Ecosystems haveEcosystems have complex dynamicscomplex dynamics

• By Sven Erik Jørgensen• By Sven Erik Jørgensen• Environmental Chemistry• University Park 2• 2100 Copenhagen Øg• Denmark• sej@farma ku dk or• sej@farma.ku.dk or

msijapan@hotmail.comHome page: www ecologicalmodel net• Home page: www.ecologicalmodel.net

Outlines

• Ecotones• Two types of ecology• Properties of ecosystems included the p y

application of the thermodynamic laws on ecosystemsy

• Description of ecosystem development by the use of gradients, as complex,the use of gradients, as complex, adaptive, hierarchical, self-organizing systemssystems

Importance of ecotones

• Buffer zones between two ecosystems - the transition zones are for ecosystems as theare for ecosystems as the membrane for cells

• Often high biodiversity, representing both the adjacentrepresenting both the adjacent ecosystems

Ecosystems have not clear borders - the transition zone is

d t d tdenoted an ecotone

Two types of ecology:

• Reductionistic ecology: organism or populations are examined one by one -provide data about organisms p gand populations

• Holistic ecology named system• Holistic ecology named system ecology: examine entire ecosystems - consider many interacting processes simultaneously - data p yfrom reductionistic ecology is often needed

Properties of ecosystems:• Complex: middle number systems• Adaptable• Hierarchical organization • Self-organizingSelf organizing • History• A common biochemistry for all organisms• A common biochemistry for all organisms,

which implies the concept of one or more limiting factorsg ac o s

• Recycling

In addition ecosystemsIn addition, ecosystems follow:follow:

• The first law of thermodynamics –the conservation of matter and energygy

• The second law of thermodynamicsTh t l ti f D i ‘ th t• The translation of Darwin ‘s theory to

Thermodynamics, sometimes denotedThermodynamics, sometimes denoted ELT.

Structure Function

Interactions and Dynamics

Organization

Gradients

• Structure has gradients• Pattern has gradients• Energy forms that can perform workEnergy forms that can perform work

have gradients: pressure energy: pressure; electrical energy: voltage;pressure; electrical energy: voltage; chemical energy chemical potential, heat energy: temperature potentialheat energy: temperature, potential energy: height and so on

What is exergy?

• Exergy is work capacity - energy that can d k It th f b f d thdo work. It can therefore be found as the gradient (= difference in potential)x

t i d i t d d t thextensive descriptor , dependent on the energy form, for instance

• Chemical energy= (µ1-µ2) N or • Pressure energy= (p1-p2)(-V)gy (p1 p2)( )• Potential energy= (h1 - h2) m g• Electrical energy= (V V ) Q• Electrical energy= (V1- V2) Q

Air Pressure Gradients(Potential)(Potential)

Wi dWind(Flux)

Wind Utilisation

Gradients in Ecoystems

G di t (X)Gradients (X)Provide the

P t ti l (V)Potentials (V)for all

Fl (J)Flows (J)

G di tGradientDissipation

i R l t dis Regulatedby Site-SpecificR sist n s (R)Resistences (R)

ElevationGradientsGradients

Forest CatenaCatena

lAgricultureCatena

What are Gradients?Figure from Schleuss 1992, Dilly 1994

Gradientsare

• ConcentrationProfiles;

High Values

Profiles;

• PatternsLow Valuesof ParameterDistribution

Low Values

Distance

AgricultureCatena

What are Gradients?

GradientsExhibitP i lPotentialsfor their DegradationDegradationbyFlows

What are Gradients?

Structural GradientsBi ti G di t- Biotic Gradients

Structural Gradients- Biotic Gradients

LakeBelau

Terrestrial-AquaticAquaticEcotone

Phragmites australis

30

35

gRubus idaeusLysimachia spp.Carex spp.

15

20

25

L Chl

(n/m

)2

5

10

15

P R

C

Anza

h

5 7,5 10 12,5 15 17,5 200

Entfernung vom Ufer (m)

Data after Dittert (1994)

Distributionof Someof Some

Plant Species

Biocenotic Heterogeneity

100

Variations-Koeffizient

Dominanten-Identität

&

Data from

U.Ir

40

60

80

Dom

inantenide

& Variationskoe

rmler and J.Sc

A k Uf id H id B h ld Uf b h E l b h0

20

40

entität

effizient

hrautzer

Acker Uferweide Hangweide Buchenwald Uferbrache Erlenbruch

DistributionPatternPattern

of Species „Yellow“

Low Heterogeneity High HeterogeneityLow HeterogeneityNo Gradients

High HeterogeneityVisible Gradients

2 What is complexity?2. What is complexity?the minimal amount of information that is needed to describe a system's …the minimal amount of information that is needed to describe a system s structure (Kolmorogow, 1965)

...the observer's ignorance of a system (Salthe, 1993)...the observer s ignorance of a system (Salthe, 993)

Rheingau Cairns, Australia

2. What is complexity?… something complicated, that is difficult to understand and hard to explain

(Hornby, 1974) h l i l d h f bj (B 1988)… the logical depth of an object (Bennett, 1988)

2. What is complexity?… the consequence of multiple interrelationships between the elements of … the consequence of multiple interrelationships between the elements of

systems, increasing with the number of interacting units and the intensity of their interactions (Nicolis, 1986)

… a property of an entity which makes long messages necessary to describe the system and which demands long time periods for their development (Salthe, 1993)

Complexity rises withComplexity rises with…

… the number of elements in a system the differences between the elements… the differences between the elements

… the number of relations between the elements the differences between the relations… the differences between the relations

… the non-linearity of the interactions

… information and connectedness

Complexity can be indicated byComplexity can be indicated by…

… the number of elements in a system the differences between the elements… the differences between the elements

… the number of relations between the elements the differences between the relations… the differences between the relations

… the non-linearity of the interactions

… information and connectedness

… the length of the shortest possible programme to describe a systemto describe a system

Complexity influences control…

Input signal Output signal

Ecosystem

Complexity influences control…

… While characterising complexity, the role of distributed control in ecosystems, in contrast with human systems, was discussed.

Control in ecosystem is network driven being diffuse and Control in ecosystem is network driven, being diffuse and distributed (Schramski et al, 2004). This decentralized control structure promotes ecosystem complexity as an control structure promotes ecosystem complexity as an attribute useful in buffering inputs by smoothing external uneven signals. As a result, ecosystems receiving very complex signals from the environment, are able to smoothen the respective output.Thus the high internal ecosystemic complexity is able to Thus, the high internal ecosystemic complexity is able to “de-complexify” complicated input signals, buffering them and producing un-complex, resilient outputs…and producing un complex, resilient outputs…

…from a workshop protocol from Slowenia

…complexity can rise throughout ecosystem development

…one task of systems sciences is to reduce complexity

…understanding nature as a hierarchical entity can be helpful

Werden – developing

Sein - being

Vergehen - passing

Giovanni Segantini: Alpen-Triptychon: Sein, 1898-1899, Öl auf Leinwand, 235 × 400 cm, St. Moritz, Segantini-Museum, Land: Italien, Stil: Symbolismus.

4. What is „Adaptation“?

- In general: Adjustment to a changing environment.

- Evolutionary: An alteration or adjustment in structure or habits often hereditary by which a species or individual habits, often hereditary, by which a species or individual improves its condition in relationship to its environment.

S t b d Ch i b h i f t ( - Systems based: Change in behavior of a system (or person or group) in response to new or modified surroundings.

- Ecosystem based: Self-organized modification of structural and functional features as a reaction on changes in

nst ints constraints.

Structurized Systems StateStructurized Systems Statewith Concentration Gradients

Structurized Systems StateStructurized Systems Statewith Concentration Gradients

Structurized Systems StateStructurized Systems Statewith Concentration Gradients

Diffusion, Dispersion,Dissipation

Thermodynamic Equilibrium(no Gradients)(no Gradients)

Structurized Systems StateStructurized Systems Statewith Concentration Gradients

Diffusion, Dispersion,Dissipation

Thermodynamic Equilibrium(no Gradients)(no Gradients)

DissipativeDissipativeSelf - Organization

Structurized Systems Statewith New Gradients

Self - Organization:g

SSpontaneous

Creation of

Macroskopic

Structures from

Microskopicc os op c

Disorder

Gradient Formation

Principles of Self - Organization:

nt m

ironm

en

Sys

tem

Env

Ope

n

Principles of Self - Organization:

Exergy Import

nt m

ironm

en

Sys

tem

Exergy Degradation

Env

Ope

n

Entropy Export

Principles of Self - Organization:

Exergy Import Convertible Energye.g. Radiation

CO2-Input

nt m

Energy Transformatione.g. Physiological Processes

G thironm

en

Sys

tem

Exergy Degradation

GrowthRespirationE

nv

Ope

n

Entropy Export Energy Outpute.g. Heat

CO2-Output

Principles of Self - Organization:

Exergy Import Convertible Energye.g. Radiation

CO2-Input

nt m

Energy Transformatione.g. Physiological Processes

G thironm

en

Sys

tem

Exergy Degradation

GrowthRespirationE

nv

Ope

n

Entropy Export Energy Outpute.g. Heat

CO2-Output

Gradient Formation

Principles of Self - Organization:

Exergy Import Systems State Far from Equilibrium

Internal Control / Regulation

Symmetry Breaking OrganizationExergy Degradation Cooperativity of Subsystems

Th d i O f S t

Symmetry Breaking Organization

Thermodynamic Openess of System

Constraints in Hierarchies

Entropy ExportMeta Stability after Small Impulses

Fluctuations in Phase Transitions

Gradient Formation

Historicity and Irreversibility

Hierarchical properties:Hierarchical properties:le

1 MyL

og T

ime

scal

1 ky

Ecosystem dyna- mics

Ecosphere dynamics

Climate

1 year

Population dyna- mics

Pesticide Use

changeEffluent

1 day

Microorga- nism popu- lations

Macroorgan- Oil spills

1h

1 day gism physiolo- gy

Microorgan- ism physiology

spills

µm mm m km Mm

Log Spatial scale

1s

ism physiology

Disturbances: scale and recovery

Vitousek, P.M. 1994. Ecology 75: 1861-1876.

Hierarchy of regulationHierarchy of regulation mechanisms

Level Rate regulated by: Illustration phytoplanktonphytoplankton

_________________________________________________

1 Concentrations Uptake of nutrients2 Needs Uptake of nutrients2 Needs Uptake of nutrients3 Solar radiation Chlorophyll a conc4 Biochem Adaptation Size selection4 Biochem. Adaptation Size selection5 Survival Shift of species6 Survival ecol Netw Shift of ecological6 Survival ecol. Netw. Shift of ecological

netw.7 Mutation/ sexual rec Genome is

External factors Forcing functions

Ecosystem structureEcosystem structure at time t

N biNew recombina- tions of genes / mutations

Gene pool Selection

E t t t

Gene pool

Ecosystem structure at time t +1

S ti l E t t Hi hHolon Level (+1) Spatial Extent: High

Minor Interactions

Holon Level (0)Multiple Interactions

Holon Level Spatial Extent: Small

(-1)

Hierarchies and the problems of scale

S ti l E t t Hi hHolon Level (+1) Spatial Extent: HighFrequencies: Low

Minor Interactions

Holon Level (0)Multiple Interactions

Holon Level Spatial Extent: Small

(-1) Frequencies: High

Hierarchies and the problems of scale

S ti l E t t Hi hHolon Level (+1) Spatial Extent: HighFrequencies: Low

Minor Interactions

Holon Level (0)Multiple Interactions

Filter

Constraints

Holon Level Spatial Extent: Small

(-1) Frequencies: High

Hierarchies and the problems of scale

Geological featuresConstraints

ale

Constraints

Soil features

Vegetation features

pora

l sc

a

Microbiota features

Temp

Biotic potential

Spatial scaleSpatial scale

Region

Watershed

Landscape

Ecosystemy

SpatialExtent

Region

Watershed

Landscape

Ecosystemy

Time for Change

SpatialExtent

Region Broad

extents

SlowprocessesConstraints WatershedpConstraints

LandscapeSmall

Extents Potentials

EcosystemRapid

processesy p

Hypotheses for (natural) hierarchical systems in steady state

Time for Change

SpatialExtent

Region

Pressure:Nitrogen

fertilisationWatershed

fertilisation

Landscape

Ecosystemy

Time for Change

SpatialExtent

Region

Pressure:Nitrogen

fertilisation

Nitrate concentrationin drinking water

fertilisationNitrate concentration

in river waterWatershed

Nitrate concentrationin groundwater

Landscape

Nitrate concentration

g

State:Ecosystemin soil solution Nitrate

concentrations

y

Time for Change

SpatialExtent Ecological

Region

gimpact

Quality of drinking water

Eutrophication of freshwater

Watershed

Landscape

Nitrate discharge in groundwater

Ecosystem

Nitrate leaching

y

Time for Change

SpatialExtent Ecological Economic

Region

gimpact impact

Quality of drinking water

Watershed Human health

Eutrophication of freshwater

LandscapeFishing reduction

Ecosystem Water treatment costs

Nitrate discharge in groundwater

Nitrate leaching

y

Fertiliser loss (capital loss)

Time for Change

SpatialExtent Ecological Economic

Region

gimpact impact

S i t lQuality of

drinking water

Societal perception

Watershed Human health

Eutrophication of freshwater

Decision Process

LandscapeFishing reduction

Ecosystem Water treatment costs

Nitrate discharge in groundwater

Nitrate leaching

y

Fertiliser loss (capital loss)

Time for Change

SpatialExtent Targets of

regional Drivers

Region Regional

regional development

Drivers

Water framework directive

Regional agricultural

policy targets

Watershed

Economic programmes

Social state in rural

Settlement

Landscapefor rural areas

Settlement

Economic situation

Ecosystem

Information, consultation and training

of the farmer

y

Local restrictions or nitrogen taxes

Response

Time for Change

g

3

varia

ble 2

Stat

e v

1

Forcing function

Ecosystems have history:

• They response to external factors (f i f ti )(forcing functions)

• The same combination of numerous forcing functions will never appear again

• All the biological components areAll the biological components are currently changing their properties due to biochemical and biological adaptationto biochemical and biological adaptation - and shifts in species composition

(Eutrophication, (measured by phyto plankton conc. or primary production)

Range, where biomanipu- lation can be applied

Range, where biomanipulation

Range, where biomanipula- tion not is

li d

Faster reco- very obtain- ed by bio- biomanipulation

hardly can be applied

applied ed by bio maipula- tion

Nutrient concentration

Two structures areTwo structures are competing:competing:

• Below 60 µg P/l zooplankton and carnivorous fish dominance Above 125µg P/l phytoplankton andAbove 125µg P/l phytoplankton and planktivorous fishBetween 60 and 125µg P/l both structures are possible dependentstructures are possible dependent on the history

Shallow lakes have twoShallow lakes have two competing structures, too:competing structures, too:

• Below about100 µg P /l submergedBelow about100 µg P /l submerged vegetation is dominant

/• Above about 250µg P/l phytoplankton is dominant

• Between about 100 and 250 µg P/l the two structures are both possibletwo structures are both possible, dependent on the history

Results obtained by aResults obtained by a structurally dynamic model:structurally dynamic model:

14

10

12

^2)

4

6

8

P_SP

(g/m

^

0

2

4

0 0.1 0.2 0.3 0.4 0.5 0.6TP (mg P/l)

Caspar David Friedrich: Einsamer Baum (Dorflandschaft bei Morgenbeleuchtung, Harzlandschaft), 1822, Öl auf Leinwand, 55 × 71 cm, Berlin, Alte Nationalgalerie, Romantik.

Systems are .....

.....Complexes of

Element

.....Complexes of elements which areinterrelated,Purposeful.... Purposefulabstractions (models),

.... Units in spaceand time with

e.g. Organ

e g Nutrition

Sub system

and time withinteracting sub units,e.g. Organisms

e.g. Nutrition

Relation

y

System.... Irreduceable,.... Hierarchical,

e.g. Population

System boundary

Super system

.... Self regulated,

.... Self-organized.

Ecosystems are .....Inputs

.....Networks of organisms and their

ProduzentenProducers

Consumers o ga s s a d t enon-living environment

Consumers

R l t fDestructors

.....Real parts of space

.....Functional models of interacting structural Soil (Pool)units

.....Interacting entities built up by gradients

Soil (Pool)

p y gof materials and energy Bedrock

Outputs

Structure Function

- Which are the „processors“ of the system?- which elements?which elements?- which components?- which subsystems?

- Which is the relevant pattern of these elements?which species composition?- which species composition?

- which habitat features?- which abiotic patterns in the landscape?which abiotic patterns in the landscape?

- How does this pattern change / develop?

The composition of most elements in all livingelements in all living

organisms are surprisinglyorganisms are surprisingly similar:

• C:N:P = 106:16:1 (by moles)C N P 42 7 1 (b i ht)• C:N:P= 42:7:1 (by weight)

• About 20 elements are common in almost the same ratio for all living organismsorganisms

• The basic biochemical processes are the same in all organisms

µ

µ-max

ckm

Michaelis-Menten’s equation describe the effect of the limitingdescribe the effect of the limiting

element(s):• µ = µ-max*NS / (NS + kn), where kn is

the half saturation constantthe half saturation constant• If two or more limiting factors are

working simultaneously,• µ = µ-max*(NS / (NS + kn))*(PS / PS +µ µ max (NS / (NS + kn)) (PS / PS +

kp)or• µ = µ-max* min(NS / (NS + kn)), (PS / µ µ a ( S / ( S )), ( S /

(PS + kp))

Solar radiation2

3

1719

Nitrate Phytoplankton - N1

315

1414

520

Ammonium Zooplankton - N8

12

16

913721

Sediment- N Detritus-N11

6

18 1022

Fish-N

Mass Conservation:• Applied in all biogeochemical models

dT / dt d il t k k*T t d• dTox/ dt = daily uptake - k*Tox -> steady state concentration: Tox = daily uptake /kBi l ti 10 000 k f h t l kt• Bioaccumulation: 10 000 kg of phytoplankton -> 1000 kg of grazing zooplankton-> 100 kg of carnivorous zooplankton > 10 kg of fish >of carnivorous zooplankton -> 10 kg of fish - > 1 kg of otter or bold eagle - but as the toxic substance is not decomposed through thesubstance is not decomposed through the food chain as the food is, the toxic substance may have up to 10 000 times higher y p gconcentration in bold eagles or otters

• Streeter-Phelps famous model

Energy conservation:9

1.95*109

Reflection and evaporation

Sunlight 1.97 * 10

evaporation

9

Respiration 0.4*10 Net Production 2.0*107 7

Gross Production 2.4*107

2. Law of thermodynamics has many different

f l tiformulations:• All processes are irreversible

E t i b ll l• Entropy increases by all real processes• We loose energy that can do work to energy

( d ) th t t d k b(named exergy) that cannot do work because the energy is in form of heat at the temperature of the environmenttemperature of the environment

• It costs exergy to maintain ecological systems far from equilibrium which is delivered by thefar from equilibrium which is delivered by the solar radiation

Solar ra- diation Plants

Energy for mainte- nance of ani- mals

Animals

mals

Inorganic compounds Animalscompounds

f Detritus Energy for main- tenance of other animals through the food chain

Energy for detrivo- rous animals and microorganims

the food chain

System at temperature T, pressure p and the chemical potential µ(1)p µ( )

Exergy difference or gradient= work produced bygradient= work produced by the gradient in chemical potential

Reference environment at same tempera- ture T and pressure p but by a chemicalture T and pressure p, but by a chemical potential at thermodynamic equilibrium (no free energy available, no gradients):µ[0)gradients):µ[0)

ProteinsProteins P l tid

Exergy

Polypeptides

Amino acids

Urea

Ammonia

Nitrite

Nitrate

Ecosystem growth andEcosystem growth and developmentdevelopment

• 3 growth formsg• E.P. Odum’s attributes• Ecosystem growth and development canEcosystem growth and development can

be described by different thermodynamic variables: maximum power, maximum p ,emergy power, maximum exergy storage, ascendency - they cover different aspects, but are consistent

- complex system can be described correctly by different view points

Three Growth Forms

• 1) Biomass physical structure• 1) Biomass - physical structure• 2) The network2) The network• 3) The information)Notice that they summarize the

E.P. Odum’s attributes

E.P. Odum’s attributes: development of an ecosystem

means that:means that:The biomass increases (GF1)( )Biodiversity increases (GF3)The network becomes more complex (GF2)The network becomes more complex (GF2)More feed backs - more self regulation (GF2)The ratio respiration / biomass decreases (GFThe ratio respiration / biomass decreases (GF

1+3)More and more narrow ecological niches (GF3)More and more narrow ecological niches (GF3)Higher total buffer capacity (resistance) (mainly

GF 1+2)GF 1+2)

100

80

** * * *

60

**

40

20

*0 20 40 60

0**

Exergy (MJ / m 2 )

Incoming exergy (solar radiation)Incoming exergy (solar radiation)An ecosystem at an early stage

Reflected

Utilised for maintenance

Added to exergy storage

maintenance

Incoming exergy (solar radiation)A mature g gy ( )

Reflected

A mature ecosystem

Reflected

Added to exergy Utilised for maintenance

gy storage

structure 2

resp2

structure 1

precycling

solar rad food chain

resp1 resp : struicture

exergy

informationdevelopment Graph 1prod: structure

Table 1

1: information 2: structure 1 3: structure 2 4: solar rad 5: exergy1:2:3:

6000.00500.00

3000.00

1: information 2: structure 1 3: structure 2 4: solar rad 5: exergy

2 2 2

4 4 4 4

4:5:

100.0030000.00

1

2 2 2

5

1:2:

3000.00250 00

1

2

3

3 3 3

5

55 5

2:3:4:5:

250.001500.00

50.0015000.00

1

11:2:3:4:5

0.000.000.000.000 00 1

20:11 fretor 2810sep05 200101

0.00 625.00 1250.00 1875.00 2500.00

Time

5: 0.00 1

Graph 1: p1 (Untitled)

Table 2: Relationships between growth forms and goal functions

Growth FormI II III

____________________________________________________________

Exergystorage up up upgy g p p pPower / throughflow up up upAscendency up up upAscendency up up upExergy dissipation up equal equalRetention time equal up upEntropy up equal equalEntropy up equal equalExergy / Biomass=specific exergy equal up upentropy/biomass=entropy /biomass=spec. entropy prod. equal down downRatio indirect /di-

t ff t lrect effects equal up up______________________________________________________________

III. CONSERVATION

II EXPLOITATION

Trend of each further cycle

I. RENEWALIV CREATIVE DESTRUCTION

Specific exergy = exergy / biomass

A hypothetic trajectory of the adaptive cycle

Maturity / Conservation

Exergystored

Renewal /Reorganization

Release / Creative destruction

Pioneer stage / Exploitation

Connectedness

The original adaptive cycle after Holling (2001)

© Resilience Alliance

Potential for developmental directions in the adaptive cycle

Develop-mental

Potential

Connectedness

We have an ecosystem theory -

consequences:

• It should be applied to ppexplain ecological observations and applied i l i l i i !!in ecological engineering!!

A WORLD EQUATION ????

EINSTEIN’S

THE BASIC LAWS

NEWTON’S LAWS

MAXWELL’S LAW OF E- LECTRICITY

EINSTEIN’S RELATIVITY THEORIES

QUANTUM THEORY

THERMODYNA- MIC LAWS

LAWS OF RADIATION

A number of laws, cove- ring many physical rela- tionships

ALL POSSIBLE OBSERVATIONS

THANK YOU FOR YOURTHANK YOU FOR YOUR ATTENTIONATTENTION