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Volume 4. Physiological Ecology of Forest Production

Published: 22nd October 2010 Authors: J. Landsberg Peter Sands
1  Introduction
1.1  Some background on forests
a)  Goods and services
b)  Wood products
c)  Water
d)  CO2 sequestration
1.2  Models and physiology
a)  Importance of physiology
b)  The nature of models
c)  Complexity and uncertainty
d)  Mathematics
e)  Statistical analyses
f)  Importance of physiological modelling
1.3  Outline
2  Weather and Energy Balance
2.1  Process rates at different levels
2.2  Weather factors that affect plant growth
2.2.1  Solar radiation
a)  Types of radiation
b)  Irradiance
c)  Insolation
d)  Determination of insolation in the absence of direct observations
2.2.2  Temperature
a)  Air temperature
b)  Leaf temperatures
c)  Stem and soil temperatures
d)  Diurnal variation of temperature
2.2.3  Humidity and vapour pressure deficit
a)  Calculating vapour pressure and humidity
b)  Vapour pressure deficit
c)  Calculating average vapour pressure deficits
2.2.4  Wind
2.3  Variation of climatic factors within a canopy
2.4  Energy Balance, Evaporation and Transpiration
2.4.1  Radiant energy
a)  Net radiation
b)  Albedo of forest canopies
2.4.2  Energy balance and flux equations
2.4.3  Resistances and conductances
a)  Boundary layer conductance
b)  Stomatal conductance
2.4.4  Heat and vapour fluxes
2.4.5  Energy balance of a surface
2.5  Canopy energy balance and transpiration
2.5.1  Wind and transfer processes
2.5.2  Partitioning Absorbed Energy
2.5.3  Canopy transpiration
a)  Canopy conductance
b)  Penman-Monteith equation for canopy transpiration
2.5.4  Eddy correlation
3  Physiological Processes
3.1  Photosynthesis
3.1.1  Overview of biochemistry of photosynthesis
3.1.2  Gas analysis and the observation of photosynthetic data
3.1.3  Empirical relationships for CO2 supply and demand
3.1.4  Farquhar and von Caemmerer model of leaf photosynthesis
a)  Assimilation
b)  Temperature effects
c)  Parameterisation
3.2  Stomatal conductance
3.2.1  Stomatal response to irradiance
3.2.2  Stomatal response to vapour pressure deficit
3.2.3  Stomatal response to CO2
3.2.4  Stomatal response to leaf water potential
3.2.5  Phenomenological models of stomatal conductance
a)  The Jarvis model
b)  The Ball and Berry model
c)  Parameterisation of Ball and Berry model
3.3  Coupled model of photosynthesis and stomatal function
3.3.1  The supply and demand curves
3.3.2  Solution of the supply and demand equations
3.3.3  Results from applying the coupled model
a)  Parameter sensitivity of coupled model
b)  A-Ci and light response curves from the coupled model
3.4  Respiration
3.4.1  Temperature dependence of respiration
3.4.2  Dark respiration of leaves
3.5  Allocation of biomass
3.5.1  Principles underlying models of allocation
a)  Functional balance
b)  Local determination of growth
c)  Optimality principles
d)  Coordination theory
3.5.2  Mechanistic approaches to modelling allocation
a)  Transport-resistance and allocation
b)  Pipe model
4  Stand Structure and Dynamics
4.1  Stem population dynamics
4.1.1  Representing mortality
4.1.2  Environmental affects on mortality
a)  Frost hardiness
b)  Drought induced mortality
4.1.3  Self thinning
4.2  Height and diameter relations and distributions
4.2.1  Height and diameter relations
4.2.2  Predicting stem mass from height and diameter
4.2.3  Height and diameter distributions
4.3  Allometric scaling and its implications
4.3.1  Allometry – how objects scale
4.3.2  Allometric relationships between biomass pools
4.3.3  Biomass as a function of diameter
a)  Allometry of stem biomass
b)  Age effects on stem allometry
c)  Allometry of other biomass pools
d)  Clonal or species effects on allometry
e)  Effects of stem numbers, fertility and water status on allometric relationships
4.4  Leaf area of trees and canopies
4.4.1  Specific leaf area
4.4.2  Estimating leaf areas
a)  Tree foliage area
b)  Canopy leaf area index
4.4.3  Modelling closed canopy LAI
a)  Closed canopy LAI in terms of site factors
b)  Closed canopy LAI and physiological parameters
4.4.4  Foliage distribution
4.4.5  Foliage dynamics 
a)  Environmental effects on foliage dynamics
b)  Modelling litterfall
4.5  Roots
4.5.1  Estimation of root mass
4.5.2  Root dynamics
4.5.3  Fine roots
5  The Carbon Balance of Trees and Stands
5.1  Radiation interception
5.1.1  Beer’s Law.
5.1.2  Sun-shade or two-stream model
5.1.3  Single tree and rows
a)  MAESTRA, a general light interception model
b)  Single tree and hedgerow light interception models
c)  Summary models of light interception
5.2  Modelling canopy photosynthetic production
5.2.1  Scaling of canopy processes
5.2.2  Structure of whole-canopy models
a)  Plant-environment models
b)  Multilayer models
c)  Two-leaf or sun-shade models
d)  Big-leaf models
5.2.3  Examples of whole-canopy models
5.2.4  Analytical models of gross canopy photosynthesis
a)  The basic analytical model
b)  Sun and shade leaves
c)  Inclusion of frost effects
5.3  Light-use efficiency and canopy photosynthetic production
5.3.1  Observations of 
5.3.2  Dependence of  on physiological and environmental factors
5.3.3  Growth modifiers
5.4  Non-homogeneous canopies
5.4.1  Mixed-species stands
5.4.2  Edge effects for block or strip plantings
5.5  Stand respiration
5.5.1  Growth and maintenance respiration
5.5.2  Observations of respiration
5.5.3  Carbon use efficiency
5.6  Allocation of biomass
5.6.1  A generic tree growth model
5.6.2  Taking allometry into account
5.6.3  Determination of allocation ratios
6  Nutrient Dynamics and Tree Growth
6.1  Nutrient cycling
6.1.1  The Geochemical Cycle
6.1.2  The Biogeochemical Cycle
a)  Nutrient addition to soil
b)  Nitrogen
c)  Losses: fire
6.2  Forest nutritional requirements
6.2.1  Nutrient uptake
a)  Movement of nutrients towards roots
b)  Factors affecting nutrient uptake
c)  Calculating nutrient uptake
6.2.2  Nutrient retranslocation
6.2.3  Growth in relation to nutrition
a)  Growth and relative nutrient addition rate
b)  Nitrogen productivity and growth
c)  Comprehensive forest nutrition trials
6.3  Modelling soil nutrient dynamics
6.3.1  CENTURY
6.3.2  SNAP
6.3.3  Modelling nitrogen uptake rate
6.4   A pragmatic fertility index
6.4.1  A fertility index based on closed canopy leaf area index
6.4.2  The 3-PG fertility rating
7  Hydrology and Plant Water Relations
7.1  The Hydrological Balance
7.1.1  Equation of hydraulic balance
7.1.2  Quantifying soil water content
7.2  Components of the hydrological balance
7.2.1  Transpiration
7.2.2  Rainfall interception
a)  Observations of rainfall interception
b)  Models of rainfall interception
7.2.3  Redistribution of Rainfall
7.2.4  Soil evaporation
7.2.5  Run-off and drainage
7.3  Water in soils and the root zone
7.3.1  The soil water potential
7.3.2  Root distribution and soil-root resistance
7.3.3  Movement of water in soil
7.4  Water Movement Through Trees
7.4.1  Water movement and water potential
7.4.2  The hydraulic hypothesis
7.4.3  Representation of effects on conductance
7.4.4  Stem water storage
7.5  Models including storage
7.5.1  Tissue water storage
7.5.2  Models based on pools and resistances
7.5.3  Stem hydraulic conductivity and its implications
7.6  Water relations of stands
7.6.1  Quantitative measures of water stress
7.6.2  Consequences of Water Stress
a)  Effects of water stress on foliage
b)  Water use efficiency
c)  Acclimation to drought
7.7  Concluding Remarks
8  Modelling tree growth: concepts and review
8.1  Concepts and principles
8.2  Types of model in forest ecophysiology
8.2.1  Empirical models
a)  NitGro – empirical growth model for Eucalyptus nitens
b)  CanSPBL, a stand-level growth and yield model for Pinus radiata
8.2.2  Process-based or mechanistic models
a)  Light-use models
d)  CenW
e)  The ITE Edinburgh model
f)  ProMod
g)  Cabala
h)  Modular-hierarchical forest growth models
8.2.3  Hybrid models
a)  PROMOD-NITGRO hybrid model
b)  Forest 5
c)  Triplex
d)  General comments
8.3  Discussion arising from empirical, process-based and hybrid models
8.3.1  Reinventing the wheel
8.3.2  Parameterisation and calibration
8.3.3  Why use process-based models?
8.4  Model evaluation: testing and sensitivity analyses
8.4.1  Model testing
8.4.2  Sensitivity analysis
9  The 3-PG Process-Based Model
9.1  An overview of 3-PG
9.1.1  The basis for the structure of 3 PG
9.1.2  Input data
a)  Weather data
b)  Site-specific factors
c)  Stand-initialisation data
d)  Species-specific parameters
9.1.3  Structure of 3 PG
9.1.4  Typical 3 PG output data
9.1.5  Spatially explicit versions of 3-PG
9.2  Biological sub-models of 3 PG
9.2.1  The basic stand-level model
9.2.2  Determination of NPP
9.2.3  Growth modifiers for site and environmental effects
9.2.4  Biomass allocation and turnover
9.2.5  Stem numbers and mortality
9.2.6  Age-dependent variables
9.2.7  Soil water balance
9.2.8  Stand management outputs and interventions
9.3  Calibration, performance and validation
9.3.1  Calibration and parameter estimation
9.3.2  Performance
9.3.3  Validation
9.4  Applications
9.4.1  Analysis and prediction of plantation growth
a)  Temperate eucalypts
b)  Sub-tropical eucalypts in South America
c)  Sub-tropical eucalypts in South Africa
d)  Conifers
e)  New Zealand native species
9.4.2  Use of the model as an analytical tool
9.4.3  Spatial applications
a)  A “proof of concept” application
b)  Comparison with BIOME-BGC
c)  GIS applications using 3-PG-Spatial
d)  Other applications
9.5  Possible improvements
9.5.1  Light interception
9.5.2  Open canopies
9.5.3  Edge effects
9.5.4  Growth modifiers
a)  Temperature-dependent growth modifiers
b)  VPD-dependent growth modifier
c)  Age-dependent growth modifier
9.5.5  Biomass allocation
9.5.6  Water balance
9.5.7  Site fertility
9.5.8  Thinning and pruning
9.6  Concluding remarks
10  Future developments
10.1  Measurement and instrumentation
10.2  Remote sensing
10.3  Meta-analyses
10.4  Respiration
10.5  Stomatal control and hydraulic limitation
10.6  Soil fertility
10.7  Models
10.8  Concluding remarks
Appendix 1    Determining solar direction and radiation
A 1.1  Solar direction
a)  Path of the sun through the sky
b)  Solar transit, day length, and direction of sunrise and sunset
A 1.2  Extra-terrestrial radiation
A 1.3  Transmittance
a)  Optical air mass
b)  Direct beam transmittance
c)  Diffuse beam transmittance
d)  Vertical transmittance
A 1.4  Calculating insolation
Appendix 2   Some mathematical details of 3 PG
A 2.1  Equations for the 3 PG growth modifiers
A 2.2  Equations for explicitly age-dependent parameters
Appendix 3   Further reading

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