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A Modelling Approach to Explore the Impact of Root
Distribution and Citrate Release on Phosphorus
Use Efficiency of Crops
Sustainable Agriculture Flagship


 Enli Wang, Brad Ridoutt
 Zhongkui Luo, Ulrike Bende-Michl
Outline of the presentation


1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
Outline of the presentation


1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
1.1 APSIM

• APSIM: Agricultural
  Production Systems
  Simulator                                                      System Control
                                                          Manager

• A modular modelling                          Clock                 Report

                                          Canopy                          Met
  framework                              Wheat                           SoilWat
                               Crops                         E                       SWIM
• Simulates biophysical                  Maize               N
                                                             G
                                                                        Soil C & N
                                       Sorghum                            SoilPH
                                                             I
  processes in farming                  Legume               N            SoilP
                                                             E                       Soil
                                       Other Crops                      New Module
  systems                                                                Erosion
                                         Irrigate

• Links to economic &                        Fertiliz                  Manure
                                Management              Economics   Residue
  ecological outcomes of
  management practices under
  climate variation
1.1 APSIM

 APSIM is a farming systems model, able to simulate
 • >20 crops, including pastures & trees
 • complex rotation patterns
 • water, carbon, nitrogen and phosphorus cycling
1.2 Crop Modules



  • Potential growth of crops


  • P demand based on critical P
    concentrations, biomass & new growth


  • Reduction in growth due to P deficiency
1.3 SoilP Module



                                                                                  APSIM-SoilP:
               P fertilizer        P fertilizer
               broadcast             banded                                       • does not simulate
                                       Placement effect
                                                                                  • precipitation/dissolution of
                                                                                     tightly bonded soil P
          Dissolution                       Mineralization          Unavailable   • may not work in acidic soils
 Rock P                 Labile P            Immobilization          Organic P        (pH<5.5) and alkaline soils
                                               L                                     (pH>7.3)
                                              Gaoss o
                          Desorption
                          Adsorption




                                                in f av
                                                  of ai
                                                      ava lab                     • new development is on-going
                                                         ilab ility
                                                             ilit                    for P modeling in those soils
                                                                  y
                                                                                  • original version does not
 Crop P                 P in Soil                               Unavailable          consider effect of root
 Uptake                 solution                                Inorganic P          exudation on PUE
Outline of the presentation



1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
2.1 Process understanding and data availability


• Organic anions can enhance P mobilisation into soil solution
  (Jones, 1998; Khademi et al, 2010)
• Release of citrate, malate & oxalate from roots increases with P
  deficiency (Vance et al, 2003; Ryan et al, 2001)




          Table 1 citrate efflux from plant roots
          Crop          Citrate Efflux   References
                        (nmol/gFW/h)
          Rice             155~360       Kirk et al (1999)
          Wheat              5~185       Ryan et al (2009)
          White lupin     1656~2373      Roelofs et al (2001)
          Proteaceae      3600~9000      Roelofs et al (2001)
2.2 Modelling


• Only one modeling study to simulate rice P uptake
  in controlled laboratory experiment
   • Rice (Kirk et al, 1999) –
     diffusion, decomposition and P solubilisation of
     citrate, only a few mm around roots, no zero
     efflux treatment
• No field scale modelling studies so far
Outline of the presentation



1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
3.1 APSIM enhancements

1.   Original Model: crop P uptake is linked to solution P in rooted soil layers
2.   Enhancement #1: need to link crop P uptake to solution P and root length density (RLD)
3.   Enhancement #2: need to link crop P uptake & solution P to citrate efflux from roots (Fc)


                           P fertilizer        P fertilizer
                           broadcast             banded
                                                   Placement effect


                     Dissolution                        Mineralization       Unavailable
           Rock P                   Labile P            Immobilization
                                      Desorption                             Organic P
                                      Adsorption




                                                     P released by citrate
           Crop P                   P in Soil                                Unavailable
           Uptake                   solution                                 Inorganic P
Outline of the presentation



1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
4.1 Biomass
                       1. Original Model: crop P uptake                                 2. Enhancement #1: need to link
                          is linked to solution P in rooted                                crop P uptake to solution P and
                          soil layers                                                      root length density (RLD)



                                  7000       (a) APSIM-SoilP                                                       7000       (b) APSIM-SoilP&RLD




                                                                                   Simulated wheat yield (kg/ha)
  Simulated wheat yield (kg/ha)




                                  6000        y = 1.1256x + 10.593                                                 6000       y = 1.1742x - 259.46
                                                   R2 = 0.8891                                                                     R2 = 0.896
                                  5000                                                                             5000

                                  4000                                                                             4000

                                  3000                                                                             3000

                                  2000                                                                             2000

                                  1000                                                                             1000

                                    0                                                                                0
                                         0    1000 2000 3000 4000 5000 6000 7000                                          0    1000 2000 3000 4000 5000 6000 7000
                                              Observed wheat yield (kg/ha)                                                     Observed wheat yield (kg/ha)

 Comparison of simulated and observed wheat grain yields under different levels of P
 fertiliser inputs at one QLD site and two NSW sites.



 The modified model performed slightly better than the original one!
4.2 Scenario analysis

•   Study site: Kingaroy, QLD (1957-2009)
•   Cropping system: Continuous wheat, rainfed condition
•   Citrate efficiency = 0.4
•   Five levels of soil P sorption capacity:
     •   Low         50, 100
     •   Medium      200
     •   High        500
     •   Very high   1000
• Seven levels of citrate efflux (nmol/gFW/h): 0, 50, 100, 200,
  500, 1000, 2000
• 11 levels of P application rates (kg P/ha): 0~200 kg P/ha with
  increase of 20 kg P/ha.
4.3 Biomass responses: short-term vs long-term


                                  First year 1957                                                                                      Average 1957~2009
         (a) Citrate efflux = 0 nmol/gFW/h          (b) Citrate efflux = 200 nmol/gFW/h                        (a) Citrate efflux = 0 nmol/gFW/h          (b) Citrate efflux = 200 nmol/gFW/h
          20                                                     20                                             20                                                     20




                                                                                                       Biomass (ton/ha)




                                                                                                                                                          Biomass (ton/ha)
 Biomass (ton/ha)




                                                    Biomass (kg/ha)
                16                                               16                                                   16                                               16

                12                                               12                                                   12                     Sorption:                 12                         Sorption:
                                                                                                                                                   50                                                   50
                    8                                                  8                                                  8                        100                       8                          100
                                                                                                                                                   200                                                  200
                    4                                                  4                                                  4                        500                       4                          500
                                                                                                                                                   1000                                                 1000
                    0                                                  0                                                  0                                                  0
                        0   40   80   120 160 200                          0   40   80   120 160 200                          0   40   80   120 160 200                          0   40     80    120 160 200
                             P Rate (kg/ha)                                     P Rate (kg/ha)                                     P Rate (kg/ha)                                         P Rate (kg/ha)

(c) Citrate efflux = 1000 nmol/gFW/h                (d) Citrate efflux = 2000 nmol/gFW/h               (c) Citrate efflux = 1000 nmol/gFW/h               (d) Citrate efflux = 2000 nmol/gFW/h
              20                                       20                                                           20                                                 20




                                                                                                                                                          Biomass (ton/ha)
                                                                                                       Biomass (ton/ha)
                                                    Biomass (ton/ha)
 Biomass (ton/ha)




              16                                                 16                                                 16                                                 16

              12                                                 12                                                 12                       Sorption:                 12                          Sorption:
                                                                                          Sorption:                                                50                                                    50
                    8                                                  8                        50                        8                        100                       8                           100
                                                                                                100                                                200                                                   200
                                                                       4                        200                       4                        500                       4                           500
                    4                                                                           500                                                1000                                                  1000
                                                                                                1000                                                                         0
                    0                                                  0                                                  0
                        0   40   80   120 160 200                          0   40   80   120 160 200                          0   40   80   120 160 200                          0   40      80   120 160 200
                             P Rate (kg/ha)                                     P Rate (kg/ha)                                     P Rate (kg/ha)                                         P Rate (kg/ha)
4.4 Biomass, yield and inorganic soil P

                                                (a) Citrate Efflux = 0                                                 (b) Citrate Efflux = 200
                                        25                                                                      25


      Biomass or grain yield (ton/ha)




                                                                              Biomass or grain yield (ton/ha)
                                                     nmol/gFW/h                                                              nmol/gFW/h
                                                                    Biomass                                                                  Biomass
                                        20                         Yield                                        20                          Yield

                                        15                                                                      15

                                        10                                                                      10

                                         5                                                                       5

                                         0                                                                       0
                                         1950 1960 1970 1980 1990 2000 2010                                      1950 1960 1970 1980 1990 2000 2010
                                                        Year                                                                    Year

                                                (c) Citrate Efflux = 0                                                 (d) Citrate Efflux = 200
                                        200                                                                     200
      Total inorganic soil P (kg/ha)




                                                                              Total inorganic soil P (kg/ha)
                                                                    Layer 1                                                                 Layer 1
                                                                    Layer 2                                                                 Layer 2
                                        150                         Layer 3                                     150                         Layer 3


                                        100                                                                     100


                                        50                                                                       50


                                         0                                                                       0
                                         1950 1960 1970 1980 1990 2000 2010                                      1950 1960 1970 1980 1990 2000 2010
                                                        Year                                                                    Year
4.5 P rate required for 90% max biomass


     P rate required for 90% max biomass (kg/ha)   160
                                                                                                   SP50
                                                   140                                             SP100
                                                                                                   SP200
                                                   120                                             SP500
                                                                                                   SP1000
                                                   100

                                                    80

                                                    60

                                                    40

                                                    20

                                                     0
                                                         0   200   400   600 800 1000 1200 1400 1600 1800 2000
                                                                          Citrate efflux (nmol/gFW/h)
4.6 P recovery


                       (a) P Rate = 20 kg/ha               (b) P Rate = 40 kg/ha               (c) P Rate = 60 kg/ha
                 180                                 180                                 180
                                                                                                          Sorption = 50
                 160                                 160                                 160              Sorption = 200
                                                                                                          Sorption = 500
                 140                                 140                                 140              Sorption = 1000
P recovery (%)




                 120                                 120                                 120
                 100                                 100                                 100
                  80                                  80                                  80
                  60                                  60                                  60
                  40                                  40                                  40
                  20                                  20                                  20
                   0                                   0                                   0
                       0       500 1000 1500 2000          0       500 1000 1500 2000          0       500 1000 1500 2000
                           Citrate efflux (nm/g/h)             Citrate efflux (nm/g/h)             Citrate efflux (nm/g/h)
Outline of the presentation



1. APSIM and its capability to simulate crop response to soil P


2. Current understanding and available data on citrate release from
   plant roots and its impact on PUE


3. New model development to enable APSIM to simulate impact of
   citrate efflux on PUE


4. Simulated response of wheat growth and P uptake to citrate
   effluxes


5. Summary
5. Summary


     Long-term effect is different from short-term
      effect, due to residual effect of applied P.

     Major impact is to increase PUE of applied P
      fertilisers, because soil P reserve can be depleted
      in a relatively short time period.

     P rate required to achieve maximum crop yield
      decreases with increasing citrate efflux from the
      roots.

     The impact increases with soil P sorption
      capacity, decreases with P application rate.
Thank you

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A modelling approach to explore the impact of root distribution and citrate release on phosphorus use efficiency of crops. Zhongkui Luo

  • 1. A Modelling Approach to Explore the Impact of Root Distribution and Citrate Release on Phosphorus Use Efficiency of Crops Sustainable Agriculture Flagship Enli Wang, Brad Ridoutt Zhongkui Luo, Ulrike Bende-Michl
  • 2. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 3. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 4. 1.1 APSIM • APSIM: Agricultural Production Systems Simulator System Control Manager • A modular modelling Clock Report Canopy Met framework Wheat SoilWat Crops E SWIM • Simulates biophysical Maize N G Soil C & N Sorghum SoilPH I processes in farming Legume N SoilP E Soil Other Crops New Module systems Erosion Irrigate • Links to economic & Fertiliz Manure Management Economics Residue ecological outcomes of management practices under climate variation
  • 5. 1.1 APSIM APSIM is a farming systems model, able to simulate • >20 crops, including pastures & trees • complex rotation patterns • water, carbon, nitrogen and phosphorus cycling
  • 6. 1.2 Crop Modules • Potential growth of crops • P demand based on critical P concentrations, biomass & new growth • Reduction in growth due to P deficiency
  • 7. 1.3 SoilP Module APSIM-SoilP: P fertilizer P fertilizer broadcast banded • does not simulate Placement effect • precipitation/dissolution of tightly bonded soil P Dissolution Mineralization Unavailable • may not work in acidic soils Rock P Labile P Immobilization Organic P (pH<5.5) and alkaline soils L (pH>7.3) Gaoss o Desorption Adsorption in f av of ai ava lab • new development is on-going ilab ility ilit for P modeling in those soils y • original version does not Crop P P in Soil Unavailable consider effect of root Uptake solution Inorganic P exudation on PUE
  • 8. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 9. 2.1 Process understanding and data availability • Organic anions can enhance P mobilisation into soil solution (Jones, 1998; Khademi et al, 2010) • Release of citrate, malate & oxalate from roots increases with P deficiency (Vance et al, 2003; Ryan et al, 2001) Table 1 citrate efflux from plant roots Crop Citrate Efflux References (nmol/gFW/h) Rice 155~360 Kirk et al (1999) Wheat 5~185 Ryan et al (2009) White lupin 1656~2373 Roelofs et al (2001) Proteaceae 3600~9000 Roelofs et al (2001)
  • 10. 2.2 Modelling • Only one modeling study to simulate rice P uptake in controlled laboratory experiment • Rice (Kirk et al, 1999) – diffusion, decomposition and P solubilisation of citrate, only a few mm around roots, no zero efflux treatment • No field scale modelling studies so far
  • 11. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 12. 3.1 APSIM enhancements 1. Original Model: crop P uptake is linked to solution P in rooted soil layers 2. Enhancement #1: need to link crop P uptake to solution P and root length density (RLD) 3. Enhancement #2: need to link crop P uptake & solution P to citrate efflux from roots (Fc) P fertilizer P fertilizer broadcast banded Placement effect Dissolution Mineralization Unavailable Rock P Labile P Immobilization Desorption Organic P Adsorption P released by citrate Crop P P in Soil Unavailable Uptake solution Inorganic P
  • 13. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 14. 4.1 Biomass 1. Original Model: crop P uptake 2. Enhancement #1: need to link is linked to solution P in rooted crop P uptake to solution P and soil layers root length density (RLD) 7000 (a) APSIM-SoilP 7000 (b) APSIM-SoilP&RLD Simulated wheat yield (kg/ha) Simulated wheat yield (kg/ha) 6000 y = 1.1256x + 10.593 6000 y = 1.1742x - 259.46 R2 = 0.8891 R2 = 0.896 5000 5000 4000 4000 3000 3000 2000 2000 1000 1000 0 0 0 1000 2000 3000 4000 5000 6000 7000 0 1000 2000 3000 4000 5000 6000 7000 Observed wheat yield (kg/ha) Observed wheat yield (kg/ha) Comparison of simulated and observed wheat grain yields under different levels of P fertiliser inputs at one QLD site and two NSW sites. The modified model performed slightly better than the original one!
  • 15. 4.2 Scenario analysis • Study site: Kingaroy, QLD (1957-2009) • Cropping system: Continuous wheat, rainfed condition • Citrate efficiency = 0.4 • Five levels of soil P sorption capacity: • Low 50, 100 • Medium 200 • High 500 • Very high 1000 • Seven levels of citrate efflux (nmol/gFW/h): 0, 50, 100, 200, 500, 1000, 2000 • 11 levels of P application rates (kg P/ha): 0~200 kg P/ha with increase of 20 kg P/ha.
  • 16. 4.3 Biomass responses: short-term vs long-term First year 1957 Average 1957~2009 (a) Citrate efflux = 0 nmol/gFW/h (b) Citrate efflux = 200 nmol/gFW/h (a) Citrate efflux = 0 nmol/gFW/h (b) Citrate efflux = 200 nmol/gFW/h 20 20 20 20 Biomass (ton/ha) Biomass (ton/ha) Biomass (ton/ha) Biomass (kg/ha) 16 16 16 16 12 12 12 Sorption: 12 Sorption: 50 50 8 8 8 100 8 100 200 200 4 4 4 500 4 500 1000 1000 0 0 0 0 0 40 80 120 160 200 0 40 80 120 160 200 0 40 80 120 160 200 0 40 80 120 160 200 P Rate (kg/ha) P Rate (kg/ha) P Rate (kg/ha) P Rate (kg/ha) (c) Citrate efflux = 1000 nmol/gFW/h (d) Citrate efflux = 2000 nmol/gFW/h (c) Citrate efflux = 1000 nmol/gFW/h (d) Citrate efflux = 2000 nmol/gFW/h 20 20 20 20 Biomass (ton/ha) Biomass (ton/ha) Biomass (ton/ha) Biomass (ton/ha) 16 16 16 16 12 12 12 Sorption: 12 Sorption: Sorption: 50 50 8 8 50 8 100 8 100 100 200 200 4 200 4 500 4 500 4 500 1000 1000 1000 0 0 0 0 0 40 80 120 160 200 0 40 80 120 160 200 0 40 80 120 160 200 0 40 80 120 160 200 P Rate (kg/ha) P Rate (kg/ha) P Rate (kg/ha) P Rate (kg/ha)
  • 17. 4.4 Biomass, yield and inorganic soil P (a) Citrate Efflux = 0 (b) Citrate Efflux = 200 25 25 Biomass or grain yield (ton/ha) Biomass or grain yield (ton/ha) nmol/gFW/h nmol/gFW/h Biomass Biomass 20 Yield 20 Yield 15 15 10 10 5 5 0 0 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 Year Year (c) Citrate Efflux = 0 (d) Citrate Efflux = 200 200 200 Total inorganic soil P (kg/ha) Total inorganic soil P (kg/ha) Layer 1 Layer 1 Layer 2 Layer 2 150 Layer 3 150 Layer 3 100 100 50 50 0 0 1950 1960 1970 1980 1990 2000 2010 1950 1960 1970 1980 1990 2000 2010 Year Year
  • 18. 4.5 P rate required for 90% max biomass P rate required for 90% max biomass (kg/ha) 160 SP50 140 SP100 SP200 120 SP500 SP1000 100 80 60 40 20 0 0 200 400 600 800 1000 1200 1400 1600 1800 2000 Citrate efflux (nmol/gFW/h)
  • 19. 4.6 P recovery (a) P Rate = 20 kg/ha (b) P Rate = 40 kg/ha (c) P Rate = 60 kg/ha 180 180 180 Sorption = 50 160 160 160 Sorption = 200 Sorption = 500 140 140 140 Sorption = 1000 P recovery (%) 120 120 120 100 100 100 80 80 80 60 60 60 40 40 40 20 20 20 0 0 0 0 500 1000 1500 2000 0 500 1000 1500 2000 0 500 1000 1500 2000 Citrate efflux (nm/g/h) Citrate efflux (nm/g/h) Citrate efflux (nm/g/h)
  • 20. Outline of the presentation 1. APSIM and its capability to simulate crop response to soil P 2. Current understanding and available data on citrate release from plant roots and its impact on PUE 3. New model development to enable APSIM to simulate impact of citrate efflux on PUE 4. Simulated response of wheat growth and P uptake to citrate effluxes 5. Summary
  • 21. 5. Summary  Long-term effect is different from short-term effect, due to residual effect of applied P.  Major impact is to increase PUE of applied P fertilisers, because soil P reserve can be depleted in a relatively short time period.  P rate required to achieve maximum crop yield decreases with increasing citrate efflux from the roots.  The impact increases with soil P sorption capacity, decreases with P application rate.

Notes de l'éditeur

  1. For the start year (single year 1957), the results show that: 1) biomass increased with citrate efflux levels at all P application rates, 2) the biomass increase is the highest at medium level of soil P sorption capacity (a=200), and 3) the biomass increase becomes smaller with increased P application rates. If no P fertiliser was applied, a citrate efflux of 200 nmol/gFW/h (similar to the level of Carazinho Wheat) lead to a biomass increase by 78-116% on soils with sorption capacity between 50-200, and by 16 and 31% on soil with sorption capacity of 500 and 1000, respectively (Fig 6a). For long-term (1957-2009) averaged biomass, the difference in biomass responses to P application rates under different citrate effluxes and soil sorption capacities are much smaller than that in a single year (Fig 5). The reduced difference in the response curves at lower rates of P applications is due to the depletion of soil P reserve in the first 15-20 years (Fig 7b, 7d), leading to near zero biomass production thereafter. The reduced difference in the response curves at high rates of P applications is due to the accumulation of P in the soil to reach a point where P supply could meet crop P demand on soils with any sorption capacity. As a result, the potential increase in biomass production at long term increases with citrate efflux levels and becomes increasingly higher on soils with higher sorption capacity (Fig 6b). If no P fertiliser was applied, a citrate efflux of 200 nmol/gFW/h (similar to the level of Carazinho Wheat) lead to a biomass increase by 30%,81%, 216%, 210% on soils with sorption capacity of 50, 100, 200 and 500, respectively (Fig 6b). Fig 7b, 7d show the simulated time courses of biomass, grain yield, and the total inorganic P in the top 30cm layers on a soil with sorption capacity of 200.
  2. The reduced difference in the response curves at lower rates of P applications is due to the depletion of soil P reserve in the first 15-20 years (Fig 7b, 7d), leading to near zero biomass production thereafter. The reduced difference in the response curves at high rates of P applications is due to the accumulation of P in the soil to reach a point where P supply could meet crop P demand on soils with any sorption capacity. As a result, the potential increase in biomass production at long term increases with citrate efflux levels and becomes increasingly higher on soils with higher sorption capacity (Fig 6b). If no P fertiliser was applied, a citrate efflux of 200 nmol/gFW/h (similar to the level of Carazinho Wheat) lead to a biomass increase by 30%,81%, 216%, 210% on soils with sorption capacity of 50, 100, 200 and 500, respectively (Fig 6b). Fig 7b, 7d show the simulated time courses of biomass, grain yield, and the total inorganic P in the top 30cm layers on a soil with sorption capacity of 200.
  3. P application rates required to achieve 90% of the potential biomass under different levels of citrate effluxes on a given soil (Fig 9). With increased level of citrate efflux from the roots, the P application rates required to achieve 90% of the potential biomass is significantly decreased, particularly on soils with high P sorption capacity. Fig 8 shows the simulated impact of citrate efflux on average P recovery (calculated as crop P uptake divided by P application rate) on different soils at three P application rates. The results show that: 1) P recovery increases with citrate efflux till certain efflux rate is reached, 2) the increase in P recovery is more significant on higher sorption soils, and 3) the increase in P recovery becomes much smaller with increased P application rates.
  4. P application rates required to achieve 90% of the potential biomass under different levels of citrate effluxes on a given soil (Fig 9). With increased level of citrate efflux from the roots, the P application rates required to achieve 90% of the potential biomass is significantly decreased, particularly on soils with high P sorption capacity. Fig 8 shows the simulated impact of citrate efflux on average P recovery (calculated as crop P uptake divided by P application rate) on different soils at three P application rates. The results show that: 1) P recovery increases with citrate efflux till certain efflux rate is reached, 2) the increase in P recovery is more significant on higher sorption soils, and 3) the increase in P recovery becomes much smaller with increased P application rates.