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NATIONAL UNIVERSTITY OF RWANDA FACULTY OF SCIENCES DEPARTEMENT OF BIOLOGY TH 4 YEAR BOTANY AND CONSERVATION

INTERNSHIP REPORT

SUMMARY REVIEW on PROCESS BASED MODEL (3PG MODEL)

By Honore Hubert UWIZEYE

Supervisor: Dr Donat NSABIMANA

August4,

2011

ACKNOWLEDGEMENT I would like to express my sincere gratitude to the government of Rwanda for the institutional strengthening of undergraduate studies and for its financial support without which my studies would have not been possible at the national university of Rwanda. I am highly indebted to my supervisors Dr Donat Nsabimana for his tireless guidance, critical comments, his valuable scientific advice, and his endeavor to help me along this internship. Your great academic strength, devotion, competence and kindness provided me the inspiration I needed most. I have learned a lot through this internship. Thank you very much. I would like to thank all those who made valuable contributions to this work in various ways. I cannot end this acknowledgement without saying a word to my class mates who supported me in various ways.

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Table of content
ACKNOWLEDGEMENT .......................................................... ERROR! BOOKMARK NOT DEFINED. TABLE OF CONTENT ............................................................. ERROR! BOOKMARK NOT DEFINED. 1. GENERAL INTRODUCTION ............................................. ERROR! BOOKMARK NOT DEFINED.
1.1. OBJECTIVES: .........................................................................................................Error! Bookmark not defined. 2.1. Introduction .........................................................................................................Error! Bookmark not defined. 2.2. Flow diagram outlining the model inputs and what it calculate ...........................Error! Bookmark not defined. 2.3. Principles underlying the model ...........................................................................Error! Bookmark not defined. 2.3.2. Soil water modifier ........................................................................................... Error! Bookmark not defined. 2.3.3. Temperature frost modifier (fT)........................................................................ Error! Bookmark not defined. 2.3.4. Nutrition ........................................................................................................... Error! Bookmark not defined. 2.3.5. Age effect (fage) ................................................................................................. Error! Bookmark not defined. 2.3.5. Calculation of p.a.u .......................................................................................... Error! Bookmark not defined. 2.3.6. Dry mass production (radiation conversion efficiency) ................................... Error! Bookmark not defined. 2.3.7. Litterfall ............................................................................................................ Error! Bookmark not defined. 2.3.8. Stem population ............................................................................................... Error! Bookmark not defined. 2.3.9. Respiration and root turnover ......................................................................... Error! Bookmark not defined. 2.3.10. Carbon allocation ........................................................................................... Error! Bookmark not defined. 2.3.11. Effect of Nutrition .......................................................................................... Error! Bookmark not defined. 2.4. Results .................................................................................................................Error! Bookmark not defined. 2.5. Discussion and recommendations ........................................................................Error! Bookmark not defined.

3. MODELING FOREST ECOSYSTEMS: STATE OF THE ART, CHALLENGES AND FUTURE DIRECTIONS ............................................................................ ERROR! BOOKMARK NOT DEFINED.
3.1. Introduction .........................................................................................................Error! Bookmark not defined. 3.2. Model user and their needs .................................................................................Error! Bookmark not defined. Forest industries ................................................................................................ Error! Bookmark not defined. Broader public community ................................................................................ Error! Bookmark not defined. The academic and scientific communities ......................................................... Error! Bookmark not defined. 3.3. Characteristics of some currently available process-based and hybrid models . Error! Bookmark not defined. 3.3.1. Model relating to industry: .............................................................................. Error! Bookmark not defined. 3.3.2. Model related to broader public community ................................................... Error! Bookmark not defined. 3.3.3. Academia .......................................................................................................... Error! Bookmark not defined.

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3.4. State of the art .....................................................................................................Error! Bookmark not defined. 3.5. Challenge and future ............................................................................................Error! Bookmark not defined. 3.6. Conclusion and remarks .......................................................................................Error! Bookmark not defined.

4. PHYSIOLOGY IN FOREST MODELS: HISTORY AND THE FUTURE ...ERROR! BOOKMARK NOT DEFINED.
4.1. Introduction .........................................................................................................Error! Bookmark not defined. 4.2. The physiological processes that govern forest growth ........................................Error! Bookmark not defined. 4.2.1. Photosynthesis ................................................................................................. Error! Bookmark not defined. 4.2.2. Stomatal conductance ...................................................................................... Error! Bookmark not defined. 4.2.3. Water relations ................................................................................................ Error! Bookmark not defined. 4.2.4. Nutrition ........................................................................................................... Error! Bookmark not defined. 4.3. The evolution of process-based models ...............................................................Error! Bookmark not defined. 4.4. Hybrid models and the future: linking process-based models with statistical descriptions of stands ....... Error! Bookmark not defined.

5. PERFORMANCE OF THE FOREST PRODUCTIVITY MODEL 3- PG APPLIED TO A WIDE RANGE OF FOREST TYPES................................................... ERROR! BOOKMARK NOT DEFINED.
5.1. Introduction .........................................................................................................Error! Bookmark not defined. 5.2. Model structure and outputs ...............................................................................Error! Bookmark not defined. 5.2.1. Structure........................................................................................................... Error! Bookmark not defined. 5.2.2. Outputs............................................................................................................. Error! Bookmark not defined. 5.3. Inputs...................................................................................................................Error! Bookmark not defined. 5.4. Parameters ..........................................................................................................Error! Bookmark not defined. 5.4.1. Canopy quantum efficiency and fertility .......................................................... Error! Bookmark not defined. 5.5. Calibration procedures and results ......................................................................Error! Bookmark not defined. 5.5.1. Calibration ........................................................................................................ Error! Bookmark not defined. 5.5.2. Results for a range of species ........................................................................... Error! Bookmark not defined. 5.5.3. Tests against independent data ....................................................................... Error! Bookmark not defined. 5.6. Discussion ............................................................................................................Error! Bookmark not defined.

6. UNDERSTANDING 3-PG USING A SENSITIVITY ANALYSIS ...... ERROR! BOOKMARK NOT DEFINED.

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6.1. Introduction .........................................................................................................Error! Bookmark not defined. 6.2. Overview of 3-PG .................................................................................................Error! Bookmark not defined. 6.3. Method ................................................................................................................Error! Bookmark not defined. 6.3.1. Data used in sensitivity analysis ....................................................................... Error! Bookmark not defined. 6.3.2. Calculation of sensitivity analysis ..................................................................... Error! Bookmark not defined. 6.4. Parameter sensitivity analysis ..............................................................................Error! Bookmark not defined. 6.5. Model output as function of site factors ..............................................................Error! Bookmark not defined. 6.6. Results and discussions ........................................................................................Error! Bookmark not defined. 6.7. Discussion ............................................................................................................Error! Bookmark not defined.

7. ANALYSIS OF BIOMASS ACCUMULATION AND STEM SIZE DISTRIBUTIONS OVER LONG PERIOD IN MANAGED STANDS OF PINUS SYLVESTRIS IN FINLAND USING THE 3PG MODEL ................................................................................ ERROR! BOOKMARK NOT DEFINED.
7.1. Introduction .........................................................................................................Error! Bookmark not defined. 7.2. Outline of 3-PG model..........................................................................................Error! Bookmark not defined. 7.3. Summary of stand dynamics in the model ...........................................................Error! Bookmark not defined. 7.4. Data availablility ..................................................................................................Error! Bookmark not defined. 7.5. Estimation of model parameters ..........................................................................Error! Bookmark not defined. 7.6. Results .................................................................................................................Error! Bookmark not defined. 7.7. Discussion ............................................................................................................Error! Bookmark not defined.

8. GENERAL DISCUSSION ..................................................... ERROR! BOOKMARK NOT DEFINED. 9. GENERAL CONCLUSION ................................................... ERROR! BOOKMARK NOT DEFINED. REFERENCES............................................................................ ERROR! BOOKMARK NOT DEFINED.

1. GENERAL INTRODUCTION Climate change Predictions indicated profound effects on the distribution, function, and productivity of forests in different parts of the world (Lenihan et al. 2003, Battles et al. 2008). Although forests cover approximately 30% of the Earths land surface and provide critical ecosystem goods and services, including food, fodder, water, shelter, nutrient cycling, and cultural and recreational value. Forests also store carbon, provide habitat for a wide range of species and help alleviate land degradation and desertification. The goal is a compelling one forests should be managed to meet current needs without compromising the ability of future generations to meet their needs. However any plan designed to strike a balance between current and future demands depends on the ability to make reliable projections of forest dynamics. Thus there is a pressing need to provide managers with the means to predict forest responses under a range of expected climate scenarios. Equally important, anticipated changes in productivity over the next century must be considered in policies being designed to promote the use of forestry projects for climate mitigation (John et al. 2009). From a forest management point of view, predicted climate change (CC) implies that current practices may need to be adapted for future conditions (Kellomki et al. 1997). By forest management it would be possible to respond to and adapt forests to the environmental changes and following changes in forest dynamics. Planning of adaptive forest management would require forestry modeling tools that are usable under climate change conditions. They should be at their best both responsive to changing environmental conditions and be able to use traditional forest inventory information as inputs to produce reliable predictions on forest growth and yield for application scales of practical forestry. The models currently used in forest management and as research tools can be classified into statistical (empirical) growth and yield models and process-based models (Mohren and Burkhart 1994). Generally both types have been developed independently one another. However, both these approaches aim at providing tools for analyses on forest resource management and related research (Korzukhin et al. 1996). So far, statistical models of forest growth and yield were thought to be preferable whenever locally focused and site-specific predictions have been required to support decision-making in practical forestry (Mohren and Burkhart 1994). However, their predictions are based on the assumption that the future environmental conditions are those which prevailed in the past (Korzukhin et al. 1996). Therefore, these models fail to tackle the

impacts of the changing environment on the tree growth as opposed to the physiological models, in which climatic and edaphic factors interact with the growth processes of trees. Accordingly, forest growth models based on physiological processes (e.g. photosynthesis, transpiration and respiration) with hydrological and nutrient cycles as controlled by climatic factors have been generally preferred when predicting stand productivity under changing climatic conditions (Landsberg and Waring 1997). Thus as far as environmental fluctuation due to climate change is concerned process based models appear to be useful in predicting impacts on forests. The current work will provide the state of knowledge about one process based model: 3-PG model (Physiological Principals for Predicting Growth).

1.1. OBJECTIVES: To provide a critical review of one process based model (PBMs) by mean of a critical summary of six publications on these forest growth predicting models. In this review, the following papers were used: 1. Generalized model of forest productivity using carbon balance and partitioning. 2. Modeling forest ecosystems: state of the art, challenges and future directions. 3. Physiology in forest models: history and the future. 4. Performance of the forest productivity model 3- PG applied to a wide range of forest types. 5. Understanding 3-PG using a sensitivity analysis. 6. Analysis of biomass accumulation and stem size distributions over long period in managed stands of Pinus sylvestris in Finland using the 3-PG model. concepts of radiation-use efficiency,

2. Generalized model of forest productivity using

concepts of radiation-use efficiency,

carbon balance and partitioning (by landsberg J.J. and Waring .R.H.) 2.1. Introduction This paper describes a model called 3-PG (the acronyms derived from the use of physiological principles in predicting growth) built in the stream of model development of the last 10-15 year. This model was built in response to the questions of complication great deal of information, strict and careful parameterization required for running its contemporary. Estimate of carbon fixed and biomass produced as only results of these models are of limited value for those concerned with forest growth in the convention sense of tree mass and its distribution to trees and their component parts. The model is based on well established principles recently confirmed constant and simplified calculations. It requires little adjustment to obtain realistic forest growth estimate and is easy to parameterize for a particular forest types. The model can also be applied to ground based forests inventory maps incorporated in a GIS or maps derived from analysis of satellite images to extend estimate of forest growth over large heterogeneous Area. The following section provide description of the simplifying concepts used to run it and some results of sensitivity analysis to show the performance of 3-PG. The model uses dimensionless factors (fi) called modifiers with value varying between zero and unity to account for the environmental constraints on growth. These are high atmospheric vapor pressure deficit (D), soil drought, defined by ratio of the amount of water in the root zone to the maximum possible amount (), or the effects of sub-freezing temperature (T).

2.2. Flow diagram outlining the model inputs and what it calculate Inputs Weather data (for representative year or each year considered) monthly average values of radiation, frost days per month, humidity. Total precipitation Initial biomass values: foliage: (wf) stem( ws), roots (wr) Variables: maximum available soil water, initial stem number, stand age, Parameter values: canopy quantum efficiency (c), ratio

Cpp=

maximum stomatal conductance (gs.max), maximum canopy

conductance (gc.max), Parameter of allometric equations: soil type parameters, maximum litterfall rate, root turnover rate. Calculate
Monthly time step of: Leaf area index L* from foliage mass, p.a Vapour pressure deficit modifier to gs.max, (stomatal conductance) gc.max (max. canopy conductance) Monthly transpiration from the Penman-Monteith equation Soil water balance, moisture ratio, soil water modifier Stem mass/tree = ws/stem number Utilizable p.a (p.a.u) from p.a and modifiers PG= c p.a.u, PN = PG Cpp Carbon allocation coefficient (nr, ns, nf ) Component mass increments : wf = nfPNetc. Update component mass : wf(t)= wf(t-1) + wf

2.3. Principles underlying the model 2.3.1. Vapor pressure deficit modifier (fD): used to reflects the constraints imposed on utilization of absorbed radiation by leaves because of stomatal closure caused by high atmospheric vapor pressure deficit, soil drought and effect of sub-freezing to.

FD=

with kg: empirical coefficient describing the relationship between stomatal

and canopy conductance and D. And this is used to get the canopy conductance gc and

gc= gcmax exp (kg D)

This was proved by Landsberg et al (1996) showing that monthly growth increment of Eucalyptus plantations were negatively correlated with monthly average of D 2.3.2. Soil water modifier The moisture ratio and water balance have first to be found Water balance: monthly transpiration (calculated using Penman-Montheith equation) monthly precipitation The moisture ratio (ro): [current soil water content + water balance] / available water (, mm) The soil water modifier is calculated as follow:

f =

the relationship shows that as r increases f decreases.

2.3.3. Temperature frost modifier (fT) It is assumed that there is no photosynthesis on any day that to falls below zero.

fT= 1- (
2.3.4. Nutrition Nutrition is dealt through the mechanism of carbon allocation rather than dealing with complicated relationship between leaf photosynthesis and nitrogen concentration

2.3.5. Age effect (fage) As a forest ages, its above ground net primary production decreases. Paired young and old tree comparisons made on the same site showed that older trees had similar maximum rate of photosynthesis in early morning, but during the day these rates fall 25-30% more than those observed on foliage of young trees. These differences are attributed to the differences in relative sensitivity of stomatal to atmospheric VPD associated with hydraulic limitations to the flow of water through an increasingly long and torturous path as trees age. Because of reduction in stomatal conductance and hydraulic conductivity as a tree ages, photosynthesis is reduced and less carbon is available to maintain previously acquired leaf area as result the canopy opens. Then this is accounted for through the age modifier (fage).

fage =

Fa : relative stand age; nage: empirical power term

2.3.5. Calculation of p.a.u Utilizable, absorbed photosynthetically active radiation is calculated by applying the modifier for D (fD), soil water (fo), temperature (fT), and age (fage) to p.a.u 2.3.6. Dry mass production (radiation conversion efficiency) GP (gross primary production is calculated by applying p.a.u the canopy quantum efficient coefficient c, and PN follows. 3-PG uses a universal canopy quantum efficiency coefficient c, with a value of 0.03 mol C (mol photon)-1 equivalent to 1.8 g C MJ-1, a value that was proved to be constant. In a number of studies (Mc Murtrie et al 1994;Warring 1995) this maximum value does not vary widely about that value except in severely nutrient deficient soils, or where atmospheric pollutant causes chlorosis of foliage, where it slightly decreases. The standing biomass at any time W(t) Dry mass production PG

= PG R

= p.a.u c

Canopy quantum efficient coefficient is universal: 0.03 mol.C/mol photon

The ration

Cpp=

was proved to be constant, so PN is deduced without need to

calculate respiration. 2.3.7. Litterfall A monthly litter fall rate is used (fmax) of about 0.02 but has been made a function of age for young stand

f =

with

f:

foliage litterfal rate,

:empirical constant =15;

:empirical

coefficient = 0.12; t: time in month 2.3.8. Stem population The model requires stem number because the allometric ratios used to partition carbon are invariably determined for a single tree. Also forest managers are not interested in the total biomass produced by a forest; they want to know stem growth rates and final stem volumes. 3PG use the relationship between stem population and maximum achievable individual stem mass (Ws.max) to calculate change in stem population (p)

Ws.max = ksp-3/2

Averages stem mass 1 2 Ws= ksp-3/2

Ws: mean stem mass

Stem population (ha-1)

If Ws < Ws.max (point 1) the population remains unchanged, if Ws > Ws.max, reduces by the number necessary to conform to the relationship. Ks value has to be provided, calculated from empirical data, or from stem mass value got by running the model for 120 years using initial stem population of 150 ha-1 with normal environmental constraints by using the model.

Mass of stems that die must be discarded from standing biomass Leaf mass does not reduce on the assumption that dying stems would have few leaves and stand foliage mass is distributed among the living stems.

2.3.9. Respiration and root turnover The ratio

Cpp=

has been proved to be constant averaging about (0.45 0.05) for a wide

range of forests. It also has been proved that: 1o there is a strong relationship between growth and fraction of PN allocated below ground. 2o as environmental conditions become harsher , the fraction of PN allocated to fine roots growth increases from about 25 to nearly 60%. The measure of harsherness is provided by environmental modifiers. Under zero stress: fD, f, and fT respectively VPD, Soil water and temperature modifiers are unit and harsherness can be deduced in the ratio

p.a.u

p.a

hence

2.3.10. Carbon allocation Mechanism of carbon allocation to component part of plant is not well understood. In model carbon allocation is sometimes estimated using allocation coefficient ni derived from allometric equation describing observed relationship between the mass or size of different parts of plants. Allometric relationships for single trees can, almost invariably be described by equations of the forms: Wi

= aiWni

with W: total mass of the plant; i: any component part of the plant.

Foresters normally use stem diameter at breast height as a measure of stem size. This can be used as a surrogate for W. for instance the following equation shows how nr (allocation coefficient to root) is expected to vary with growing conditions.

r=

Making nr dependent on the relative harshness of the growing condition defined by the ratio , reflects the accumulating evidence that trees allocate increased amounts of carbon to their roots when growing conditions deteriorate. In 3-PG model average stem mass is calculated form total (stand) stem mass in PN is reduced by poor growing condition, nr is increased and stem and foliage growth are reduced in a way that conserves the allometric balance of the trees. 2.3.11. Effect of Nutrition Nutrition affects the amount of carbon allocated to roots with greater proportion going to roots of trees on infertile sites than those on fertile sites. Deterministic relationship is not yet found. A simple scaling procedure minimizing carbon below ground of fertile sites and increases as fertility decreases.

with a new variable (m) that has a maximum value 1 on highly

fertile site, reducing towards zero as site fertility decreases. 2.4. Results The model requires weather data (monthly values for as many years as required using the same set of data). It works with monthly time step and has been run for a period of 120 years producing realistic pattern of stem growth and stem diameter increments, as well as realistic time curse of leaf area index for a range of soil conditions and atmospheric constrains. 2.5. Discussion and recommendations The use of is reliable and keeps the model relatively simple. But the

procedure need to be tested against long-term detailed flux measurements Hydrology is extremely simple and need to be reconsidered. The use of monthly water balance through assuming that rainfall is evenly distributed over the month is not

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obviously the case and soil recharge and depletion patterns are distorted. The use of daily precipitation and water balance could be a solution but it is a high data demand and increased complexity process. The constancy of ratio eliminates the need to calculate respiration in various tissues

and components which would leads to large errors. But certainly the errors are smaller than those that would result from calculating respiration. Allometric ratios used in the model to deal with carbon allocation, which are results of carbon allocation up to the times they are measured, are assumed to be genetically determined and tree tend to conserve them. But these can undoubtedly be modified by environmental conditions. The matter of age effect remarquable in the reduction of PN as a forest ages is linked with reduction in hydraulic conductivity which feeds back to stomata finally photosynthesis, is consistent and should be reinvestigated. Parameterization of the model to estimates derived from the satellite which monitor seasonal and yearly variation of L* (leaf area index), would enable the model to scale up stand prediction of growth to region wide estimate of PN. In a word, the model can confidently be used as a practical tool.

3. Modeling forest ecosystems: state of the art, challenges and future directions (by Landsberg J.J.)
3.1. Introduction
This paper was a contribution to the conference that discussed how current and future modeling efforts will address the information needs for ecosystem management, forest certification and sustainable management. Challenges faced by forest modelers and the most useful future directions for their works are discussed. For the identification of information needs of forests managers, challenges and future directions for forest models, they considered who they are and what they are trying to achieve. Forest managers concerned with wood production will require information on the present status of the resource (e.g. number of trees by species and sizes etc.) forecasts of the nature and timing of future harvest. Other concerned with non-woody values may require different information. Models of any sort are abstractions that should encapsulate the essential features of the system being modeled. Conventional empirical

11 growth and yield models are developed from large amount of field data to meet the need of managers. They provide statistical descriptive accuracy; process based models (PBMs) provide flexibility, generality and predictive power. A process based model implies a model of a particular process (e.g. leaf photosynthesis, carbohydrate allocation,). A system is modeled at whatever level of complexity based on sub-models that handle its constituent processes. The need to develop PBMs was first recognized by Mkel et al (1986) and PBMs were also stated to have significantly more advantages over empirical models in increasing our understanding of, and predicting forest behavior Korzukhin et al (1996). Reliability of future on hybrid model (combination of PGMs and relationship based on stand level measurements), the close interaction between ecophysiological and empirical modeler and forest practitioners, potential valuable role of PBMs in forest in forest management and involvement of prospective client in model development have all been recommended in different reviews. 3.2. Model user and their needs Forest managers are not a homogeneous group, and there is no universal model that can provide for all users needs. Forest managers can be grouped into forest industries, the broader public community, and the academic and scientific communities. These entire groups may use models as tools to aid decision-making by serving as guides and mean of exploring options and alternatives, evaluation of the consequences of particular actions, or examination of the sensitivity of the system to specific disturbance or lack of it. Forest industries: may be represented by state organizations or by private companies who own or lease tracts of forested land. Operational managers (or manager on the ground) of these, in charge of operations like logging schedules, thinning, re-establishment, disease and weed control, may need growth model to provide better information for forest management but these should not threaten his own experience and judgments. Manager at high level need PBMs while dealing with larger scale questions such as wood flow and market requirements or the economics of alternative practices such as thinning or fertilization. Commercial companies also may use PBMs in their need to know the probable productivity of new land to make decisions about purchase and development. Plantation, primarily for pulpwood are increasing rapidly. Seeking to develop new resources to replace the logging of natural forests involve the use of technology and silvicultural practices for which rate of change is outstripping the capacity to gather measurement data thus need PBMs. Broader public community: is concerned with the use and sustainability of natural forests. Exclusion of areas of natural forests from commercial exploitation will involve considerations of

12 the potential wood yield of forests. Assessment of potential yield in any given area essentially involves estimation of total aboveground biomass production; estimates of the economic value of that biomass involves prediction of stem size distribution and wood quality, all these information could be provided by PBMs. Questions of carbon sequestration by forests is becoming of increasing importance in relation to climate change and its consequences. Reports about it depend heavily on modeling, and they are crucial factors in raising political awareness and concern about climate change and its implications. Environmental groups may need proof made of concrete to fight some economical activities by logging bans, the use of reliable model may help in these discussions to reach a rational conclusion. PBMs may also be used to assess the impact of plantations on the water balance of an area since it has been shown that deep-rooted evergreen trees can cause sufficient disruption to the local hydrology. The academic and scientific communities: PBMs are done by scientists who are either in universities or in institutions such as government research organizations. The objective of their modeling exercise is to improve understanding of the processes and the factors affecting forest growth. The development of models of any type in the academic community also contributes to the ability of people concerned to teach effectively. Model developed in academic communities are not intended as practical management tools. They provide a vital source of understanding and techniques that, directly and indirectly contribute to the modeling exercises carried out by people more concerned with management. 3.3. Characteristics of some currently available process-based and hybrid models Models considered in this section are categorized as relating to the user groups and have characteristics that would allow them, often with some modifications, to meet the requirements and objectives of the clients. 3.3.1. Model relating to industry: They are carbon balance models, combined with some means of allocating carbon to stem to provide results useful to forest managers. Pathfinding model by Mohren et al (1984), Makel and Hari (1986) hybrid models describing stand growth in terms of simplified physiological processes and assimilate distribution on leading to the calculation of potential productivity. The model worked with average weather data on a yearly time step and did not include the dynamics of

13 water relations and nutrient supply. Stand structure was represented by equations very similar to those used in conventional modeling FORCYTE model developed by Kimmins et al. (1990), a comprehensive forest ecosystem model intended to predict the consequences of action such as forest harvesting and fire. This model aim to simulate the light climate and the availability and dynamics of up to five nutrients and their effects on tree growth and secondary succession over the management cycle, as well as interaction between herbs, shrubs and trees. Immensely complicated since it represents an excellent summary of

current knowledge about forest ecosystems and requires a considerable amount of input information. Its complexity makes it difficult to envisage how it could be rigorously tested in any quantitative sense Sievnen (1993) produced a model of the dimensional growth of even-age stands in which biomass production was calculated from intercepted radiation and a canopy conversion efficiency (, biomass produced per unit intercepted light). Respiration and senescence terms gave the carbon balance. The model could be used for the assessment of timber quality. 3-PG (Landsberg and Waring 1997) is a generalized stand model (i.e., it is not site or species specific but needs to be parameterized for individual species) applicable to plantations or even-aged , relatively homogeneous forests that was developed in a deliberate attempt to bridge the gap between conventional, mensuration- based growth and yield and process-based carbon- balance. More details are provided in the article published on the model. FOREST-BGC (Running and coughland 1988; Running ans Gower 1991) is the best known process-based model of forest growth. It includes hydrologic, photosynthetic, and maintenance respiration processes computed daily and carbon allocation and nitrogen processes computed annually. Outputs are in terms of biomass pools (t.ha1

.year-1-) which are of little interest to forest managers. The model allocated carbon to

individual trees on the basis of their relative sizes and allometric relationships. It produces reasonable estimates of NPP available for growth and high correlation between measured and modeled stand volume increment and basal area increment. PROMOD by Battaglia and Sands (1997) that estimates site productivity in terms of mean annually increment of Eucalyptus globulus. It contains simple empirical relationships for closed-canopy leaf area index in terms of site climate factors. Soils information includes water-holding capacity and a fertility index. It includes a

14 canopy photosynthesis production model and respiration calculations, a standard water balance model and respiration calculations, a standard water balance model and carbon allocating to stems and foliage on the basis of empirical relationship. 3.3.2. Model related to broader public community The succession or gap models that derived from the work of Botkin et al. (1993) were developed for regional scale analysis of forest dynamics. In general, they simulate establishment as a stochastic process constrained by environment filters. The growth of trees is simulated in terms of species-specific empirical relationships describing height and diameter growth, constrained by temperature, drought, and nitrogen dynamics formulated in various ways. Completion is simulated by shading based on height and canopy size relationships. Formix 3-Q model formulated by Ditzer et al (2000) to estimate the timber yield. The growth of trees was described by form factors and allometric relationships, with dry mass production estimated from light interception and a standard value of . Respiration is estimated by manipulation of the allometric equations to give trees size increments. Site quality was estimated from soil maps giving nutrient concentrations and plant available water. Succession development, above ground biomass, and the impacts of logging are simulated using GIS and stratification procedures. Information got from the model could otherwise be difficult to obtain and can contribute to management decisions GDAY model (generic decomposition and yield) that describes how photosynthesis and nutritional factors interact to determine the productivity of forests growing under Nlimited conditions. The model is used to estimate N-mineralization rate from (assumed) soil organic matter composition, and the C:N ratios of plant and soil pools are specified. It provides insights into the probable interactions between increased photosynthetic rates that might result from increased atmospheric CO2 concentrations with nitrogen supply constraints and the likely equilibrium situations. The model does not provide results relevant to any particular practical situation but give perception of the relationship that exist between factors, such as N concentration in tissues, or biomass growth in relation to N-uptake rates and retranslocation. It is a tool that can bring insight into discussions about the effect of climate change on forest and sustainability in terms of N supply and cycling.

15 3.3.3. Academia These are model developed primarily for the purpose of understand mechanisms and the interaction between processes. Some already mentioned would also appear for they have some practical aspects. Thornleys (1991) transport-resistance model of forest growth and partitioning applies the transport-resistance approach of crop plants, to forests. This is coupled with representation of growth in terms of the size and activity of meristems. The model has five compartments and is based on carbon and nitrogen and nitrogen pools and fluxes. But due to the fact that the model use a large number of parameter, state variables and equations, it is very difficult to reproduce and work with it without the help from Thornley. Although, the model produces plausible curves of the time course of the growth of various components of the simulated forest and interesting and informative response to stimuli. The pipe model also called PIPESTEM, by Shinozaki et al. (1964) measured growth in terms of mean stem length (from leaves to feeder roots), basal area, woody volume, and total (carbon equivalent) mass. The rate of biomass production is estimated as the rate of production of substrate minus the rate of maintenance respiration. Cen W, produced by Kirschbaum (1999), is a comprehensive model of the growth of monospecific, even aged stands that attempts to deal with all the major processes involved in CO2, water and nitrogen fluxes, foliage dynamics, wood production, and stand architecture. It has been used to make simulations of wide-scale forest growth and response to environmental conditions. Cen W has not been designed as a tool for management but is useful as an attempt to link in one model all the biophysical processes considered important in the growth of trees and timber production. HYBRID immensely detailed model of ecosystem dynamics produced by Friend el al. (1997). It aims to couple carbon, water, and nutrient cycles in the soil-plant-atmosphere system, prediction of the consequences of climate change in terms of GPP and NPP, heterotrophic (soil) respiration, latent heat flux, total carbon biomass, and annual maximum L*. It also predicts the probable dominance of particular plant types. The model is not intended to be used in any operational or decision-making context in relation to forestry but it may contribute to evaluation of the probable impacts of climate change.

16 3.4. State of the art The brief survey on some PBMs leads to the conclusion that the use of mixed process-based and empirical models will lead to significant improvements in model flexibility and predictive power. Whatever the improvement to simulated ecosystem processes, models have to be massively simplified to be of any value as a practical tool. 3.5. Challenge and future Carbon allocation, nutrition, communication between scientists and managers to see the need of the end user, the accuracy required of output variables, number and availability of parameter values are still challenges in the field of PBMs development. 3.6. Conclusion and remarks PBMs are at the stage where they can be used as management tools to predict growth and yield of forest stands. The optimization of effectiveness and to create confidence in users, the scientists who develop and work with these models must consults, and work closely, with the prospective model users. PBMs can provide quantitative estimates of the effects of climate change. They are important tools for heuristic analysis to answer the what if questions in relation to factors such as management actions, insect attack, or drought. Complex, multiparameter models tend to remain as research tools within the scientific community, and even they tend to be used only by their creators and perhaps, a small circle of colleagues. They need to develop simplified versions that might be more widely used. Scientists are often hesitant to develop models in the form suitable for release and use by others. They know that they are imperfect and are reluctant to commit themselves, and they do not want to allow time to provide technical support for users.

4. Physiology in forest models: history and the future (by Joe Landsberg) 4.1. Introduction Foresters usually concerned with wood production have been using empirical model to get information such as volume of timber, how forest is growing, and expected volume of timber in the future. These models have been potential sources of information for management and planning that is accurate within limits of sampling and measurement accuracy. However these

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models are site specific, lack flexibility, the capacity to simulate the results of environmental stress, and they cannot be driven by data obtained by remote sensing. Therefore, they cannot take advantages of technology and that is a limitation to their spatial applicability. On the other hand, it is believed that model based on physiology that governs tree growth could be a response to the challenges faced by empirical models. Tree physiology seemed to be irrelevant to forest managers and process based models (PBMs) tend to be ignored for they contain too may poorly known parameters for their projection to be as reliable in practice as those of empirical models (EM), and their outputs have not been of any interest to foresters. Despite the perceived problems and dichotomy of views between physiological and empirical modelers, the view that we should move towards mixed or hybrid model that incorporate elements of PBMs and EM and exploit the strength of each, is gaining ground. 4.2. The physiological processes that govern forest growth Using radiant energy and CO2 captured by foliage of plants, carbohydrates are formed, respired to produce energy needed for protein synthesis and formation of new tissues, and the rest is translocated to components parts of a tree. This process called photosynthesis is affected by nutrient status of the foliage, uptake of CO2 through stomata and temperature. Carbohydrate that becomes wood is the interest of commercial forester. EM based on measurement of the product describe it statistically while PBMs predict the product by describing the process that lead to them, their response to external driving variables and interaction between them. The problem comes while forming relationships on which the models are based. Pn = (, N, T) abs R (1) With : coefficient describing the conversion of absorbed photosynthetically active radiant energy (abs) to carbohydrates, , N, T denote water relations, nutrition and temperature respectively, R: respiration. Photosynthesis is driven by photosynthetically active radiation absorbed by the canopy. To calculate light absorbed by the canopy, a description of amount and distribution of the foliage that makes up the canopy in terms of Leaf area index (L*), is required. Light penetration into, and absorption by plant stand has been described by Beers Law: in the horizontal plane light intensity decreases exponentially projected foliage area index generally referred to as L*. Some

18

models like MESTRO by Wang and Jarvis (1990) have been developed on this principles but the amount of information need about canopy structure combined with hourly time step needed to run it, preclude the model use as tool for predicting canopy photosynthesis. 4.2.1. Photosynthesis Early research in 1960s and 70s lead to the establishment of basic characteristics of photosynthesis, the most important of which was the fact that the rate of leaf photosynthesis, as measured by CO2 uptake, invariably shows a hyperbolic response to light intensity, with negative rates of uptake, indicating respiration, at low intensities. The process is also strongly dependent on ambient CO2 concentration. This was followed by establishment of a set of light response curves for shoots at various depths through the canopy by Jarvis et al. (1976). Recently Farquhar et al. (1980) came up with a biochemically-based model, with parameters that can be associated with ambient CO2 concentration, leaf nutritional status and temperature. 4.2.2. Stomatal conductance They are interface between leaf surface and the air. They tend to open in light and close in the dark and they are very responsive to water status of leaves and CO2 concentration. The extent to which stomata are opened or closed and the rate of gas flow through them, is described by stomatal conductance (gs) while gas exchange of forest canopies depend on canopy conductance (gc= gsL* ), which is function of canopy leaf area and stomatal conductance. A semi- empirical model for stomatal conductance produced by Leuning, 1990, 1995 now provides a single equation, a general description of stomatal responses. 4.2.3. Water relations Enormous work have been carried out aiming at providing an understanding about physics of water movement through plants and interaction between plant water status, soil water content and the effects of water stress on plants. In general soil water content in the root zone has a nonlinear influence on plant growth. When soil is wet, water is non-limiting and other factors will determine growth rates. As the soil dries, combined with the influence of VPD, it becomes more and more important, reducing the ability of trees to absorb nutrients until it becomes dominant determinant of growth. Different factors are taken into account to calculate water holding

19

capacity of the soil exploited by trees: the proportion of rainfall lost by interception by canopy, surface runoff, drainage, then calculating transpiration by trees and the water balance. Water content in the root zone of trees is variable at any time. But a 2-layer model was proved to be sufficient to describe the situation with useful accuracy for practical purpose. 4.2.4. Nutrition Yet nutrient uptake and its effects on tree growth cannot be predicted due to several factors including complex process of mass flow, diffusion to mychorhizea and fine roots that are almost impossible to measure, nutrient and their chemistry complexity, the poor knowledge of nutrient concentration in plant tissues, its influence on growth and mobility. Nutrient uptake rates by trees at any site are still lacking, prospect has only been done on Nitrogen where it is assumed that uptake rate is equal to N-mineralization rate. But for practical purpose some model use site fertility index like fertility rating (FR) in 3-PG model where values are given based on local knowledge about soil fertility. 4.3. The evolution of process-based models The evolution of PBMs starts with the publication of Pathfinding (Montheith 1977) showing that the growth rate of crops increases linearly with seasonal abs. The production of model that can serve as practical tools for forest managers like 3-PG shows that understanding of physiological process underlying the growth is good enough at levels ranging from plant organ to stands, and can allow to establish Models that can predict biomass proclivity accurately and reflect the effect of environmental variables on growth. But some problems like carbon partitioning remain problematic. 4.4. Hybrid models and the future: linking process-based models with statistical descriptions of stands The future modeling is expected to lie on hybrid models combining the strengths and flexibility of these PBMs with the strong point of measurement-based, EM. This will require a change in mind set among the practionners who will need to recognize that the new tools can offer more that the old ones the determination of model structure appropriate to particular applications must start with the end user, whose needs determine the model context, the question to be addressed,

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the accuracy required of estimates and the range of situation or environment over which the model is to be applied. The use of hybrid model will reduce the measurement costs while increasing the information available to management. The final point should be made about the future of modeling concerns the role of remote sensing (RS), which allows estimate of L* so that, linked to weather data, growth rate can be calculated.

5. Performance of the forest productivity model 3- PG applied to a wide range of forest types (by J.J> Landsberg, R.H. Waring, N.C. Coops)
In the current paper, the structure of the 3 PG is outlined indicating the input data and parameter values required to run it and changes introduced since the model description was originally published. 5.1. Introduction The model called 3-PG (Physiological principles Predicting Growth), developed by Landsberg and Waring (1997), is a simple process- based model requiring few parameter values and only readily available data as inputs. It is a generalized stand model (not site- specific, but need to be parameterized for individual species) applicable to plantations or even- aged, relatively homogeneous forests. It has been developed in a deliberate attempt to bridge the gap between conventional, measurement- based growth and yield, and process- based carbon balance model. The output variables are of direct interest and applicability to forest managers. Parameter values that provide reasonable predictions of the growth of particular forest types can be estimated from relatively limited data. Different analysis of the model performance and increasing experience with the model have allowed the evaluation of parameter values that are most critical and variation in parameter values derived from fitting 3-PG to different set of measurements made on the same species subjected to different treatments. The strength of the model is that it produce values of a number of output variables that correspond to those observed , these include the time course of stand biomass, average stem diameter and hence stand basal area and leaf area index (L*), as well as reasonable values of monthly transpiration rates. Growth, in terms of carbon fixed by the stand, and water use, are linked through the stomata, which are explicitly dealt with in the model.

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5.2. Model structure and outputs

5.2.1. Structure

Here general outline indicating changes introduced since the model description was originally published are provided. The model consists of 2 sets of calculations: Those that leads to biomass values Those that allocate biomass between various components of the trees, and hence determine growth pattern of the stand. 5.2.2. Outputs These are monthly or annually values of: Leaf area index Stem mass and volume Stem growth rate Mean annual (volume) increment (MAI) Stem number The model can also provide estimate of stand height.

3 PG can run for any number of years, and uses actual weather for each month or monthly averages for many years. Like other model it is based on the calculation of: Radiation interception Canopy photosynthesis or gross primary production(GPP, GP) obtained by applying a canopy quantum efficiency (QE, c) value to the photosynthetically active radiation(PAR, p) absorbed by the stand (APAR, p.a) Net primary production (NPP, PN) Allocation of carbohydrates to component parts of the trees

Maximum value of c is constrained by vapor pressure deficit (VPD) through its effect on stomatal conductance (gs) and hence canopy conductance (gc), air temperature, frost, water balance and nutrition to get effective value of c.

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Soil nutritional status is represented by an index: the fertility rating (FR) that takes valued between o and 1.

The allocation equation is fully developed by Landsberg and Waring (2003). The allocation of carbohydrate is on a single tree basis. Allocation to roots is influence by moisture relation and soil nutrition and symbolized by the coefficient (nr). Allocation to stems and foliage relies on the ratio of the derivatives (Pf,s) of the allometric equations ( Wi= aiBin) describing Leaf (wf) and stem (ws) mass in terms of stem (bole) diameter at breast height (B).

(1)

The model calculates monthly values of the average mass of individual stem; and the relationship between B and ws determines stand basal area and volume at any time. The use of the Pf.s ratios provides a convenient way to estimate the rate of change of foliage mass with stem diameter.

Initial tree populations are specified, its change is caused by natural mortality and thinning. Soil water balance is calculated using Penman-Monteith equation to calculate transpiration. Canopy conductance is estimated from gs and L* The limiting factor is one that applies, i.e. if soil water is more limiting than VPD, growth is assumed to be constrained by soil water during that Period.

5.3. Inputs Even if 3 PG model can be started at any stand age, normally it simulates stand growth from year 2. The inputs data required by the model are: Weather data (monthly average values of solar radiation, atmospheric VPD, rainfall, frost days per month and average temperature). All these are routinely available except in some cases where solar radiation can be calculated from temperature data (Bristow and compbell, 1984) Soil water holding capacity in the root zone (, mm depth equivalent)

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Initial stem number (nst) Foliage and root mass (ws, wf, and wr , t ha-1) appropriate to the stand at the starting age A value of fertility rate (FR): this is used due to the fact that soil nutrient status in terms of usable in quantitative models of plant growth is extremely limited because good quality information about soil physical and chemical properties of plantation and forested areas are scarce, moreover no simple relationship between state measure of soil fertility and tree growth. Nutrient availability depends on biogeochemical cycling (Waring and Schlesinger, 1985) particularly in relation to nitrogen. There is also need of expert knowledge in chemical analysis that lead to FR 5.4. Parameters Parameter values required are the constants (as) and coefficients (ns) of the allometric equations for stem mass (including balk) in terms of diameter. 5.4.1. Canopy quantum efficiency and fertility This is a change from the original Landsberg & Waring (1997) formulation of the model. The equation used is:

c = 0(fNo + (1- fNo) FR)

0: is the maximum value of QE, at FR = 1 and 0 fNo gives the minimum value of QE at a very low FR Based on experience with the model a value of 0.5 for fNo has proved suitable when calibrating the model against growth data from a range of soil types (range of FR). Temperature response function (fT) has proved to be an essential part of the model. Since when it was not included in the model, modeled GPP values were generally more than twice those measured, but when applied, it resulted in reasonable correspondence between measured and modeled values of GPP. This shows that in some part of the world (Sweden, Finland...) temperatures completely determine the growing season.

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5.5. Calibration procedures and results 5.5.1. Calibration 3 PG is calibrated by fitting to individual sets of observational data: biomass production is determined by QE, the FR, weather conditions and soil moisture holding capacity at the site. The calibration procedure involves running the model and comparing output with observed values. Parameter values are then adjusted to improve the fit and the model re-run. Successive adjustments lead, in most cases, to good fits between observed and simulated variable values. The more observed/ measured data available, the more precise will be the calibration. In the rare cases that root mass values are also available these provide an additional constraint. Calibration of soil water balance against measurements of soil water content provides the opportunity to adjust the parameter controlling the rate of change of stomatal conductance with VPD (Ewers et al., 2000). A number of parameter values can be varied to alter the output of the model but the normal procedure is to use standard default or the best available empirical values for as many parameters as possible. Values of constant (as) and power (ns) of allometric equation provide the estimates of stem diameter when the equation (1) is resolved for B from mean stem mass they should be based on experimental data. The non linearity of the equations cause significant differences on the values of B, basal area and stand volume for small variations in the parameter values therefore as and ns should not be altered once established. Uncertainty of FR is treated by adjusting parameter within limits of 0.1 or 0.2 units to match observed and simulated biomass production. 5.5.2. Results for a range of species In all cases it was possible to simulate observed data with useful accuracy. 3-PG was fitted to the data for each species. For significant differences between region and species sensitivity analysis was used to optimize parameter values. Data from sites across environmental range from sub-tropical Africa and Australia to the northern Europe were analyzed to evaluate 3- PG model generality and parameter variation. They are divided into two sets:

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Experimental data: including spacing experiment and experiment designed to evaluate effect of fertilization, irrigation, elimination of constraint to tree growth Data derived from measurement made in commercial plantation. All data set from experiment were time series of varying length. Plantation data were much less detailed than experimental data, consisting of single set of measurement (with doubtful accuracy). Standard management of experimental plantation is greater than that of commercial plantation. Climatic condition experienced are shown in tables, showing weather data, average to, total annually short wave radiant energy income. Parameter values for data sets like FR were based on physical characteristics of soil chemical analysis when available and advice from forest officer. SLA value was obtained from various sources in literature. Litter fall values were obtained from best information available. Stem number were available for all experiments, but initials were missing for commercial plantations. 5.5.3. Tests against independent data Independent set of measurements were used to test whether the model can predict stand growth when it has not been fitted to the data. Model performance was evaluated at three level of severity. Test against data from the same site: different treatment were applied; (fertilization / irrigation) and (no fertilization / no irrigation). Results are evaluated in terms of r2 (denoting the proportion of variance in the observed values accounted for by simulated value) and bias. Test on data from different site: a similar experiment is carried out on the same species plantation in different climatic condition. In these first two tests r2 0.9 showing high correspondence of simulation to observation. Test against results from spacing experiment: data from spacing thinning experiment at a different location to the fertilization/irrigation experiment were tested. Here r2 is lower but results are not biased.

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5.6. Discussion Species considered in this paper grow in a wide range of environmental condition. Good data sets of periodical measurements are necessary for 3- PG calibration, since once well calibrated it can produce observed data measurement accurately. Test against data prove that 3-PG is robust and reliable and can be used with confidence to predict growth where tree have not been grown (evaluation of site productivity). The model can be used to explore the influence of climatic parameters (VPD, to limits, drought) on the distribution of productivity of a tree species. Wide spread inadequacy of soil survey, poor understanding of relationship between soil chemical properties and growth lead to the absence of clear guideline for FR selection. 6. Understanding 3-PG using a sensitivity analysis (by L.J. Esprey, P.J. Sand, C.W. Smith) 6.1. Introduction The paper reports results of a sensitivity analysis of 3-PG using the methodology where sensitivity of several model outputs of variations of each parameter were calculated and used to assess the sensitivity of 3-PG to changes in its parameter values. The analysis helps to understand individual parameter, how processes are modeled, identify parameters that need to be accurately determined, provide insight in the limitation of the model, most importantly build confidence in the model users. Data used come from 31 Eucalyptus glandis plantations in 2 provinces in South Africa. 6.2. Overview of 3-PG The model overview principals underlying it (Landsberg 2003), however 3-PG had already been parameterized for Eucalyptus glandis in Kwa- zulu natal, South Africa using data from two contrasting trial sites. Full details of this preliminary parameterization of 3-PG for E.glandis are outlined in Gush 1999.

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6.3. Method 6.3.1. Data used in sensitivity analysis Data used came from 31 n of each trail were used in the model E.glandis research trials. Each received high quality silvicultural management comprising full weed control, adequate nutrition and blacking (replant of area where seedling had died). Long term monthly mean temperature and solar radiant were obtained from Shulze et al. (1994). Actual monthly rainfall data for the duration of each trial were used in the model prediction and selected using the driver station approach. Site factors, latitude and soil texture were directly available for each site. Maximum available soil water (sx) was derived as a product of soil water per unit soil depth. The assignment of fertility rating to site was base on whether the trial had received fertilizer or not. 6.3.2. Calculation of sensitivity analysis The relative sensitivity 1(X,p) of a given model output X with respect to a parameter, site factor and model input p is defined as: (1)

This is the change x in X produced by a change p relative to the original values of X is independent of p, and is positive or negative depending on whether an increase in p results in an increase or a decrease , respectively, in X. The relative non linearity 2 of X with respect to p by:

and this is zero if the variable X depends linearly on p. 6.4. Parameter sensitivity analysis The relative sensitivity analysis and non-linearity of stand volume (SV) and leaf area index (L) were determined for each site by running the model with reference values of each parameter (p) and with each p varied 30% at either side of its reference value. The effects of fertility rating (FR) on canopy quantum efficient (c) were taken into account in this sensitivity analysis by fNo: (parameter that controls the effects of FR on cx). The contrasting characteristics of the 2 sites

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considered are: 1o. High productivity, deep soil, law water stress and high temperature; 2o low productivity, high altitude, high water stress, and cooler temperature. 6.5. Model output as function of site factors Rainfall, temperature and maximum available soil water (sx) were varied by 30% to assess model out as a function of site factors. 6.6. Results and discussions The model performance was evaluated by comparing predicted and observed stand volume. Quality of fit is very similar in each case and the slope of the regression line was close to unity. Result of model outputs as a function of parameter values and site factors shows that: Increasing in FR increases biomass portioning to stem and decreases partitioning to roots. There is only a slight response in SV and WR to change in sx. Increasing rainfall increases both SV and WR (root biomass) at both site but increase in SV is greater on more productive site. Increasing the mean temperature reduces both SV and WR and the reduction is greater at warmer sites. The finding that variation of stand biomass data could strongly affect early stand development and had little effects on predicted growth rates was confirmed. The little effect of variation in variable soil water on SV, WR, and L at the time of initialization was found. Sensitivity measures are relatively insensitive to initial stand conditions A potential applicability of 3-PG model is the prediction of the growth at site for which there may be no preexisting assessments of site quality. The accuracy with which parameter need to be determined with is as follow: 1. Parameters with a low ranking in sensitivity can be assigned generic values. 2. Parameters with a high ranking in sensitivity must be assigned species-specific values.

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3. Parameter with a moderate sensitivity ranking but with a high non-linearity or site variation require current parameter special attention as particular circumstances of the current parameter values or site factors might be hiding high sensitivity. 6.7. Discussion Parameter sensitivity analysis highlighted the existence of 3 classes of parameters: Insensitive parameters: those that can be varied widely without affecting the studied output studied. Sensitive parameters: those whose value strongly affect the predicted outputs Non-linear parameter and linear parameters. The following parameter will require accurate determination since they have a moderate or high sensitivity ranking for stand volume SV or leaf area index L: fNo: parameter that shows the effect of site fertility on canopy quantum efficiency as: constant in stem allometric relationship ns: power in stem allometric relationship nRn: minimum fraction of net primary production to roots nRx: maximum fraction of net primary production to roots cx: maximum canopy conductance : basic wood density The ratio (Y) of net primary production gcx : maximum canopy conductance 1: specific leaf area (1) for mature trees p20: stem partitioning ratio maximum litter fall rate Tmax (maximum temperature for growth) and Topt (optimum temperature for growth) require close attention since they are highly non-linear and have a moderate sensitivity ranking. Extinction coefficient k can probably be assigned a generic value. There is also recognition of parameters in the list requiring close attention essentially affecting primary production and stem growth multiplicatively, and hence

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uncertainties in one affect the determination of values for others. The only resolution to this problem is to estimate parameters by fitting observed time-series data for foliage, stem and root biomass to data predicted by 3-PG. The relationship between SV and L as a function of selected parameter can be explained by considering how each parameter affects the calculation of SV and L. For instance increasing the biomass portioned to roots reduce both SV and L. So L and SV are sensitive to nRx and nRn. In relation to climatic factors, both SV and wR increase with increasing rainfall, but at different rates since water stress increases partitioning to root but this effect are much less than the effects of increased rainfall on PN. However increase in monthly mean temperature results in reduction of SV and wR . This paper identified parameter that need to be accurately determined in 3-PG is to be used with confidence as a management tool. It also provides information for a sound parameterization of the model for other species under other conditions. 7. Analysis of biomass accumulation and stem size distributions over long period in managed stands of Pinus sylvestris in Finland using the 3-PG model (by Joe Landsberg, Annikki Mkel, Risto Sievnen and Mikko Kukkola) 7.1. Introduction This paper presents test result of a process based model (PBM) based on 3-PG developed by Landsberg and Warring (1997) in which modification in biomass allocation routine have been done. A key assertion in new application of the model is to what extent parameters can be estimated from independent data generally available for the species and regional application, and what part of the model need to be calibrated against the actual test data. The objective of the paper was to determine the parameter values as independently as possible of the data from the site where the simulation were carried out.

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7.2. Outline of 3-PG model An outline of the model and detailed principle underlying the model are found in Landsberg and Warring (1993) but the current model application modifies some aspect of the allocation routine on the basis of empirical results while retaining the following model basic characteristics: The ratio of allocation to foliage and stem decreases with increase mean tree diameter Allocation to root depends on site quality. Allocation to roots (nr), is assumed constant over time and depends on site fertility (Vannien 2003). Allocation to foliage relative to wood (pf:w) is assumed to decreases with increasing mean breast height diameter (B) as follows:

pf:w= aBn (1)

To calculate growth, it is assumed that the fraction s of biomass allocated to wood is directed to stems, where 1- s is directed to braches and coarse roots. The carbon allocation coefficient for foliage (nf), wood (nw), and fine roots (nr) must sum to unity. They are obtained as follows: (2a) and (2b) .

Allocation to stem growth is obtained from allocation to wood growth as:

The model uses mean stand breast diameter (B) got from the mean mass of individual stems with and allometric equation between mean stem mass and B , where as, ns are empirical

coefficients. The relationship between B and ws combined with stocking density determines stand basal area at any time. 7.3. Summary of stand dynamics in the model The dynamics of stem number are calculated from removal during thinning as follows: (4) Where T(t) is stem number removed during thinning at time step t. the biomass state variables, wi (kgDMha-1) (i=s for stem mass, i=f for foliage mass and i=r for fine root mass) are calculated from three dynamic equations of the following form:

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(5) When ni is allocation to component I, si is turnover in I, and mi is a coefficient dependent on the size distribution of the stand, relating to the proportion of removed stem number to the proportion of removed mass of component i. mi is determined from data as for NPP(t) is calculated monthly as usual in the model. 7.4. Data availablility Model predictions were compared with long-term measurement data from two well managed sites in Finland. These data include stem diameter (B), height, number of stems in each stem size class, stem mass and wood density. 7.5. Estimation of model parameters Most parameter values required in calculating NPP are derived from knowledge of the behavior of species in question or default value that have been found suitable for a wide range of tree species. Fertility rating (FR) was set to a value that gives satisfactory model performance due to lack of empirical information from plots under study. Conductance parameters, specific leaf area (SLA), canopy quantum efficiency coefficient was given default values for conifers. Due to high slow growth rate of Boreal pine compared to temperate ones, the maximum age parameter related to photosynthetic efficiency and stomatal conductance was set to 500 years, hence no age effects on GPP within the age range considered. Allocation coefficient and turn over rates were estimated from data and calculations from Vannien (2003). Foliage litter fall rate are estimated according to the time the species under study hold its foliages. No natural tree mortality was assumed. Mean tree volume and basal area were calculated from stand volume, stand basal area, and sticking density given by growth and yield table from Scot pine in Finland Koivisto (1959) to fit parameters of equation (3). The effects of size distribution and removals is taken into account by means of coefficient mi dependent on the size distribution of the stand, relating the proportion of removed stem number to the proportion of removed mass of component i.

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7.6. Results The potential productivity of both sites depends only on the mean annually weather conditions and hence remained constant over time in the simulation because there was no age effect and leaf area was not limiting. Stand development in terms of Basal area, volume and mean B closely followed the measured stand characteristics in all stand. Discrepancies between basal area and volume in some case, suggest that the allometry derived from the yield table did not accurately describe the stand. The foliage mass prediction from the model although tending toward lower values than the empirical estimates, was close to them. For the analysis of stem size distribution , quantitative measure of stem size distribution are derived from fitting the Weibull probability density function to the stem size distribution data from each plot at each measurement time i.e. after thinning was carried out. This method is well described by Nanang (1998). 7.7. Discussion The simulations were generally in good agreement with the data. However, the overestimation of stem growth at the later stage of stand development observed is accompanied by foliage mass estimate that is closest to the measured of all stands. We may infer that the discrepancy is increasing as the stands age. A limitation of the 3-PG model as a practical management tool is that it produces, as output, B, whereas the diameters of stems in stands vary. In addition to simulating variables thing practices, stem size distributions are important determinant of the commercial value of a given volume of timber. 8. GENERAL DISCUSSION An understanding of the physiological processes underlying the growth of forests stands is good enough at the levels ranging from plant organs to stands, to allow scientists to write models that can predict biomass productivity accurately and reflect the effects of environmental variables.

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3-PG model is robust and is built on well established principals and can be used with confidence as a practical tool to produce results that show excellent correspondence between stand growth measurements and simulations. The major scientific challenges faced by scientists in the improvement of forest models are the questions of carbon allocation and the nutritional problem. It is important to handle these questions through experimentation under controlled conditions, using detailed physiological techniques and chemical analysis. There is a need for detailed research on uptake mechanisms, in controlled environments, with measurements designed specifically to test particular hypotheses as expressed in models. For the matter of communication between scientists and forest managers, they have to sit down and search for common ground so that model development starts with the needs of the end users 9. General conclusion A good process-based model will provide estimates of the potential productivity of sites for which no mensuration data exits, calculate realized growth and evaluate the efforts of actions such as fertilization or thinning, or the impact of pests and diseases on productivity. It should also be possible for models to make an assessment of the impacts of rising atmospheric CO2 concentrations and climate change.

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