DdeI+ and DdeI+/DdeI- values. For the combined
AluI and DdeI genotypes only weaning weight showed sig-
nificant differences (p < 0.05) between genotypes, with the
difference between the highest and lowest value being
greater than 8.5 kg (Table 5). Regarding the average effects
of haplotype substitution, the substitution of L plus DdeI-
for L plus DdeI+ produced a significant difference
(p < 0.05) for weaning weight only (+3.56 kg), which also
presented a significant difference between genotypes (Ta-
ble 6).
876 GH1 gene polymorphisms in beef cattle
Table 2 - Genotype and allele frequencies of the AluI and DdeI poly-
morphisms in exon 5 of the growth hormone 1 gene (GH1) of Canchim
calves.
Polymorphism, genotype and allele Frequency
AluI polymorphism (n = 203 animals)
L/L genotype 0.818
L/V genotype 0.182
V/V genotype 0.000
L allele 0.909
V allele 0.091
DdeI polymorphism (n = 200 animals)
DdeI+/DdeI+ genotype 0.240
DdeI+/DdeI- genotype 0.495
DdeI-/DdeI- genotype 0.265
DdeI+ allele 0.488
DdeI- allele 0.512
Table 3 - Observed and expected frequencies of each genotype in Can-
chim cattle considering the combined distribution of the AluI and DdeI
polymorphisms in the amplified fragments of the growth hormone 1 gene
(GH1) gene of Canchim calves
Frequency1
Genotype Observed Expected
L plus DdeI+/L plus DdeI+ 0.175 (35) 0.158 (31.6)
L plus DdeI+/L plus DdeI- 0.380 (76) 0.405 (81.1)
L plus DdeI+/V plus DdeI+ 0.065 (13) 0.074 (14.7)
L plus DdeI-/L plus DdeI- 0.260 (52) 0.260 (52)
L plus DdeI-/V plus DdeI+ 0.120 (24) 0.094 (18.9)
V plus DdeI+/V plus DdeI+ 0.000 (0) 0.009 (1.7)
1The number of observed and expected frequencies in the animals is
shown in parentheses.
Kemenes et al. (1999) reported an AluI V allele fre-
quency of 0.28 for Charolais taurine cattle, which contrib-
uted 5/8 of the Canchim genome. However, for Zebu
indicine cattle, which account for the remaining proportion
of the Canchim genome, almost all studies have shown fix-
ation of the AluI L allele (Kemenes et al., 1999; Tambasco
et al., 2000; Curi et al., 2006), except for the study by
Unanian et al. (2000) of Nelore indicine cattle, which the
reported frequencies of 0.85 for the L/L genotype and 0.15
for the L/V genotypes. Thus, as expected, the allelic fre-
quencies observed by us for Canchim calves were interme-
diate between those of the two foundation breeds. The
genotype and allele frequencies obtained by us for the DdeI
polymorphism differ from the standard reported for Euro-
pean breeds (B. taurus). For taurine breeds, Yao et al.
(1996) reported Holstein cattle DdeI genotype frequencies
of 0.74 (DdeI+/DdeI+), 0.24 (DdeI+/DdeI-) and 0.02
(DdeI-/DdeI-), whereas Ferraz ALJ (2001, MSc Disserta-
tion, São Paulo State University, Jaboticabal, SP, Brazil)
reported Simmental DdeI genotype frequencies of 0.81
(DdeI+/DdeI+), 0.14 (DdeI+/DdeI-) and 0.05
(DdeI-/DdeI-). No studies on the haplotypes were found in
the literature. Thus, few inferences can be made regarding
the haplotypes in relation to the two genetic groups that
were used to produce the Canchim breed. However, our re-
sults indicate that the V plus DdeI+ haplotype can only be
derived from the Charolais breed.
The higher weaning and yearling weights obtained
for the AluI polymorphism L/L genotype differed from
those reported by Schlee et al. (1994a), who reported an as-
sociation between higher weights and the heterozygous ge-
notype. However, Di Stasio et al. (2002) found no evidence
of an association between this polymorphism and growth or
carcass traits in the same breed. Tambasco et al. (2003),
Silveira et al. 877
Table 4 - Mean estimated breeding values (EBV) for birth weight (BW), weaning weight (WW), yearling weight (YW) and mean plasma insulin-like
growth factor-1 (IGF-1) concentration for different genotypes of the growth hormone 1 gene (GH1) polymorphisms AluI and DdeI of Canchim calves.
Polymorphism and genotype EBV-BW EBV-WW (*) EBV-YW IGF-1
AluI
L/L -0.0114 ±2.04 4.6527 ±8.80 7.3141 ±9.08 141.07 ±79.72
L/V 0.1998 ±1.65 1.8626 ±7.80 4.1700 ±8.07 128.50 ±87.70
DdeI
DdeI+/DdeI+ 0.4718 ±2.29 7.8330 ±9.86 8.6167 ±7.72 140.24 ±82.00
DdeI+/DdeI- 0.0199 ±1.78 2.8291 ±7.96 5.5099 ±9.41 136.42 ±80.20
DdeI-/DdeI- - 0.3694 ±1.97 3.3481 ±8.05 7.4510 ±8.95 141.84 ±83.69
* Significant difference between genotype means obtained for the Dde I polymorphism (F test, p < 0.05).
Table 5 - Mean estimated breeding values (EBV) for birth weight (BW), weaning weight (WW), yearling weight (YW) and mean plasma insulin-like
growth factor-1 (IGF-1) concentration for the combined growth hormone 1 gene (GH1) polymorphisms AluI and DdeI genotypes of Canchim calves.
Genotype EBV-BW EBV-WW (*) EBV-YW IGF-1
L plus DdeI+/L plus DdeI+ 0.5336 ±2.43 8.6995 ±10.13 8.8681 ±8.06 144.62 ±84.46
L plus DdeI+/L plus DdeI- -0.0198 ±1.84 3.6879 ±8.18 6.5183 ±9.61 138.92 ±75.61
L plus DdeI+/V plus DdeI+ 0.2966 ±1.84 5.3780 ±8.89 7.9042 ±6.97 128.45 ±76.90
L plus DdeI-/L plus DdeI- -0.3694 ±1.96 3.3481 ±8.05 7.4510 ±8.91 141.84 ±83.69
L plus DdeI-)/V plus DdeI+ 0.1492 ±1.59 0.0285 ±6.58 2.2218 ±8.04 128.52 ±94.62
* Significant difference between means (F test, p < 0.05).
Table 6 - Average effects of haplotype substitution according to the combined distribution of the growth hormone 1 gene (GH1)AluI and DdeI
polymorphisms on birth weight (BW), weaning weight (WW), yearling weight (YW) and mean plasma insulin-like growth factor-1 (IGF-1) concentra-
tion of Canchim calves.
Haplotype BW WW (*) YW IGF-1
V plus DdeI+ with L plus DdeI+ 0.52 ±0.76 2.32 ±5.91 4.26 ±7.42 13.51 ±15.59
V plus DdeI+ with L plus DdeI- -0.67 ±0.79 -1.24 ±5.60 2.57 ±7.47 16.29 ±15.91
L plus DdeI- with L plus DdeI+ 1.20 ±0.44 3.56 ±3.35 1.69 ±4.37 -2.78 ±9.06
* Significant effect (F test, p < 0.05) of substitution of L plus DdeI- haplotype with L plus DdeI+ haplotype.
studying Canchim x Nelore, Simental x Nelore and Angus
x Nelore crossbred beef cattle, observed a higher weight
gain between birth and weaning in animals with the L/L ge-
notype compared to those with the L/V genotype, while the
opposite was noted between weaning and one year of age.
In a review, Switonski (2002) concluded that most studies
found lower growth rates in V/V cattle compared to those
with the other two genotypes (Chrenek et al., 1998;
Oprzadek et al., 1999; Sirotkin et al., 2000; Grochowska et
al., 2001). These conflicting results suggest that the AluI
polymorphism is not directly responsible for the phenotype
variations and that the contradictory data can be explained
by differences in the linkage disequilibrium between mark-
ers and quantitative trait loci (QTL) between the various
populations studied, or by different epistatic interactions
between the genetic bases of these populations and QTL.
On the other hand, it is interesting to note that in beef cattle,
in addition to the direct effect of GH on growth, its effect on
milk yield can also affect the phenotype of the animals. If
the effects of the alleles on milk yield and growth were an-
tagonistic, this factor would explain part of these controver-
sial results, especially those regarding weaning and
postweaning weight. Since the V allele of the AluI poly-
morphism is rare, or does not exist, in Zebu cattle, the chro-
mosome region corresponding to this restriction site in
Canchim cattle carrying this allele probably originated
from the Charolais breed. Canchim cattle heterozygous at
the AluI locus are therefore also expected to be heavier be-
cause Charolais taurine cattle are normally heavier than
Nelore indicine cattle, especially when the production sys-
tem favors the expression of these differences.
Although the DdeI polymorphism does not cause an
amino acid substitution in the protein sequence, Yao et al.
(1996) observed a highly positive association between the
DdeI+ allele and milk, fat and protein yield in Holstein cat-
tle. With respect to growth traits, no data are available in the
literature to permit comparison with our present results,
probably because the polymorphism is silent, thus not
arousing the interest of researchers.
The results of the effect of the haplotypes on the traits
analyzed, in contrast to those observed for the separate ge-
notype distributions, seem to better explain the variation
observed by us in the estimated breeding values.
It is interesting to note that the differences in weaning
weight were not accompanied by differences in yearling
weight as would be expected. The effect of substitution
with the L plus DdeI+ haplotype was positive for the three
production traits analyzed, although the standard error
found in our analysis did not indicate that the values ob-
tained were significantly different from zero, except for the
effect of substitution of L plus DdeI- with L plus DdeI+ on
birth weight. The effect of substituting haplotype V plus
DdeI+ with haplotype L plus DdeI- was negative for wean-
ing weight and positive for yearling weight. These results
might be explained by antagonistic effects of some
haplotype on growth and maternal ability.
The lack of effect of the GH1 genotypes on plasma
IGF-1 concentration at weaning was due to the wide varia-
tion observed in this trait, as demonstrated by its coefficient
of variation (58.72%), indicating that a very large number
of animals would be necessary to observe significant ef-
fects. In addition, collection of a single blood sample for the
determination of IGF-1 does not seem to be an appropriate
strategy since it does not permit assessment of the release
profile of this growth factor. Schlee et al. (1994b) reported
higher plasma GH levels in L/L cattle, a finding also re-
ported by Sorensen et al. (2002), whereas heterozygous
L/V cattle presented more elevated plasma IGF-1 levels. In
contrast, Grochowska et al. (2001) found a positive associ-
ation between the V/V genotype and peak GH, whereas
higher IGF-1 levels were associated with the L/L genotype.
These controversial results might be attributed to the lack
of uniformity of the population studied in terms of breed,
sex, age and productive capacity, as well as to the experi-
mental method used.
In conclusion, based on the analysis of the genotype
frequencies and of the effects of the polymorphisms on the
different weight parameters it is evident that the restriction
enzymes analyzed determine haplotypes that should be
considered in the study of growth hormone 1 gene poly-
morphisms. In addition, the results indicate that the GH1
polymorphisms can be used for the selection for growth
traits, with the haplotype containing the two restriction sites
provoking a greater increase in weaning weight in the
Canchim herd studied.
References
Boldman KG, Kriese LA, Van Vleck LD, Van Tassel CP and
Kachman SD (1995) A Manual for Use of MTDFREML: A
Set of Programs to Obtain Estimates of Variance and Co-
Variance. Agricultural Research Service, Lincoln, 125 pp.
Chrenek P, Kmet J, Sakowski T, Vasicek T, Huba J and Chrenek J
(1998) Relation-ships of growth hormone genotypes with
meat production traits of Slovak Pied bulls. Czech J Anim
Sci 43:541-544.
Curi RA, Palmieri DA, Suguisawa L, de Oliveira HN, Silveira AC
and Lopes CR (2006) Growth and carcass traits associated
with GH1/Alu I and POU1F1/Hinf I gene polymorphisms in
Zebu and crossbred beef cattle. Genet Mol Biol 29:56-61.
Davis ME, Boyles SL, Moeller SJ and Simmen RC (2003) Ge-
netic parameter estimates for serum insulin-like growth fac-
tor-I concentration and ultrasound measurements of backfat
thickness and longissimus muscle area in Angus beef cattle.
J Anim Sci 81:2164-2170.
Di Stasio L, Sartore S and Alberta A (2002) Lack of association of
GH1 and POU1F1 gene variants with meat production traits
in Piedmontese cattle. Anim Genet 33:61-64.
Goodman HM (1993) Growth hormone and metabolism. In:
Schreibman MP, Scanes CG and Pang PKT (eds) The Endo-
crinology of Growth, Development and Metabolism in Ver-
tebrates. Academic Press, San Diego, pp 93-115.
878 GH1 gene polymorphisms in beef cattle
Gordon DF, Quick DP, Erwin CR, Donelson JE and Maurer RA
(1983) Nucleotide sequence of the bovine growth hormone
chromosomal gene. Mol Cell Endocrinol 33:81-95.
Grochowska R, Sorensen P, Zwierzchowski L, Snochowski M
and Lovendahl P (2001) Genetic variation in stimulated GH
release and in IGF-I of young dairy cattle and their associa-
tions with the leucine/valine polymorphism in the GH gene.
J Anim Sci 79:470-476.
Kemenes PA, Regitano LCA, Rosa AJM, Packer IU, Razook AG,
Figueiredo LA, Silva NA, Tchegaray MA and Coutinho LL
(1999) k-casein, b-lactoglobulin and growth hormone allele
frequencies and genetic distances in Nelore, Gyr, Guzerá,
Caracu, Charolais, Canchim and Santa Gertrudis Cattle.
Genet Mol Biol 22:539-541.
Nicoll CS, Mayer GL and Russell SM (1986) Structural features
of prolactins and growth hormones that can be related to
their biological properties. Endocr Rev 7:169-203.
Oprzadek J, Dymnicki E, Zwierzchowski L and Lukaszewicz M
(1999) The effect of growth hormone (GH), κ-casein
(CASK) and β-lactoglobulin (BLG) genotypes on carcass
trait in Friesian bulls. Anim Sci Pap Rep 17:85-92.
SAS (1999) Statistical Analysis System, Systems for Windows.
SAS Institute Inc., Cary, NC.
Schlee P, Graml R, Rottmann O and Pirchner F (1994a) Influence
of growth hormone genotypes on breeding values of Sim-
mental bulls. J Anim Breed Genet 111:253-256.
Schlee P, Graml R, Schallenberger E, Schams D, Rottmann R,
Olbrich-Bludau A and Pirchner F (1994b) Growth hormone
and insulin-like growth factor 1 concentrations in bulls of
various growth hormone genotypes. Theor Appl Genet
88:497-500.
Sirotkin AV, Chrenek P, Makarevich AV, Huba J and Bulla J
(2000) Interrelation-ships between breed, growth hormone
genotype, plasma IGF-I level and meat performance in bulls
of different ages. Arch Anim Breed 43:591-596.
Sorensen P, Grochowska R, Holm L, Henryon M and Lovendahl
P (2002) Polymorphism in the bovine growth hormone gene
affects endocrine release in dairy calves. J Dairy Sci
85:1887-1893.
Switonski M (2002) Molecular genetics in beef cattle breeding - A
review. Anim Sci Pap Rep 20:7-18.
Tambasco DD, Alencar MM, Coutinho LL, Tambasco AJ, Tam-
basco MD and Regitano LCA (2000) Caracterização molec-
ular de animais da raça Nelore utilizando microssatélites e
genes candidatos. Rev Bras Zootec 29:1044-1049.
Tambasco DD, Paz CCP, Tambasco-Studart MD, Pereira AP,
Alencar MM, Freitas AR, Coutinho LL, Packer IU and
Regitano LCA (2003) Candidate genes for growth traits in
beef cattle crosses Bos taurus xBos indicus. J Anim Breed
Genet 120:51-56.
Unanian MM, Barreto CC, Freitas AR, Cordeiro CMT and Josah-
kian LA (2000) Associação do polimorfismo do gene do
hormônio de crescimento com a característica peso em bovi-
nos da raça Nelore. Rev Bras Zootec 29:1380-1386.
Yao J, Aggrey SE, Zadworny D, Hayes JF and Kuhnlein U (1996)
Sequence variations in the bovine growth hormone gene
characterized by single-strand conformation polymorphins
(SSCP) analysis and their association with milk production
traits in Holsteins. Genetics 144:1809-1816.
Associate Editor: Luiz Lehmann Coutinho
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