Multivariate random regression analysis for body weight and main morphological traits in genetically improved farmed tilapia (Oreochromis niloticus)
- Jie He^{1, 2},
- Yunfeng Zhao^{2},
- Jingli Zhao^{1, 2},
- Jin Gao^{1, 2},
- Dandan Han^{3},
- Pao Xu^{1} and
- Runqing Yang^{2}Email author
Received: 16 September 2016
Accepted: 24 October 2017
Published: 2 November 2017
Abstract
Background
Because of their high economic importance, growth traits in fish are under continuous improvement. For growth traits that are recorded at multiple time-points in life, the use of univariate and multivariate animal models is limited because of the variable and irregular timing of these measures. Thus, the univariate random regression model (RRM) was introduced for the genetic analysis of dynamic growth traits in fish breeding.
Methods
We used a multivariate random regression model (MRRM) to analyze genetic changes in growth traits recorded at multiple time-point of genetically-improved farmed tilapia. Legendre polynomials of different orders were applied to characterize the influences of fixed and random effects on growth trajectories. The final MRRM was determined by optimizing the univariate RRM for the analyzed traits separately via penalizing adaptively the likelihood statistical criterion, which is superior to both the Akaike information criterion and the Bayesian information criterion.
Conclusions
In the selected MRRM, the additive genetic effects were modeled by Legendre polynomials of three orders for body weight (BWE) and body length (BL) and of two orders for body depth (BD). By using the covariance functions of the MRRM, estimated heritabilities were between 0.086 and 0.628 for BWE, 0.155 and 0.556 for BL, and 0.056 and 0.607 for BD. Only heritabilities for BD measured from 60 to 140 days of age were consistently higher than those estimated by the univariate RRM. All genetic correlations between growth time-points exceeded 0.5 for either single or pairwise time-points. Moreover, correlations between early and late growth time-points were lower. Thus, for phenotypes that are measured repeatedly in aquaculture, an MRRM can enhance the efficiency of the comprehensive selection for BWE and the main morphological traits.
Background
From an economical point of view, growth and developmental characters are the most important traits in farmed fish species, and persistent efforts are made to genetically improve these traits in fish breeding. Growth and developmental traits, such as body weight (BWE) and morphological traits, are measured at different times. Faster BWE growth shortens time to market and selection on morphological traits allows fish shape and size to be standardized. Growth and developmental traits are dynamic quantitative traits because they vary spatially and temporally [1]. They are also called infinite-dimensional traits, since they are expressed on a continuous time and space scale [2]. In tilapia breeding, growth and developmental traits at a specific age (in day, week, or year), such as the time to market are the main targets of genetic improvement. As such, genetic analysis of the growth and developmental process can increase the efficiency of selection compared to that of traits measured at specific ages [3].
Considerable attention has been paid to the genetic analysis of BWE and morphological traits at a specific age in breeding tilapia. In earlier studies, growth traits at specific ages were considered as separate traits and analyzed by using univariate animal models [4–13]. Subsequently, multivariate animal models were applied to estimate both the heritabilities of growth traits at specific ages and the genetic correlations between traits measured at different specific ages [8, 11, 12, 14–22]. Although multivariate genetic analysis of growth traits has improved the accuracy of parameter estimation by using more records, at multiple ages, such an analysis is strongly limited by the variable and irregular timing of these measurements [23].
For growth and developmental traits that are recorded at multiple time-points during growth, the patterns of growth curves have been shown to be heritable [2, 3, 24]. The genetic analysis of dynamic quantitative traits was initially conducted by first fitting individual growth curves and then analyzing their estimated parameters within the framework of a multivariate animal model. However, with such genetic analyses, it was not possible to determine whether some individuals had too few records to fit the growth curves. Since then, random regression models (RRM) [3, 25] were developed to dissect dynamic phenotypes into different functions that describe the effects of various genetic and environmental factors. Only three papers on the application of RRM for the genetic analysis of growth curves have been published, in rainbow trout [26] and tilapia [9, 27]. The use of RRM allows heritabilities of growth traits at any age to be estimated, as well as genetic correlations between pairwise traits measured at different ages. Moreover, multiple trait RRM (MRRM) can estimate genetic correlations between pairs of traits measured at different ages and improve heritability estimates for each trait. This not only facilitates the genetic analysis of dynamic traits, but also improves the prediction of breeding values [3, 28].
The objective of this study was to construct and implement a MRRM to simultaneously model the genetic changes in growth traits and estimate the heritability of each growth trait, as well as the genetic correlations between pairs of growth traits measured at specific ages, such as the time to market for a population of genetically improved farmed tilapia in order to formulate a selection criterion for simultaneously selecting BWE and morphological traits. For this purpose, BWE, body length (BL), and body depth (BD) were measured six times on 1451 fish from 45 mixed families of full and half-sibs.
Methods
Experimental population
The base population of 1800 one-month old tilapia fingerlings (sex ratio of 1:1) was imported from WorldFish, Malaysia, in 2006, and was derived from 60 families with complete pedigree. The fish were systematically selected for seven generations at the experimental station of the Freshwater Fisheries Research Center in Wuxi, China. Based on breeding values for BWE at 120 days of age estimated using an animal model, 40 broodstocks per family were chosen at each generation, with a sex ratio of 1:1. The selection intensity in males and females was ~ 5%. Broodstocks from different families were randomly mated with each other and 100 to 125 families were retained in each generation. There was one generation of selection per year. In the seventh generation, in 2014, 120 males and 120 females from different families (one male and one female from each family) were selected as experimental parents. In May of that year, each male and two females ready to spawn were maintained in a 1-m^{3} fiberglass tank for one week. Next, 120 females with fertilized eggs in their mouths were separately placed into multiple hapas (1 m × 1 m × 1 m) in a large concrete pond (50 m × 7 m × 1 m) for a one-week incubation period and were then isolated from their progenies. At 50 days of age, 50 families, each consisting of no less than 1000 surviving progenies above 30 g, were used to construct the experimental population. At the same time, 32 progenies that were randomly chosen from each family were tagged with passive integrated transponder tags, and 1600 tagged fish were mixed in a larger concrete pond (50 m × 20 m × 1.8 m). Because of this, we did not include any effect of the rearing facilities on the measurements in the statistical model. In the subsequent experiment, all fish were fed on a standard commercial diet (crude protein: 28%, crude lipids: 4%, crude fiber: 15%, ash: 18%, total phosphorus: 1%, lysine: 1.2%) manufactured by the Feedstuff Incorporated company (Ningbo, China). Dissolved oxygen was maintained at 8.23 to 8.67 mg/L by air pumps and the water temperature naturally fluctuated between 24.3 and 26.7 °C during the entire experiment.
The tagged fish were weighed on electronic scales to measure BWE, while BL and BD were measured with calipers. Before each measurement, the fish were anesthetized with clove oil at a density of 100 mg/L. The fish were measured once every 15 days during the experiment, with the first recording at the time of tagging. A maximum of six records for each fish and each trait were available, since 4.6% of the individuals died before the end of the experiment. Although experimental fish were measured only six times, the traits were recorded on 20 different days of age because of different spawning times. A total of 7560 records were extracted for BWE, BL and BD.
Averages and standard deviations (in parentheses) for body weight (g), body length (mm) and body depth (mm) at 20 days of age
Age | Sample size | BWE | BL | BD |
---|---|---|---|---|
52 | 473 | 36.3 (8.5) | 88.19 (9.68) | 31.74 (3.42) |
59 | 354 | 49.94 (3.67) | 99.28 (3.92) | 36.62 (2.00) |
65 | 304 | 62.47 (12.47) | 107.13 (11.46) | 41.30 (3.90) |
68 | 469 | 82.99 (11.00) | 116.86 (9.79) | 43.71 (2.87) |
72 | 320 | 95.98 (23.40) | 125.40 (12.26) | 48.25 (3.52) |
75 | 352 | 106.19 (10.56) | 129.58 (5.23) | 49.32 (2.68) |
81 | 302 | 124.03 (31.99) | 133.49 (15.32) | 53.66 (4.54) |
85 | 470 | 138.50 (27.12) | 139.96 (11.65) | 56.56 (3.74) |
88 | 319 | 153.00 (40.52) | 141.67 (15.28) | 60.09 (3.87) |
92 | 352 | 173.97 (19.17) | 149.15 (6.48) | 62.43 (2.26) |
98 | 303 | 200.39 (51.45) | 159.64 (19.21) | 63.70 (4.61) |
102 | 469 | 221.12 (44.36) | 161.13 (13.83) | 64.04 (3.73) |
105 | 319 | 243.64 (63.82) | 164.32 (17.93) | 66.17 (4.46) |
109 | 354 | 275.08 (33.08) | 172.25 (8.63) | 67.19 (3.04) |
115 | 304 | 313.77 (77.68) | 180.04 (23.21) | 69.58 (5.59) |
121 | 473 | 384.56 (65.23) | 185.63 (15.63) | 70.32 (4.46) |
122 | 320 | 391.92 (89.49) | 187.85 (19.95) | 71.87 (5.06) |
128 | 354 | 427.41 (105.57) | 190.47 (26.18) | 72.09 (6.45) |
134 | 304 | 480.17 (50.97) | 194.43 (10.70) | 73.15 (3.76) |
141 | 320 | 538.13 (87.26) | 197.54 (17.46) | 74.97 (5.21) |
Pedigree data were collected by tracing back three generations, thus including 1604 individuals from 77 sires and 88 dams. Although maternal effects on growth traits may be large in the early growth period for tilapia, they were not considered in the analyses, because observations from only a single generation and from such a small number of dams without records render the maternal effects unidentifiable.
Random regression model
To estimate the genetic correlations between growth traits measured at multiple time-points, the RRM for those traits must be solved simultaneously by a multivariate genetic analysis. The covariance matrices in the MRRM were estimated by using restricted maximum likelihood (REML), as implemented in DMU Version 6 [29]. The starting values were set to 0 for each fixed effect, to the identity matrix for each random effect covariance matrix and to 1 for the residual variance. The convergence criterion for REML was set to 10^{−6}.
After estimating the covariance matrices, heritabilities for the traits at any age and the genetic correlations between traits measured at different ages were estimated by the covariance functions [30]. The covariances for a trait between the \(i\)th and \(j\)th days of age can be calculated as \({\mathbf{z}}_{i} {\mathbf{Fz}}_{j}^{\text{T}}\) for the family effect, \({\mathbf{w}}_{i} {\mathbf{Aw}}_{j}^{\text{T}}\) for the additive genetic effect, and \({\mathbf{s}}_{i} {\mathbf{PEs}}_{j}^{\text{T}}\) for the permanent environmental effect. Therefore, genetic correlations between the \(i\)th and \(j\)th days of age for a trait were estimated by \(\frac{{{\mathbf{w}}_{i} {\mathbf{Aw}}_{j}^{\text{T}} }}{{\sqrt {({\mathbf{w}}_{i} {\mathbf{Aw}}_{i}^{\text{T}} )({\mathbf{w}}_{j} {\mathbf{Aw}}_{j}^{\text{T}} )} }}\), phenotypic correlations by \(\frac{{{\mathbf{P}}_{ij} }}{{\sqrt {{\mathbf{P}}_{ii} {\mathbf{P}}_{jj} } }}\) with phenotypic covariance \({\mathbf{P}}_{ij} = {\mathbf{z}}_{i} {\mathbf{Fz}}_{j}^{\text{T}} + {\mathbf{w}}_{i} {\mathbf{Aw}}_{j}^{\text{T}} + {\mathbf{s}}_{i} {\mathbf{PEs}}_{j}^{\text{T}} + \sigma_{e}^{2} {\mathbf{I}}(i = j)\), and heritability at the \(i\)th day of age by \(\frac{{{\mathbf{w}}_{i} {\mathbf{Aw}}_{i}^{\text{T}} }}{{{\mathbf{P}}_{ii} }}\).
Choice of model
Results
Choice of MRRM
The best MRRM for BWE, BL, and BD was selected by separately optimizing the univariate RRM for growth traits. The Legendre polynomial for the effect of sex was generally modeled by the population’s growth curve, chosen as the Legendre polynomial of three orders according to the highest goodness-of-fit for the three analyzed traits. The three random effects, i.e. family, additive genetic, and permanent environmental effects, was characterized by using Legendre polynomials of different orders that ranged from 0 to 3 [3]. The models were designated as \({\text{LPm}}_{1} {\text{m}}_{2} {\text{m}}_{3}\), e.g., \({\text{LP}}121\) is a model with a Legendre polynomial of order 1 for the family effect, order 2 for the additive genetic effect, and order 1 for the permanent environmental effect. A total of 64 RRM for each analyzed trait were compared based on the PAL criterion to determine the best combination of orders of the Legendre polynomials for the three random effects.
Choice of RRM for BWE, BL and BD based on the PAL criterion
Model | Parameters | BWE | BL | BD | |||
---|---|---|---|---|---|---|---|
− 2log(ML) | PAL | − 2log(ML) | PAL | − 2log(ML) | PAL | ||
LP00 | 3 | 59,206.2 | 36,641.6 | 28,386.1 | |||
LP11 | 11 | 47,172.4 | 47,172.4 | 30,293.0 | 30,293.0 | 23,486.4 | 23,486.4 |
LP12 | 16 | 45,006.1 | 45,035.6 | 29,667.0 | 29,700.4 | 22,765.9 | 22,797.9 |
LP13 | 22 | 44,259.8 | 44,309.2 | 29,583.0 | 29,650.3 | 22,726.0 | 22,798.5 |
LP21 | 16 | 46,821.6 | 46,884.8 | 30,162.3 | 30,260.6 | 23,263.7 | 23,334.8 |
LP22 | 22 | 44,991.3 | 45,032.6 | 29,657.8 | 29,705.3 | 22,745.5 | 22,792.1 |
LP23 | 29 | 44,228.7 | 44,294.0 | 29,580.0 | 29,670.8 | 22,706.3 | 22,812.4 |
LP31 | 22 | 46,752.8 | 46,877.5 | 30,149.3 | 30,311.3 | 23,256.0 | 24,142.4 |
LP32 | 29 | 44,827.3 | 44,882.0 | 29,642.1 | 29,704.9 | 22,736.9 | 22,798.5 |
LP33 | 37 | 44,215.4 | 44,301.9 | 29,574.4 | 29,695.9 | 22,705.9 | 22,850.5 |
Genetic parameters for growth traits
Covariance estimates
Heritability estimates
Estimates of ratios of family to phenotypic variances (f ^{2}) and of heritabilities (h ^{2}) for BWE, BL and BD at selected days of age based on the univariate and multivariate RRM
Age | Univariate | Multivariate | ||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
BWE | BL | BD | BWE | BL | BD | |||||||
f ^{2} | h ^{2} | f ^{2} | h ^{2} | f ^{2} | h ^{2} | f ^{2} | h ^{2} | f ^{2} | h ^{2} | f ^{2} | h ^{2} | |
60 | 0.259 | 0.108 | 0.732 | 0.163 | 0.698 | 0.035 | 0.242 | 0.080 | 0.728 | 0.173 | 0.653 | 0.098 |
65 | 0.334 | 0.169 | 0.654 | 0.256 | 0.628 | 0.141 | 0.356 | 0.187 | 0.656 | 0.256 | 0.611 | 0.177 |
70 | 0.392 | 0.308 | 0.588 | 0.335 | 0.566 | 0.249 | 0.428 | 0.299 | 0.592 | 0.331 | 0.558 | 0.270 |
75 | 0.421 | 0.401 | 0.538 | 0.396 | 0.527 | 0.331 | 0.456 | 0.378 | 0.542 | 0.391 | 0.514 | 0.348 |
80 | 0.428 | 0.462 | 0.500 | 0.442 | 0.505 | 0.386 | 0.458 | 0.438 | 0.504 | 0.438 | 0.482 | 0.405 |
85 | 0.422 | 0.507 | 0.472 | 0.478 | 0.492 | 0.422 | 0.447 | 0.485 | 0.476 | 0.474 | 0.460 | 0.446 |
90 | 0.412 | 0.541 | 0.451 | 0.505 | 0.483 | 0.445 | 0.433 | 0.521 | 0.454 | 0.502 | 0.445 | 0.474 |
95 | 0.403 | 0.565 | 0.435 | 0.527 | 0.476 | 0.461 | 0.421 | 0.547 | 0.438 | 0.523 | 0.436 | 0.494 |
100 | 0.395 | 0.581 | 0.422 | 0.544 | 0.469 | 0.472 | 0.412 | 0.565 | 0.427 | 0.539 | 0.430 | 0.508 |
105 | 0.392 | 0.590 | 0.413 | 0.556 | 0.463 | 0.480 | 0.406 | 0.576 | 0.420 | 0.550 | 0.426 | 0.518 |
110 | 0.391 | 0.595 | 0.408 | 0.565 | 0.456 | 0.488 | 0.403 | 0.583 | 0.416 | 0.557 | 0.422 | 0.527 |
115 | 0.392 | 0.597 | 0.406 | 0.570 | 0.449 | 0.496 | 0.402 | 0.587 | 0.415 | 0.561 | 0.418 | 0.535 |
120 | 0.393 | 0.598 | 0.406 | 0.571 | 0.442 | 0.506 | 0.401 | 0.590 | 0.416 | 0.562 | 0.412 | 0.545 |
125 | 0.393 | 0.600 | 0.409 | 0.570 | 0.433 | 0.520 | 0.398 | 0.595 | 0.418 | 0.561 | 0.404 | 0.556 |
130 | 0.389 | 0.605 | 0.414 | 0.566 | 0.421 | 0.538 | 0.391 | 0.603 | 0.422 | 0.560 | 0.392 | 0.570 |
135 | 0.380 | 0.615 | 0.420 | 0.563 | 0.404 | 0.560 | 0.379 | 0.616 | 0.424 | 0.559 | 0.377 | 0.588 |
140 | 0.365 | 0.632 | 0.423 | 0.560 | 0.379 | 0.588 | 0.362 | 0.635 | 0.423 | 0.562 | 0.358 | 0.610 |
Correlation estimates
Estimates of phenotypic (lower triangle) and family (upper triangle) correlations for body weight between pairwise measurements at selected days of age
Age | 60 | 65 | 70 | 75 | 80 | 85 | 90 | 95 | 100 | 105 | 110 | 115 | 120 | 125 | 130 | 135 | 140 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
60 | 1.00 | 0.98 | 0.97 | 0.93 | 0.89 | 0.86 | 0.83 | 0.80 | 0.77 | 0.75 | 0.72 | 0.69 | 0.67 | 0.64 | 0.62 | 0.59 | 0.57 |
65 | 1.00 | 1.00 | 0.99 | 0.98 | 0.96 | 0.94 | 0.91 | 0.89 | 0.87 | 0.85 | 0.82 | 0.80 | 0.78 | 0.76 | 0.74 | 0.72 | 0.70 |
70 | 0.98 | 0.99 | 1.00 | 1.00 | 0.99 | 0.97 | 0.96 | 0.94 | 0.92 | 0.90 | 0.88 | 0.87 | 0.85 | 0.83 | 0.81 | 0.79 | 0.77 |
75 | 0.95 | 0.97 | 0.99 | 1.00 | 1.00 | 0.99 | 0.98 | 0.97 | 0.95 | 0.94 | 0.92 | 0.91 | 0.89 | 0.88 | 0.86 | 0.84 | 0.83 |
80 | 0.92 | 0.94 | 0.97 | 0.99 | 1.00 | 1.00 | 0.99 | 0.98 | 0.97 | 0.96 | 0.95 | 0.94 | 0.92 | 0.91 | 0.90 | 0.88 | 0.87 |
85 | 0.88 | 0.90 | 0.95 | 0.98 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.98 | 0.97 | 0.96 | 0.95 | 0.93 | 0.92 | 0.91 | 0.90 |
90 | 0.84 | 0.87 | 0.92 | 0.96 | 0.99 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.98 | 0.97 | 0.96 | 0.95 | 0.94 | 0.93 | 0.92 |
95 | 0.81 | 0.84 | 0.90 | 0.94 | 0.97 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 | 0.98 | 0.98 | 0.97 | 0.96 | 0.95 | 0.94 |
100 | 0.79 | 0.82 | 0.87 | 0.92 | 0.96 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.98 | 0.97 | 0.97 | 0.96 |
105 | 0.76 | 0.79 | 0.85 | 0.90 | 0.94 | 0.96 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.98 | 0.98 | 0.97 |
110 | 0.74 | 0.77 | 0.83 | 0.88 | 0.92 | 0.95 | 0.97 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.98 | 0.98 |
115 | 0.72 | 0.75 | 0.80 | 0.85 | 0.89 | 0.92 | 0.94 | 0.96 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 | 0.99 | 0.99 |
120 | 0.70 | 0.72 | 0.77 | 0.82 | 0.86 | 0.89 | 0.91 | 0.94 | 0.95 | 0.97 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 | 1.00 | 0.99 |
125 | 0.67 | 0.69 | 0.74 | 0.78 | 0.82 | 0.85 | 0.87 | 0.90 | 0.92 | 0.94 | 0.96 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 | 1.00 |
130 | 0.64 | 0.66 | 0.70 | 0.74 | 0.77 | 0.80 | 0.82 | 0.85 | 0.87 | 0.90 | 0.93 | 0.96 | 0.98 | 0.99 | 1.00 | 1.00 | 1.00 |
135 | 0.61 | 0.62 | 0.65 | 0.68 | 0.70 | 0.73 | 0.76 | 0.78 | 0.81 | 0.85 | 0.88 | 0.91 | 0.95 | 0.97 | 0.99 | 1.00 | 1.00 |
140 | 0.57 | 0.57 | 0.59 | 0.61 | 0.63 | 0.65 | 0.68 | 0.71 | 0.74 | 0.78 | 0.82 | 0.86 | 0.90 | 0.94 | 0.97 | 0.99 | 1.00 |
Discussion
In this study, we used an MRRM to estimate heritabilities of BWE, BL, and BD based on repeated measurements during growth period, as well as genetic correlations between pairwise growth traits at specific days of age. Legendre polynomials were chosen to characterize the influence of fixed and random effects, i.e. family, additive genetic, and permanent environmental effects, on growth curves. The best MRRM was established by separately optimizing univariate RRM for each growth trait, according to the PAL. Based on the final MRRM, we found that heritabilities increased with age for all traits and that they were slightly lower than those obtained by using univariate RRM. In addition, for each trait, genetic correlations between measurements decreased monotonously with increased lag in days of age. For measurements at the same days of age, estimates of heritabilities and genetic correlations were close to those previously reported for tilapia [4–6, 8, 10–16, 18, 20–22, 33, 34].
For the selected RRM, changes in the genetic effects with day of age were optimally modeled for all analyzed traits by using Legendre polynomials of three orders. In contrast, Rutten et al. [9] and Turra et al. [27] used polynomials of two orders to fit the fixed and random effects, without selecting the orders of the polynomials for the different fixed and random effects. The non-significant variance for the permanent environmental effects showed that they have no impact on growth traits. Ratios of family variance [9] and of permanent environmental variance [27] to phenotypic variance are largely underestimated in a population with multiple families, which may be caused by the collinearity between the family and permanent environmental effects in the RRM used.
Repeated measurements during growth are required to estimate changes in both the fixed and the random effects with age when applying the RRM to genetic analysis of growth traits. More longitudinal measurements per individual would help to model growth curves with a higher goodness-of-fit and to more robustly estimate genetic parameters for random regression. However, more longitudinal measurements not only increase the experimental costs, but also affect fish growth, especially when all experimental individuals are measured together. In our experiment, the simultaneous measurement of 1600 tilapia fish incurred high labor costs and a measuring frequency of once every 15 days influenced fish growth to some extent. In future trials, the experimental population could be divided into several separately reared subpopulations, and each subpopulation could be observed in turn over more growth time-points. The extension of such measurements to more families from multiple generations would also help to identify maternal effects on early growth in tilapia within the MRRM framework.
Conclusions
Using repeated records of growth duration from multiple families in one generation, first we introduced MRRM to genetically analyze growth curves of body weight and main morphological traits in genetically improved farmed tilapia. The optimal MRRM was chosen by comparing the univariate RRM for the analyzed traits separately via PAL. By using the covariance functions of the MRRM, changes in heritabilities with days of age and genetic correlations between lags in days of age were estimated, which could be used to carry out early selection for each trait analyzed. More importantly, for phenotypes that are measured repeatedly in aquaculture, an MRRM can enhance the efficiency of the comprehensive selection for BWE and the main morphological traits at a specific age.
Declarations
Authors’ contributions
RY proposed the method and supervised JH, JZ and JG for writing the manuscript. JH, YZ, JZ and JG wrote computer codes and conducted real data analysis. DH and PX took part in the interpretation of results. JH and PX constructed the breeding population and observed traits. All authors read and approved the final manuscript.
Acknowledgements
This study was funded by Special Scientific Research Funds for Central Non-profit Institutes, Chinese Academy of Fishery Sciences (2014B004 and 2014A04XK01) and the National Natural Science Foundations of China (31201989).
Competing interests
The authors declare that they have no competing interests.
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Authors’ Affiliations
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