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Poult Sci 2007. 86:2461-2465. doi:10.3382/ps.2007-00086
© 2007 Poultry Science Association
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PRODUCTION, MODELING, AND EDUCATION

Hyperbolastic Models as a New Powerful Tool to Describe Broiler Growth Kinetics

H. Ahmadi and M. Mottaghitalab1

Department of Animal Science, Faculty of Agricultural Science, University of Guilan, PO Box 41635-1314, Rasht, Iran

1 Corresponding author: m_mottaghi{at}gstp.ir


    ABSTRACT
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
The mathematical models for describing growth kinetics are important tools to examine biological parameters, such as BW at specific time, maximum growth response, and growth rates. Classical growth models such as Gompertz and Richards have been extensively used in broiler studies. To obtain a more intuitive understanding of growth, a class of flexible growth models containing 3 and 4 parameters were developed as hyperbolastic models. These models provide description of growth behaviors for continuous output in different fields, for instance cancer and stem cell growth. This study was conducted to find out if 3 flexible new hyperbolastic growth models, called type 1, 2, and 3 (H1, H2, and H3), may be used to illustrate relationship between live weight and age in broilers and also how effective such models would be compared with 2 classical growth models, namely Gompertz and Richards. A set of growth data over 70 d, obtained from 18 male broilers, was used to fit growth models. Goodness of fit of the models was determined using mean square error, R2, and residual standard deviation. It was revealed that the 3 new models may be used to fit broiler growth data successfully and could be implanted in SAS PROC NLIN. Goodness of fit criteria refers better fit with H3, presumably due to its greater flexibility, followed by the Richards, Gompertz, H2, and H1. In conclusion, it seems that the H3 can be considered as a more useful tool for modeling broiler growth. However, it is reasonable to compare models for fit before selection of the more accurate one.

Key Words: broiler • growth model • Gompertz • Richards • hyperbolastic


    INTRODUCTION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
In animal production models, growth curves deal with genetic potential evaluation and estimation of daily nutrient requirements for growth. These estimates could be used for calculation of total feed requirements, which sets an upper limit to feed intake when animals are given ad libitum access to high-quality feeds. An appropriate growth model conveniently simulates information observed on an animal in terms of a small set of parameters that can be interpreted biologically and used to derive other relevant growth traits.

A number of nonlinear functions have been used to describe growth in fish, poultry, and mammals (France et al., 1996; Lopez et al., 2000). Despite numerous reported growth models, derivatives of the commonly used functions failed to describe the corresponding mean growth rate curve (Taylor, 1980), emphasizing the need to examine obtained growth models with similar situations that may have appeared in field observations.

The use of growth models is usually empirical, and the form of the model is chosen by its ability to fit the data. A growth model can be characterized by some underlying physiological or biochemical mechanism or constraint (France and Thornley, 1984). Such models can be defined as rate as a function of state, in which the instantaneous growth rate is a function of the organism’s size. Unlike equations in which growth rate is purely an empirical function, an equation in this form can usually be interpreted biologically and can be ascribed by its parameters (Lopez et al., 2000).

The Gompertz and Richards models have been the most well known models for describing growth in broilers (Aggrey, 2002; Darmani Kuhi et al., 2003). In these models, the growth curves are asymmetric around the point of maximum growth rate. However, differences between them are points of inflection, which in the Gompertz model is a fixed and in the Richards model is a variable (flexible) proportion of their asymptotic growth values (Tabatabai et al., 2005).

Tabatabai et al. (2005) introduced a class of 3 and 4 parameter models, namely hyperbolastic models, to predict self-limited growth behavior including growth in tumors as well as stem cells. These models are called hyperbolastic because the outcome is a function of inverse hyperbolic sine function (arcsinh). These models are a family of flexible growth models that can predict variety of growth behaviors for continuous outputs in many fields of biological research. They introduced these hyperbolastic growth models in 3 types, including hyperbolastic growth model type 1 or H1 (generalizes logistic growth model); hyperbolastic growth model type 2 or H2 (stand alone); and hyperbolastic growth model type 3 or H3 (generalizes Weibull growth model).

The aim of present study is evaluation of 3 new flexible hyperbolastic growth models, the H1, H2, and H3, and their ability to describe relationship between live weight and age in broiler chickens. In the meantime, model outputs will be compared with the Gompertz and Richards growth models.


    MATERIALS AND METHODS
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Data Source
The BW data used in present study was first reported by Roush et al. (2006). Growth data are average values of 18 male broilers (Ross x Ross 308) BW for a 70-d period. Details of growth experiment with broiler chickens are described in the paper.

Growth Models
Five growth models were fitted to data using NLIN procedure (Marquart algorithm) of the SAS (SAS Institute Inc., 1999). The models are as below:


Formula


Formula


Formula


Formula


Formula


Formula

Gompertz model (Gompertz, 1825),


Formula


Formula

and Richards model (Richards, 1959),


Formula


Formula

In all models, W(t) is referred to live weight (g) at age t, β is the intrinsic growth rate, {theta} and {gamma} are parameters, and M represents the asymptotic or maximum growth response (final weight), which is assumed to be constant, though final weight may usually change over time.

In each model {alpha} is defined as a function of the other parameters (M, β, and initial observed value W0 at time t0), which allows reduction of the number of parameters to be estimated and also anchors the first predicted value to the original value observed at the initial time point.

A quantitative verifying of the fit of the predictive models was made using error measurement indices commonly used to evaluate forecasting models. The accuracy of models (goodness of fit) was determined by mean squared error (MSE) and R2 value as well as residual standard deviation (RSD). Forecasting error measurements were based on the value differences between model predicted and observed BW (Oberstone, 1990).


    RESULTS
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Behavior of 5 growth models and related residuals are shown in Figure 1Go. As Figure 1Go indicates, all growth models could be fitted to the data without difficulty by nonlinear regression, although they were sensitive to choice of initial values; however, they provided an excellent fit.


Figure 1
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Figure 1. Observed and growth models predicted BW vs. age (predicted values using 5 growth models, namely hyperbolastic 1, 2, 3, Gompertz, and Richards).

 
Estimated parameters and their standard error obtained with H1, H2, H3, Gompertz, and Richards growth models are summarized in Table 1Go. As the table shows, the lowest estimate of M or final weight is obtained with H1 (5,220 g for male broiler chicken). It is followed by the H3 (5,367 g), H2 (5,491 g), Richards (5,934 g), and Gompertz (6,038 g). Based on M, remarkable differences were observed among prediction of introduced models.


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Table 1. Estimated growth parameters (with standard error in parentheses) using different growth models
 
Table 2Go shows observed and predicted values of broiler BW using 5 models, as well as values of model error measurements and determined residuals. The results show that the H3 (MSE = 364, R2 = 0.99995, RSD = 19.2) and Richards (MSE = 603.3, R2 = 0.999919, RSD = 24.7) models provide the best fit to broiler growth data. It is followed by Gompertz (MSE = 627.7, R2 = 0.999916, RSD = 25.2), H2 (MSE = 1,294, R2 = 0.99983, RSD = 34.9), and H1 (MSE = 2,533, R2 = 0.99966, RSD = 48.8). All growth models are powerful to describe the broiler growth pattern (see Figure 1Go).


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Table 2. Observed and predicted broiler BW using 6 models with model error measurements and determined residuals1
 

    DISCUSSION
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
The overall calculated statistical values (MSE, R2, and RSD) have shown that the H3 and Richards model provide higher accuracy fit to data set, followed by the Gompertz, H2, and H1. Similar results were observed with other flexible and generalized models used to evaluate growth data fit (Brown et al., 1976; Gille and Salomon, 1995; France et al., 1996; Lopez et al., 2000).

Although models are closely related, the parameter values may have appeared quite different when these models were fitted to a single set of data. The Gompertz model has 2 parameters (M and β) and can fit asymmetric growth, though it is not very flexible when compared with the H3 and Richards model. The Richards model used in this study was formed with 3 parameters (M, β, and {gamma}). It is an asymmetric model and also more flexible than the Gompertz model. The H1 has one more parameter ({theta}) than the Gompertz functions, which allows more flexibility and can fit asymmetric growth patterns with enough ability to detect increasing and decreasing growth kinetics. The H2 has the same number of parameters as H1 and can fit asymmetric curves, but it cannot fit decreasing growth patterns, so it seems to be less flexible. The H3 model has the same flexibility as the H1 function at the expense of one more parameter ({gamma}) similar to the Richards equations. More flexibility of H3 and Richards may lead to more accurate prediction and better fit to the broiler growth data set than other models (Darmani Kuhi et al., 2003). This means that the fixed point of inflection (less flexibility) in the growth models may act as a limitation for data fitting. Consequently, consideration of flexible growth models (such as the H3 and Richards) as an alternative to the Gompertz is worthwhile due to the following reasons: 1) they are easy to fit and fitting them to growth data leads to biologically meaningful parameters; 2) they have more flexibility, i.e., very often give a closer fit to data points and therefore a smaller MSE and RSD and higher R2 values than the simpler models; and 3) they encompass simpler models for additional parameters. This is especially important when the behavior of a particular data set is not defined previously (Darmani Kuhi et al., 2003).

Despite unstable value, the asymptote can be regarded as the final weight. In this study, to predict final weight (M) some bias was observed among models (see Table 1Go). It has been established in different species that estimation of final weight is a function of algorithm fitting, and its accuracy judging is possible when a precise final weight is available (Ricklefs, 1985; Lopez et al., 2000).

The new growth models, and H3 in particular, clearly demonstrate with valid data fitting for broiler and better goodness of fit than others introduced in the present study. Based on calculated R2 for the same data set (Roush et al., 2006), the results revealed that the H3 model (R2 = 0.99995) has more accurate prediction than artificial neural network of broiler growth model (R2 = 0.999938), though in H3 model higher MSE value was observed (364 vs. 163.1).

In the present study, the H3 model showed lower residuals distribution (in terms of RSD) than that of Gompertz and Richards. This is in agreement with Tabatabai et al. (2005) when classical and hyperbolastic models were compared, to evaluate cancer, craniofacial, and stem cell growth data fitting. Their suggestion was that choosing a flexible and highly accurate predictive model such as hyperbolastic can significantly improve the outcome of a study, and the accuracy of a model determines its utility.

In conclusion, hyperbolastic growth models can be used to fit a broiler growth data set, and the results indicate higher prediction accuracy for the H3. However, it is reasonable to compare growth models before making a decision to select the more accurate one.

Received for publication February 20, 2007. Accepted for publication May 9, 2007.


    REFERENCES
 TOP
 ABSTRACT
 INTRODUCTION
 MATERIALS AND METHODS
 RESULTS
 DISCUSSION
 REFERENCES
 
Aggrey, S. E. 2002. Comparison of three nonlinear and spline regression models for describing chicken growth curves. Poult. Sci. 81:1782–1788.[Abstract/Free Full Text]

Brown, J. E., H. A. Fitzhugh, Jr., and T. C. Cartwright. 1976. A comparison of nonlinear models for describing weight-age relationships in cattle. J. Anim. Sci. 42:810–818.[Abstract/Free Full Text]

Darmani Kuhi, H., E. Kebreab, S. Lopez, and J. France. 2003. An evaluation of different growth functions for describing the profile of live weight with time (Age) in meat and egg strains of chicken. Poult. Sci. 82:1536–1543.[Abstract/Free Full Text]

France, J., J. Dijkstra, and M. S. Dhanoa. 1996. Growth functions and their application in animal sciences. Ann. Zootech. 45:165–174.[CrossRef]

France, J., and J. H. M. Thornley. 1984. Mathematical Models in Agriculture. Butterworths, London, UK.

Gille, U., and F. V. Salomon. 1995. Bone growth in ducks through mathematical models with special reference to the Janoscheck growth curve. Growth Dev. Aging 59:207–214.[ISI][Medline]

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. Phil. Trans. Royal Soc. 182:513–585.

Lopez, S., J. France, M. S. Dhanoa, F. Mould, and J. Dijkstra. 2000. A generalized Michaelis-Menten equation for the analysis of growth. J. Anim. Sci. 78:1816–1828.[Abstract/Free Full Text]

Oberstone, J. 1990. Management Science—Concepts, Insights, and Applications. West Publ. Co., New York, NY.

Richards, F. J. 1959. A flexible growth function for empirical use. J. Exp. Bot. 10:290–300.[Abstract/Free Full Text]

Ricklefs, R. E. 1985. Modification of growth and development of muscles in poultry. Poult. Sci. 64:1563–1576.[ISI][Medline]

Roush, W. B., W. A. Dozier III, and S. L. Branton. 2006. Comparision of Gompertz and neural networks models of broiler growth. Poult. Sci. 85:794–797.[Abstract/Free Full Text]

SAS Institute Inc. 1999. SAS/STAT User’s Guide. Version 8. SAS Institute Inc., Cary, NC.

Tabatabai, M., D. K. Williams, and Z. Bursac. 2005. Hyperbolastic growth models: Theory and application. Theor. Biol. Med. Model. 2:14.[CrossRef][Medline]

Taylor, St C. S. 1980. Genetically standardized growth equations. Anim. Prod. 30:167–175.[ISI]





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