Zed Rengel

Univ Western Australia Perth

Ukupna kolicina nitrata u tlu

- u 1.5-m profilu tla - u 1.5-m profilu tla -

Lupina Kolicina nitrata 21 srpnja 27 rujna 29 studenog

psenica

pasnjak

------------------------------ kg/ha -----------------------------

74 33 44

46 13 24

35 4 17

Ukupna kolicina ispranog nitrata, kg/ha kg/ha

- dolje do 1 m u profilu tla - dolje do 1 m u profilu tla -

21 lipnja 26 rujna

24

20

Dolling et al. 1995

Heterogenost concentracije nitrata u tlu

- blok 5 m 5 m 5 m, mali blok od 25 cm x 25 cm x 25 cm mg/kg

mg/kg

Nakon 41 dana rasta

Lupinus pilosus

Lupinus angustifolius

Kompenzacija u primanju nitrata, L. angustifolius

- rata primanja plitkim korijenima opskrbljenih visokom koncentracijom nitrata -

0.3

Rata primanja nitrata (mol m-2 s-1)

L. angustifolius

L. pilosus

Uniformno visoka opskrba nitratima (1500 / 1500 M) Razlicita opskrba nitratima (1500 / 250 M)

*
0.2

0.1

0.0

Plitki korijeni

Dunbabin, Rengel & Diggle 2001 Aust J Agric Res 52, 505-512.

Kompenzacija u primanju nitrata, L. angustifolius

- rata primanja dubokim korijenima opskrbljenih visokom koncentracijom nitrata -

Rata primanja nitrata (mol m-2 s-1)

0.3

L. angustifolius

L. pilosus

Uniformno visoka opskrba nitratima (1500 / 1500 M) Razlicita opskrba nitratima (250 / 1500 M)

**
0.2

0.1

0.0

Duboki korijeni

Dunbabin, Rengel & Diggle 2001 Aust J Agric Res 52, 505-512.

Kompenzacija u primanju nitrata, L. angustifolius

- rata primanja plitkim korijenima opskrbljenih niskom koncentracijom nitrata -

Rata primanja nitrata (mol m-2 s-1)

0.06

L. angustifolius

L. pilosus

0.05

Uniformno niska opskrba nitratima (250 / 250 M) Razlicita opskrba nitratima (250 / 1500 M)

0.04

0.03

0.02

0.01

0.00

Plitki korijeni
Dunbabin, Rengel & Diggle 2001 Aust J Agric Res 52, 505-512.

Kompenzacija rasta korijena, L. pilosus

U odnosu na bilje opskrbljene uniformnin nitratima
Duboki korijeni
2.2

Plitki korijeni

Visoka opskrbljenost plitkih korijena nitratima, niska opskrbljenost dubokih korijena nitratima

L. angustifolius
2

L. pilosus

1.8

1.6

Smanjenje Povecanje

1.4

1.2

0.8

Rata rasta

Broj novih laterala

Rata rasta

Broj novih laterala

Biljke su bile stare 21 dan na pocetku tretmana, Rast nakon 9 dana

Dunbabin, Rengel & Diggle 2001 Aust J Agric Res 52, 495-503.

Kratak opis modela Komponenta modela za reakciju korijena Koristenje modela u simulaciji poljskih pokusa Koristenje modela za istrazivanje rasta korijena i primanja nitrata u heterogenom tlu

Modul za distribuciju resursa

Kontrolira rast korijena i primanja hraniva na bazi isplativosti ulaganja

Modul za vodu
Drenaza, redistribucija, evaporacija, primanje

ROOTMAP 3-D model arhitekture korijena

Modul za hraniva
Ispiranje, masovni tok, difuzija, primanje

Model koji simulira reakciju pojedinacnih korijena, kao i cijelog korijenovog sistema, u odnosu na opskrbu asimilatima koja varira u prostoru i vremenu
Dunbabin, Diggle, Rengel & Van Hughten 2002. Plant Soil 239, 19-38.

Potraznja na nivou cijele biljke

(bazirana na resursima koji su na raspolaganju za koristenje u biljci)

Lokalni signali koji pokazuju potencijalnu opskrbu resursima

Opskrba asimilatima za rast i odrzavanje, ovisno o lokalnoj akviziciji resursa i prioritetima

Lokalni interni zahtjevi za resursima

Lokalni opskrbni sistem

(korijeni, korijenove kvrzice, stabljika)

Vanjska opskrba resursima (npr. nitrati, voda)

Osnovna metodologija modela

Stvarna reakcija kroz primanje hraniva ili rast biljke (ovisno o resursima) bazirana na opskrbi i potrebama
Dunbabin, Diggle, Rengel & Van Hughten 2002. Plant Soil 239, 19-38.

Kratak opis modela Komponenta modela za reakciju korijena Koristenje modela u simulaciji poljskih pokusa Koristenje modela za istrazivanje rasta korijena i primanja nitrata u heterogenom tlu

nitrat = ljubicasta

voda = svjetlo plava

nitrati + voda

lokalni volumen s visokim nitratima

Lokalni volumen suhog tla

Lokalni volumen s visokim nitratima

Lokalni volumen suhog tla

Pocetni profil tla

Ne-reagirajuci korijenov sistem koji raste na osnovu predefiniranih pravila

Korijenovi sistem koji reagira na lokalni okolis

Lokalni volumen s visokim nitratima

Lokalni volumen suhog tla

Ispiranje nitrata u profilu tla nakon uzastopnih kisa

Kratak opis modela Komponenta modela za reakciju korijena Koristenje modela u simulaciji poljskih pokusa Koristenje modela za istrazivanje rasta korijena i primanja nitrata u heterogenom tlu

Modelirani podaci (cm/cm3)

3.5 3 2.5 2 1.5 1 0.5 0 0.0

Gustoca korijena

Korelacija izmedju
y = 0.97x

r2 = 0.90

modeliranih i poljskih podataka

L. angustifolius je rasla na polju 1995 i 1996

0.5

1.0

1.5

2.0

Poljski pokusi
Modelirani podaci (kg nitrate/ha)
14 12 10 8 6 4 2 0 0 2 4 6 8

3.0 (cm/cm3)

2.5

3.5

Sadrzaj nitrata u tlu

y = 0.96x

r2 = 0.86

10

12

14
Dunbabin, Diggle & Rengel 2002. Plant Soil 239, 39-54.

Poljski pokusi (kg nitrate/ha)

Rast korijena simuliran modelom

- L. angustifolius (rasla na polju 1996) -

Gustoca korijena (cm/cm3)

0
0

2
0

4
0

20

20

20

40

40

40

Dubina (cm)

60

60

60

Field Data
80 80 80

Modelled data
100 100 100

120

120

120

79 DAS
140 DAS=dana nakon sijanja 140

108 DAS
140

146 DAS
Dunbabin, Diggle & Rengel 2002. Plant Soil 239, 39-54.

Nitratni profili simulirani modelom

- L. angustifolius (rasla na polju) -

Sadrzaj nitrata (kg NO3--N/ha)

0 0 5 10 15 0 0 5 10 15 0 0 5 10 15

20

20

20

40

Poljski podaci Modelirani podaci

40

40

Dubina (cm)

60

60

60

80

80

80

100

100

100

120

120

120

86 DAS
140 140

106 DAS
140

217 DAS

DAS=dani nakon sijanja

Dunbabin, Diggle & Rengel 2002. Plant Soil 239, 39-54.

Profil vode tla simularog modelom

- L. angustifolius (rasla na polju 1996) 0.16

0 - 10 cm dubina

Poljski podaci Modelirani podaci

0.12 0.08 0.04

25 75 125 175 225

Kolicina vode u tlu (v/v)

0.16 0.12 0.08 0.04 0.16 0.12 0.08 0.04 25 75 125

30 - 50 cm dubina

25

75

125

175

225

120 - 150 cm dubina

175

225

Dunbabin, Diggle & Rengel 2002. Plant Soil 239, 39-54.

Dani nakon sijanja

Nitratni budget simuliran modelom

- u 1.5 m simuliranog profila -

1995 Izvor/sink za nitrate pocetni Lupina: primanje

1996

------ kg nitrata / ha ------

32 -21

57 -26 -28 69 -26 (-23) 46

Capeweed: primanje -0 mineralizirano ispiranje zavrsno 60 -32 (-35) 39

Izmjereno i objavljeno: Anderson et al. 1988

Dunbabin, Diggle & Rengel 2002. Plant Soil 239, 39-54.

Kratak opis modela Komponenta modela za reakciju korijena Koristenje modela u simulaciji poljskih pokusa Koristenje modela za istrazivanje rasta korijena i primanja nitrata u heterogenom tlu

Riblja kost

razgranati

Simulirana efikasnost primanja nitrata za razne arhitekture korijena

- plasticitet primanje nitrata ili plasticitet rasta korijena je bio onemogucen -

Riblja kost
Plasticnost rasta korijena
Staticna opskrba nitratima

razgranat

Plasticnost primanja nitrata Plasticnost rasta korijena

Dinamicna opskrba nitratima

Plasticnost primanja nitrata

1 1.1 1.2 1.3 1.4 1.5 1.6 1.7 1 1.1 1.2 1.3 1.4 1.5 1.6 1.7

Relativna efikasnost primanja nitrata

Dunbabin, Rengel & Diggle (2004). Func. Ecol. 18, 204-211

Modelirane arhitekture korijena

(s istim volumenom korijena) Razne arhitekture su identicne po troskovima (tj. ista kolicina asimilata je koristena) L. angustifolius riblja kost (fina) L. pilosus (duboka) L. pilosus (plitka)

Riblja kost (fina) 0 0.2 0.4

highly branched

Dubina (m)

0.6 0.8 1.0 1.2 1.4

Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Izmjerene padaline u Moora 1995 (Anderson et al., 1998)

40 30 20 10 0 0 7 14 21 28 35 42 49 56 63 70 77 84 91 98 105

Ista ukupna kolicina kao i u primjeru abore, ali pljuskovi su odgodjeni za kasnije u sezoni
40 30 20 10 0 0 7 14 21 28 35 42 49 56 63 70 77 84 91 98 105

Dani nakon sijanja

Primanje nitrata raznim arhitekturama korijena

Totalna primljena kolicina
L=pljuskovi kasnije u sezoni E=rani pljuskovi u sezoni

29

-1

26

23

Primanje nitrata (L)

20

Primanje nitrata (E)

17

14

11 Riblja kost debela

Riblja kost fina

L. angustifolius L. angustifolius L. pilosus debela fina duboka

L. pilosus plitka

Highly branched

Smanjenje dubine zakorijenjivanja, povecanje gustoce korijena

108 dana nakon sijanja Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Ispiranje nitrata - razne arhitekture korijena Totalna isprana kolicina

L=pljuskovi kasnije u sezoni E=rani pljuskovi u sezoni

29

26

-1
23

Isprani nitrati (L)

20

Isprani nitrati (E)

17

14

11 Riblja kost debela

Riblja kost fina

L. angustifolius L. angustifolius L. pilosus debela fina duboka

L. pilosus plitka

Highly branched

Smanjenje dubine zakorijenjivanja, povecanje gustoce korijena

108 dana nakon sijanja Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Primanje nitrata raznim arhitekturama korijena

Rata primanja po povrsini korijena

-1

L. pilosus (plitak)
-2

L. pilosus (dubok)

0 0
E=rani pljuskovi u sezoni (izmjereno u Anderson et al 1988)

20

40

60

80

100

Dani nakon sijanja

Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Primanje nitrata raznim arhitekturama korijena

Rata primanja po povrsini korijena

-2

-1

6 Riblja kost (fina) Highly branched 4

0 0
E=rani pljuskovi u sezoni (izmjereno u Anderson et al 1988)

20

40

60

80

100

Dani nakon sijanja

Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Zakljucci
Sistemi opskrbe/potreba (supply/demand) se mogu koristiti za simuliranje reakcija korijena na okolis Rast korijena, primanje nitrata i sadrzaj vode su uspjesno modelirani u sistemu lupina-tlo Kapacitet simuliranja reagirajuceg korijenovog sistema omogucava studiranje interakcija izmedju korijena i heterogenog okolisa Uspjeh u primanju nitrata je balans izmedju brzine akvizicije i velicine ukupnog volumena eksploatiranog tla s jedne strane I brzine ispiranja nitrata s druge

Acknowledgements

Dr Vanessa Dunbabin (UWA, now Univ Tasmania) Dr Art Diggle (State Dept of Agric)

Root length density and cumulative nitrate leached after the main rainfall event
- four L. angustifolius plants grown in the field in 1996 -

2-2.5 1.5-2 1-1.5 0.5-1 0-0.5

*
Root length density (cm/cm3)

12-15 9-12 6-9 3-6

0-3

42 days after sowing

92 days after sowing

50-60 40-50 30-40 20-30 10-20 0-10

*
Nitrate leached (kgNO3--N/ha)

50-60 40-50 30-40 20-30 10-20

0-10

Root growth as influenced by the nitrate supply

- first order laterals -

L. angustifolius

L. pilosus
35 30 25 20 15 10 5 0

Root growth (cm day-1)

35 30 25 20 15 10 5 0 750/750 250/750

shallow roots deep roots

250/250

750/250

750/750

250/750

250/250

750/250

Nitrate concentration in upper/lower compartment, M

21-d-old plants at t=0, growth after 9 days. Dunbabin et al. 2001. Aust J Agric Res 52, 495.

Root growth as influenced by the nitrate supply

- second order laterals -

L. angustifolius
16 16 shallow roots deep roots 14 12 10 8 6 4 2 0 750/750 250/750 250/250 750/250 750/750

L. pilosus

Root growth (cm day-1)

14 12 10 8 6 4 2 0

250/750

250/250

750/250

Nitrate concentration in upper/lower compartment, M

21-d-old plants at t=0, growth after 9 days.

Dunbabin et al. 2001. Aust J Agric Res 52, 495.

Modelling root growth of L. pilosus

- first order laterals 40 35 upper root system

Experimental data Modelled data lower root system

Root growth (cm day-1)

30 25 20 15 10 5 0
250/750 250/250 750/250 750/750 750/750

250/750

250/250

750/250

Nitrate supply in upper/lower compartment (M)

Dunbabin et al. 2002. Plant Soil 239, 19.

Modelling nitrate uptake by L. angustifolius

- in the high nitrate patch when split-nitrate supply Experimental data Modelled data

0.3

Nitrate uptake rate (mol m-2 s-1)

0.25 0.2 0.15 0.1 0.05 0

750/750

upper root system

lower root system

750/250 1500/1500 1500/250

750/750

250/750 1500/1500 250/1500

Nitrate supply in upper/lower compartment (M)

Dunbabin et al. 2002. Plant Soil 239, 39.

Parameters used in the model for simulation of lupin root growth and nutrient uptake in nutrient solution (determined by parameter fitting and from experimental data) for three orders of branching: tap root (0), primary laterals (1), secondary laterals (2). The branch lag time = the time lag
before a branch will grow (eg. a primary lateral) from a node on the previous branching order (eg. the tap root).
Parameter description root radius 0 root radius 1 root radius 2 unit N growth rate 0 unit N growth rate 1 unit N growth rate 2
* * * * * * * * * *

Symbol a0 a1 a2 gN0 gN1 gN2 gnN0 gnN1 gnN2 bN1 bN2 bnN1 bnN2 Tbl0 Tbl1 D fix nodm

L. angustifolius 1.0 0.5 0.3 0.08 0.12 0.14 3.5 0.77 0.54 5.9 2.1 0.42 0.42 192 528 90 0.25 0.05 0.075 9 56 1.6 5.3 0.119 2.55*10
-8

L. pilosus 0.8 0.4 0.2 0.08 0.14 0.96 4.2 0.73 0.73 3.5 3.5 1.2 0.42 216 291 90 0.25 0.05 0.075 9 265 1.1 2.6 0.115 1.89*10
-8

Units mm mm mm m mol s m mol s m mol s m mol s m mol s m mol s

-1 -1 -1 -1 -1 -1 -1 -1 -1 -1 -1 -1

unit nN growth rate 0 unit nN growth rate 1 unit nN growth rate 2

unit N branch density 1 unit N branch density 2

branches m -1 mol -1 branches m -1 mol -1 branches m -1 mol -1 branches m -1 mol -1

unit nN branch density 1 unit nN branch density 2 branch lag time 0 branch lag time 1 initial branch angle deflection index unit N fix
* * *

h h deg unitless mol nodule s mol nodule nodule mol mol unitless nmol m s mol m
3 -2 -3 -1 -2 -1 -1 -1 -1 -1

unit nodule cost

unit nodule growth rate total seed reserve

nodg (N+nN)see
d

N down regulation factor max. N net influx rate kinetic constant poten. transpiration rate

Fmax Km Tmax

m m s

Parameters used in the model for simulation of lupin root growth and nutrient uptake in soil [4 orders of branching: tap root (0), primary laterals (1), secondary laterals (2) and tertiary laterals (3)]. The branch lag time represents the time lag before a branch will grow (eg. a primary lateral) from a node on the previous branching order (eg. the tap root).
Parameter description Symbol L. angustifolius Units

root radius 0 root radius 1 root radius 2 root radius 3 * unit N growth rate 0 * unit N growth rate 1 * unit N growth rate 2 * unit N growth rate 3 * unit nN growth rate 0 * unit nN growth rate 1 * unit nN growth rate 2 * unit nN growth rate 3 branch spacing 0 branch spacing 1 branch spacing 2 branch spacing 3 branch lag time 0 branch lag time 1 branch lag time 2 initial branch angle Field data were used only deflection* index unit N fix * to initialise the unit nodule cost * environmental variables unit nodule growth rate total seed reserve required by the model,

* represents those parameters previously calibrated in the model through simulating nutrient solution experiments.

a0 a1 a2 a3 gN0 gN1 gN2 gN3 gnN0 gnN1 gnN2 gnN3 bN0 bN1 bnN0 bnN1 Tbl0 Tbl1 Tbl1 D fix nodm nodg (N+nN)se
ed

1.0 0.5 0.3 0.2 0.08 0.12 0.14 0.14 3.5 0.77 0.54 0.54 8 8 5 5 192 528 528 90 0.25 0.05 0.075 9 56 5.3 0.119 -8 2.55*10 30 1.9*10
-9

mm mm mm mm -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s -1 -1 m mol s mm mm mm mm h h h deg unitless -1 -1 mol nodule s -1 mol nodule -1 nodule mol mol nmol m s -3 mol m 3 -2 -1 m m s mm m s
2 -1 -2 -1

eg. soil bulk density, initial soil water contents, drained upper limit and wilting point parameters and rainfall

maximum N net influx rate kinetic constant potential transpiration rate displacement dependent dispersivity nitrate diffusion coefficient

Fmax Km Tmax D

Model Output The intensity of the blue background shows the water content and the black dots represent the nitrate content in soil

Model Output L. angustifolius plants growing in sandy soil The intensity of the blue background shows the water content and the black dots represent the nitrate content in soil

Model Output L. angustifolius plants growing in sandy soil The intensity of the blue background shows the water content and the intensity of pink colour (also black dots) dots represent the nitrate content in soil

initial soil profile

root systems responding to their local environment

local high nitrate patch

local dry patch

nitrate leaching through soil profile after successive rain events

- root elongation rate - branching density - duration of apical non-branching

- soil resistance - time of appearance of each branching order - axis initiation parameters - direction of branching

Root branching
o ld dir ec tio n

tion

new c dire

= angle of deflection = orientation of deflection

Extensible tree:
data structure to store information on root architecture

node pointers

null

If the influx, In, is known, the driving force for the diffusive flux can be calculated

1 1 CL = CL CL0 = 4 DL f 1 r0 2 RLv
In

r0 2

RLv

CL = average soil solution concentration of the bulk soil CL0 = solution concentration at the root surface DL = diffusion coefficient in water

= volumetric soil water content

f = tortuosity factor r0 = root radius RLv = root length density

Transport equation
incorporating diffusion and mass flow

C = - 1 rD b C + v r C b t r r r
L L e 0 0 L
b = buffer power CL = soil solution concentration in the bulk soil r = radial distance from the root axis De = effective diffusion coefficient in soil v0 = water flux across the soil-root interface r0 = root radius

Uptake of nitrate by various root architectures

L=high leaching events late in the season E=high leaching events early in the season

29

-1

26

23

nitrate uptake (L) nitrate leached (L)

20

nitrate uptake (E) nitrate leached (E)

17

14

11 herringbone coarse

herringbone L. angustifolius L. angustifolius L. pilosus fine coarse fine deep decreasing rooting depth, increasing rooting density

L. pilosus shallow

dichotomous

at 108 days after sowing

Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Static Environment Processes

Bulk Density Module Manager Module

Wilting Point

Model structure
Nitrate Module
Dynamic Processes

External Control Module

ENGINE

Weather Module

Clock Plant Module Graphics Display Module

Plant1

Engine interacts with modules, with modelling activities synchronised by the clock.

Plant2 Plant3

Scoreboard

The scoreboard represents the simulation volume, containing all 3-D parameter values.

1 2 3 4 5 6 7

A B C DE F

Scoreboard Display Module (Spreadsheet-like)

Uptake of nitrate by various root architectures

Uptake rate per surface area

-1

-2

L. pilosus (shallow) 6 L. pilosus (deep) . Herringbone (fine) x dichotomous 4

0 0 20 40 60 80 100

E=high leaching events early in the season (as measured by Anderson et al 1988)

Dunbabin, Diggle & Rengel (2003). Plant Cell Environ. 26, 835-844

Nitrate leaching from topsoil causes topsoil acidification

Change in soil pH
-0.6 0 -0.4 -0.2 0.0 0.2 -0.6 -0.4 -0.2 0.0 0.2

20

Depth (cm)

40

60

80

Lupinus angustifolius

Trifolium subterraneum
Tang, Raphael, Rengel & Bowden 2001 Aust J Soil Res 38, 837-849.

Simulating nitrate uptake efficiency of root architectures

Relative nitrate uptake efficiency

(dynamic/static nitrate supply)

herringbone
1.8
(0.11)

highly branched
0.78
(0.21)

Averaged over 30 simulations and 5 replicates

Dunbabin, Rengel & Diggle (2003). In: Innovative Soil-Plant Systems for Sustainable Agricultural Practices, pp.2-16.

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