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Find out moreof models P. Huhtanen, A. Seppälä, S. Ahvenjärvi and M. Rinne J Anim Sci 2008.86:2657-2669. doi: 10.2527/jas.2008-0894 originally published online Jun 6, 2008;

The online version of this article, along with updated information and services, is located on the World Wide Web at: http://jas.fass.org/cgi/content/full/86/10/2657

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suggesting that silage pdNDF is not a homogenous substrate. 1993. Huhtanen et al. Further. (1999) de- 1 Corresponding author: pjh87@cornell. The fit improved with increasing number of pools. the digestion rate (kd) is variable during fermentation]. Pitt et al. 2008. Furthermore. Generally. However. seven 2-pool. and without the mechanisms of selective retention. rived equations to estimate the effective first-order kd from non-first-order models assuming that rumen is a 1-compartment system with random passage of feed particles. the comparison of feedstuffs on the basis of parameter values is more unequivocal than in the case of multiple-pool models. Animal Production Research..6 between the nonescapable and escapable pools.† and M. fiber. and three 3-pool models were compared in fitting gas production data and predicting in vivo NDF digestibility and effective first-order digestion rate of potentially digestible NDF (pdNDF). Schofield et al.† S. Seppälä. The concentration of pdNDF was estimated by a 12-d in situ incubation. Cornell University. Cone et al.*1 A. All rights reserved. it is impossible to attain observed in vivo digestibility of potentially digestible NDF (pdNDF) with realistic passage kinetic parameter values (Ellis et al. J. gas production. Huhtanen. they proposed a method for estimating the effective first-order kd from 2657 Downloaded from jas. 1994. the models predicted in vivo NDF digestibility and digestion rate accurately. Most of the new models are non-first-order models [i. There were marked differences between the models in fitting the gas production data.. The effective first-order digestion rate was computed both from observed in vivo and model-predicted pdNDF digestibility assum- ing the passage kinetic model described above. 2008. 2008. 2000. 1996). 1994. Relatively simple models such as the France model or even a single exponential model with discrete lag period satisfied the minimum criteria for a good model. rumen digestion ©2008 American Society of Animal Science. the feed particles are retained selectively in the rumen (Allen and Mertens.. Ahvenjärvi. Rinne† *Department of Animal Science. Ithaca.2527/jas. The parameter values from gas production profiles and pdNDF were used in a 2-compartment rumen model to predict pdNDF digestibility using 50 h of rumen residence time distributed in a ratio of 0. (2008) demonstrated that using digestion kinetic parameters derived from gas production data fitted with a 2-pool Gompertz model and indigestible NDF (iNDF) concentration estimated by 12-d in situ incubation in a 2-compartment rumen model predicted the in vivo NDF digestibility of grass silages accurately and precisely.edu Received January 22.org by on September 21. Investigating the time-related residuals during the later phases of fermentation is important when the data are used to estimate the first-order digestion rate of pdNDF. Huhtanen et al. . Finland ABSTRACT: Eleven 1-pool. Isolated NDF from 15 grass silages harvested at different stages of maturity was incubated in triplicate in rumen fluid-buffer solution for 72 h to estimate the digestion kinetics from cumulative gas production profiles.e. Mathematical description of the data has been accomplished by fitting a variety of nonlinear models (France et al. 86:2657–2669 doi:10.. The models overestimating the asymptotic gas volumes tended to underestimate the in vivo digestibility.. 2006). Sci. Anim. a good fit of gas production data was not necessarily translated into improved predictions of the in vivo data. Accepted May 30.. A limitation of the non-first-order kinetic parameters is that they cannot be used in steady-state rumen models. FI-31600 Jokioinen. To overcome these problems. 2010.Prediction of in vivo neutral detergent fiber digestibility and digestion rate of potentially digestible neutral detergent fiber: Comparison of models P.4:0. and †MTT Agrifood Research Finland.fass. mathematical model. However. 1988). NY 14853. Key words: effective digestion rate.2008-0894 INTRODUCTION The in vitro gas production technique has widely been used during the last decade among ruminant nutritionists to study feed degradation. In vivo digestibility was estimated by the total fecal collection method in sheep fed at a maintenance level of feeding.

Gompertz (GOMP2S and GOMP2F). lacked robustness. discrete lag time. resulting in 288 recordings per sample. (2000).2658 Huhtanen et al. (1996) proposed an empirical multiphase model... Gompertz (GOMP) and Richards (RICH) models are extensively used to describe growth functions. and logistic (LOG3) models. Three-pool models were used for Gompertz (GOMP3S). (1993) equation is a generalization of the Mitscherlich model. DLag. and the European convention for the protection of vertebrate animals used for experimental and other scientific purposes. The France et al. 2010. Models and Curve-Fitting For each NDF residue. The generalized Michaelis-Menten (Gen M-M) model allows the rate of digestion to decrease continuously or to increase first and then decrease and includes a lag time.8 kPa to Downloaded from jas. Hollis. appendix A and B. g/kg of DM) was calculated as follows: [NDF intake (g/d) − fecal NDF (g/d)]/DMI (kg/d). which will decrease the relative contribution of gas from the inoculum and decrease the random errors associated with a small sample size (e.. and in vivo digestibility of the silages are described in more detail by Huhtanen et al. and three 3-pool models were fitted to the data by the NLIN procedure (SAS Inst. The objective of the present study was to evaluate the importance of choice of models in fitting gas production data to estimate the in vivo NDF digestibility and effective first-order digestion rate of grass silages. the parameter values of these models do not always permit unequivocal comparison of feedstuffs. the model-predicted pdNDF digestibility. in vivo digestibility of pdNDF was calculated as follows: [NDF intake (g/d) − NDF output (g/d)]/pdNDF intake (g/d). Groot (Groot3). Cary. (1994. GOMPF) were used. The improvement in goodness of fit was marginal for this model. 1976. For the Gen M-M. and lag pool. MATERIALS AND METHODS All sheep in the digestibility trials and ruminally fistulated cattle as rumen fluid donors for the in situ incubations were managed according to legislation documented in the Finnish Animal Welfare Act (247/96). and EXPLPool. The concentration of pdNDF (g/kg of DM) was calculated as NDF (g/kg of DM) − iNDF (g/kg of DM). and in many cases. The concepts of discrete lag and lag pool are explained in detail by Mertens (1993a). The logistic (LOG) model assumes that gas production is proportional both to the microbial mass and the digestible substrate (Schofield et al. Inc. . h) describes the time at which 50% of the asymptotic value of gas was produced and C is a dimensionless switching parameter that together with B determines the shape of the curve. implemented under the auspices of the local animal use and care committee. 1994). those related to attachment of feed particles on the walls of fermentation bottles). The models EXPDLag (EXPDLag2). and LPool denote absence of lag. or resulted in local optima or biologically meaningless parameter values. and Weibull (WEIB2) were also extended to accommodate 2 cell wall pools with fast and slow degradation rates. NH). The concentration of digestible NDF (dNDF. estimation of iNDF concentration.g. (2008). In Vitro Gas Production Measurements In vitro gas production measurements were made by an automated system described in detail by Huhtanen et al.. the rates were similar for both pools. in which B (units. EXPDLag.1 mL). Solenoid valves are adjusted to open when the differential pressure in the bottle reaches 2. Gas production from each sample was determined in 3 replicated runs. (2000. The compartmental interpretation of most of the functions is described in detail by France et al. The goodness of fit was markedly improved for the other 2-pool models except for the EXPDLag2 model. Cumulative gas production was recorded every 15 min for the 72-h incubation periods. The system is based on pressure transducers (142PC05D. Because iNDF by definition is indigestible. which allows accommodating either a sigmoidal or nonsigmoidal shape. The 1-pool models are presented in Table 1. The model predicted the in vivo digestion rate accurately and precisely. The models included 3 exponential equations EXP0. (2003a). The system allows using a relatively large sample size (500 mg). GOMPS) and France et al. Groot et al. MN) and solenoid valves (11–15–1-SV24Q70. logistic (LOG2). Thus. Honeywell Inc. isolation of NDF. The problem of the 2-compartment Gompertz model and other non-first-order models is that although they fit the data better than simpler models such as the first-order exponential model. Details of silages.fass. the order of using vertebrate animals for scientific purposes (1076/85). The contents of fermentation bottles are stirred intermittently with 15 s of stirring and 30-s pauses using a magnetic stirring plate. A probability function Weibull (Ross. Two different forms of the Gompertz equations as derived by Schofield et al.org by on September 21. Eleven different 1-pool. the system consists of 39 serum bottles with a volume of 120 mL. (2008) and chemical analyses by Nousiainen et al. Feeds In vitro gas production was measured from isolated neutral detergent residue of 15 primary growth timothy (Phleum pratense)-meadow fescue (Festuca pratensis) silages harvested at different stages of maturity over 3 yr. amount of iNDF ingested in feeds was also used for output. Groot (Groot2). 7 different 2-pool. release accumulated gas (approximately 1. the fit was not markedly improved by including the second pool or the model did not converge. In brief. WEIB) was also included in the analysis. Subscripts 0. Richards (RICH2). NC). Minneapolis. Pneutronics. a mean gas curve was calculated from the 3 replicates.

Digestibility of cell wall carbohydrates in the rumen is competition between the rates of digestion and passage.. 2006). Downloaded from jas. Pond et al. b. respectively. c > 0 t≥L t ≥0 1 Gen M-M = generalized Michaelis-Menten.6 of rumen residence time between the nonescapable and escapable pools was assumed (Huhtanen et al.5}/2. GOMP = Gompertz. RICH = Richards. With the models including discrete lag time..0625-h integration step and using the Euler integration method. 1988). The passage rate from the escapable pool was assumed to be a first-order process. The amount of dNDF predicted by the model was calculated as pdNDF (g/kg of DM) × model-predicted pdNDF digestibility. WEIB = Weibull. The rate constant (kr) was calculated as: kr = (t × λ2)/[1 + (t × λ)]. Specific gravity has also been shown to influence the probability of particles to escape from the rumen. the digestibility (D) with a kd and 1-compartment rumen model with first-order passage (kp) can be evaluated by a simple algebraic equation [D = kd/(kd + kp)].. Lund et al.. L = lag parameter. Model-predicted pdNDF digestibility was used to calculate the effective first-order digestion rate (effective kd) using the equation described by Huhtanen et al.fass. and n = adjustable parameters. B. kL = rate parameter from lag pool. Mathematical models to describe the cumulative gas production from in vitro fermentation Model1 EXP0 EXPDLag EXPLPool Groot France Gen M-M LOG GOMP GOMPFR WEIB RICH Equation2 Vt = V [1 − e ] Vt = V {1 − e−[k⋅ (t − L)]} Vt = V {1 − 1/[kd − kL] ⋅ [kd⋅e(−kL⋅t) − kL⋅e(−kd⋅t)]} Vt = V/[1 + (BC/tC)] Vt = V⋅ (1 − e −{k⋅[t − L] − [c⋅ (√ t − √L)]}) Vt = V (t − T)c/[(t − L)c + Kc] Vt = V/[1 + e [2 − 4⋅k⋅ (t − L)]] Vt = V1 × exp {−exp[1 − k1 × (t − L1)]} Vt = V(1 − exp{−b[ec(t − L) − 1]/c}) Vt = V⋅{1 − e−[k⋅(t − L)]}n Vt = V⋅{1 − e −[k⋅(t − L)]}1/n −(k⋅t) Domain t≥0 t≥L t≥0 t≥0 t≥L t≥L t≥0 t≥0 t ≥ L. Total NDF digestibility was calculated as model-predicted dNDF/NDF. . and EXPLPool models. The mean residence time in the rumen was assumed to be 50 h.4:0. a gamma time-related function (G2G1) was used to describe this process. V = asymptotic gas volume.0333 were used for kp and kr. but the release from the nonescapable to the escapable pool commenced at time t = 0. the digestion began at t = lag time. Except for the EXP0. Estimated gas production parameters from different models were used to compute the value of F at different time points. and total extent of digestion is defined as the total amount degraded divided by the total amount entering the system. (2008) to predict ruminal pdNDF digestibility for a single batch of feed. Simulations were run for 120 h with a 0. the goodness of fit was compared using the proportion of variance accounted for (R2) and residual mean squares (RMS) and ranking the models according to RMS. where λ = the rate constant (= 2/residence time in the compartment) and t = time.05 and 0.. Statistical Analysis To evaluate the performance of the models. If the function of cumulative gas production is described as F(t) and the derivative of the function F’(t) = dF/dt describes the mass disappearing at time = t.Gas production models 2659 Table 1. (2006): effective kd = {−[kp + kr] + [(kp + kr)2 + 4 × pdNDF digestibility × krkp/(1 − pdNDF digestibility)]0. Norway). r. For the simple exponential model. Values of 0. The amount of pdNDF digested in the rumen was estimated using a model incorporating the selective retention of feed particles in the rumen (Allen and Mertens. Lund et al. 2006) have demonstrated that the release of feed particles from the nonescapable pool to the escapable pool is a time-dependent process. 2000) and must be evaluated numerically. Rumen Digestion Models Digestion kinetic parameters derived from the gas production data were used in a dynamic mechanistic model described by Huhtanen et al. c.g. The total mass disappearing by digestion is the integral over time from t > lag to ∞. Single effective first-order digestion rate was calculated from in vivo pdNDF digestibility similarly as from model-predicted pdNDF digestibility described earlier. EXPDLag. C. 2006. 1988. therefore. Large forage particles entering the rumen need to be comminuted below a threshold size before they can escape from the rumen. The rumen models were constructed using Powersim software (Bergen. In the model ExpDLag. 2010. the retention in the lag pool was included in the rumen nonescapable pool. Marker kinetic studies (e. LOG = logistic. (1999) demonstrated that the function derived to estimate effective kd is independent of intake. The kd at each time point can be estimated as the mass digested as a proportion of the mass in the rumen. but for most models. which allows using this parameter in continuous rumen models. k and kd = rate parameters. 2 Vt = volume of gas at time t. the integrals are nonanalytical (France et al.. the kd at each time point (t) can be described as follows: kd = F’(t)/F(t).org by on September 21. kd is or can be time-dependent. Distribution of 0. Pitt et al.

Although the mean RMS was greater for the Groot model compared with Gen M-M.99730 0.8 3.26 3.47 9.24 3. The GOMPS model had a smaller RMS than LOG and EXP0.073 0. the silages showed a large variation in all parameters. as for all 15 gas production curves. Root mean squared errors (RMSE) between the observed in vivo and model-predicted dNDF and NDF digestibility values and in vivo derived and modelpredicted effective kd values were calculated as follows: RMSE = √∑ (observed − predicted)2/n.34 2.34 Median 19.99 1.997 (Table 3).30 0. The silages were well-preserved as indicated by low pH (4.6 38.613 0.085 0.72 1 Gen M-M = generalized Michaelis-Menten. The differences between the models were much greater when evaluated on the basis of RMS.2 7.94 1. concentrations of VFA (22 g/kg of DM) and ammonia N (47 g/kg of total N). whereas LOG was the worst according to this criterion. .99967 Mean 23.99979 0.fass. Goodness of fit of the 1-pool models Residual mean squares Model1 EXP0 EXPDLag EXPLPool France Gen M-M GOMPS GOMPF Groot LOG RICH WEIB R2 0.10) and low Table 3.85 17.6 10. The Gen M-M and Groot models showed the greatest average R2 values.597 0. Only the fit to models LOG (11/15) and EXP0 (4/15) resulted in the largest RMS.99982 0.84 6.53 3.2660 Huhtanen et al.83 10. and the 3 three-pool models.52 2. Table 2. and random variation around the regression line (Bibby and Toutenburg.5 7.13 1. 1977).3 137 93 50 0. it was greater than 0.11 34.99771 0.3 82 47. The average RMS across the 15 curves was smallest with the Gen M-M model followed by the Groot model. LOG = logistic. RESULTS Feeds The chemical and morphological composition and in vivo digestibility of the silages are shown in Table 2.725 0. the 6 two-pool models.99966 0. These evaluations were made separately for the 11 one-pool models.840 0.907 Indigestible NDF. Downloaded from jas.41 2.99960 0.99968 0.76 7. Mean square prediction error (MSPE) was divided to components resulting from mean bias.99966 0. g/kg of DM Leaves Stems Heads Digestibility OM NDF pdNDF2 1 2 Mean 927 161 586 81 36 346 596 57 0. The RMSE were 1.035 Minimum 909 112 402 17 20 192 375 0 0.0 1.1 to 4.28 0.3 3.99893 0.83 4.12 0. Due to differences in maturity.20 1. the smallest RMS was observed more often with the Groot model (8/15) than with Gen M-M (4/15).35 15.737 0.43 1.46 4. slope bias. The significance of the deviation of the intercept from 0 and the slope from 1 was analyzed by t-test.46 2. RICH = Richards.6 7. WEIB = Weibull.99969 0.8 2.83 11.66 25.13 25. but it was always greater than for the other models.org by on September 21. Fitting of the Gas Production Profiles The proportion of variance explained by the models was in general very high.98 3. Potentially digestible NDF.844 SD 8. GOMP = Gompertz. 2010.6 2.3% of the final gas volume. g/kg of DM OM CP NDF iNDF1 Lignin Morphological composition.8 3.4 6.21 Maximum 57. Models LOG and EXP0 clearly had the greatest RMS. Composition and digestibility of the silages Item Composition.15 1.83 1.791 Maximum 940 239 669 158 55 625 714 129 0.75 Minimum 6.869 0.

RICH = Richards.001).94 0. The residuals were usually greatest during the first 20 h of incubation. especially with multiple pool models. Downloaded from jas. Asymptotic Gas Volume Model-predicted asymptotic gas volumes together with observed endpoints are shown in Figure 1. The models EXP0. intercept close to 0.99996 0.99997 0. slope close to 1. ranging from 116.02 0. and WEIB2 performed well according to these criteria. a disadvantage of this and other models with a time-dependent fractional digestion rate.org by on September 21.17 0.80. One. is that parameter values are strongly correlated and do not permit unequivocal comparison of feedstuffs. RICH. However.30 0. The model described the cumulative gas production profiles very well as indicated by small mean square errors and confidence intervals of the parameter values.g.47 0. LOG2..38 0.99999 1. The differences between the models were highly significant (P < 0. and LOG further decreased RMS. whereas the LOG model slightly overpredicted it.1 mL (Gen M-M).99992 0. In most cases.11 Median 2. and Groot2 underpredicted in vivo NDF digestibility. WEIB = Weibull.22 0.005) and precisely R2 > 0. the RMSE was only 0. Most models predicted the in vivo-derived effective kd accurately (bias from −0.99999 Average 3.2 mL (LOG) to 130.22 0.49 0. GOMP2S.Gas production models 2661 Extending the models for a second pool assuming that the 2 pools describe the gas produced from rapidly and slowly digestible NDF decreased RMS markedly except for EXPDLag (Table 4).16 0. For the best models. In many cases. The R2 values were greater than 0. Table 4.70 0.32 to 0. The relationship between the effective kd calculated from model-predicted and in vivo-determined pdNDF digestibility is shown in Table 7.and 3-pool models Residual mean squares Model1 Two-pool ExpDLag2 GOMP2S GOMP2F Groot2 RICH2 LOG2 WEIB2 Three-pool GOMP3S Groot3 LOG3 1 R2 0.47 0. Goodness of fit of the 2. 2010.99994 0. with R2 values ranging from 0. Danfær et al. and RMS was below 1. but the models France. .99969 0.15 0.20 0. which preclude the use of kinetic parameters estimated using models includ- Prediction of NDF Digestibility The relationships between the predicted and observed NDF digestibility are shown in Table 6.9999 with all other 2-pool models.05 Maximum 7.61 0. Steady-state rumen models (e. The variation between the models in predicted gas volume was relatively large.00000 0. LOG = logistic.69 0.70% of the final gas volume.757 to 0. the variation resulting from mean bias or random variation across the regression line had the greatest contribution to the total MSPE.05 0.02 0. fitting the gas production profiles to the 2-pool EXPDLag model resulted in similar rates for the both pools. 2008) demonstrated that in vivo NDF digestibility could be predicted accurately and precisely by a model in which digestion kinetic parameters were estimated from gas production profiles by the 2-pool Gompertz equation and iNDF concentration by a 12-d in situ incubation. but there were differences also during the later phases of incubation. GOMP = Gompertz.95 0. GOMP2F.15 0.15 0. Including a third pool into the Groot.99995 0. Groot.70 0.00 in all cases.04 0.04 0. The R2 values were above 0.14 Gen M-M = generalized Michaelis-Menten. Our previous study (Huhtanen et al. DISCUSSION The main objective of the present study was to investigate the importance of curve-fitting to the gas production data.005 to 0.62 0.07 0. Including the third pool in the Groot model clearly improved the relationship between potential NDF digestibility and asymptotic gas volume.98 0. and high R2).940 (Table 5).96 for all other models except for the WEIB model.02 0.99999 0.and 2-pool Groot and Gen M-M models had the lowest R2 values between potential NDF digestibility and asymptotic gas volume.. Gen M-M. GOMPS. Some examples of time-related residuals are shown in Figure 2.24 0.12 0. Asymptotic extent of gas production was closely associated with potential NDF digestibility (pdNDF/NDF).39 0.32 0.. The parameter values of gas production kinetics were used in a mechanistic rumen model to estimate in vivo NDF digestibility and effective firstorder digestion rate of pdNDF. indicating that the precision of this model was less than that of the other models.26 0.fass.03 0. None of the models was the best according to all criteria (small bias.10 0. 2006) need first-order digestion rates.71 1.12 Minimum 1.

(1999) proposed equations to estimate effective first-order digestion rates for models including digestion lag and time-dependent digestion rate for 1-compartment rumen models. this often results in negative values at zero time. Pitt et al. Allen and Mertens (1988) proposed a model that describes digestive processes by a 2-step mechanism: the first step is a first-order process involving a change of substrate from unavailable to available form (lag pool). Gen M-M = generalized Michaelis-Menten. GOMP = Gompertz. similarly as for the in situ data (Ørskov and McDonald. 1993a). or attachment of microbes with the substrate. The discrete lag time can be attributed to hydration. Model-predicted asymptotic gas volumes for 15 gas production curves (SE = 1. (2008) for 2-compartment rumen models.fass. by France et al. This approach was extended by Huhtanen et al. (2000). Sometimes a parameter describing a rapidly or immediately degradable fraction has been included in models describing cumulative gas production. Observed value is the mean of the 5 data points during the last hour of incubation. As discussed Downloaded from jas. .org by on September 21. This model assumes that digestion begins instantaneously after the lag time.2662 Huhtanen et al. ing time-dependent digestion rates. 2010. LOG = logistic. RICH = Richards. Although the discrete lag time does not completely describe the lag phenomena. and the Gas Production Models Different versions of simple exponential equations have been applied to gas production studies. Figure 1. WEIB = Weibull. negative gas volumes at zero time are biologically meaningless. However. 1979). Including a discrete lag time in the simple firstorder equation improved the fit of models markedly.74 mL). and the usefulness of such parameter values in rumen models is questionable. it provides a simple quantitative measure of lag effects (Mertens. The effective first-order kd was computed from the modelpredicted pdNDF digestibility by solving the equation proposed by Allen and Mertens (1988). removal of digestion inhibitors.

003. and France. and exhaustion of the rapidly degradable substrate toward the end of fermentation). it did not fit the data better than the discrete lag model. 1.Gas production models 2663 Figure 2. Biologically. In the WEIB model. Groot et al.0) or first decreases rapidly and thereafter slowly (n < 1. the kd increases first and decreases later during fermentation (c >1.31. substrate accessibility. 1996. the fractional digestion rate can increase or decrease. the kd increased first and declined thereafter. Although the lag pool model is theoretically more correct than the discrete lag model. An increase in digestion rate with time may reflect an increase in substrate availability.. The average value for the c parameter across the 15 gas production curves was 1. the parameter n influences the pattern of changes in kd with time.0) with increased digestion time. 1997). . probably reflecting increased microbial number. and consequently. respectively. the model is exponential).023. In the RICH model. the kd either increases (n > −1. but the reductions in RMS were rather small. Time-related residuals after fitting of the 2-pool Gompertz (GOMP2S). (1993) proposed a generalized Mitscherlich model. decreased availability of growth factors) that could result in a decreased fractional digestion rate as the degradation proceeds. and France models to a single gas production curve. MichaelisMenten (Gen M-M). France et al.. and increased microbial numbers. 23. Compared with other gas production models.0 (WEIB). the first step can be interpreted to be related to the same processes as the discrete lag. The biases (observed − predicted) of NDF digestibility predictions were 0.79. The France model fit the data consistently better than the ExpDLag model.46 to 2. The parameter n increases the flexibility in curve-fitting with both WEIB Downloaded from jas. i. This equation has been extensively used by the Dutch workers (Cone et al.0) or decreases (n < −1. which can accommodate both sigmoid and nonsigmoid shapes. 2000). In the RICH and WEIB models. and 2. Gen M-M.fass. and microbial attachment at the beginning of incubation. but it was still much smaller than the RMS of the other 1-pool models. microbial attachment.30. Omitting the lag parameter from the model increased the RMS.0). 0. LOG1. second step represents the first-order digestion of the substrate.0 (RICH) or 1. the changes are more rapid at the early stages of incubation. the Gen M-M is more flexible (France et al. France et al.69 (range 1.16.995). The effects of parameters n and c on the gas production curve in the WEIB and France models are very similar. Mertens (1993a) postulated that a discrete lag time was a necessary addition to the model to adequately describe the digestion processes.046. When the parameter value is −1. suggesting that the France model was not able to describe the digestion kinetics of isolated NDF very accurately.g. build-up of fermentation products.org by on September 21... (1996) proposed a modified Gen M-M equation without a discrete lag.17 for the models GOMP2S. Further. −0. in all cases. and 0.e. (2000) discussed several factors (e. Residual mean squares for fitting the gas production curves were 0.e. the kd either first increases rapidly and thereafter slowly (n > 1. 2010..000 units.0) or decreases continuously (c ≤ 1.0). logistic (LOG1). and it decreased RMS compared with the previous models. the kd is constant (i. the RMS values of the 15 gas curves were strongly correlated (R2 = 0. In the Gen M-M model.

7 11. RICH = Richards. including the second pool markedly decreased the positive intercept.2 11.22 in WEIB2). (1994) have an inherent error in that predicted gas volumes are positive at time zero.15 in Groot2. Gompertz and logistic equations as given by Schofield et al.fass. 0.5 9.62 2.6 9.68 3.70 0.6 12.8 10. (2000) presented some biological problems despite the fact that the fit was markedly improved.7 RMSE2 3. the division between the rapidly and slowly degradable pools can be rather arbitrary.23 2.3 P-value 0.20 per hour toward the end of fermentation period. 0.9 12. GOMP = Gompertz. LOG = logistic.3 9. (1994) fit all 15 cumulative gas production profiles better than the model derived by France et al.02 3.6 9.9 10. the fractional digestion rate increases with time when the parameter value of c >0.0 14. the fractional digestion rates of the slow pool increased up to 0. Predicted asymptotic gas volume was smaller than observed gas volume at 70 to 72 h in many cases (9/15). The RMS with 1-pool models was positively (R2 = 0.53 0. in the present study using isolated NDF. WEIB = Weibull.4 17. because the initial gas volume should be zero. (1994). microbial attachment and microbial growth.3 10.73 0. and it took only 1.922 0.933 0. the value of parameter c was only marginally different from zero. which is not biologically possible.927 0.1 15.936 0. This was probably due to high values of the parameter c.002 0.96 2.85) correlated with the in vivo NDF digestibility.26 in GOMP2FR. However.39 0. The 2-pool models assume that the pdNDF is comprised of rapidly and slowly degradable fractions as described by Schofield et al.757 0. Large variation between the models in the proportions of the rapidly and slowly degradable pools despite very good fit to the data may indicate that there are not 2 easily distinguishable pdNDF pools described by the cumulative gas production profiles but rather several cell wall fractions differing in the rate of digestion.909 0.8 11.7 9. Table 5. 0.23 in RICH2. but the proportion of the rapidly degradable pool varied markedly between the models (0.0 9.004 0.88 2.913 0.70 R2 0.38 0.85 0.0 10.73 3.60 2.5 12.94 0. Depending on the model structure.74 3. The proportion of the rapidly degradable pool of the total gas produced increased with improved in vivo NDF digestibility.789 0.6 10.917 0. For example. and the model approached ExpDLag.66 0. or to chemical and structural changes in cell walls with the advancing fermentation time.65 0.82 2. (2000).16 0.9 10.10 0.934 Gen M-M = generalized Michaelis-Menten. which was interpreted as increased microbial activity per unit of feed.16 0. The effects of the n parameter may be attributed to the changes in hydration.871 0.69 2.890 0.0 9. Including the second pool to the Gompertz model derived by France et al. The 1-pool Gompertz model derived by France et al. The 2-pool models fit the data markedly better than the corresponding 1-pool models except the ExpDLag model. RMSE = root mean square error. (1994) model.8 13.41 0.5 10.840 0.916 0.37 0. suggesting that 1-pool models did not adequately describe the NDF digestion kinetics of early harvested silages.62 4.3 10. (2000) fit the data better than the model derived by Schofield et al.2 10.5 12. In the Gompertz model derived by France et al. and RICH models improving the fit compared with the EXPDLag model.4 10. However.09 2.88 2. especially for the rapidly degradable pool but also for the slowly degradable pool.21 0.4 10.5 h to reach zero residual. 2010.3 9.15 to 0.937 0.65 to 0. . Differences in kd have been reported between Downloaded from jas. The Gompertz model derived by Schofield et al.39 Slope 125 125 131 140 101 146 130 94 150 133 135 144 151 154 154 150 142 145 149 151 SE 12.4 10.929 0.909 0.2664 Huhtanen et al.org by on September 21.3 10.1 9.9 11. Relationships between the potential NDF digestibility (pdNDF/NDF) and model-predicted asymptotic gas production (mL) Model1 One-pool EXP0 EXPDLag EXPLPool France Gen M-M GOMPS GOMPF Groot LOG RICH WEIB Two-pool GOMP2S GOMP2FR Groot2 LOG2 RICH2 WEIB2 Three-pool GOMP3 Groot3 LOG3 1 2 Intercept 20 14 8 1 42 −9 9 47 −14 6 5 −3 −11 −6 −14 −8 −2 −4 −5 −10 SE 11.74 0.53 2.39 in GOMP2.50 2.25 0. (1994) and Schofield and Pell (1995).3 9.940 0.2 9. (2000) when 2-pool models were used despite the positive intercept at time zero with the Schofield et al.1 9.8 10. 0.67 2. (1994).55 in LOG2.892 0.933 0.49 2. and 0.74 2. in agreement with the study of Schofield et al.

152a 0.082 1.810 0.108 0.988 0.919b 0.928b 0.005 0.562 Slope 0.933 0.008 Mean bias 0.168 0.180 0.856 0.019 0.030 0.840 0.847 0.338 0.962 0.182 1.005 0. Downloaded from jas.005 0.720 0.013 0.011 0.019 0.004 0. mean bias (observed − predicted).854 0.986 0.834b 0.005 0.063 0.107 0.882 0.985 0.502 0.212 0.org by on September 21.002 0.013a 0.178 0.150 0.017 0.009 0.060 0.001 0.007 0. mean bias (observed − predicted).233 0.254 0.079 0.020 0.014 0.048 0.166 0.721 0.630 Slope 0.905 0.493 0.872 0.833 0.832 0.033 −0.036 −0. Table 7.fass.001 −0.006 0.002 0.112 0. 2010.006 0.002 0.988 0.020 0.143a 0.461 0.011 −0.966 0.004 0.846 0.005 0.005 Bias 0.928 0.264 0.012 0.682 0. root mean square error (RMSE).561 0. .05) different from 1.010 0.727 0.957 0.019 0.005 0.003 −0.012 0.993 1.989 0.002 0.030 0.001 0.092 0.027 1.009 0.009 0.491b 1.010 0.042 0.998 0. Relationships between the digestion rates of potential NDF digestibility [pdNDF (1/h)] calculated from the model-predicted and in vivo-determined digestibility of pdNDF.050 0. WEIB = Weibull.005 0.356 Intercept significantly (P < 0.023 0.954 0.014 0.011 0.897 0.006 0.004 −0.002 0.083 0.009 0.472 0.Gas production models 2665 Table 6.001 0.864 0.930 0.039 0.032 0. LOG = logistic.038 −0.013 0.023 0.012 0.795b 0.003 0.001 0.858b 0.015 0.000 0.001 0.003 0.041a 0.015 0.093 0.974 0.012 0. 1 Gen M-M = generalized Michaelis-Menten.147 0.002 0.989 0. and distribution of the mean squared prediction error (MSPE) Distribution of MSPE Model1 One-pool EXP0 EXPDLag EXPLPool France Gen M-M GOMPS GOMPF Groot LOG RICH WEIB Two-pool GOMP2S GOMP2F Groot2 LOG2 RICH2 WEIB2 Three-pool GOMP3S LOG3 Groot3 a b Intercept 0.015 Random 0.989 0.010 0.007 0.059a 0.044 −0.941 0.269 0.975 R2 0.989 0.967 0.017a 0.399 0. Slope significantly (P < 0.854 0.107 0.005 Slope 1.014 0.934 0.004 0.048 0.800 0.013 0. Slope significantly (P < 0.902 0.864 0. RICH = Richards.722 0.989 0.823b 0.820 0.027 0.933 0.003 −0.420 0.004 −0.05) different from 0.000 0.989 0.036 0.010 −0.013 0.002 0.065a 0.858 0.144 0.995 1.984 0.206 0.873 0.095 0.939 0.032a −0.025 1.075 1.742 0.018 0.022 0. GOMP = Gompertz.064a 0.012 0.005 0.302 0.740 0.063 0.668 0.903 0.850 0.011 0.769 0.05) different from 1.040 0.039 0.004 0.858 0.011 R2 0.505 0.016 −0.012 0.018 Mean bias 0.160 0.003 0.012 0.010 0.879 0.486 0.001 −0.008 0.438 Intercept significantly (P < 0.021 0.036 0.042 0. LOG = logistic.856 0.025 1.033 Slope 0.018 −0.186 0.013 0.005 0.002 0.818 0.235 0.823 0.003 0.042 0.790 0.048 0.160 0.984 0.861 0.003 0.014 0.814 RMSE 0.078 0.114 0.268 0.961 0.006 0.013 0.05) different from 0.005 0.835 0.014 −0.840 0.986 0.766 0.049 0.020a 0.732 0.303 0.986 1.006 0.015 0.178 0.007 0.851 0.010 0.579 0.065 0.146 0.001 −0.990 0.984 RMSE 0.066 0.271 0.005 0. and distribution of the mean squared prediction error (MSPE) Distribution MSPE Model1 One-pool EXP0 EXPDLag EXPLPool France Gen M-M GOMPS GOMPF Groot LOG RICH WEIB Two-pool GOMP2S GOMP2F Groot2 LOG2 RICH2 WEIB2 Three-pool GOMP3S LOG3 Groot3 a b Intercept 0.006 0.862 0.153 0.014 −0. Relationships between the predicted and observed in vivo NDF digestibility. GOMP = Gompertz.986 0.047 0.008 1.758 0. root mean square error (RMSE).025 0.014 Bias 0.284 0.006 0.140 0.872 0.000 0.055 1.985 0.005 0.862 0.391 0.883 0.000 0.000 Random 0.010 0.100 0.005 0.037 0.005 0.065 0. WEIB = Weibull.007 0.953 1.001 0.983 0.040 0.063 0. RICH = Richards.986 0.633 0.004 0. 1 Gen M-M = generalized Michaelis-Menten.

The pools estimated by the multipool models should therefore be viewed as purely mathematical constructs that may or may not correspond to chemical entities (Schofield.. In their study with sugars. extended ruminal in situ incubation is likely to be a more reliable method as indicated by the close empirical relationship between iNDF concentration in grass or legume silages and in vivo OM digestibility (Nousiainen et al.1 mL) and Groot2 (128. 2006) and also between individual cell wall polysaccharides (Mertens. This is in contrast to what could be assumed from the greater energy supply and increased microbial growth from early-harvested silages. Assuming a digestion rate of 0. Groot3. Our data indicate that despite the very good fit of the models.2666 Huhtanen et al. the possibility of encountering a local minimum increases. 2008). Prediction of NDF Digestibility As indicated earlier.2 mL) compared with Groot1 (128. However. Cone et al. the small improvement in the fit is likely because the data generated by summing 2 exponential functions produce a curve shaped very much like a single-pool exponential function (Schofield et al.e.8 mL) indicates. and 0.06 per hour. On contrary. Because the digestible neutral detergent solubles can be estimated accurately and precisely by the Lucas test (Huhtanen et al.and a slow-digesting pool (Van Soest et al. Because some gas production still occurred at 72 h (mean 0. . The proportion of the third gas pool characterized with the longest lag time and slow rate decreased with improved in vivo NDF digestibility.05 or 0. Furthermore.15. However. 2000). the length of the incubation period may be extended. 2005)... Although the use of kinetic parameters estimated by the 1-pool LOG Relationship Between Potential NDF Digestibility and Gas Production The close relationship between the extent of potential NDF digestibility and total volume of gas produced indicates that isolated NDF used in the present study was a relatively homogenous substrate. rather than attempting to estimate the potential extent of NDF digestion from asymptotic gas production. the gas production from the pool assumed to represent microbial turnover was below 10% of the total gas volume (i.. this model can be used to estimate both the D-value (concentration of digestible OM in DM) and kinetic parameters needed in the mechanistic dynamic models.938 of that predicted by the models EXP0. especially if forage-type specific equations are used. Groot.12 mL/h). 2003b.org by on September 21. 1994). these models had a small MSPE and bias. Including a second pool for the exponential models did not improve the fit much. GOMP. and LOG2 predicted lower asymptotic gas production than the observed values (mean 70 to 72 h). (2) using digesta passage models including selective retention of feed particles in the rumen. Including a third pool in the model further decreased the RMS.987 of pdNDF will be digested during 72 h. none of the models were able to identify a gas pool that could result from the microbial turnover. and microbial turnover. the intercept was not significantly different from 0. multipool models do not accurately and consistently identify feed entities characterized by differences in degradation kinetics. the measured value at 70 to 72 h was proportionally only 0. Another solution is to increase the number of pools as lower asymptotic gas volume with the model Groot3 (123. Rinne et al. 2006). (1997) suggested that the 3 phases of gas production can be related to the fermentation of soluble and insoluble substrates. and the slope was not different from 1. The models LOG. the declining robustness with increased number of pools is inherent in the nonlinear curve-fitting procedure. The good performance of the model can be attributed to the following 3 factors: (1) separation of NDF into iNDF and pdNDF. 0. Differences in VFA pattern in fermentation or changes in partitioning of carbon between VFA production and microbial synthesis would influence the amount of gas produced per unit of fermented substrate. 2006).. these models were not able to describe the gas production profiles correctly. respectively. Most likely.973 to 0. Downloaded from jas. and when the number of parameters is increased.. the large variation between the models in the predicted asymptotic gas volumes suggests that it is not possible to estimate pdNDF from the gas volume. this pool represents the gas production both from slowly digestible fiber and microbial turnover. However. much less than the gas production from the third pool in the present study).27% of the mean asymptotic gas production for models GOMP3S. Good performance in the curve-fitting was not always translated to accurate and precise predictions of the in vivo NDF digestibility values.. With the exponential model. and LOG3. proportionally 0. The kd decreases at later stages of fermentation in the Groot and Gen M-M models..927 to 0. In vitro NDF digestion of various forages also demonstrated a fast.20. The France and RICH models were the best 1-pool models in predicting NDF digestibility. using the gas production kinetic data in a rumen model predicted the in vivo NDF digestibility both accurately and precisely. To at least partly overcome this problem. which although theoretically sound may lead to overestimation of the asymptotic gas volume as the time-related pattern of residuals suggests. in contrast to the other 2-pool models.fass. This together with the distinct time-related pattern of changes in the residuals after 40 to 50 h of incubation (discussed later) suggests that these models overestimated the asymptotic value of gas production. 2010. stems and leaves (Chaves et al. and (3) an accurate and precise prediction of kd by a gas production system (Huhtanen et al. with the average RMSE being 0. and Groot2. 1993b). Overestimation of the asymptotic gas volume by the EXP0 model is related to its inability to describe the lag phenomenon. and the fit was almost perfect. Gen M-M.

The 1-pool RICH model was the best model in predicting in vivo first-order kd of NDF in terms of small RMSE and bias. and random variation increased. biased digestibility estimates. which resulted in underestimated extent and overestimated rate of digestion. Dhanoa et al. Both the Gen M-M and Groot1 models resulted in a significant slope bias (i. Also. consequently. Huhtanen et al. However. and WEIB2 models were almost as good. which also suggests that in vitro techniques may allow a more accurate estimation of the true intrinsic kd (ultimate limit to digestion imposed by cell wall characteristics) than the in situ technique. The studies comparing particle-associated enzyme activities within the in situ bags and in the surrounding rumen digesta (Nozière and Michalet-Doreau.fass. whereas time-related patterns during later stages of fermentation markedly influence the asymptotic value and consequently the kd. EXPDLag.78) between the residual of dNDF estimates with the value of parameter n. . with advancing incubation time. GOMP2S. the residuals decreased consistently. (1994) demonstrated that the effect of lag time on digestibility is smaller when estimated using an appropriate 2-compartment rumen model rather than a 1-pool model. and the slope close to 1. data not shown). With parameter values above 1. 1998) showed much lower activities within the bags. but they were less accurate in predicting the NDF digestibility than the other 1-pool models except for EXP0. It is possible that the compartmental retention time varied between the feeds. the France. and with values below 1. This appears to be related to the strong relationship (R2 = 0.Gas production models 2667 and GOMP models resulted in accurate and precise estimates of NDF digestibility. Accurate predictions of the in vivo values by the Groot equation appeared to require 3 pools to be used. GOMP2S. but this variation would have the same influence on both model-predicted and Downloaded from jas. much poorer fit and underestimation of the asymptotic gas volume preclude acceptance of these models. see Huhtanen et al. the EXP0 model was as accurate in predicting the in vivo NDF digestibility as the models Gen M-M and Groot despite the fact that the fit of gas production data was much better with the latter 2 models. It is possible that the assumption of decreased kd with increased fermentation time with these models was not correct. if the sum of the negative and positive residuals is equal. and consequently underestimation of the parameters describing kd. or that the effect was too strong. This may result from the time-related patterns of the residuals (observed − predicted) of gas production curves. This may be related to the time-related pattern of residuals during late stages of incubation with the Gen M-M and Groot models. 2006). biased estimates of the extent of gas production lead to biased rate estimates and. The LOG model showed an opposite pattern of time-related residual. Surprisingly.. The Gen M-M and Groot1 models fit the data much better than the other 1-pool models. high R2 value. Investigating the residuals from approximately 50 h of incubation onwards showed a clear time-related pattern (Figure 2). dNDF was underestimated. 2010. During early stages of fermentation when both under. The third pool in the LOG and GOMP models had minimal effects on the accuracy and precision of NDF digestibility estimates. and RICH2 models. Prediction of the Effective First-Order Digestion Rate The ultimate goal of using the in vitro gas production system is to produce reliable parameter values for the effective first-order kd of pdNDF to be used in mechanistic dynamic rumen models.and overfitting occur.. and the model performed similarly to the France.. 1996. which is consistent with lower kd with the in situ compared with in vitro or in vivo techniques. Ellis et al. suggesting an overestimation of the asymptotic gas volume. we used kd estimates calculated from in vivo pdNDF digestibility to validate the method.org by on September 21. In agreement with our data.0.e. the mean extent and rate of gas production of the 15 silages were strongly and negatively correlated (R2 = 0. the digestibility of low digestibility silages was underestimated more than that of high-digestibility silages). 2008). (2000) compared several models in estimating the extent of rumen digestion from gas production profiles using measured values for potentially digestible fraction but assumed the rumen as a 1-compartment system. Consistent with this. it can be calculated from the duodenal excretion pattern of internally labeled lucerne ADF-15N that only 10% of the cumulative marker excretion at the duodenum was recov- ered from dairy cows during the first 10 h after dosing (Huhtanen and Hristov. The residuals of the gas production curve showed a similar pattern as the corresponding 1-pool model. It seems that large residuals during early stages of fermentation (<20 h) have a relatively small effect on the predicted dNDF. Including the second pool in the Groot model did not improve the accuracy. Because of many methodological problems of the in situ technique and strong evidence of underestimation of the in vivo rates of pdNDF digestion (for a review. (2000) observed a negative bias with the model Gen M-M compared with the France model in the extent of rumen digestion. Dhanoa et al. Digestion rates estimated by a gravimetric in vitro method were greater than those determined in situ (Bossen et al. it was overestimated. the feed particles are mainly in the rumen nonescapable pool and not eligible for passage.95.0. all 3-pool models performed well. The second pool in the WEIB model decreased the random variation markedly. For example. including a third pool in the Groot model decreased the bias markedly.. The dNDF estimates of the WEIB model had lower correlations with the other models and also a greater random error. 2001). whereas LOG tended to overestimate the extent of digestion. However. Because the extent of digestion was determined separately using 12-d in situ incubation. In the present study.

Mertens. 1994). Kebreab... and H. H. Huhtanen. D. The estimated effective kd can be used in steady-state rumen models. and they predicted the in vivo NDF digestibility and digestion rate accurately and precisely. J. M. C. A. Visscher. Hill. Feed Sci. In vitro gas production profiles to estimate extent and effective first-order rate of neutral detergent fiber digestion in the rumen. indicating that pdNDF in grass silage cannot be considered as a homogenous substrate. D. D. D. the model EXPDLag predicted the in vivo data accurately despite greater RMS in curve-fitting. Feed Sci. Pitt et al. 61:113–128. 2000. and both passage and lag time had an influence on the estimated effective kd. Huhtanen. Estimating passage kinetics using fibre-bound 15N as an internal marker. C. Hatfield. H. J. France. Technol. Technol. J. Hvelpund. and J. France. R.org by on September 21. Nørgaard. 83:143–150. Conclusions Two-pool models could be fit better to gas production data than 1-pool models. J. G. Danfær. D. M. G. 1996. A. P. J. and F. France. 2006. P. Chaves. S. Digestion and passage of fibre in ruminants.. Lopez. Zileshi. and M. no time-related pattern or residuals. Matis. the proportion of pdNDF disappearing by passage is smaller during the early residence time compared with a 1-compartment system. Oudshoorn. Feed Sci. and M. Bibby. A. B. Pages 407–415 in Nutrient Digestion and Utilization in Farm Animals: Modelling Approaches.. M. UK.. A. Brookes. 64:77–89. Weisbjerg. H. 2008. Huhtanen. and it results in the same pdNDF digestibility as estimated for a single feed batch using time-related digestion rates estimated from the gas production curves. Forbes and J. Br. There was no evidence of benefits from using 2-pool gas production models in terms of predicting the in vivo parameters despite markedly better fit of the data. R. 1994.2668 Huhtanen et al. Wallingford. Estimating the extent of degradation of ruminant feeds from a description of their gas production profiles observed in vitro: Comparison of models. Anim. Mertens. M. van Gelder. Dijkstra. C. B. WI. Technol. 1998. UK. Williams. They assumed rumen as a single-compartment system without selective retention of particles. V. However. S. D. Nutr. 2006. J. and WEIB had good fit of data. Downloaded from jas. H. J. J. Theodorou. Madison. Dijkstra. Enzyme activities of rumen particles and feed samples incubated in situ with differing types of cloth. The best fit was not always translated into improved accuracy of the in vivo predictions. Davies. Fahey Jr. J. Anim. 2001.. WI. J. T.. R. When the first-order digestion rate for mechanistic rumen models is to be estimated. M. Nutr. E. in the 2-compartment system. C. and T. Digestion kinetics of leaf. Bannink. Dhanoa. Moser. M. J. (1999) presented a formal derivation for several models with variable digestion rates for estimating the effective kd including the effects of digestion lag. Influence of fermentation method on NDF degradation parameters. S. and D. 91:1464–1476. 128:229–252. Lister. Davies. LITERATURE CITED Allen.. a digestion lag time would have a smaller effect on the digestibility in the 2-compartment compared with 1-compartment system (Ellis et al. M. stem and inflorescence from five species of mature grasses. and J. Sci. M. W. Anim. S. M. but for later residence times. J. Cone. Nutr. R.. Lund. ed. A. and E. 163:99–111. UK. Nutr. 118:261–270. London. Pages 682–756 in Forage Quality. Ahvenjärvi. Williams. J.. R. Huhtanen. K. S. Feed Sci. the Netherlands. 2000. Hellämäki. M. Weisbjerg. 79:161–168. and M. Seppälä. and P. R. Wallingford. an investigation of the pattern of residuals during later stages of fermentation is important in addition to fitting criteria. J. Dairy Sci.. CAB International. E.. 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