Keywords:

Creole bovine

Molecular markers

Precocity

Productivity

Zoogenetic resource

Effect of IGF-1 and LEP/ob SNPs on growth parameters of Blanco Orejinegro cattle

Efecto de SNPs de IGF-1 y LEP/ob sobre parámetros de crecimiento del ganado Blanco Orejinegro

Efeito dos SNPs de IGF-1 e LEP/ob nos parâmetros de crescimento do gado Blanco Orejinegro

Marisol Londoño-Gil

Luis Gabriel González-Herrera

Albeiro López-Herrera

Juan Carlos Rincón Flórez

Rev. Fac. Agron. (LUZ). 2022, 39(2): e223933

ISSN 2477-9407

DOI: https://doi.org/10.47280/RevFacAgron(LUZ).v39.n2.11

Animal Production

Associate editor: Dra. Rosa Razz

University of Zulia, Faculty of Agronomy

Bolivarian Republic of Venezuela

Abstract

The aim of this work was to identify the effect of some SNPs of the

IGF-1 and LEP/ob genes, on the growth in animals of the Blanco Orejinegro

creole breed (BON) and to evaluate the relationship of age at rst calving

(AFC) with the curve trajectory. For this, 1217 phenotypic and 439 genomic

records of pure BON bovines were used. The Gompertz, Logistic, Von

Bertalanffy and Brody models were evaluated. The individual growth curve

parameters were estimated and the effect of SNPs of the IGF-1 and LEP/

ob genes on the curve parameters was estimated through a linear model.

Finally, the association of the curve parameters and the AFC was analyzed

trough a linear model. The model that represented the best t to the growth

trajectory was Brody’s. On average, the BON animals presented an adult

weight (β

) of 479.9±7.4 kg and a growth rate expressing the daily weight

gain as a proportion of the total weight (β

) of 0.002±0.00004. The SNPs

rs110654613 (nucleotide change A/G) and rs110959643 (A/G), within the

IGF-1 gene, showed a signicant effect (p<0.05) on the parameters of the

BON cattle growth curve. There were no associations of LEP/ob gene SNPs

on the parameters of the BON cattle growth curve. AFC was signicantly

associated (p<0.05) with the parameters β

and β

. It is concluded that the use

of genomic information for the IGF-1 gene can lead to higher growth rates

and earlier AFC.

Universidad Nacional de Colombia sede Medellín,

Departamento de Producción Animal, Grupo de investigación

Biodiversidad y Genética Molecular (BIOGEM), Carrera 65

N 59A-110, CP 050034, Medellín, Colombia.

Universidad Nacional de Colombia sede Palmira,

Departamento de Ciencia Animal, Facultad de Ciencias

Agropecuarias, Carrera 32 N 12 - 00, CP 763352, Palmira,

Colombia.

Received:

30-10-2021

Accepted: 30-05-2022

Published: 21-06-2022

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2022, 39(2): e223932. April - June. ISSN 2477-9407.2-7 |

Resumen

El objetivo de este trabajo fue identicar el efecto de algunos SNPs

de los genes de IGF-1 y LEP/ob, sobre el crecimiento de animales de

la raza criolla Blanco Orejinegro (BON) y evaluar la relación de la

edad al primer parto (EPP) con la trayectoria de la curva. Para ello

se utilizaron 1.217 registros fenotípicos y 439 registros genómicos

de bovinos puros BON. Los modelos de Gompertz, Logístico, Von

Bertalanffy y Brody fueron evaluados y se estimaron los parámetros

individuales de la curva de crecimiento. Se estimó el efecto de SNPs

en los genes IGF-1 y LEP/ob sobre los parámetros de la curva, a

través de un modelo lineal. Finalmente, se analizó la asociación de

los parámetros de la curva y su trayectoria con la EPP, mediante

un modelo lineal. El modelo que representó un mejor ajuste a la

trayectoria de crecimiento fue el de Brody. En promedio los animales

BON presentaron un peso adulto (β

) de 479,9±7,4 kg y una tasa de

crecimiento expresando la ganancia diaria de peso como proporción

del peso total (β

) de 0,002±0,00004. Los SNP rs110654613 (cambio

nucleotídico A/G) y rs110959643 (A/G), dentro del gen IGF-1,

mostraron efecto signicativo sobre los parámetros de la curva de

crecimiento de ganado BON. No hubo asociaciones de SNPs del gen

LEP/ob con la curva de crecimiento de ganado BON. La EPP estuvo

asociada signicativamente (p<0,05) con los parámetros β

. Se

concluye que el uso de información genómica para el gen IGF-1

puede llevar a mayores tasas de crecimiento y EPP más tempranos.

Palabras clave: bovino criollo, marcadores moleculares, precocidad,

productividad, recurso zoogenético.

Resumo

O objetivo deste trabalho foi identicar os efeitos de alguns

SNPs dos genes IGF-1 e LEP/ob, sobre a curva de crescimento de

animais da raça Blanco Orejinegro (BON) e avaliar a relação da idade

ao primeiro parto (IPP) com a trajetória da curva. Para isso, foram

utilizados 1.217 registros fenotípicos e 439 registros genômicos de

bovinos BON puros. Foram avaliados os modelos de Gompertz,

Logístico, Von Bertalanffy e Brody. Os parâmetros individuais da

curva de crescimento foram estimados e o efeito dos SNPs dos genes

IGF-1 e LEP/ob sobre os parâmetros da curva foi estimado por meio

de um modelo linear. Finalmente, a associação dos parâmetros da

curva e sua trajetória com a IPP foi analisada por meio de um modelo

linear. O modelo que mais se adequou à trajetória de crescimento foi

o de Brody. Em média, os animais BON apresentaram peso adulto

(β

) de 479,9±7,4 kg e uma taxa de crescimento expressando o ganho

de peso diário em proporção ao peso total (β

) de 0,002±0,00004.

Os SNPs rs110654613 (mudança de nucleotídeo A/G) e rs110959643

(A/G), dentro do gene IGF-1, mostraram um efeito signicativo

(p<0,05) nos parâmetros da curva de crescimento de bovinos BON.

Não houve associações com SNPs do gene LEP/ob com os parâmetros

da curva de crescimento de gado BON. A IPP foi signicativamente

associada (p<0,05) aos parâmetros β

e β

. Conclui-se que o uso de

informações genômicas para o gene IGF-1 pode levar a maiores taxas

de crescimento e IPP mais precoce.

Palavras-chave: polimorsmos, precocidade, produtividade, raça

crioula, recurso zoogenético.

Introduction

Growth is dened as the increase in weight of animals from

birth to its stabilization in adulthood (Ramírez et al., 2009), where

both biological and environmental aspects interfere (Rincón and

Quintero, 2015). Some of the biological aspects are hormonal that

regulate various productive and reproductive characteristics in the

body. For example, insulin-like growth factor 1 (IGF-1), which is a

protein hormone that plays an important role in various metabolic and

physiological processes and in growth with anabolic effects (Castrelln

et al., 2010). The bovine IGF-1 gene is located on chromosome 5

(Bos taurus autosome – BTA5) and has been proposed as a candidate

gene for growth traits in cattle (Rogberg-Muñoz et al., 2013). Another

important hormone involved in energy metabolism is leptin (Saleem,

2015), a protein that regulates appetite and metabolism (Lusk, 2007)

and has been used as a biological marker that reects the degree of fat

in the body. Leptin is encoded by the LEP/ob gene, which in bovines

is located on chromosome 4 (BTA4).

In cattle, multiple genetic variants have been reported within the

IGF-1 gene, many of them single nucleotide polymorphisms (SNPs),

some signicantly associated with phenotypic variation and breeding

values in the early growth phase (Andrade et al., 2008; Castrelln

et al., 2010). It has also been proposed that the LEP/ob gene is

associated with parameters of the growth curve and backfat in cattle

(Lusk, 2007), and that variations in the SNP UASMS2 (C/T) of this

gene have been signicantly inuential in the variability found for

parameters of the growth curve (Lusk, 2007). However, the study

of the association of markers of some genes with parameters of the

growth curve has not been carried out in any of the Colombian Creole

breeds, not even in the Blanco Orejinegro (BON), one of the most

widely reported.

The BON is a breed that has undergone a process of natural

selection close to 500 years in the conditions of the Colombian

tropics (López-Herrera et al., 2001), belongs to the Bos taurus species

(Rincón and Quintero, 2015) and is characterized phenotypically due

to its white fur, black ears, skin and nose (López-Herrera et al. 2001).

Its economic importance lies in its rusticity, ability to efciently take

advantage of low-quality forages and to be in steep terrain, ability to

reproduce, survive, be long-lived and because it is a triple-purpose

animal (milk, meat and work) (Bedoya et al., 2001; López-Herrera

et al., 2001).

In cattle studies, Inoue et al. (2020) reported a negative genetic

association between the adult weight of Japanese Black cattle and

the age at rst calving (AFC), where females with a lower adult

weight would show a higher AFC. Likewise, these authors found a

positive but low association between the maturation rate and AFC. In

Holstein cattle in Brazil, Coelho et al. (2009) found positive, but not

signicant, correlations between AFC and adult weight; and negative

and signicant between AFC and the maturation rate, suggesting that

cows with higher maturation rates have a better productive efciency.

In BON cattle, there is no type of study associating AFC with growth

curve parameters, and the information available on BON growth at

the molecular level is scarce (Caivio-Nasner et al., 2021; Cañas et al.

, 2008; Londoño-Gil et al., 2021), so it is important to study the allelic

variants of certain genes or regions of the genome that are related

to growth variation in this breed and, in turn, to AFC (productive

ability and reproductive). Therefore, the objective of this work was to

identify the effect of some SNPs of the IGF-1 and LEP/ob genes on





= 



−1

− 2

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Londoño-Gil et al. Rev. Fac. Agron. (LUZ). 2022, 39(2): e2239323-7 |

the growth curve of animals of the Creole breed Blanco Orejinegro

(BON) and to evaluate the relationship between age at rst calving

(AFC) with the trajectory of the curve.

Materials and methods

Data and animals of the BON breed

To model the growth of the animals, phenotypic and genotypic

information was used from a BON cattle database belonging to 7

Colombian herds located in the departments of Antioquia, Caldas,

Meta, Risaralda and Tolima. Management and environmental

conditions varied according to the herd and its location. Historical

production data from 1993 to 2018 were used. The information was

analyzed and rened in the R program (R Core Team, 2020), taking

into account the weight at birth as a starting point for modeling the

growth curve, guaranteeing a minimum of 4 weighings per animal

and that the last weight measurement had been recorded at least at

800 days (26.31 months) of age. After purication, the information

from 11,140 consecutive weighings in the life of 1,217 animals (9.15

weighings per animal) was used to estimate the parameters of the

growth curves.

Growth curve tting models

To estimate the curve that best t the data, the models were used:

Gompertz (1825) ; Logistical (Verhulst, 1838):

Von Bertalanffy (1938):

and Brody (1945): ; where: corresponds

to the j-th weight of the i-th animal at time “t”, β

: corresponds to

the asymptotic weight when “t” tends to innity and is commonly

interpreted as the adult weight, β

: corresponds to a parameter of t

when Y ≠ 0 and when t ≠ 0, and indicates the proportion of asymptotic

mature weight that is gained after birth, β

: corresponds to the maturity

index represented as a percentage proportion of maximum growth

with respect to the adult weight of the animal; t

is the age in days

at the j-th weighing of the i-th animal (Dominguez-Viveros et al.,

2017). The t of the models to the data was determined by the criteria

of lowest sums of squares of the error (SQE), Akaike information

criterion (AIC), Bayesian information criterion (BIC), and highest

percentages of convergence (%C) and coefcient of determination

). Finally, the individual parameters of the growth curve of each of

the animals were estimated.

SNPs within LEP/ob and IGF-1 genes

Genotype information was obtained for 439 individuals, each with

genotype information for 118,116 SNPs, which were part of the BON

cattle data bank. These genotypes were coded as 0: homozygous for

the rst allele, 1: heterozygous, and 2: homozygous for the second

allele. From the GenBank assembly ARS-UCD1.2 (https://www.

ncbi.nlm.nih.gov/assembly/GCA_002263795.2; GenBank accession

number NKLS00000000.2), where mapped SNPs located within

IGF-1 genes (AC_000162.1; 66523798…66604881) and LEP/

ob (AC_000161.1; 93249803…93266625), located in BTAs 5 and

4, respectively. Five SNPs were identied within the IGF-1 gene:

rs133675145 change (A/G), rs110654613 (A/G), rs110959643 (A/G),

rs109297640 (A/G) and rs109148720 (T/G); and two within the LEP/

ob gene: rs136588988 (T/C) and rs110559656 (A/G).

Association of SNPs with the variation of the growth curve

format

Once the parameters of the curve were estimated, individuals

that presented a value for the parameter β

(adult weight) between

300 and 600 kg were kept, considering them biologically possible

in the breed; records of animals that were well represented by herd

(minimum 6 individuals per herd), sex, season (summer 1: January,

February, March; summer 2: July, August, September; winter 1:

April, May, June; and winter 2: October, November, December),

year (from 2009 to 2017, minimum 3 observations per year) and also

had a genotype. Finally, 90 animals (69 females and 21 males) had

consistent phenotypic (growth curves) and genotypic information.

The linear model used to evaluate the effect of the variations of the

SNPs on the parameters of the growth curve of these animals was:

Where, is the parameter evaluated for growth in

animal i; μ is the general mean of the evaluated traits, and

are the xed effects of herd, season

of birth, year of birth and sex of the individuals, respectively. is

the effect of the marker variant of the individually evaluated

SNPs and is the random error associated with each

observation (Cardona et al., 2015). The lm function was used and the

means of the genotypes were compared using the lsmeans package

(Lenth, 2016), through the Tukey test in the R software (R Core

Team, 2020). The curves of the animals with signicant SNPs were

plotted in the R software, to visually observe the differences in the

growth curves according to the genotype.

Association of growth curve parameters with AFC

To make this association, information from 140 females that had

sufcient and consistent information on growth curve parameters and

AFC was used. The linear model used was:

Where, is the AFC in animal i; μ is the overall mean of the

AFC, and are the xed effects of herd and year of birth,

respectively; are the curve parameters of the model that

was chosen as the best and is the random error associated with

each observation. This analysis was performed using the lm function

and the means of signicant effects were compared by Tukey’s test

using the lsmeans package using R software (R Core Team, 2020).

Results and discussion

Table 1 shows the results of the t evaluation of the models used

and the means of the parameters obtained for each model.

The model that presented a better t for the description of the

growth curve of BON animals was the Brody model, with the lowest

AIC, BIC and SQE, while obtaining the highest R

. This model has

been widely used to describe the growth curve in cattle, since it

usually presents the best ts (Agudelo et al., 2008: Ramírez et al.,

2009; Rincón and Quintero, 2015), especially at ages after 6 months,

since for rst months of life it tends to underestimate the weight

(Agudelo et al., 2008). However, for authors such as Dominguez-

Viveros et al. (2017) the model with the best performance for grazing

zebu cattle was the logistic one.

The average parameters with biological interpretation in the Brody

model were: β

: 479.9 ± 7.4 kg (adult weight) and β

: 0.002 ± 0.00004

(maturity index or growth rate expressing the daily gain of weight as a

proportion of the total weight), large numbers in this index indicate an

early maturation of the animals (Hojjati and Hossein-Zadeh, 2018).

These results are consistent with those previously reported in the

BON breed by Rincón and Quintero (2015), under the Brody model,

the data t better, with higher adult weights (590 kg) and a maturity





= 



−1

− 2





= 

+ (1 − 

−1

)

−1

;





= 

(1 − 



−2

)





= 

+ (1 − 



−2

)





ijklmn

= μ + 

+ 

+ 

+ Sex



+ SNP



+ 

ijklmn

ijklm



, 

, 

, Sex



SNP





ijklmn

ijk

= μ + 

Year + 

+ 

+ 

ijk



and 



, β

and β



ijk

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Rev. Fac. Agron. (LUZ). 2022, 39(2): e223932. April - June. ISSN 2477-9407.4-7 |

Table 1. Evaluation of t of non-linear models used in the description of the growth curve of animals of the Blanco Orejinegro (BON)

breed from Colombia and estimated mean parameters.

Model AIC BIC SQE R

% C β

Bertalanfy 119238.2 119267.5 29014728 0.79 50.53 389.4 0.5155 0.003

Logistic 119944.2 119973.5 30912991 0.78 84.96 347.0 4.95 0.006

Brody 118939.9 118969.2 28248116 0.80 76.83 474.9 0.924 0.002

Gompertz 119414.9 119444.1 29478432 0.79 85.30 372.0 2.042 0.004

AIC: Akaike Information Criterion, BIC: Bayesian Information Criterion, SQE: Error Sum of Squares, R

: Determination Coefcient, %C: Convergence Percentage.

index of 0.00153. The small differences may be due to the fact that

the parameters of the growth curves are probably inuenced either

by the genetic component of the animals studied, by management,

by their ability to adapt to their environment (Rincón and Quintero,

2015), or because in the previous BON study information from a

single herd was used, so there was probably greater homogeneity.

Association of the SNPs with the variation of the growth

curve format

Of the 7 SNPs under study (2 within the LEP/ob gene and 5 SNPs

within the IGF-1 gene), only 2, within the IGF-1 gene, showed a

signicant association with the growth curve parameters (table 2),

corresponding to markers rs110654613 (β

) and rs110959643 (β

and β

In table 2, it is observed that the herd and the year of birth inuence

all the parameters of the curve, thus affecting the complete growth

curve of the BON animals. This is important, since differences are

generated in the growth of the animals that can be attributed to

changes in forage availability, selection and management processes

in the different herds, as well as climatic variations between years

(Martins et al., 2000). Similarly, in studies of growth traits in BON

cattle, Martínez et al. (2012) found that herd, year of birth and sex

were important sources of variation. On the other hand, the SNP

rs110959643 seems to be almost xed in this population sample,

since of the 90 animals under study, 85 animals presented genotype

0, ve 1 and none 2 (0: homozygous for the rst allele - GG, 1:

heterozygous - GA, 2: homozygous for the second allele - AA). This

situation may be because the animals in the population have been

selected, naturally or articially, towards genotype 0 (GG), due to

the effects of drift, increased inbreeding, structuring or imbalance

in the model, which could generate a spurious association and affect

the robustness of the result. Gui et al. (2018) in a study evaluating

the inuence of IGF-1 gene polymorphisms on growth traits,

showed that the low sample number can lead to certain genotypes

not being observed. However, in the present study, the individuals

included came from a database where there were several selection

criteria for genotyping, such as being the most representative of

the population, having several phenotypic records, among others,

which does not allow the sample was not representative.

The mean difference for each of the parameters of the growth

curve according to the genotype of the SNPs is shown in table 3. For

the parameter β

, related to the asymptotic adult weight, the animals

with genotype 1 (heterozygous - GA) for SNP rs110959643 will

have a lighter adult weight (427 kg) compared to the homozygous

genotype 0 (GG - 534 kg), which is consistent with what has been

previously reported in other beef cattle breeds.

Table 2. SNPs and factors associated with each of the parameters

of the growth curve under the Brody model in a

population of Blanco Orejinegro (BON) cattle from

Colombia.

Parameter SNP Year Sex Herd Season R

rs110959643* ** ** ** NS 0.45

rs110959643** ** NS ** NS 0.45

rs110654613* ** NS ** NS 0,48

**p<0,01, *p<0,05, NS p>0,05. rs110959643 y rs110654613: Intronic variants

of the IGF-1 gene. R

: Coefcient of determination.

Table 3. Adjusted means of the parameters of the growth curve

(β

y β

) according to the genotype of the associated

SNPs in a population of BON cattle from Colombia.

SNP/Genotype

0 1 2

rs110959643 - β

534 ± 20.3

427 ± 48.3

rs110959643 - β

0.937 ± 0.004

0.906 ± 0.010

rs110654613 - β

0.0015 ± .0001

0.0009 ± 0.0002

rs110959643 and rs110654613: Intronic variants of the IGF-1 gene. Same letters

on the same line do not differ statistically. For rs110959643, 0 = GG, 1 = GA, 2

= AA; for rs110654613 0 = GG, 1 = GA, 2 = AA.

Rogberg-Muñoz et al. (2011) found that the SNP rs110959643

has important implications in the productivity of animals, since this

gene has been reported to have a role in the growth and development

of bovines and its polymorphisms have been associated with

growth traits (weight at birth, daily weight gain, adult weight and

other body weights) and development of mammals. The effect of

these SNPs on the growth curve of BON animals can be veried

graphically in gure 1.

Figure 1. Average growth curve according to the genotype of the

SNP rs110959643 associated with the parameters β0

and β1 of the Brody model.

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Londoño-Gil et al. Rev. Fac. Agron. (LUZ). 2022, 39(2): e2239325-7 |

Regarding the parameter β

, associated with the maturity index,

in table 3 a dominance effect is observed for the SNP rs110654613

(IGF-1 SNP), where animals with genotypes 0 (GG) or 1 (GA)

(i.e., homozygous for the rst allele or heterozygotes) will have a

higher maturity index and therefore will reach maturity earlier

than homozygous 2 (AA) (0.001567, 0.001564 and 0.000942,

respectively) (gure 2). These results disagree with reported in

Brahman cattle from Brazil by Crispim et al. (2015), who found that

for the maturity index (β

) the most signicant SNP was found in BTA

20 and was rs42482169. For this SNP the mean of the homozygous

dominant genotype was 0.0063, for the heterozygous 0.0064 and for

the homozygous recessive 0.0068.

Figure 2. Average growth curve according to the genotype of the

SNP rs110654613 associated with the parameter β

of the

Brody model (1945).

The SNPs rs110959643 (5:66228391) and rs110654613

(5:66224046), important for the growth curve of Colombian Creole

cattle BON, are located within introns of the IGF-1 gene, with a

difference of 4345 bp between each other and with a high degree

of linkage with the possible causal variant, given the linkage

disequilibrium that exists in the BON breed, of 0.39 at distances

less than 25 Kb (Caivio-Nasner et al., 2021). The IGF-1 gene has

previously been related to myogenic factors directly involved in

growth and sexual precocity, due to the effect of this gene on different

metabolic and endocrine pathways, which have shown a connection

between growth and the beginning of reproductive life (Mota et al.,

2020). This has been shown in Nelore females from Brazil, where

the higher concentration of the hormone IGF-1 in the blood has been

associated with greater growth and, therefore, a lower age at puberty

and a lower AFC (Ferraz et al., 2018) and in Canchim animals

(Gui et al., 2018), where its importance in growth traits has been

demonstrated. This gene is important in the modulation of a variety of

physiological activities such as cell development, differentiation and

proliferation, leading to increased muscle growth (Gui et al., 2018);

therefore, by nding an association of SNPs of the IGF-1 gene with

the parameters of the growth curve in this study, it can be inferred that

it can be used as a candidate gene for early selection of BON animals

for future work.

Finally, the SNPs within the LEP/ob gene did not show a

signicant association with growth or AFC. Previously, the LEP/

ob gene has been associated with parameters of the growth curve

and backfat of cattle (Lusk, 2007) and it has been reported that the

variations in the SNP UASMS2 of this gene have been signicant for

the body weight growth curve (p<0.001). This is in contrast to what

was reported by Ferraz et al. (2018), who stated that this gene has no

implications for age at puberty, and that hormone secretion after 18

months was very low, which would help to understand why in the

present study the LEP/ob gene had no signicant effect.

Association of the growth curve parameters with AFC

In this study, the mean AFC for the BON females analyzed was

found to be 1,044.04 ± 177.04 days or 34.23 ± 5.80 months. This AFC

is lower than that reported by M-Rocha et al. (2012) in 22 populations

of BON cattle, of 1,104 ± 141 days (36.8 months).

The AFC under study was signicantly associated with the

parameters β

(p = 0.0124), β

(p = 0.0117) and with the herd of origin

(p = 0.0010), which can be evidenced in the table 4, but not with the

year of birth (P = 0.0654), nor the parameter β

(p = 0.2414) (data not

shown). These ndings differ from those reported by M-Rocha et al.,

(2012), who found no effect of herd or year on AFC (p>0.05).

Table 4. Adjusted means of AFC according to the birth herd of

BON females from Colombia.

Herd PPE adjusted mean

1 830 ± 91.7

2 978 ± 35.6

3 991± 31.8

4 1029 ± 54.6

5 1061 ± 38.9

6 1140 ± 47.3

7 1308 ± 77.3c

a, b, c

Means with different letters differ signicantly (p<0.05) from each other.

The parameter β

, associated with adult age, showed that for each

kg more of weight at adult age in BON cattle, the AFC of BON females

decreased by an average of 0.538 ± 0.212 days. In addition, for one

unit of change in the parameter β

, the AFC decreases by an average

of 0.000719 ± 0.0003 days in BON females from Colombia; showing

that animals that grow faster and have a higher adult weight, will be

those that possibly have an earlier AFC. These animals will continue

to grow after the rst service and will end up with a considerable

adult weight. These results differ from those found by Bayram et al.

(2004), in Brown Swiss cattle using the Richards model, who reported

a positive correlation between age at rst calving and estimated adult

weight (β

). For these authors, an increase in weight at the age at rst

calving meant an increase in adult weight.

Similarly, Gaviolli et al. (2012), reported positive phenotypic

correlations between AFC and the parameters β

and β

, indicating

that animals with higher adult weight and higher growth rate will have

a higher AFC, which is not desirable from the productive point of

view. The difference may be due to the differential management given

to BON females in each herd and effects of the breed. According to

the information provided by the producers of their herds, they often

prefer to wait for a specic age to serve their animals, instead of

doing it by weight, so animals that present a high growth rate can

be served late in his life. In each herd, a different handling of the

rearing of these BON females is carried out, and this should be taken

into consideration in future works because the present results show

that it is necessary to try to standardize the AFC according to the

weight, ensuring that this traits do not vary signicantly. Likewise,

given that the genotype for genes such as IGF-1 has an impact on the

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Londoño-Gil et al. Rev. Fac. Agron. (LUZ). 2022, 39(2): e2239326-7 |

growth rate of animals, it is important to take it into consideration as

a candidate gene within the improvement and management plans of

BON herds to select animals. more efcient both productively and

reproductively.

Conclusions

Of the methods evaluated in this research, Brody’s method

yielded the best estimate of the parameters studied, which makes

it a practical tool to evaluate the growth curves of BON animals in

Colombia, which presented an adequate adult weight and growth

rate, which allows them to be economically and environmentally

protable. In this study, it was found that two SNPs in the IGF-1 gene

inuenced the growth of BON cattle; however, no association was

found between the growth curve parameters and the LEP/ob gene. In

addition, the growth parameters, according to the Brody model, are

associated with the age at rst calving in the BON breed. This will

allow studies to be carried out and the IGF-1 gene to be applied as a

candidate gene for growth in the selection of these animals, which in

turn will allow improving AFC.

Funding source

This study was funded by the Universidad Nacional de Colombia,

Medellín, and the Ministerio de Ciencia Tecnología e Innovación of

Colombia - Minciencias, through a call for Young Researchers and

Innovators 812 - 2018, contract 289 - 2019.

Literature cited

Agudelo, D., Cerón, M., y Restrepo, L. (2008). Modelación de las funciones de

crecimiento aplicadas a la producción animal. Revista Colombiana de

Ciencias Pecuarias, 21(1), 39–58. https://doi.org/10.17533/udea.rccp

Andrade, P. C., Grossi, D. A., Paz, C. C. P., Alencar, M. M., Regitano, L. C. A., &

Munari, D. P. (2008). Association of an insulin-like growth factor 1 gene

microsatellite with phenotypic variation and estimated breeding values of

growth traits in Canchim cattle. Animal Genetics, 39(5), 480–485. https://

doi.org/10.1111/j.1365-2052.2008.01755.x

Bayram, B., Yanar, M., & Akbulut, Ö. (2004). Relationships among richards

growth curve parameters, reproductive and milk production traits in

brown swiss cattle sag feisfcfoh. Journal of Applied Animal Research,

26(1), 29–32. https://doi.org/10.1080/09712119.2004.9706500

Bedoya, G., Carvajal, L. G., Moreno, F. L., Davies, S., Derr, J., Ossa, J., y

Ruiz, A. (2001). Estructura molecular y poblacional del ganado criollo

Colombiano (GCC). Revista Colombiana de Ciencias Pecuarias, 14(2),

109–120. https://doi.org/10.17533/udea.rccp

Brody, S. (1945). Bioenergetics and growth with special reference to the efciency

in domestic animals. Hafner Publishing Company, INC. 1033 p. https://

onesearch.library.rice.edu

Caivio-Nasner, S., López-Herrera, A., González-Herrera, L. G., & Rincón, J. C.

(2021). Diversity analysis, runs of homozygosity and genomic inbreeding

reveal recent selection in Blanco Orejinegro cattle. Journal of Animal

Breeding and Genetics, 138(5), 613–627. https://doi.org/10.1111/

jbg.12549

Cañas, J. J., Ramírez, J., Arboleda, O., Ochoa, J., Vergara, O., y Cerón, M. (2008).

Estimación de parámetros genéticos para peso al destete en ganado blanco

orejinegro (BON) en el noroccidente colombiano. Revista MVZ Córdoba,

13(1), 1138–1145. https://doi.org/10.21897/RMVZ.405

Cardona, S. J. C., Álvarez, J. D. C., Sarmento, J. L. R., Herrera, L. G. G., y

Cadavid, H. C. (2015). Associação de SNPs nos genes para κ-caseína e

β-lactoglobulina com curvas de lactação em cabras leiteiras. Pesquisa

Agropecuaria Brasileira, 50(3), 224–232. https://doi.org/10.1590/S0100-

204X2015000300006

Castrelln, V. V., Vera, W. A., Bracamonte, M. P., Rincn, A. M. S., Montoya, H.

M., & De la Rosa Reyna, X. F. (2010). Polymorphisms in the IGF1

gene and their effect on growth traits in Mexican beef cattle. Genetics

and Molecular Research, 9(2), 875–883. https://doi.org/10.4238/vol9-

2gmr745

Coelho, J. G., Barbosa, P. F., Tonhati, H., & Freitas, M. A. R. de. (2009). Análise

das relações da curva de crescimento e eciência produtiva de vacas da

raça Holandesa. Revista Brasileira de Zootecnia, 38(12), 2346–2353.

https://doi.org/10.1590/S1516-35982009001200008

Crispim, A. C., Kelly, M. J., Guimarães, S. E. F., e Silva, F. F., Fortes, M. R.

S., Wenceslau, R. R., & Moore, S. (2015). Multi-Trait GWAS and New

Candidate Genes Annotation for Growth Curve Parameters in Brahman

Cattle. PLOS ONE, 10(10), 1–19. https://doi.org/10.1371/journal.

pone.0139906

Dominguez-Viveros, J., Urbina-Valenzuela, A. R., Palacios-Espinoza, A.,

Callejas-Juárez, N., Ortega-Gutiérrez, J. Á., Espinoza-Villavicencio, J. L.,

Padrón-Quintero, Y., y Rodríguez-Castro, M. (2017). Caracterización del

crecimiento de bovinos cebú en pruebas de comportamiento en pastoreo.

Ecosistemas y Recursos Agropecuarios, 4(11), 341–348. https://doi.

org/10.19136/era.a4n11.1149

Ferraz, M. V. C., Pires, A. V., Santos, M. H., Silva, R. G., Oliveira, G. B., D. M.,

Polizel, D. M., Biehl, M. V., Sartori, R., & Nogueira, G. P. (2018). A

combination of nutrition and genetics is able to reduce age at puberty in

Nelore heifers to below 18 months. Animal, 12(3), 569–574. https://doi.

org/10.1017/S1751731117002464

Gaviolli, V. R. N., Buzanskas, M. E., Cruz, V. A. R., Savegnago, R. P., Munari, D.

P., Freitas, A. R., & Alencar, M. M. (2012). Genetic associations between

weight at maturity and maturation rate with ages and weights at rst and

second calving in Canchim beef cattle. Journal of Applied Genetics,

53(3), 331–335. https://doi.org/10.1007/s13353-012-0100-6

Gompertz, B. (1825). On the nature of the function expressive of the law of human

mortality, and on a new mode of determining the value of life Contingencies.

Philosophical Transactions ofthe Royal Society of London, 115, 513–583.

http://www.jstor.org/journals/rsl.html

Gui, L., Wang, Z., Jia, J., Zhang, C., Chen, Y., & Hou, S. (2018). IGF-1 Gen

Polimorzmi Çin Qinchuan Sığırında Büyüme Özelliklerini Etkiler.

Kafkas Universitesi Veteriner Fakultesi Dergisi, 24(3), 387–392. https://

doi.org/10.9775/kvfd.2017.19036

Hojjati, F., & Hossein-Zadeh, N. G. (2018). Comparison of non-linear growth

models to describe the growth curve of Mehraban sheep. Journal of

Applied Animal Research, 46(1), 499–504. https://doi.org/10.1080/0971

2119.2017.1348949

Inoue, K., Hosono, M., Oyama, H., & Hirooka, H. (2020). Genetic associations

between reproductive traits for rst calving and growth curve

characteristics of Japanese Black cattle. Animal Science Journal, 91(1),

1–8. https://doi.org/10.1111/asj.13467

Lenth, R. V. (2016). Least-Squares Means: The R Package lsmeans. Journal of

Statistical Software, 69(1), 1–33. https://doi.org/10.18637/jss.v069.i01

López-Herrera, A., Saldarriaga, O. A., Arango, A. E., Fabio, N., Olivera, M.,

Bedoya, G., y Ossa, J. E. (2001). Ganado Blanco Orejinegro (BON):

Una alternativa para la producción en Colombia. Revista Colombiana de

Ciencias Pecuarias, 14(2), 121–128. https://doi.org/10.17533/udea.rccp

Londoño-Gil, M., Rincón Flórez, J. C., Lopez-Herrera, A., y Gonzalez-Herrera,

L. G. (2021). Genome-wide association study for growth traits in Blanco

Orejinero (BON) cattle from Colombia. Livestock Science, 243(104366),

104366. https://doi.org/10.1016/j.livsci.2020.104366

Lusk, J. L. (2007). Association of single nucleotide polymorphisms in the leptin

gene with body weight and backfat growth curve parameters for beef

cattle 1. American Society of Animal Science, 85, 1865–1872. https://doi.

org/10.2527/jas.2006-665

M-Rocha, J. F., Gallego, J. L., Vásquez, R. F., Pedraza, J. A., Echeverri, J., Cerón-

Muñoz, M. F., y Martínez, R. (2012). Estimación de parámetros genéticos

para edad al primer parto e intervalo entre partos en poblaciones bovinas

de la raza Blanco Orejinegro (BON) en Colombia. Revista Colombiana

de Ciencias Pecuarias, 25, 220–228. https://doi.org/10.17533/udea.rccp

Martínez, R., Vásquez, R., y Gallego, J. L. (2012). Eciencia productiva de la

Corporación Colombiana de Investigación Agropecuaria, Corpoica. 215

Martins, G. A., Martins Filho, R., Lima, F. de A. M., y Lôbo, R. N. B. (2000).

Inuência de fatores genéticos e de meio sobre o crescimento de bovinos

da raça Nelore no Estado do Maranhão. Revista Brasileira de Zootecnia,

29(1), 103–107. https://doi.org/10.1590/S1516-35982000000100014

Mota, L. F. M., Lopes, F. B., Fernandes Júnior, G. A., Rosa, G. J. M., Magalhães,

A. F. B., Carvalheiro, R., y Albuquerque, L. G. (2020). Genome-wide

scan highlights the role of candidate genes on phenotypic plasticity for

age at rst calving in Nellore heifers. Scientic Reports, 10(1), 6481.

https://doi.org/10.1038/s41598-020-63516-4

R Core Team. (2020). R: A Language and Environment for Statistical Computing

(4.0.0). R Foundation for Statistical Computing. https://www.r-project.

org

Ramírez, E., Cerón-Muñoz, M., Herrera, A. C., Vergara, O. D., Arboleda, E. M.,

y Restrepo, L. F. (2009). Crecimiento de hembras cruzadas en el trópico

colombiano. Revista Colombiana de Ciencias Pecuarias, 22, 642–647.

https://doi.org/10.17533/udea.rccp

Rincón, J., y Quintero, J. (2015). Comparación de modelos no lineales para

describir el crecimiento en ganado Blanco Orejinegro (BON). Revista

CES Medicina Veterinaria y Zootecnia, 10(1), 31–37. https://doi.

org/10.21615/cesmvz

This scientic publication in digital format is a continuation of the Printed Review: Legal Deposit pp 196802ZU42, ISSN 0378-7818.

Londoño-Gil et al. Rev. Fac. Agron. (LUZ). 2022, 39(2): e2239327-7 |

Rogberg-Muñoz, A., Melucci, L., Prando, A., Villegas-Castagnasso, E. E., Ripoli,

M. V., Peral-García, P., Baldo, A., Añon, M. C., & Giovambattista, G.

(2011). Association of bovine chromosome 5 markers with birth and

weaning weight in Hereford cattle raised under extensive conditions.

Livestock Science, 135(2–3), 124–130. https://doi.org/10.1016/j.

livsci.2010.06.160

Rogberg-Muñoz, A., Cantet, R. J. C., Elena, M., Pedro, J., Prando, A., Nélida,

A., Verónica, M., y Giovambattista, G. (2013). Longitudinal analysis of

the effects of IGF1 - SnaBI genotypes on the growth curve of Angus bull

calves. Livestock Science, 154(1–3), 55–59. https://doi.org/10.1016/j.

livsci.2013.03.016

Saleem, A. H. (2015). Role of Leptin in Growth, Reproduction and Milk Production

in Farm Animals: A Review. Advances in Animal and Veterinary Sciences,

3(5), 302–307. https://doi.org/10.14737/journal.aavs/2015/3.5.302.307

Verhulst, P. (1838). Notice sur la loi que la population pursuit dans son

accroissement. Correspondance Mathématique et Physique, 10,113-

121. http://www.neo-classical-physics.info/uploads/3/4/3/6/34363841/

verhulst_-_law_of_population.pdf

Von Bertalanffy, L. (1938). A quantitative theory of organic growth (inquires

on growth laws. Human Biology, 10(2), 181–213. https://doi.

org/10.2307/41447359