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Lecture 2 Standard DEB model. defecation. feeding. food. faeces. assimilation. reserve. somatic maintenance. maturity maintenance. . 1- . maturation reproduction. growth. maturity offspring. structure. Standard DEB model. One type of food faeces reserve structure
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defecation feeding food faeces assimilation reserve somatic maintenance maturity maintenance 1- maturation reproduction growth maturity offspring structure Standard DEB model One type of food faeces reserve structure Isomorphy
Topological alternatives 11.1c From Lika & Kooijman 2011 J. Sea Res 66: 381-391
Test of properties 11.1d From Lika & Kooijman 2011 J. Sea Res, 66: 381-391
Feeding • Definition: • Disappearance of food from environment • Embryo’s do not feed • Comprises: • searching of food (stochastic) • handling of food
Busy periods not only include handling but also digestion and other metabolic processing Feeding arrival events of food items fast SU binding prob. 0 time slow SU binding prob. 0 time
Assimilation Definition: Conversion of substrate(s) (food, nutrients, light) into reserve(s) Transformation: food + O2 reserve + excreted products (e.g. faeces, CO2, NH3)
Reserve dynamics & allocation Increase: assimilation structural surface area Decrease: mobilisation reserve-structure interface Change in reserve density structural length-1 Reserve dynamics follows from weak homeostasis of biomass = structure + reserve -rule for allocation of mobilised reserve to soma: constant fraction of mobilisation rate
Reserve dynamics in starving active sludge PHB density, mol/mol Data from Beun, 2001 time, h
Yield of biomass on substrate reserve maintenance Data from Russel & Cook, 1995 1/spec growth rate, h-1
-rule for allocation Ingestion Respiration Ingestion rate, 105 cells/h O2 consumption, g/h Length, mm Length, mm Length, mm Reproduction Cum # of young • large part of adult budget • to reproduction in daphnids • puberty at 2.5 mm • No change in • ingest., resp., or growth • Where do resources for • reprod. come from? Or: • What is fate of resources • in juveniles? Growth: Von Bertalanffy Age, d Age, d
Somatic maintenance • Definition: • Collection of processes required to • maintain current amount of structure • Transformation: • reserve + O2 excreted products (e.g. CO2, NH3) • Comprises: • protein turnover (synthesis, but no net synthesis) • maintaining conc gradients across membranes (proton leak) • (some) product formation (leaves, hairs, skin flakes, moults) • movement (usually less than 10% of maintenance costs)
Maturity maintenance • Definition: • Collection of processes required to • maintain current state of maturity • Transformation: • reserve + O2 excreted products (e.g. CO2, NH3) • Comprises: • maintaining defence systems (immune system)
Maintenance first Chlorella-fed batch cultures of Daphnia magna, 20°C neonates at 0 d: 10 winter eggs at 37 d: 0, 0, 1, 3, 1, 38 Kooijman, 1985 Toxicity at population level. In: Cairns, J. (ed) Multispecies toxicity testing. Pergamon Press, New York, pp 143 - 164 30106 cells.day-1 400 Maitenance requirements: 6 cells.sec-1.daphnid-1 300 300 number of daphnids max number of daphnids 200 200 100 100 106 cells.day-1 0 0 6 12 30 60 120 8 11 15 18 21 24 28 32 35 37 30 time, d
Growth Definition: Conversion of reserve(s) to structure(s) Transformation: reserve + O2 structure + excreted products (e.g. CO2, NH3) Allocation to growth: Consequence of strong homeostasis:
Growth at constant food length, mm Von Bert growth rate -1, d time, d ultimate length, mm Von Bertalanffy growth curve:
Isomorphic growth 2.6c diameter, m Weight1/3, g1/3 Amoeba proteus Prescott 1957 Saccharomyces carlsbergensis Berg & Ljunggren 1922 time, h time, h Weight1/3, g1/3 Toxostoma recurvirostre Ricklefs 1968 length, mm Pleurobrachia pileus Greve 1971 time, d time, d
Mixtures of V0 & V1 morphs volume, m3 hyphal length, mm Bacillus = 0.2 Collins & Richmond 1962 Fusarium = 0 Trinci 1990 time, min time, h volume, m3 volume, m3 Escherichia = 0.28 Kubitschek 1990 Streptococcus = 0.6 Mitchison 1961 time, min time, min
Reproduction Definition: Conversion of adult reserve(s) into excreted embryonic reserve(s) Transformation : reserve + O2 reserve + excreted products (e.g. CO2, NH3) Involves: reproduction buffer + handling rules Allocation to reproduction in adults: Strong homeostasis: Fixed conversion efficiency Weak homeostasis: Reserve density at birth equals that of mother Reproduction rate: follows from maintenance + growth costs, given amounts of structure, reserve and maturity at birth
Reproduction at constant food 103 eggs 103 eggs Rana esculenta Data Günther, 1990 Gobius paganellus Data Miller, 1961 length, mm length, mm
Embryonic development 2.6.2e Carettochelys insculpta Data from Web et al 1986 embryo yolk O2 consumption, ml/h weight, g time, d time, d
General assumptions • State variables: structural body mass & reserve & maturity • structure reserve do not change in composition; maturity is information • Food is converted into faeces • Assimilates derived from food are added to reserve • Mobilised reserve fuels all other metabolic processes: • somatic & maturity maintenance, growth, maturation or reproduction • Basic life stage patterns • dividers (correspond with juvenile stage) • reproducers • embryo (no feeding • initial structural body mass is negligibly small • initial amount of reserves is substantial) • juvenile (feeding, but no reproduction) • adult (feeding & male/female reproduction)
Specific assumptions • Reserve density hatchling = mother at egg formation (maternal effect) • foetuses: embryos unrestricted by energy reserves • Stage transitions: cumulated investment in maturation > threshold • embryo juvenile initiates feeding • juvenile adult initiates reproduction & ceases maturation • Somatic maintenance structure volume & maturity maintenance maturity • (but some somatic maintenance costs surface area) • maturity maintenance does not increase • after a given cumulated investment in maturation • Feeding rate surface area; fixed food handling time • Body mass does not change at steady state (weak homeostasis) • Fixed fraction of mobilised reserve is spent on soma: • somatic maintenance + growth (-rule) • Starving individuals: • can shrink to pay somatic maintenance till some threshold • can rejuvenate to pay maturity maintenance, but this increases the hazard
1E,1V isomorph 2.9b All powers are cubic polynomials in l
1E,1V isomorph 2.9c all quantities scaled dimensionless
1E,1V isomorph 2.9d length l, survival S reserve density, e maturity, vH time, time, time, cum. feeding,10 reprod. acceleration, q hazards, h, hH time, time, time,
1E,1V isomorph 2.9D, continued scaled flux of CO2 scaled flux of H2O time, time, scaled flux of NH3 scaled flux of O2 time, time,
Primary DEB parameters 2.8a time-length-energy time-length-mass