Nematode Population Dynamics and Economic Thresholds Dinâmica das Populações de Nematóides e Níveis de Dano Econômico 23o CONGRESSO BRASILEIRO DE NEMATOLOGIA March 14, 2001 Howard Ferris Department of Nematology University of California, Davis Basic components of the dynamics of populations: • • • • Birth and death rates Development and senescence rates Population size Density dependence – resource availability • Predator pressure Birth Rates • Intrinsic factors – oocytes and sperm – age effects • Extrinsic factors – resource availability – mate availability – temperature Consequences of Multiple Mating •C. elegans produces 4x more eggs when multiple-mated than by hermaproditism. Sex Ratios and Multiple Mating Effects 4000 Pf •Females of Heterodera attract and are mated by several males 2000 Eggs fertilized •R. pellio male does not supply sufficient sperm to fertilize all oocytes from a single female 0 Rhabditis pellio 400 350 300 250 200 150 100 50 0 Total sperm = 884 Female produces 600 oocytes Only 150 fertilized at a single mating 3 5 7 Male Age 1:1 F:M 0 9 50 0.3:0.7 F:M Pi 0.7:0.3 F:M 100 •Probability that female genes are perpetuated is increased •Population may increase at a greater rate when there are fewer females and more males 120 Caenorhabditis elegans Age-specific Reproduction N2 100 clk-1 age-1 80 60 40 20 0 3 4 5 6 7 Age (days) Chen, Carey and Ferris (2001), Expt. Gerontology 36:431-440 8 9 10 wild type 400 300 200 Lifetime egg production 100 0 0 8 16 24 32 40 48 age-1 400 300 200 100 0 0 8 16 24 32 LIFESPAN (days) Chen, Carey and Ferris (2001), Expt. Gerontology 36:431-440 40 48 Death Rates • Intrinsic factors – natural longevity – relationships of fecundity and longevity • Extrinsic factors – resource availability – environmental extremes – predation – management NUMBER ALIVE 48 C. elegans wild type ³80 eggs/day 40-79 eggs/day 1-39 eggs/day 0 eggs/day 24 lx 1 .0 w i l d ty p e c lk -1 age-1 0 .8 0 .6 0 .4 0 .2 0 .0 0 0 0 8 16 24 AGE (days) Chen, Carey and Ferris (2001), Expt. Gerontology 36:431-440 32 8 16 24 32 A G E (d ays ) 40 40 48 48 Many types of models represent our understanding of the dynamics of populations…. • Continuous and discrete time models – differential equations and time steps – understand behavior through calculus or sensitivity analysis • Age and stage structured models • Deterministic and stochastic models • Individual and event-based models – time steps or event steps Models with parameters related to properties of the organisms are usually more satisfying to biologists than equations that draw lines through points on a graph Continuous time models Nt=N0 t Nt=N0ert, dN/dt=rN 6000 r=dNt/Ntdt (growth rate/indiv.) 4000 Nt =er (pop. growth/unit time) 2000 0 0 800 Nt 600 400 200 0 0 20 40 60 Time 80 100 200 400 N0 600 800 Continuous time models Nt=N0 t Nt=N0ert, dN/dt=rN 6000 r=dNt/Ntdt (growth rate/indiv.) 4000 Nt =er (pop. growth/unit time) 2000 Seasonal Multiplication: 0 Nt/N0=ert 0 Nt=aN0(b+1) 800 10 600 8 Nt/N0 Nt Nt/N0=aN0b, 400 4 0 2 20 40 60 Time 80 100 400 N0 600 800 6 200 0 200 0 200 N0 400 600 Ln Final Population (Pf) 10 dN/dt=rN(1-N/K) Nt=K/(1+((K/N0-1)(e-rt)) 5 dP/dt=aP(1-P/E) Pf=aEPi/((a-1)Pi+E) Pf=(75/-Ln0.993)(1-0.993Pi) 0 0 2 4 6 8 Ln Initial Population (Pi) Sep 99 10 Pf=(a/-Lnq)(1-qPi) 500 Multiplication Rate Pf/Pi=((a/-Lnq)(1-qPi))/Pi Multiplication Rate (Pf/Pi) 450 400 350 Pf/Pi=1018 Pi-0.71, r2=0.71 Pf/Pi=(400/-Ln0.90)(1-0.90Pi)/Pi 300 250 200 150 100 50 0 0 2 4 6 Ln(Pi) Sep 99 8 10 Multiplication Rate (Pf/Pi) Seasonal population change 500 Meloidogyne arenaria - oriental melon 400 for Pi<5, Pf/Pi=325, else: Pf/Pi=1018 Pi 300 -0.71 2 , r =0.71 200 100 0 0 2 4 6 Ln(Pi) Sep 99 Kim and Ferris (2001) 8 10 Discrete time models 1.2 1 0.8 0.6 0.4 0.2 1.2 0 1 10 C 15 C 20 C 25 C 30 C 35 C 0.8 Soil Temperature Rate Rate Discrete time models 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 0.96 1.92 3.84 1.2 1 0.8 0.6 0.4 0.2 1.2 0 1 20 C 25 C 30 C 35 C 0.8 Soil Temperature 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 0.96 1.92 3.84 Soil Moisture (bars) 1.2 1 0.8 Rate 15 C 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C 30 C 35 C Soil Temperature 1.2 1 0.8 Rate 10 C Rate Rate Discrete time models 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 0.96 1.92 3.84 1.2 1 0.8 Rate Discrete time models 0.6 0.4 0.2 1.2 0 1 10 C 15 C 20 C 25 C 30 C 35 C 0.8 Rate Soil Temperature 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 0.96 1.92 3.84 Soil Moisture (bars) 1.2 1 Rate 0.8 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C 30 C 35 C Soil Temperature 1.2 1 Rate 0.8 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 1.2 1 Rate 0.8 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C Temperature 30 C 35 C 0.96 1.92 3.84 1.2 1 0.8 Rate Discrete time models 0.6 0.4 0.2 1.2 0 1 10 C 15 C 20 C 25 C 30 C 35 C 0.8 Rate Soil Temperature 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 0.96 1.92 3.84 Soil Moisture (bars) 1.2 1 Rate 0.8 1.2 0.6 0.4 1 0.2 0.8 0 0.6 10 C 15 C 0.4 20 C 25 C 30 C 35 C Soil Temperature 0.2 1.2 0 1 20 C 25 C 30 C 35 C 0.8 Temperature Rate 15 C 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 1.2 1 0.8 Rate 10 C 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C Temperature 30 C 35 C 0.96 1.92 3.84 1.2 1.2 0.6 0.4 1 0.2 0.8 Rate 1 0.8 Rate Discrete time models 1.2 0 1 10 C 0.6 15 C 20 C 25 C 30 C 35 C 0.8 Soil Temperature Rate 0.4 0.6 0.2 0.4 0 10 C 15 C 20 C 25 C 30 C 0.2 35 C Temperature 0 0.015 0.03 0.06 0.12 0.24 0.48 0.96 1.92 3.84 Soil Moisture (bars) 1.2 1 Rate 0.8 1.2 0.6 0.4 1 0.2 0.8 0 0.6 10 C 15 C 0.4 20 C 25 C 30 C 35 C Soil Temperature 0.2 1.2 0 1 20 C 25 C 30 C 35 C 0.8 Temperature Rate 15 C 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 1.2 1 0.8 Rate 10 C 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C Temperature 30 C 35 C 0.96 1.92 3.84 1.2 Discrete time models 1 Rate 0.8 1.2 0.4 1 0.2 0.8 Rate 0.6 1.2 0 1 10 C 0.6 15 C 20 C 25 C 30 C 35 C 0.8 Soil Temperature Rate 0.4 0.6 0.2 0.4 0 10 C 15 C 20 C 25 C 30 C 0.2 35 C Temperature 0 0.015 0.03 0.06 0.12 0.24 0.48 0.96 1.92 3.84 Soil Moisture (bars) 250 1.2 1 200 150 1 0.8 100 0.6 0.4 0.2 Rate Eggs J2 J3 J4 Ad 1.2 0.8 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C 30 C 35 C Soil Temperature 50 1.2 0 1 20 C 25 C 30 C 35 C Temperature 0.8 0 Rate 15 C 0 10 20 30 Days 40 50 0.6 0.4 0.2 0 0.015 0.03 0.06 0.12 0.24 0.48 Soil Moisture (bars) 1.2 1 0.8 Rate 10 C 0.6 0.4 0.2 0 10 C 15 C 20 C 25 C Temperature 30 C 35 C 0.96 1.92 3.84 250 200 Eggs J2 J3 J4 Ad 150 100 50 0 0 10 20 30 Days 40 50 Total (all stages) 600 500 400 300 200 100 0 0 10 20 30 Days 40 50 Statistical Models Total (all stages) 600 500 400 300 200 100 0 0 10 20 30 Days 40 50 Crop Yield in Relation to Nematode Population Density Oriental melon - Meloidogyne arenaria 1.2 1 0.8 0.6 0.4 0.2 0 0 2 C Relative Yield B Early season Relative Yield Relative Yield A 4 6 Ln (Pi+1) 8 10 Total harvest Late season 1.2 1 0.8 0.6 0.4 0.2 0 0 2 4 6 Ln (Pi+1) 8 A: Early season Y = 0.43+0.57*0.998Pi, ym=19743 1.2 1 0.8 0.6 0.4 0.2 0 B: Late season Y = 0.03+0.97*0.998Pi, ym=10170 0 2 Kim and Ferris (2001) 4 6 Ln (Pi+1) 8 10 C: Total harvest Y = 0.50+0.50*0.999Pi, ym=12312 10 Value Loss (WON) Early Late Total Crop Value Early Harvest Late Harvest Panel A 2019 won/kg 967 won/kg Panel B 967 won/kg 2019 won/kg 0 10 20 30 40 50 60 Pi Sep 99 Value Loss (WON) A 300000 250000 200000 150000 100000 50000 0 B Kim and Ferris (2001) 400000 300000 Early Late 200000 Total 100000 0 0 10 20 30 40 50 60 Pi Sep 99 The Economic Threshold That initial population at which the loss in value due to nematode damage is equal to the cost of nematode management The Economic Threshold amended That initial population at which the difference in crop value with and without management is equal to the cost of the management Profitability Limit constraint That initial population level at which net returns become zero Management Efficacy = 100% ET = 63 PL1 = 245 600 400 200 4.15 5.5 0 0 2 4 Ln (Pi+1) 6 8 Management Efficacy = 90% 800 Net Returns Net Returns 800 ET = 74 PL1 = 245 PL2 = 1153 600 400 200 4.3 5.5 7.05 0 0 2 4 Ln (Pi+1) 6 8 Fixed Cost Economic Threshold Net Returns ($) 600 500 400 300 200 100 0 0 2 4 6 Nematode Population (Ln) 8 10 Continuous Model Optimization 1600 1400 1200 a b Pi m T z $max E.T. = = = = = = = = 15 50 550 0.1 50 0.999 1000 110 1000 $ 800 600 400 200 0 0 2 4 6 log2 Pi 8 10 a= 600 Pi = 200 m= 0.1 T= 20 z= 0.99 $max = 1000 E.T. = 78.48428 Discrete Model 1200 1000 800 $ 600 400 200 0 0 2 4 log2 Pi 6 8 10 Optimized Discrete Model Seasonal Multiplication Rates (Host Crop) 500 Pf/Pi 400 a= b= amax = p= q= s= 500 -0.2 500 1 -0.1 0.65 300 200 100 0 0 500 1000 Pi 1500 2000 Overwinter Survival Rates 1 Pi2/Pf1 0.8 a= b= amax = p= q= s= 500 -0.2 500 1 -0.1 0.65 0.6 0.4 0.2 0 0 500 1000 Pf1 1500 2000 Annual Population Change (Host Crop) 120000 a= b= amax = p= q= s= 500 -0.2 500 1 -0.1 0.65 Pi1 * (Pi2/Pi1) 100000 80000 60000 40000 20000 0 0 500 1000 Pi1 1500 2000 Annual Population Change (Non-host) 1400 1200 Pi(t+x) 1000 800 Pi1 Pi2 a= b= amax = p= q= s= 500 -0.2 500 1 -0.1 0.65 Pi3 600 400 200 0 0 500 1000 Pi(t) 1500 2000 1600 a= b= s= Pi(0) = 1400 Pi(t+x) 1200 300 0.6 0.4 70 1000 800 600 400 200 0 0 1 2 3 4 5 6 7 Years After Planting Host Crop 8 0NHR Population Convergence Population Level 3000 2NHR 4NHR 6NHR 2000 1000 0 0 5 Year 10 15 Ave. Annual Returns ($) Optim um Rotation Length 300 200 100 0 -100 -200 0 1 2 3 4 5 6 7 Years of Non-host 8 9 10 Perennial Crop Considerations 12000 Mesocriconema xenoplax 10000 Lovell Nemaguard 8000 6000 4000 2000 0 0 200 400 600 800 1000 1200 1400 1600 1800 2000 Days 12000 Mesocriconema xenoplax 10000 8000 6000 4000 2000 0 0 2000 4000 6000 8000 Degree-Days 10000 12000 14000 2200 Year 1 Year 2 100 60 LU 40 LT 20 NU 0 NT 0 1000 DD 2000 AUC AUC 80 12000 10000 8000 6000 4000 2000 0 3000 LT NU NT 1000 2000 DD 3000 AUC AUC LU 0 LT NU NT 0 Year 3 30000 25000 20000 15000 10000 5000 0 LU 1000 DD 2000 3000 Year 4 30000 25000 20000 15000 10000 5000 0 LU LT NU NT 0 1000 DD 2000 3000 Year 1 Year 2 100 60 LU 40 LT 20 NU 0 NT 0 1000 DD 2000 3000 AUC LU LT NU NT 0 1000 2000 DD 3000 LU LT NU NT 0 Year 3 30000 25000 20000 15000 10000 5000 0 AUC AUC 80 12000 10000 8000 6000 4000 2000 0 1000 DD 2000 3000 Coefficient 12 LT-Full 10 LT-S/F 8 LU-Full 6 LU-S/F 4 NT-Full 2 NT-S/F 0 NU-Full Year 2 Year 3 Year 4 Year 5 NU-S/F Area Under Curve 80 60 Pi2170 40 Pi4 20 Pi43 Pi434 0 0 2000 4000 DD Noling and Ferris (1987) Alflafa Yield Loss 40 y=1.15+0.37x, r2=0.89 30 20 10 0 0 20 40 60 AUC 80 100 References Burt, O. R. and H. Ferris. 1996. Sequential decision rules for managing nematodes with crop rotations. J. Nematology 28:457-474. Chen, J., J.R. Carey and H. Ferris. 2001. Comparative demography of isogenic populations of Caenorhabditis elegans Expt. Gerontology 36:431-440. Ferris, H. 1978. Nematode economic thresholds: derivation, requirements and theoretical considerations. J. Nematology 10:341-350. Ferris, H. 1985. Density-dependent nematode seasonal multiplication and overwinter survivorship: a critical point model. J. Nematology 17:93-100. Hsin, H. and C. Kenyon. 1999. Signals from the reproductive system regulate the lifespan of C. elegans. Nature 399:362-366. Kim D.G. and H. Ferris. 2001. Relationship between crop losses and initial population densities of Meloidogyne arenaria in winter-grown oriental melon in Korea. J. Nematology (subm.) Noling, J.W. and H. Ferris. 1987. Nematode-degree days, a density-time model for relating epidemiology and crop losses in perennials. J. 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