In Ichthyop, there is the possibility to define temperature and salinity values below and above which the particle is killed.
If constant lethal temperatures or salinities are provided, the cold and warm lethal temperatures are provided with the cold_lethal_temperature, warm_lethal_temperature, fresh_lethal_salinity, saline_lethal_salinity, .
If a CSV file is used, the lethal temperatures and salinities are provided in a CSV (lethal_temperature_file and lethal_salinity_file parameters) which is formatted as follows:
The first column provides the class intervals, which are either the age of the particle (in days) or the length of the particle (in cm). The variable that is used to determine the class is provided by the temperature.class parameter.
DEB growth
The DEB equations are based on Kooijman (2010) and can be found in Flores-Valiente et al. (2023)
Influence of temperature is given by equation 5 of Flores-Valiente et al. (2023):
\[
c_T = \exp \left( \dfrac{T_A}{T_1} - \dfrac{T_A}{T} \right)
\left[
\dfrac{
1 +
\exp
\left(
\dfrac{T_{AL}}{T_1}
-
\dfrac{T_{AL}}{T_L}
\right)
+
\exp
\left(
\dfrac{T_{AH}}{T_H}
-
\dfrac{T_{AH}}{T_1}
\right)
}
{
1 +
\exp
\left(
\dfrac{T_{AL}}{T}
-
\dfrac{T_{AL}}{T_L}
\right)
+
\exp
\left(
\dfrac{T_{AH}}{T_H}
-
\dfrac{T_{AH}}{T}
\right)
}
\right]
\tag{6.3}\]
If the particle is an embryo (\(E_H < E_{Hb}\)), then it does not feed. Please note that, according to this definition, a larva that hatched but does not feed is considered an embryo in DEB theory. Else, the scaled functional response \(f\) is computed as:
\[
f = \dfrac{F}{F + K}
\tag{6.4}\]
with \(F\) the food vector and \(K\) the half-saturation constant.
Assimilation rates \(\{\dot{p_{Am}}\}\), energy conductance \(\dot{v}\), volume specific somatic maintenance (\([\dot{p}_M]\)) and maturity maintenances rates (\(\dot{k}_J\)) are corrected by the \(c_T\) factor.
If the accelerated DEB model is used (see e.g. Kooijman et al. (2011) and Kooijman (2014) for more details), the acceleration factor is computed as follows
\[
s_M =
\begin{cases}
1 & (E < E_{Hb})\\
\dfrac{V^{1/3}}{L_b} & (E_{Hb} \le E_H < E_{Hj}) \\
\dfrac{L_j}{L_b} & (E_{H} > E_{Hj})
\end{cases}
\tag{6.5}\]
with \(L_b\) the structural length of the particle when \(E_H = E_Hb\) (i.e the length at birth, i.e. first feeding) and \(L_j\) the structural length of the particle when \(E_H = E_Hj\) (i.e the length when metamorphosis occurs). The \(s_M\) factor is then used to multiply the assimilation rate \(\{\dot{p_{Am}}\}\) and the energy conductance \(\dot{v}\).
(NICOLAS: s’agit il de la longueur structurelle \(L = V^{1/3}\) ou de la longueur mesurée \(L_w = L / \delta_M\)). Please note that \(L_b\) and \(L_j\) are here considered as parameters but they should be in fact calculated prior any Ichthyop runs at (e.g. at \(f = 1\)) as they depend on the other parameters. If you are considering testing different parameter sets, \(L_b\) and \(L_j\) should be modified accordingly (e.g. using the DEBtool function get_tj.m).
Consequently, the assimilation (in \(J.d^{-1}\)) is computed as follows:
\[
\dot{p}_A = \begin{cases}
0 & (E_{H} < E_{Hb}) \\
c_T s_M f \{ \dot{p}_{Am}\} V^{2/3} & (E_{H} \ge E_{Hb}) \\
\end{cases}
\tag{6.6}\]
Energy loss to maintenance is given by
\[
\dot{p}_M = c_T [ \dot{p}_M ] V
\tag{6.7}\]
The mobilized energy is given by:
\[
\dot{p_C} = c_T
\dfrac{\dfrac{E}{V} \left([E_G] s_M \dot{v} V^{2/3} + \dot{p}_M\right)}
{\kappa \dfrac{E}{V} + [E_G]}
\tag{6.8}\]
Maturity maintenance is given by
\[
\dot{p}_J = c_T \dot{k}_J E_H
\tag{6.9}\]
Energy that is sent to structural growth is given by:
\[
\dot{p}_G = \kappa \dot{p}_C - \dot{p}_M
\tag{6.10}\]
Energy that goes to development (as embryo and juvenile) or to reproduction (as adults) is given by:
\[
\dot{p}_R = (1 - \kappa) \dot{p}_C - \dot{p}_J
\tag{6.11}\]
Finally, the increment of reserve \(E\) and structural volume \(V\) is given by:
\[
\dfrac{dE}{dt} = \dot{p}_A - \dot{p}_C
\tag{6.12}\]
\[
\dfrac{dV}{dt} = \dfrac{\dot{p}_G}{{[E_G]}}
\tag{6.13}\]
The increment of the energy invested into development or reproduction depends on the value of \(E_H\).
\[
\dfrac{dE_H}{dt} =
\begin{cases}
\dot{p}_R & E_H < E_{Hp}\\
0 & \text{otherwise}
\end{cases}
\tag{6.14}\]
\[
\dfrac{dE_R}{dt} =
\begin{cases}
0 & E_H < E_{Hp}\\
\dot{p}_R & \text{otherwise}
\end{cases}
\tag{6.15}\]
Finally, starvation mortality is determined whether one of the following conditions are matched:
\[
M_{starv} =
\begin{cases}
\kappa \dot{p}_C < \dot{p}_M\\
(1 - \kappa) \dot{p}_C < \dot{p}_J
\end{cases}
\tag{6.16}\]
Finally, all the states variables are then incremented:
\[
E(t + \Delta t) = E(t) + \dfrac{dE}{dt}\Delta t
\tag{6.17}\]
\[
V(t + \Delta t) = V(t) + \dfrac{dV}{dt}\Delta t
\tag{6.18}\]
\[
E_H(t + \Delta t) = E_H(t) + \dfrac{dE_H}{dt}\Delta t
\tag{6.19}\]
\[
E_R(t + \Delta t) = E_R(t) + \dfrac{dE_R}{dt}\Delta t
\tag{6.20}\]
Additional references
Kooijman, S. A. L. M. (2010). Dynamic Energy Budget theory for metabolic organisation. Cambridge Univ. Press, Cambridge. doi: 10.1098/rstb.2010. Kooijman, S. A. L. M. (2014). Metabolic acceleration in animal ontogeny: An evolutionary perspective. Journal of Sea Research, 94(Supplement C), 128–137. https://doi.org/10.1016/j.seares.2014.06.005 Kooijman, S. A. L. M., Pecquerie, L., Augustine, S., & Jusup, M. (2011). Scenarios for acceleration in fish development and the role of metamorphosis. J. Sea Res., 66, 419–423. https://doi.org/10.1016/j.seares.2011.04.016
(book?){kooijman_dynamic_2010, title = {Dynamic {Energy} {Budget} theory for metabolic organisation}, isbn = {978-0-521-13191-9}, url = {doi: 10.1098/rstb.2010.}, publisher = {Cambridge Univ. Press, Cambridge}, author = {Kooijman, S.A.L.M.}, year = {2010}, }
(article?){kooijman_scenarios_2011, title = {Scenarios for acceleration in fish development and the role of metamorphosis.}, volume = {66}, doi = {https://doi.org/10.1016/j.seares.2011.04.016}, journal = {J. Sea Res.}, author = {Kooijman, S. A. L. M. and Pecquerie, L. and Augustine, S. and Jusup, M.}, year = {2011}, pages = {419–423}, }
(article?){kooijman_metabolic_2014, series = {Dynamic {Energy} {Budget} theory: applications in marine sciences and fishery biology}, title = {Metabolic acceleration in animal ontogeny: {An} evolutionary perspective}, volume = {94}, issn = {1385-1101}, shorttitle = {Metabolic acceleration in animal ontogeny}, url = {http://www.sciencedirect.com/science/article/pii/S1385110114001105}, doi = {10.1016/j.seares.2014.06.005}, number = {Supplement C}, urldate = {2017-11-21}, journal = {Journal of Sea Research}, author = {Kooijman, S. A. L. M.}, month = nov, year = {2014}, keywords = {Acceleration, Dispersal, Maturation, Parental care, add_my_pet}, pages = {128–137}, }
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