Stochastic structure and individual-tree growth models

The majority of past and current individual-tree growth modelling methodologies have failed to characterise and incorporate structured stochastic components. Rather, they have relied on deterministic predictions or have added an unstructured random component to predictions. In particular, spatial stochastic structure has been neglected, despite being present in most applications of individual-tree growth models. Spatial stochastic structure (also called spatial dependence or spatial autocorrelation) eventuates when spatial influences such as competition and micro-site effects are not fully captured in models. Temporal stochastic structure (also called temporal dependence or temporal autocorrelation) eventuates when a sequence of measurements is taken on an individual-tree over time, and variables explaining temporal variation in these measurements are not included in the model. Nested stochastic structure eventuates when measurements are combined across sampling units and differences among the sampling units are not fully captured in the model. This review examines spatial, temporal, and nested stochastic structure and instances where each has been characterised in the forest biometry and statistical literature. Methodologies for incorporating stochastic structure in growth model estimation and prediction are described. Benefits from incorporation of stochastic structure include valid statistical inference, improved estimation efficiency, and more realistic and theoretically sound predictions. It is proposed in this review that individual-tree modelling methodologies need to characterise and include structured stochasticity. Possibilities for future research are discussed. (C) 2001 Elsevier Science B.V. All rights reserved.

Predicting the growth response to thinning for Scots pine stands using individual-tree growth models

Summary

Individual-tree basal area growth models for scots pine, pubescent birch and Norway spruce on drained peatlands in Finland

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Root growth models: towards a new generation of continuous approaches

Models of root system growth emerged in the early 1970s, and were based on mathematical representations of root length distribution in soil. The last decade has seen the development of more complex architectural models and the use of computer-intensive approaches to study developmental and environmental processes in greater detail. There is a pressing need for predictive technologies that can integrate root system knowledge, scaling from molecular to ensembles of plants. This paper makes the case for more widespread use of simpler models of root systems based on continuous descriptions of their structure. A new theoretical framework is presented that describes the dynamics of root density distributions as a function of individual root developmental parameters such as rates of lateral root initiation, elongation, mortality, and gravitropsm. The simulations resulting from such equations can be performed most efficiently in discretized domains that deform as a result of growth, and that can be used to model the growth of many interacting root systems. The modelling principles described help to bridge the gap between continuum and architectural approaches, and enhance our understanding of the spatial development of root systems. Our simulations suggest that root systems develop in travelling wave patterns of meristems, revealing order in otherwise spatially complex and heterogeneous systems. Such knowledge should assist physiologists and geneticists to appreciate how meristem dynamics contribute to the pattern of growth and functioning of root systems in the field.

Calibration and evaluation of individual-based models using Approximate Bayesian Computation

This paper investigates the feasibility of using approximate Bayesian computation (ABC) to calibrate and evaluate complex individual-based models (IBMs). As ABC evolves, various versions are emerging, but here we only explore the most accessible version, rejection-ABC. Rejection-ABC involves running models a large number of times, with parameters drawn randomly from their prior distributions, and then retaining the simulations closest to the observations. Although well-established in some fields, whether ABC will work with ecological IBMs is still uncertain. Rejection-ABC was applied to an existing 14-parameter earthworm energy budget IBM for which the available data consist of body mass growth and cocoon production in four experiments. ABC was able to narrow the posterior distributions of seven parameters, estimating credible intervals for each. ABC’s accepted values produced slightly better fits than literature values do. The accuracy of the analysis was assessed using cross-validation and coverage, currently the best available tests. Of the seven unnarrowed parameters, ABC revealed that three were correlated with other parameters, while the remaining four were found to be not estimable given the data available. It is often desirable to compare models to see whether all component modules are necessary. Here we used ABC model selection to compare the full model with a simplified version which removed the earthworm’s movement and much of the energy budget. We are able to show that inclusion of the energy budget is necessary for a good fit to the data. We show how our methodology can inform future modelling cycles, and briefly discuss how more advanced versions of ABC may be applicable to IBMs. We conclude that ABC has the potential to represent uncertainty in model structure, parameters and predictions, and to embed the often complex process of optimizing an IBM’s structure and parameters within an established statistical framework, thereby making the process more transparent and objective.

Predicting how many animals will be where: how to build, calibrate and evaluate individual-based models

Individual-based models (IBMs) can simulate the actions of individual animals as they interact with one another and the landscape in which they live. When used in spatially-explicit landscapes IBMs can show how populations change over time in response to management actions. For instance, IBMs are being used to design strategies of conservation and of the exploitation of fisheries, and for assessing the effects on populations of major construction projects and of novel agricultural chemicals. In such real world contexts, it becomes especially important to build IBMs in a principled fashion, and to approach calibration and evaluation systematically. We argue that insights from physiological and behavioural ecology offer a recipe for building realistic models, and that Approximate Bayesian Computation (ABC) is a promising technique for the calibration and evaluation of IBMs. IBMs are constructed primarily from knowledge about individuals. In ecological applications the relevant knowledge is found in physiological and behavioural ecology, and we approach these from an evolutionary perspective by taking into account how physiological and behavioural processes contribute to life histories, and how those life histories evolve. Evolutionary life history theory shows that, other things being equal, organisms should grow to sexual maturity as fast as possible, and then reproduce as fast as possible, while minimising per capita death rate. Physiological and behavioural ecology are largely built on these principles together with the laws of conservation of matter and energy. To complete construction of an IBM information is also needed on the effects of competitors, conspecifics and food scarcity; the maximum rates of ingestion, growth and reproduction, and life-history parameters. Using this knowledge about physiological and behavioural processes provides a principled way to build IBMs, but model parameters vary between species and are often difficult to measure. A common solution is to manually compare model outputs with observations from real landscapes and so to obtain parameters which produce acceptable fits of model to data. However, this procedure can be convoluted and lead to over-calibrated and thus inflexible models. Many formal statistical techniques are unsuitable for use with IBMs, but we argue that ABC offers a potential way forward. It can be used to calibrate and compare complex stochastic models and to assess the uncertainty in their predictions. We describe methods used to implement ABC in an accessible way and illustrate them with examples and discussion of recent studies. Although much progress has been made, theoretical issues remain, and some of these are outlined and discussed.

Data collapse, scaling functions, and analytical solutions of generalized growth models

We consider a nontrivial one-species population dynamics model with finite and infinite carrying capacities. Time-dependent intrinsic and extrinsic growth rates are considered in these models. Through the model per capita growth rate we obtain a heuristic general procedure to generate scaling functions to collapse data into a simple linear behavior even if an extrinsic growth rate is included. With this data collapse, all the models studied become independent from the parameters and initial condition. Analytical solutions are found when time-dependent coefficients are considered. These solutions allow us to perceive nontrivial transitions between species extinction and survival and to calculate the transition's critical exponents. Considering an extrinsic growth rate as a cancer treatment, we show that the relevant quantity depends not only on the intensity of the treatment, but also on when the cancerous cell growth is maximum.

Fluid-mobile trace element constraints on the role of slab melting and implications for Archaean crustal growth models

We use published and new trace element data to identify element ratios which discriminate between arc magmas from the supra-subduction zone mantle wedge and those formed by direct melting of subducted crust (i.e. adakites). The clearest distinction is obtained with those element ratios which are strongly fractionated during refertilisation of the depleted mantle wedge, ultimately reflecting slab dehydration. Hence, adakites have significantly lower Pb/Nd and B/Be but higher Nb/Ta than typical arc magmas and continental crust as a whole. Although Li and Be are also overenriched in continental crust, behaviour of Li/Yb and Be/Nd is more complex and these ratios do not provide unique signatures of slab melting. Archaean tonalite-trondhjemite-granodiorites (TTGs) strongly resemble ordinary mantle wedge-derived arc magmas in terms of fluid-mobile trace element content, implying that they-did not form by slab melting but that they originated from mantle which was hydrated and enriched in elements lost from slabs during prograde dehydration. We suggest that Archaean TTGs formed by extensive fractional crystallisation from a mafic precursor. It is widely claimed that the time between the creation and subduction of oceanic lithosphere was significantly shorter in the Archaean (i.e. 20 Ma) than it is today. This difference was seen as an attractive explanation for the presumed preponderance of adakitic magmas during the first half of Earth's history. However, when we consider the effects of a higher potential mantle temperature on the thickness of oceanic crust, it follows that the mean age of oceanic lithosphere has remained virtually constant. Formation of adakites has therefore always depended on local plate geometry and not on potential mantle temperature.

Dynamical behaviour on the parameter space: new populational growth models proportional to beta densities

We present new populational growth models, generalized logistic models which are proportional to beta densities with shape parameters p and 2, where p > 1, with Malthusian parameter r. The complex dynamical behaviour of these models is investigated in the parameter space (r, p), in terms of topological entropy, using explicit methods, when the Malthusian parameter r increases. This parameter space is split into different regions, according to the chaotic behaviour of the models.

Dynamical analysis in growth models: Blumberg’s equation

We present a new dynamical approach to the Blumberg's equation, a family of unimodal maps. These maps are proportional to Beta(p, q) probability densities functions. Using the symmetry of the Beta(p, q) distribution and symbolic dynamics techniques, a new concept of mirror symmetry is defined for this family of maps. The kneading theory is used to analyze the effect of such symmetry in the presented models. The main result proves that two mirror symmetric unimodal maps have the same topological entropy. Different population dynamics regimes are identified, when the intrinsic growth rate is modified: extinctions, stabilities, bifurcations, chaos and Allee effect. To illustrate our results, we present a numerical analysis, where are demonstrated: monotonicity of the topological entropy with the variation of the intrinsic growth rate, existence of isentropic sets in the parameters space and mirror symmetry.

From weak Allee effect to no Allee effect in Richards’ growth models

Population dynamics have been attracting interest since many years. Among the considered models, the Richards’ equations remain one of the most popular to describe biological growth processes. On the other hand, Allee effect is currently a major focus of ecological research, which occurs when positive density dependence dominates at low densities. In this chapter, we propose the dynamical study of classes of functions based on Richards’ models describing the existence or not of Allee effect. We investigate bifurcation structures in generalized Richards’ functions and we look for the conditions in the (β, r) parameter plane for the existence of a weak Allee effect region. We show that the existence of this region is related with the existence of a dovetail structure. When the Allee limit varies, the weak Allee effect region disappears when the dovetail structure also disappears. Consequently, we deduce the transition from the weak Allee effect to no Allee effect to this family of functions. To support our analysis, we present fold and flip bifurcation curves and numerical simulations of several bifurcation diagrams.

Symbolic dynamics and big bang bifurcation in Weibull-Gompertz-Fréchet's growth models

In this paper, motivated by the interest and relevance of the study of tumor growth models, a central point of our investigation is the study of the chaotic dynamics and the bifurcation structure of Weibull-Gompertz-Fréchet's functions: a class of continuousdefined one-dimensional maps. Using symbolic dynamics techniques and iteration theory, we established that depending on the properties of this class of functions in a neighborhood of a bifurcation point PBB, in a two-dimensional parameter space, there exists an order regarding how the infinite number of periodic orbits are born: the Sharkovsky ordering. Consequently, the corresponding symbolic sequences follow the usual unimodal kneading sequences in the topological ordered tree. We verified that under some sufficient conditions, Weibull-Gompertz-Fréchet's functions have a particular bifurcation structure: a big bang bifurcation point PBB. This fractal bifurcations structure is of the so-called "box-within-a-box" type, associated to a boxe ω1, where an infinite number of bifurcation curves issues from. This analysis is done making use of fold and flip bifurcation curves and symbolic dynamics techniques. The present paper is an original contribution in the framework of the big bang bifurcation analysis for continuous maps.

Explosion birth and extinction: Double big bang bifurcations and Allee effect in Tsoularis-Wallace's growth models

This work concerns dynamics and bifurcations properties of a new class of continuous-defined one-dimensional maps: Tsoularis-Wallace's functions. This family of functions naturally incorporates a major focus of ecological research: the Allee effect. We provide a necessary condition for the occurrence of this phenomenon of extinction. To establish this result we introduce the notions of Allee's functions, Allee's effect region and Allee's bifurcation curve. Another central point of our investigation is the study of bifurcation structures for this class of functions, in a three-dimensional parameter space. We verified that under some sufficient conditions, Tsoularis-Wallace's functions have particular bifurcation structures: the big bang and the double big bang bifurcations of the so-called "box-within-a-box" type. The double big bang bifurcations are related to the existence of flip codimension-2 points. Moreover, it is verified that these bifurcation cascades converge to different big bang bifurcation curves, where for the corresponding parameter values are associated distinct kinds of boxes. This work contributes to clarify the big bang bifurcation analysis for continuous maps and understand their relationship with explosion birth and extinction phenomena.

The Role of Institutions in Cross-Section Income and Panel Data Growth Models: A Deeper Investigation on the Weakness and Proliferation of Instruments

This paper investigates the role of institutions in determining per capita income levels and growth. It contributes to the empirical literature by using different variables as proxies for institutions and by developing a deeper analysis of the issues arising from the use of weak and too many instruments in per capita income and growth regressions. The cross-section estimation suggests that institutions seem to matter, regardless if they are the only explanatory variable or are combined with geographical and integration variables, although most models suffer from the issue of weak instruments. The results from the growth models provides some interesting results: there is mixed evidence on the role of institutions and such evidence is more likely to be associated with law and order and investment profile; government spending is an important policy variable; collapsing the number of instruments results in fewer significant coefficients for institutions.

Stability of Growth Models with Generalised Lag Structures

This paper considers the lag structures of dynamic models in economics, arguing that the standard approach is too simple to capture the complexity of actual lag structures arising, for example, from production and investment decisions. It is argued that recent (1990s) developments in the the theory of functional differential equations provide a means to analyse models with generalised lag structures. The stability and asymptotic stability of two growth models with generalised lag structures are analysed. The paper concludes with some speculative discussion of time-varying parameters.