Dynamic Energy Budget Theory - I Tânia Sousa with contributions from : Bas Kooijman
Feb 23, 2016
Dynamic Energy Budget Theory - I
Tânia Sousa with contributions from : Bas Kooijman
Metabolism in a DEB
individual. Rectangles are state
variables Arrows are flows of food
JXA, reserve JEA, JEC, JEM, JET , JEG, JER, JEJ or structure JVG.
Circles are processes The full square is a fixed
allocation rule (the kappa rule)
The full circles are the priority maintenance rule.
A DEB organism: growth
MV - Structure
Feeding
MH - Maturity
XAJ EAJ
Assimilation
ME - ReserveMobilisation
ECJ
Offspring MER
Somatic Maintenance
Growth
Maturity Maintenance
Reproduction
Maturation
ESJ
EGJEJJ
ERJ
Von Bertallanffy growth in DEB theory
DEB theory predicts: decreases with specific maintenance needs and
increases with the reserve density (food level) decreases with
𝑑𝐿𝑑𝑡 =�̇� 𝐵 (𝐿−𝐿 )
Von Bertalanffy: growth at constant food
1�̇�𝐵
=3𝐿�̇� +
(3 �̇� [𝑀𝑉 ] 𝑦𝐸𝑉+3𝐿𝑇 [ �̇� 𝐸𝑀 ] )�̇� [ �̇�𝐸𝑀 ]
Von Bertalanffy: growth at constant food
time, dultimate length, mm
leng
th, m
m
Von
Ber
t gro
wth
rate
-1, d
A lower the food level implies a smaller ultimate size and a shorter time to reach it. Empirical fact: organisms of the same species at different food levels
exhibit von Bertallanfy growth rates that are inversely proportional to ultimate length
Extremes in relative growth rate in insects
Buprestis splendens (jewel beetle)Juveniles live in wood for 20-40 a
Antheraea polyphemus (polyphemus moth)Juveniles increase weight 80000 × in 48 h
Represent dL/dt as a function of L. Represent dL/dt as a funtion of L for two
organisms of the same species living at different food densities.
Exercise
𝑑𝐿𝑑𝑡 =�̇� 𝐵 (𝐿−𝐿 )
Growth in DEB:
What happens to the reserve density in an egg?
Egg and foetal development: differences
𝑑𝐿𝑑𝑡 =1
3𝑚𝐸 �̇� [𝑀𝑉 ]− [ �̇� 𝐸𝑀 ]𝐿− { �̇�𝐸𝑇 }
𝑚𝐸 [𝑀𝑉 ]+ [𝑀𝑉 ] 𝑦𝐸𝑉
Growth in DEB:
What happens to the reserve density in an egg? It decreases in time
What happens to the reserve density in a foetus?
Egg and foetal development: differences
𝑑𝐿𝑑𝑡 =1
3𝑚𝐸 �̇� [𝑀𝑉 ]− [ �̇� 𝐸𝑀 ]𝐿− { �̇�𝐸𝑇 }
𝑚𝐸 [𝑀𝑉 ]+ [𝑀𝑉 ] 𝑦𝐸𝑉
Growth in DEB:
What happens to the reserve density in an egg? It decreases in time
What happens to the reserve density in a foetus? It tends to infinity
Obtain V(t) for a foetus
Egg and foetal development: differences
𝑑𝐿𝑑𝑡 =1
3𝑚𝐸 �̇� [𝑀𝑉 ]− [ �̇� 𝐸𝑀 ]𝐿− { �̇�𝐸𝑇 }
𝑚𝐸 [𝑀𝑉 ]+ [𝑀𝑉 ] 𝑦𝐸𝑉
Growth in DEB:
What happens to the reserve density in an egg? It decreases in time
What happens to the reserve density in a foetus? It tends to infinity
Obtain V(t) for a foetus
Empirical pattern: Volume is proportional to cubed time in a foetus
Egg and foetal development: differences
𝑑𝐿𝑑𝑡 =1
3𝑚𝐸 �̇� [𝑀𝑉 ]− [ �̇� 𝐸𝑀 ]𝐿− { �̇�𝐸𝑇 }
𝑚𝐸 [𝑀𝑉 ]+ [𝑀𝑉 ] 𝑦𝐸𝑉
V (𝑡 )=( �̇� 𝑡3 )3
Egg & Foetal development
If growth is supply driven when does growth
stops?
Competition between growth and somatic maintenance
MV - Structure
Feeding
MH - Maturity
XAJ EAJ
Assimilation
ME - ReserveMobilisation
ECJ
Offspring MER
Somatic Maintenance
Growth
Maturity Maintenance
Reproduction
Maturation
ESJ
EGJEJJ
ERJ
If growth is supply driven when does growth stops?
When all the energy that goes for somatic maintenance plus growth is used for maintenance
As the organism gets bigger it gets more food ( to V2/3) but it grows slower because somatic maintenance ( to V) is competing with growth
The higher the specific somatic maintenance needs the lower the ultimate size
Competition between growth and somatic maintenance
MV - Structure
Feeding
MH - Maturity
XAJ EAJ
Assimilation
ME - ReserveMobilisation
ECJ
Offspring MER
Somatic Maintenance
Growth
Maturity Maintenance
Reproduction
Maturation
ESJ
EGJEJJ
ERJ
�̇�𝐸𝐴=𝑦𝐸𝑋 �̇� 𝑋𝐴= 𝑓 (𝑋 ) { �̇� 𝐸𝐴𝑚}𝑉 2/3
Metabolism in a DEB
individual. Rectangles are state
variables Arrows are flows of food
JXA, reserve JEA, JEC, JEM, JET , JEG, JER, JEJ or structure JVG.
Circles are processes The full square is a fixed
allocation rule (the kappa rule)
The full circles are the priority maintenance rule.
A DEB organism: maturity maintenance
MV - Structure
Feeding
MH - Maturity
XAJ EAJ
Assimilation
ME - ReserveMobilisation
ECJ
Offspring MER
Somatic Maintenance
Growth
Maturity Maintenance
Reproduction
Maturation
ESJ
EGJEJJ
ERJ
Collection of processes that maintain the level of
maturity Defense and regulating systems
Maturity maintenance is paid from flux (1-)JE,C:
maturity level It does not increase after the onset of reproduction
Maturity maintenance
Specific maturity maintenance costs are constant because of the strong homeostasis
The complexity would decrease in the absence of energy spent in its maintenance (2nd Law of thermodynamics)
Empirical pattern: no reproduction occurs at very low food densities
�̇�𝐸 𝐽=𝑘 𝐽 𝑀𝐻
- maturity maintenance rate coefficient
Metabolism in a DEB
individual. Rectangles are state
variables Arrows are flows of food
JXA, reserve JEA, JEC, JEM, JET , JEG, JER, JEJ or structure JVG.
Circles are processes The full square is a fixed
allocation rule (the kappa rule)
The full circles are the priority maintenance rule.
A DEB organism: maturation/reproduction
MV - Structure
Feeding
MH - Maturity
XAJ EAJ
Assimilation
ME - ReserveMobilisation
ECJ
Offspring MER
Somatic Maintenance
Growth
Maturity Maintenance
Reproduction
Maturation
ESJ
EGJEJJ
ERJ
The use of reserve to increase the state of
maturity (embryo and juvenile) or to reproduce (adult)
Allocation to maturation in a juvenile (MH <MH
p) or to reproduction in na adult (MH >=MH
p) (supply driven):
Maturation/Reproduction
Empirical pattern: organisms kept at low food density never reach puberty implying that they will not reproduce
Stage transitions should not be linked with size
�̇�𝐸𝑅=(1−) �̇� 𝐸𝐶− �̇�𝐸 𝐽
MHb- threshold of maturity at birth
MHp- threshold of maturity at puberty
Extremes in relative maturity at birth in
mammals
Ommatophoca rossii (Ross Seal) ♂ 1.7-2.1 m, 129-216 kg♀ 1.3-2.2 m, 159-204 kgAt birth: 1 m, 16.5 kg; ab = 270 d
Didelphus marsupiales (Am opossum) ♂, ♀ 0.5 + 0.5 m, 6.5 kgAt birth: <2 g; ab = 8-13 d10-12 (upto 25) young/litter, 2 litters/a
Extremes in relative maturity at birth in fish
Latimeria chalumnae (coelacanth) ♂, ♀ 1.9 m, 90 kgEgg: 325 gAt birth: 30 cm; ab = 395 dFeeds on fish
Mola mola (ocean sunfish) ♂,♀ 4 m, 1500 (till 2300) kgEgg: 3 1010 eggs in bufferAt birth: 1.84 mm g; ab = ? dFeeds on jellyfish & combjellies
The amount of energy continuously invested
in reproduction is accumulated in a buffer and then it is converted into eggs providing the initial endowment of the reserve to the embryo
Initial amount of reserve follows from Initial structural vol. and maturity are negligibly
small and maturity at birth is given Empirical fact: reserve density at birth equals that of
mother at egg formation (egg size covaries with the nutritional state of the mother)
Reproduction
𝑅 �̇� 𝐸𝑅=𝑅 ((1−) �̇� 𝐸𝐶− �̇� 𝐸𝐽 )=𝑑𝑀𝐸𝑅
𝑑𝑡
- initial amount of reserve of the egg - reproduction efficiency
Rules for handling the reproduction buffer are
species-specific (different evolutionary strategies) Some species reproduce when
enough energy for a single egg has been accumulated
Some species reproduce a large clutch (some fishes have thousands of eggs)
Some species use environmental triggers for spawning (e.g., moluscs)
Reproduction: buffer handling rules
Energy flows vs. Mass flows
Fluxes Parameters�̇�𝑋= �̇� 𝑋 𝐴𝜇𝑋= 𝑓 (𝑋 ) {�̇�𝑋𝑚 }𝑉 2/ 3
�̇�𝐴= �̇� 𝐸𝐴𝜇𝐸= 𝑓 (𝑋 ) {�̇�𝐴𝑚 }𝑉 2 /3
�̇�𝑆= �̇� 𝐸𝑆𝜇𝐸=[�̇�𝑀 ]𝑉 + {�̇�𝑇 }𝑉 2 /3
=�̇�𝑅= �̇� 𝐸𝑅𝜇𝐸
{�̇�𝑋𝑚 }=𝜇𝑋 { �̇� 𝑋𝑚 }{�̇�𝐴𝑚 }=𝜇𝐸 { �̇� 𝐴𝑚}
State Variables
[�̇�𝑀 ]=𝜇𝐸 [ �̇�𝐸𝑀 ]�̇�𝐶= �̇�𝐸𝐶𝜇𝐸=𝐸( �̇�𝐿 − �̇�) {�̇�𝑇 }=𝜇𝐸 { �̇� 𝐸𝑇 }
�̇�𝐺= �̇� 𝐸𝐺𝜇𝐸=[𝐸𝐺 ] 𝑑𝑉𝑑𝑡
[𝐸𝐺 ]=𝑦𝐸𝑉 𝜇𝐸 [𝑀𝑉 ]
EHb- threshold of maturity at birth
EHp- threshold of maturity at puberty