N General, Heritability Estimates for Beef Cattle Compared to Swine Are: Slightly Lower.
Abstract
The aim of this written report was to estimate genetic parameters for superovulatory response traits in lodge to explore the possibility of genetic improvement in Japanese Black cows. Nosotros analyzed xix 155 records of the total number of embryos and oocytes (TNE) and the number of good embryos (NGE) collected from 1532 donor cows between 2008 and 2018. A two-trait repeatability animal model analysis was performed for both. Considering records of TNE and NGE did non follow a normal distribution, the records were analyzed post-obit no, logarithmic, or Anscombe transformation. Without transformation, the heritability estimates were 0.26 for TNE and 0.17 for NGE. With logarithmic transformation, they were 0.22 for TNE and 0.18 for NGE. With Anscombe transformation, they were 0.26 for TNE and 0.18 for NGE. All analyses gave similar genetic correlations between TNE and NGE, ranging from 0.60 to 0.71. Spearman's rank correlation coefficient between breeding values of cows with more 10 records was ≥0.95 with both transformations. Thus, the genetic improvement of TNE and NGE of donor cows could be possible in Japanese Black cattle.
Introduction
Japanese Blackness cattle are a Wagyu brood famous for their excellent meat quality (Gotoh et al., 2014). For Japanese Black cattle in Japan, embryo transfer is widely used in the production of commercial animals as well as convenance stocks. The value of Japanese Black calves is much higher than the other beef breeds, and therefore, information technology is assisting for dairy farmers to transfer embryos from Japanese Black cows to dairy cattle recipients. In Japan, in 2014, Japanese Blackness embryos were transferred into near 100 000 Holstein cows, and 42 000 calves were born, or 8% of the total number of Japanese Black calves built-in in 2014. Thus, embryo transfer plays an important office in efficient Wagyu production (Agriculture and Livestock Industry Promotion System, 2019). Reproductive techniques such equally multiple ovulation, embryo transfer, and ovum pickup are widely used in dairy cattle production (Jaton et al., 2016); around the globe, 470 000 bovine embryos were produced in vivo in 2018 (International Embryo Transfer Society [IETS], 2018). The number of embryos and oocytes obtained from donor dairy cows per flash is used as an indicator of the response to superovulation handling (e.m., Jaton et al., 2016; Parker Gaddis et al., 2017). Big differences in numbers produced in vivo among cows have been reported (e.g., Kafi and McGowan, 1997; Kanitz et al., 2002; Mapletoft et al., 2002). The heritabilities of in vivo embryo production traits accept been estimated for Holstein (Jaton et al., 2016; Parker Gaddis et al., 2017), Belgian Blue (Michaux et al., 2002), and Nellore (Zebu) cattle breeds (Peixoto et al., 2004), merely non so far in Japanese Black cattle. This report estimated genetic parameters of superovulatory response traits in Japanese Black cows to assess the possibility of genetic comeback for embryo production.
Materials and Methods
Animal Care and Use Committee approval was not needed because information was obtained from preexisting databases.
Phenotypic measurements
The total number of embryos and oocytes recovered (TNE) and the number of adept embryos (NGE) per flush were recorded from twenty 257 superovulation treatments of 1532 Japanese Black donor cows between 2008 and 2018 at the Zennoh Embryo Transfer middle, Hokkaido, Japan. TNE was divers as the sum of the number of embryos and unfertilized oocytes nerveless in a unmarried flush; NGE was the number of embryos morphologically classified equally grade i according to the International Embryo Transfer Society criteria (Robertson and Nelson, 1998). We analyzed 19 155 records of those with TNE ≥ i.
After their showtime calving, all cows received superovulation handling every ≥seventy d (81.4 ± 27.two d in average). Every bit a basic program, commencement, a total of xx AU of FSH (Antrin R-10, Kyoritsu Seiyaku Corp., Tokyo, Japan) was administered intramuscularly in the neck twice a solar day for 3 d. At the fifth treatment, PGF2α (cloprostenol 0.225 mg/cow, Darmajin, Kyoritsu Seiyaku Corp., Tokyo, Japan) was administered. The day after estrus was observed, cows were inseminated artificially. Ane week later on, embryos were collected by washing of the uterine horns. Because embryo production performance from the same cow declines with the number of collections performed (Donaldson and Perry, 1983), the dosage of FSH was adjusted.
Equally the distributions of TNE and NGE differed from normal distribution (Figure 1), logarithmic and Anscombe (ans) transformations were used (Jaton et al., 2016; Parker Gaddis et al., 2017) as follows:
Effigy 1.
Effigy 1.
Statistical analysis
The following two-trait repeatability brute model was used to judge genetic parameters:
where yi is the vector of phenotypic records (i = 1 for TNE, i = 2 for NGE); , , , and east i are the vectors of fixed effects (year of superovulation, calendar month of superovulation, type of superovulation program, technician, and linear and quadratic covariates of age in months at superovulation), breeding values of cows, permanent environmental furnishings of cows, and errors; , , and are the blueprint matrices relating to , , and , respectively; is the additive genetic variance of trait i, is the additive genetic covariance betwixt traits, is the permanent environmental variance of trait i; is the permanent environmental covariance betwixt traits; is the error variance for trait i; is the fault covariance between traits; A is the condiment genetic relationship matrix constructed from full-blooded data of 3,521 individuals; and I is the identity matrix. Equally preliminary cheque, we confirmed the significance of all the fixed effects in the model (P < 0.0001) by least squares assay. The issue of service sire was not included because previous studies reported the impact of service sire was estimated to be negligible (König et al., 2007; Jaton et al., 2016). We did not include the result of the stage of lactation considering our donors basically do not calve again later first calving and the effect of the number of flushes because it could be highly confounded with the age in this written report.
Variance components were estimated in AIREMLF90 software (Misztal et al., 2002) with the default convergence criterion (10−12). The SEs of the estimated heritability, repeatability, and genetic correlation were calculated by using the se_cover_function option (Houle and Meyer, 2015).
Reliabilities of estimated breeding values (EBVs) for moo-cow i ( ) were calculated every bit follows:
where is the inbreeding coefficient of cow i, and is the prediction error variance of EBV of cow i.
Results and Give-and-take
Descriptive statistics of phenotypic records
Effigy 1 shows the histogram illustrating the distributions of records of TNE and NGE. Effigy two shows the number of flushes per donor. Bones statistics of superovulatory response traits are listed in Table i. The mean TNE in this written report (16.fifty) was college than those reported in previous studies of Holstein (6.67, Asada and Terawaki, 2002; nine.21, Jaton et al., 2016; 9.27, Cornelissen et al., 2017), Belgian Bluish (6.68, Michaux et al., 2002), and Nellore (10.27, Peixoto et al., 2004). The mean NGE (6.73) was college than that of Holstein (5.11, Gaddis et al., 2017). Because beef breeds could be more responsive to superovulation treatments than dairy breeds, beefiness cattle might accept the ability to produce more than embryos per affluent (Mikkola et al., 2020). Steinhauser et al. (2018) plant that Wagyu breeds responded better to superovulation treatments than other Bos taurus and Bos indicus breeds, but they did non describe the treatment conditions. Yokoo et al. (2016) reported that Japanese Black cows responded better than Japanese Shorthorn cows. Therefore, the Japanese Black cattle could have the potential for high response to superovulation treatments, equally our results propose.
Table one.
Trait | No. of records | No. of cows | Hateful | SD | Minimum | Maximum |
---|---|---|---|---|---|---|
TNE | 19 155 | 1 532 | 16.l | 10.66 | 1 | 84 |
NGE | 6.73 | 6.12 | 0 | 56 |
Trait | No. of records | No. of cows | Mean | SD | Minimum | Maximum |
---|---|---|---|---|---|---|
TNE | 19 155 | 1 532 | 16.fifty | x.66 | 1 | 84 |
NGE | 6.73 | 6.12 | 0 | 56 |
Tabular array 1.
Trait | No. of records | No. of cows | Mean | SD | Minimum | Maximum |
---|---|---|---|---|---|---|
TNE | xix 155 | one 532 | 16.l | 10.66 | 1 | 84 |
NGE | half dozen.73 | 6.12 | 0 | 56 |
Trait | No. of records | No. of cows | Mean | SD | Minimum | Maximum |
---|---|---|---|---|---|---|
TNE | 19 155 | 1 532 | 16.fifty | 10.66 | 1 | 84 |
NGE | 6.73 | vi.12 | 0 | 56 |
Figure 2.
Figure 2.
Figure 3 shows the effects of month of superovulation obtained from the analysis using the untransformed data. The difference in TNE between the maximum and minimum values of the estimated furnishings was approximately ii embryos. NGE tended to exist lower in winter. Heat stress could bear on the estrus bike and sign in Japanese Black and Holstein cows (Sakatani et al., 2012), and common cold stress could too impact fertility and reproductive operation in Japanese Black cows (Nabenishi and Yamazaki, 2017; Kino et al., 2019). Because the data were collected in subarctic Hokkaido, cold stress might be dominant in this Japanese Blackness population.
Figure three.
Figure 3.
Figure 4 shows the event of age at superovulation obtained from the analysis using the untransformed data. The effects would be very pocket-sized (Mikkola et al., 2020), but the consequence of the number of flushes previously experienced is still unclear. Our results bespeak that the response decreased with increasing historic period of the moo-cow, maybe because responsiveness to FSH would subtract by undergoing the superovulation over and once again. In our data, mean age (mo) of superovulation was 66. Peixoto et al. (2004) using 1,036 superovulation records of 475 Nellore females, whose ages ranged from two.2 to 20.5 year onetime at the fourth dimension of superovulatory treatment, and iii%, 25%, 30%, 23%, and 19% of them were <3, three–5.9, 6–viii.9, 9–eleven.9, and >12 yr sometime, respectively. Therefore, the mean historic period of superovulation in Peixoto et al. (2004) could be older than our data. Higher means of superovulatory response traits in our report might exist due to younger historic period of superovulation than previous studies.
Figure four.
Figure 4.
Estimated genetic parameters
Genetic parameter estimates of TNE and NGE and their standard errors (SE) are listed in Tabular array 2. Without transformation, the estimated heritability was 0.26 for TNE and 0.17 for NGE. The heritability of NGE was also lower in Holstein (Parker Gaddis et al., 2017). The heritability of TNE estimated here was similar to previous estimates in Holstein (Jaton et al., 2016; Parker Gaddis et al., 2017) and Belgian Bluish (Michaux et al., 2002). The estimated repeatability was 0.37 for TNE and 0.26 for NGE, both like to previous estimates in Holstein (Jaton et al., 2016; Parker Gaddis et al., 2017).
Table 2.
Transformation | Trait | rep2 | rg | ||||
---|---|---|---|---|---|---|---|
Untransformed | TNE | 29.18 ± 5.02 | eleven.74 ± 3.53 | 69.65 ± 0.74 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.71 ± 0.08 |
NGE | half-dozen.11 ± 1.27 | 3.49 ± 0.92 | 26.85 ± 0.29 | 0.17 ± 0.03 | 0.26 ± 0.01 | ||
Logarithm | TNE | 0.16 ± 0.03 | 0.07 ± 0.02 | 0.49 ± 0.01 | 0.22 ± 0.04 | 0.32 ± 0.01 | 0.60 ± 0.09 |
NGE | 0.fifteen ± 0.03 | 0.06 ± 0.02 | 0.63 ± 0.01 | 0.18 ± 0.03 | 0.25 ± 0.01 | ||
Anscombe | TNE | 1.81 ± 0.31 | 0.73 ± 0.22 | 4.39 ± 0.05 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.65 ± 0.08 |
NGE | 0.98 ± 0.xix | 0.43 ± 0.thirteen | three.90 ± 0.04 | 0.18 ± 0.03 | 0.27 ± 0.01 |
Transformation | Trait | rep2 | rthou | ||||
---|---|---|---|---|---|---|---|
Untransformed | TNE | 29.18 ± v.02 | xi.74 ± three.53 | 69.65 ± 0.74 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.71 ± 0.08 |
NGE | 6.11 ± i.27 | three.49 ± 0.92 | 26.85 ± 0.29 | 0.17 ± 0.03 | 0.26 ± 0.01 | ||
Logarithm | TNE | 0.xvi ± 0.03 | 0.07 ± 0.02 | 0.49 ± 0.01 | 0.22 ± 0.04 | 0.32 ± 0.01 | 0.lx ± 0.09 |
NGE | 0.15 ± 0.03 | 0.06 ± 0.02 | 0.63 ± 0.01 | 0.18 ± 0.03 | 0.25 ± 0.01 | ||
Anscombe | TNE | 1.81 ± 0.31 | 0.73 ± 0.22 | iv.39 ± 0.05 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.65 ± 0.08 |
NGE | 0.98 ± 0.19 | 0.43 ± 0.xiii | iii.90 ± 0.04 | 0.18 ± 0.03 | 0.27 ± 0.01 |
, additive genetic variance , permanent ecology variance; , error variance; , heritability; reptwo, repeatability; r g, genetic correlation.
Table 2.
Transformation | Trait | repii | rg | ||||
---|---|---|---|---|---|---|---|
Untransformed | TNE | 29.eighteen ± 5.02 | 11.74 ± 3.53 | 69.65 ± 0.74 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.71 ± 0.08 |
NGE | 6.11 ± ane.27 | 3.49 ± 0.92 | 26.85 ± 0.29 | 0.17 ± 0.03 | 0.26 ± 0.01 | ||
Logarithm | TNE | 0.16 ± 0.03 | 0.07 ± 0.02 | 0.49 ± 0.01 | 0.22 ± 0.04 | 0.32 ± 0.01 | 0.60 ± 0.09 |
NGE | 0.fifteen ± 0.03 | 0.06 ± 0.02 | 0.63 ± 0.01 | 0.18 ± 0.03 | 0.25 ± 0.01 | ||
Anscombe | TNE | 1.81 ± 0.31 | 0.73 ± 0.22 | 4.39 ± 0.05 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.65 ± 0.08 |
NGE | 0.98 ± 0.19 | 0.43 ± 0.xiii | 3.90 ± 0.04 | 0.18 ± 0.03 | 0.27 ± 0.01 |
Transformation | Trait | reptwo | rone thousand | ||||
---|---|---|---|---|---|---|---|
Untransformed | TNE | 29.18 ± 5.02 | eleven.74 ± iii.53 | 69.65 ± 0.74 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.71 ± 0.08 |
NGE | six.xi ± ane.27 | 3.49 ± 0.92 | 26.85 ± 0.29 | 0.17 ± 0.03 | 0.26 ± 0.01 | ||
Logarithm | TNE | 0.16 ± 0.03 | 0.07 ± 0.02 | 0.49 ± 0.01 | 0.22 ± 0.04 | 0.32 ± 0.01 | 0.60 ± 0.09 |
NGE | 0.15 ± 0.03 | 0.06 ± 0.02 | 0.63 ± 0.01 | 0.xviii ± 0.03 | 0.25 ± 0.01 | ||
Anscombe | TNE | 1.81 ± 0.31 | 0.73 ± 0.22 | 4.39 ± 0.05 | 0.26 ± 0.04 | 0.37 ± 0.01 | 0.65 ± 0.08 |
NGE | 0.98 ± 0.19 | 0.43 ± 0.13 | 3.ninety ± 0.04 | 0.18 ± 0.03 | 0.27 ± 0.01 |
, additive genetic variance , permanent environmental variance; , error variance; , heritability; rep2, repeatability; r 1000, genetic correlation.
The estimated genetic correlation between TNE and NGE was 0.71 without transformation. Parker Gaddis et al. (2017) estimated a correlation of almost 1 in Holstein. High positive genetic correlations (0.74–0.97) betwixt TNE and the number of transferable embryos accept been estimated in Holstein (König et al., 2007; Jaton et al., 2016) and Belgian Blue (Michaux et al., 2002). Our estimated genetic correlation was slightly lower than in previous studies. Cattle breeds might touch on the genetic correlation between TNE and NGE.
Consequence of transformation of phenotypic records
With logarithmic transformation, heritability estimates were 0.22 for TNE and 0.18 for NGE, and repeatability estimates were 0.32 and 0.25 (Table 2). With Anscombe transformation, heritability estimates were 0.26 for TNE and 0.18 for NGE and repeatability estimates were 0.37 and 0.27. Estimates of genetic correlation between TNE and NGE were 0.60 with logarithmic transformation and 0.65 with Anscombe transformation. Thus, the estimates of heritability, repeatability, and genetic correlation differed little between untransformed and transformed data.
Reliabilities of estimated breeding values
Effigy 5 shows the relationship between the number of flushes and the mean reliability of estimated convenance values of cows past two-trait repeatability animal model assay using untransformed data. The mean reliability initially increased speedily with the number of records, simply and so increased more than slowly across ten flushes. Spearman'due south correlation coefficients amid the EBVs of cows (n = 805) with ≥10 records are listed in Tabular array iii. Values of both TNE and NGE were always ≥0.95. The values of the rank correlation coefficients were almost the same (≥0.95) even when all cows with records were included. This indicates that the deviation in the genetic ability of cows selected by using untransformed and transformed records was small.
Table 3.
Untransformed | Logarithm | Anscombe | |
---|---|---|---|
Untransformed | – | 0.97 | 0.99 |
Logarithm | 0.95 | – | 0.99 |
Anscombe | 0.98 | 0.99 | – |
Untransformed | Logarithm | Anscombe | |
---|---|---|---|
Untransformed | – | 0.97 | 0.99 |
Logarithm | 0.95 | – | 0.99 |
Anscombe | 0.98 | 0.99 | – |
Above the diagonal: the total number of embryos and oocytes; below the diagonal: the number of practiced embryos.
Table 3.
Untransformed | Logarithm | Anscombe | |
---|---|---|---|
Untransformed | – | 0.97 | 0.99 |
Logarithm | 0.95 | – | 0.99 |
Anscombe | 0.98 | 0.99 | – |
Untransformed | Logarithm | Anscombe | |
---|---|---|---|
Untransformed | – | 0.97 | 0.99 |
Logarithm | 0.95 | – | 0.99 |
Anscombe | 0.98 | 0.99 | – |
Above the diagonal: the total number of embryos and oocytes; below the diagonal: the number of good embryos.
Effigy 5.
Figure five.
General Discussion
In this written report, genetic parameters of TNE and NGE were estimated as traits relating to response to superovulation treatments, the start such report in Japanese Blackness cattle. The heritabilities of both were moderate and the genetic correlation between them was positive and high. Both had higher heritabilities than other female person reproductive traits such as calving interval, non-return rate, and conception charge per unit (Oyama et al., 2002; VanRaden et al., 2004; Inoue et al., 2020; Ogawa and Satoh, 2021), and the accuracy of selection for TNE and NGE could be college than those for other representative female reproductive traits. Jaton et al. (2016) and Parker Gaddis et al. (2017) reported that genetic improvement by option for superovulatory response in Holstein cows was possible. Our results betoken that it is possible also in Japanese Black.
The data analyzed here differed from those in previous studies in terms of the greater number of records. The average number of repeated records per cow in previous studies was ane to three (Michaux et al., 2002; Peixoto et al., 2004; Jaton et al., 2016; Cornelissen et al., 2017; Parker Gaddis et al., 2017), whereas that hither was 12.5. Furthermore, the total number of records analyzed here was larger than in previous studies of beef cattle (Michaux et al., 2002) and dual-purpose breeds (Peixoto et al., 2004). The reliabilities of EBV of both traits were effectually 0.half dozen when the number of repeated records was 10, but information technology increased only slowly beyond this (Figure 5). In our population, superovulation records can be collected well-nigh once every 3 mo. Equally the mean age at first calving is virtually 24 mo in Japanese Black cows, nosotros tin obtain 10 repeated records of TNE and NGE at 5 yr of age. Furthermore, the genetic correlation between them was high and positive. There were little differences in the estimated genetic parameters with and without transformation and the values of rank correlation coefficients of EBVs were very high. We concluded that the two-trait animal model assay with untransformed data would be preferable for predicting breeding values for TNE and NGE because of the ease of treatment data.
The superovulatory response decreased with age (Figure 4). The proportion of the number of high-quality embryos also tended to decrease with age (results not shown). It would be possible that some cows have good response for many times of flushes, whereas others rapidly become unresponsive to superovulation. Female reproductive traits in cattle, such as calving interval (Panetto et al., 2012; Ogawa and Satoh, 2021), could be analyzed using a random regression model, in terms of gene-environment (age) interaction. Such kinds of analysis would exist needed in the hereafter. On the other hand, improving representative carcass traits, including caste of marbling, is economically most of import in Japanese Blackness cattle (Sasaki et al., 2006; Oyama., 2011). Hirayama et al. (2019) investigated superovulatory responses among lines of Japanese Black cattle and reported that strains with better superovulatory responses had phenotypically superior growth. It is of import to select cows with meliorate TNE and NGE without expense of other economically important traits such as body weight, carcass weight, and meat quality. Hence, research to investigate the genetic relationships between carcass and embryo production traits in Japanese Blackness cattle will be needed.
Conclusion
We estimated genetic parameters for TNE and NGE in Japanese Black cattle. Estimated heritabilities were moderate and estimated genetic correlation was high. The estimates of heritability and genetic correlation differed petty between untransformed and transformed data. The values of rank correlation coefficients of EBVs between with and without transformation were very loftier. These results suggest that genetic improvement of both past selection is possible in Japanese Black cows.
Abbreviations
-
EBV
-
IETS
International Embryo Transfer Gild
-
NGE
-
TNE
full number of embryos and oocytes
Abbreviations
Conflict of involvement statement
The authors declare no real or perceived conflicts of interest.
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