Biomechanical design and long-term stability of trees: Morphological and wood traits involved in the balance between weight increase and the gravitropic reaction

Studies on tree biomechanical design usually focus on stem stiffness, resistance to breakage or uprooting, and elastic stability. Here we consider another biomechanical constraint related to the interaction between growth and gravity. Because stems are slender structures and are never perfectly symmetric, the increase in tree mass always causes bending movements. Given the current mechanical design of trees, integration of these movements over time would ultimately lead to a weeping habit unless some gravitropic correction occurs. This correction is achieved by asymmetric internal forces induced during the maturation of new wood.The long-term stability of a growing stem therefore depends on how the gravitropic correction that is generated by diameter growth balances the disturbance due to increasing self weight. General mechanical formulations based on beam theory are proposed to model these phenomena. The rates of disturbance and correction associated with a growth increment are deduced and expressed as a function of elementary traits of stem morphology, cross-section anatomy and wood properties. Evaluation of these traits using previously published data shows that the balance between the correction and the disturbance strongly depends on the efficiency of the gravitropic correction, which depends on the asymmetry of wood maturation strain, eccentric growth, and gradients in wood stiffness. By combining disturbance and correction rates, the gravitropic performance indicates the dynamics of stem bending during growth. It depends on stem biomechanical traits and dimensions. By analyzing dimensional effects, we show that the necessity for gravitropic correction might constrain stem allometric growth in the long-term. This constraint is compared to the requirement for elastic stability, showing that gravitropic performance limits the increase in height of tilted stem and branches. The performance of this function may thus limit the slenderness and lean of stems, and therefore the ability of the tree to capture light in a heterogeneous environment.     

Effect of tree size and competition on tension wood production over time in beech plantations and assessing relative gravitropic response with a biomechanical model

? Premise of the study: Gravitropic movements are unexpected mechanical processes that could disturb tree design allometries derived from the physics of nonliving bodies. We investigated whether the scaling law of gravitropic performance (power of -2 of stem diameter) derived from integrative biomechanical modeling is disturbed by ontogeny or environment, then discuss the silvicultural and dendroecological consequences. ? Methods: In a beech (Fagus sylvatica) plantation, four plots with different initial planting densities evolved without any intervention for 26 yr. Regular tree inventories and a silvicultural model were used to monitor competition over time in each plot. The radial production of tension wood was quantified using a cross-section of the stems at 1.30-m height, and an integrative biomechanical model computed the tree gravitropic performance over time. ? Key results: All trees developed tension wood over the whole period, with higher amounts at the youngest age, resulting in theoretical lean corrections of ca. 20-30° on the first 4 m of the stem over the whole period. The scaling law of gravitropic performance is slightly larger than the power of -2 of stem diameter. ? Conclusions: Gravitropic performance in forest ecosystems is mainly limited by size (diameter). Ontogenic acclimation of tension wood formation allows the youngest trees to be more reactive. No additional effect of spacing was found. However, silviculture influences size and, therefore, tree reactivity at a given age. Such results will be helpful for dendroecological approaches that use wood as a marker of environmental disturbances or a trait linked to plant strategies.     

Mechanosensing is involved in the regulation of autostress levels in tension wood

Key message

The level of stresses of tension wood changes during the gravitropic movement. These changes are induced by the perception of strains experienced by the tree during reorientation to the upright position.

Abstract

In most hardwood species, tension wood is produced to ensure tropic movements in radially growing organs. Tension wood exhibits internal tensional forces (autostresses) greater than those of normal wood, which enable the trunk to restore its verticality. During the gravitropic response, there is a first phase when the trunk curves upwards and a second phase when the trunk decurves to reach a final vertical and straight shape. Tension wood appears to be of varying strength, but the source of these variations remains partly undefined. We set out to assess the involvement of mechanosensing in the regulation of the strength of tension wood. Autostress levels characterise the strength of tension wood and can be indirectly estimated by measuring the associated residual longitudinal maturation strains (rlms) after the autostresses release. The higher the tension, the higher the measured associated shrinkage. To look for the involvement of mechanosensing in the regulation of tension wood strength, rlms were measured in different types of experiments in which the trunk mechanical state was modified. Results showed that (1) bigger trees exhibited higher levels of rlms, (2) there was a quantitative relationship between the rlms and the sum of strains experienced by the trunk, (3) artificial curving induced an increase in rlms and (4) in tilted staked trees, rlms increased towards negative values for 3 weeks and then remained constant. These findings are consistent evidence for the regulation of rlms values by mechanosensing. This brings new insight into gravitropism.     

A model to simulate the gravitropic response and internal stresses in trees,considering the progressive maturation of wood

Key message

The developed model of gravitropism takes non-instantaneous maturation of wood into account which enabled to correctly simulate different gravitropic phases and realistic internal stress profiles.

Abstract

A new biomechanical model of tree movement in relation to gravity (gravitropism) is proposed in this study. The modelling of the progressive maturation of wood cells is taken into account, as well as spatio-temporal variations in maturation strains (MS) and mechanical properties. MS were identified using an inverse method that allows the model to fit the gravitropic reaction observed experimentally. For this purpose, the curvature during righting movement, the geometry and the mass distribution of a two-year-old poplar tree was measured. The identified MS are higher than expected, which shows the underestimation of MS by usual measurements. By using the same mechanical parameters and MS as an input, the model gives satisfying results in terms of shape modelling for different trees up to 32 days after tree tilting. The model is able to simulate the latency phase observed in the tree righting movement, and the internal stress profile in the trunk is realistic (low compressive value in the central part of the trunk and zero stress in newly formed cells). The next development of the model will aim to simulate the end of the gravitropic phase in relation with the regulation of MS by the tree.     

The gravitropic response of poplar trunks: key roles of prestressed wood regulation and the relative kinetics of cambial growth versus wood maturation

In tree trunks, the motor of gravitropism involves radial growth and differentiation of reaction wood (Archer, 1986). The first aim of this study was to quantify the kinematics of gravitropic response in young poplar (Populus nigra x Populus deltoides, 'I4551') by measuring the kinematics of curvature fields along trunks. Three phases were identified, including latency, upward curving, and an anticipative autotropic decurving, which has been overlooked in research on trees. The biological and mechanical bases of these processes were investigated by assessing the biomechanical model of Fournier et al. (1994). Its application at two different time spans of integration made it possible to test hypotheses on maturation, separating the effects of radial growth and cross section size from those of wood prestressing. A significant correlation between trunk curvature and Fournier's model integrated over the growing season was found, but only explained 32% of the total variance. Moreover, over a week's time period, the model failed due to a clear out phasing of the kinetics of radial growth and curvature that the model does not take into account. This demonstrates a key role of the relative kinetics of radial growth and the maturation process during gravitropism. Moreover, the degree of maturation strains appears to differ in the tension woods produced during the upward curving and decurving phases. Cell wall maturation seems to be regulated to achieve control over the degree of prestressing of tension wood, providing effective control of trunk shape.     

Study of tension wood in the artificially inclined seedlings of <Emphasis Type="Italic">Koelreuteria henryi</Emphasis> Dummer and its biomechanical function of negative gravitropism

Key message

Stem reorientation is critical to tree survival. With anatomical observation and strain measurement, the tension wood formation and biomechanical behavior were studied to gain insights into tree uprighting process.

Abstract

Tension wood plays a role in maintaining the mechanical stability of angiosperm trees. Both biological and physical aspects of tension wood are essential in understanding the mechanism of trunk or branch reorientation. In this study, we worked on both tension wood formation and its biomechanical function in artificially inclined 2-year-old Koelreuteria henryi seedlings. The tension wood formation and reorientation process of the trunk last for about 3 months. With pinning method, we confirmed that at the beginning of inclination the cambial zone including the vascular cambium and the developing normal wood fibers on the upper side of the inclined trunk perceives the onset of mechanical change and starts to produce G-fibers that generate a strong contractile released growth strain (RGS) for gravitropic correction. Stronger contractile RGS and more tension wood were found at the trunk base than at the half-height, suggesting that the trunk base plays a key role in trunk uprighting of K. henryi seedlings. The eccentric cambial growth in the tension wood side increases the efficiency of gravitropic correction and the compressive strains measured in the opposite wood of some inclined seedlings also help the upright movement.

     

Wood mechanics, allometry, and life-history variation in a tropical rain forest tree community

Wood density plays a central role in the life-history variation of trees, and has important consequences for mechanical properties of wood, stem and branches, and tree architecture. Wood density, modulus of rupture, modulus of elasticity, and safety factors for buckling and bending were determined for saplings of 30 Bolivian rain forest tree species, and related to two important life-history axes: juvenile light demand and maximum adult stature. Wood density was strongly positively related to wood strength and stiffness. Species safety factor for buckling was positively related to wood density and stiffness, but tree architecture (height : diameter ratio) was the strongest determinant of mechanical safety. Shade-tolerant species had dense and tough wood to enhance survival in the understorey, whereas pioneer species had low-density wood and low safety margins to enhance growth in gaps. Pioneer and shade-tolerant species showed opposite relationships between species traits and adult stature. Light demand and adult stature affect wood properties, tree architecture and plant performance in different ways, contributing to the coexistence of rain forest species.     

Patterns of auxin distribution during gravitational induction of reaction wood in poplar and pine

Gravistimulation of tree stems affects wood development by unilaterally inducing wood with modified properties, called reaction wood. Commonly, it also stimulates cambial growth on the reaction wood side. Numerous experiments involving applications of indole-3-acetic acid (IAA) or IAA-transport inhibitors have suggested that reaction wood is induced by a redistribution of IAA around the stem. However, in planta proof for this model is lacking. Therefore, we have mapped endogenous IAA distribution across the cambial region tissues in both aspen (Populus tremula, denoted poplar) and Scots pine (Pinus sylvestris) trees forming reaction wood, using tangential cryosectioning combined with sensitive gas chromatography-mass spectrometry analysis. Moreover, we have documented the kinetics of IAA during reaction wood induction in these species. Our analysis of endogenous IAA demonstrates that reaction wood is formed without any obvious alterations in IAA balance. This is in contrast to gravitropic responses in roots and shoots where a redistribution of IAA has been documented. It is also of interest that cambial growth on the tension wood side was stimulated without an increase in IAA. Taken together, our results suggest a role for signals other than IAA in the reaction wood response, or that the gravitational stimulus interacts with the IAA signal transduction pathway.     

Modelling polymer interactions of the 'molecular Velcro' type in wood under mechanical stress

Trees withstand wind and snow loads by synthesising wood that varies greatly in mechanical properties: flexible in twigs and in the stem of the sapling, and rigid in the outer part of the mature stem. The ‘molecular Velcro’ model of Keckes et al. [2003. Cell-wall recovery after irreversible deformation of wood. Nat. Mater. 2, 810–814] permits the simulation of the tensile properties of water-saturated wood as found in living trees. A basic feature of this model is the presence of non-covalent interactions between hemicellulose chains attached to adjacent cellulose microfibrils, which are disrupted above a threshold level of interfibrillar shear. However, other evidence does not confirm the importance of hemicellulose–hemicellulose association in the cohesion of the interfibrillar matrix. Here, we present an alternative model in which hemicellulose chains bridging continuously from one microfibril aggregate (macrofibril) to the next provide most of the cohesion. We show that such hemicellulose bridges exist and that the stripping of the bridging chains from the cellulose surfaces under the tensile stress component normal to the macrofibrils can provide an alternative triggering mechanism for shear deformation between one macrofibril and the next. When one macrofibril then slides past another, a domain of the wood cell wall can extend but simultaneously it twists until the spacing between macrofibrils is reduced again and contact through hemicelluloses bridges is restored. Overall deformation therefore takes place through a series of local stick–slip events involving temporary twisting of small domains within the wood cell wall. Modelled load–deformation curves for this modified ‘molecular Velcro’ model are similar, although not identical, to those for the original model. However, the mechanism is different and more consistent with current views of the structure of wood cell walls, providing a framework within which the developmental control of rigidity in wood synthesised in different parts of a tree may be considered.     

Effect of circumferential heterogeneity of wood maturation strain, modulus of elasticity and radial growth on the regulation of stem orientation in trees

Active mechanisms of re-orientation are necessary to maintain the verticality of tree stems. They are achieved through the production of reaction wood, associated with circumferential variations of three factors related to cambial activity: maturation strain, longitudinal modulus of elasticity (MOE) and eccentric growth. These factors were measured on 17 mature trees from different botanical families and geographical locations. Various patterns of circumferential variation of these factors were identified. A biomechanical analysis based on beam theory was performed to quantify the individual impact of each factor. The main factor of re-orientation is the circumferential variation of maturation strains. However, this factor alone explains only 57% of the re-orientations. Other factors also have an effect through their interaction with maturation strains. Eccentric growth is generally associated with heterogeneity of maturation strains, and has an important complementary role, by increasing the width of wood with high maturation strain. Without this factor, the efficiency of re-orientations would be reduced by 31% for angiosperms and 26% for gymnosperms. In the case of angiosperms, MOE is often larger in tension wood than in normal wood. Without these variations, the efficiency of re-orientations would be reduced by 13%. In the case of gymnosperm trees, MOE of compression wood is lower than that of normal wood, so that re-orientation efficiency would be increased by 24% without this factor of variations.     

Root movements: Towards an understanding through attempts to model the processes involved

Roots have the ability to change the direction of their forward growth. Sometimes these directional changes are rapid, as in mutations, or they are slower, as in tropisms. The gravitational force is always present and roots have an efficient graviperception mechanism which enables them to initiate gravitropic movements. In trying to model and simulate the course of gravitropic root movements with a view to analyse the component processes, the following aspects of the plant's interaction with gravity have been considered: (1) The level of organization (organism, organ, cell) at which the movement process is expressed; (2) whether the gravity stimulation event is dynamic or static (i.e. whether or not physiologically significant displacements take place with respect to the gravity vector); (3) the sub-systems involved in movement and the processes which they regulate; (4) the mathematical characterization of the relevant sub-systems. A further allied topic is the nature of nutational movements and whether they are linked with gravitropic movements in some way. In considering how they can best be modelled, two types of nutational movements are proponed: stochastic nutation and circumnutation. Most, if not all, natural movements developed in response to static gravistimulation can be viewed as gravimorphisms. This applies at the levels of cell, organ and organism. However, when a system at any one of these levels experiences dynamic gravistimulation, because of its inherent homeostatic properties, it is induced to regenerate a state similar to that previously held. Thus, gravitropism is a regenerative gravimorphic process at the level of the organ.     

In the lack of extreme pioneers: trait relationships and ecological strategies of 66 subtropical tree species

Aims Despite the growing interest in the topic of functional ecology, there are still forest regions that have not been examined, as most work has been done in the tropics. Unresolved issues include the strength of a growth-mortality trade-off in trees (originally identified for seedlings) and the nature of the association between plant traits and vital rates, if any. Our objectives were to examine whether (i) ecological strategies in South American mixed forests are organized along the fast competitor × slow stress-tolerator and height gradients as the main strategy axes depicted in the overall trait and vital rate correlation structure, and (ii) a tentative path model we proposed can explain the patterns of covariation among traits and vital rates.Methods We studied a different habitat (subtropical mixed conifer-hardwood forests) and region (Brazilian Atlantic Forest) from the majority of related studies in forests, carried out in the Neotropical region. Data on total height, stem slenderness, crown depth, wood density, specific leaf area, leaf and seed length, seed dispersal mode, annual mortality, diameter relative growth rate and relative growth rate under favorable conditions were measured in southern Brazil for 66 tree species. Data were subjected to principal components analysis and path analysis. Restricted data on saplings and treelets were analyzed through correlation.Important findings Studied traits were reduced to four principal components. Principal components analysis produced axes that fit the resource acquisition versus resource-conservation and the height-mortality trade-offs, although the former was split into two distinct axes. Seed size and seed dispersal mode appeared independently of these axes. A path model showed that leaf length and specific leaf area caused direct changes in trunk slenderness and, indirectly through growth, affected mortality. Expected trade-offs between growth and survivorship and between wood density and stem slenderness trade-offs were not found. This may result from the lack of extreme pioneers and over-representation of slow-growing hardwood species found in Atlantic subtropical forests of South America. This suggests that the fastest growing species in the region do not grow so fast as to compromise wood density and survivorship, but grow fast enough to benefit from increased size. Relationships between traits and vital rates seem to be mediated by the assembly process of regional floras, and the relative importance of traits like SLA and wood density may vary between floristic regions.     

Numerical modelling of shape regulation and growth stresses in trees

The main objective of this paper is to present the results of a study of the interactions between the growth and design of a tree with regards to biomechanical factors at the plant level. A numerical incremental model dedicated to the calculation of tree mechanical behaviour has been integrated in the plant architecture simulation software AMAPpara. At any stage of tree growth, a new equilibrium was calculated considering the weight increment applied on the structure, i.e. the mass of new wood layers and vegetative elements, as well as the biomechanical reaction caused by cell maturation strains in both normal and reaction wood. The resulting incremental displacements allowed the tree shape to be modified. The field of growth stresses was calculated within the stem, using a cumulative process taking into consideration the past history of each growth ring. The simulation results of trunk and branch shape, as well as internal stresses, were examined after consideration of different growth strategies. A block of trees was also simulated in order to show the influence of spatial competition on stem curvature and the variability in growth stress.     

Wood transformation in dead-standing trees in the forest-tundra of Central Siberia

Changes in the composition of wood organic matter in dead-standing spruce and larch trees depending on the period after their death have been studied in the north of Central Siberia. The period after tree death has been estimated by means of cross-dating. The results show that changes in the composition of wood organic matter in 63% of cases are contingent on tree species. Wood decomposition in dead-standing trees is accompanied by an increase in the contents of alkali-soluble organic compounds. Lignin oxidation in larch begins approximately 80 years after tree death, whereas its transformation in spruce begins not earlier than after 100 years. In the forest-tundra of Central Siberia, the rate of wood organic matter transformation in dead-standing trees is one to two orders of magnitude lower than in fallen wood, which accounts for their role as a long-term store of carbon and mineral elements in these ecosystems.     

树木高生长限制的几个假说

树木生长到一定年龄后高生长停滞, 对这一现象的解释存在很多争议。成熟假说认为树木顶端分生组织分裂活性下降导致树木高生长减慢。营养限制假说认为土壤中营养元素(特别是氮素)在植物活体或枯落物中积累使土壤中可利用的养分含量降低, 细根生物量增加和叶片光合能力下降导致了地上部分生长的减缓。呼吸假说认为边材呼吸消耗随个体发育的增加使投入到高生长的碳减少。水力限制假说认为水分运输阻力随高度增加而增加导致了叶片总光合碳同化下降, 分配到高生长的生物量减少。树木发展假说认为植物用多种调节机制克服随个体发育增加的水力阻力, 包括叶片结构和生理特征的变化, 叶/边材面积比降低, 边材渗透性和树干储水能力的增加等。水力限制假说得到了较多的关注, 对不同高度树木的叶比导率、光合特征和树干生长量等测定结果支持这一假说。但对这一假说 也存在很多的争议, 主要表现在: 水力阻力是否确实随高度的增加而增加, 水力阻力的分布, 补偿机制的作用和生物量分配转变等。本文综述了树木高生长限制的4个假说以及争论的焦点, 并总结了目前研究的热点问题和今后的研究方向。     

树木高生长限制的几个假说

树木生长到一定年龄后高生长停滞,对这一现象的解释存在很多争议.成熟假说认为树木顶端分生组织分裂活性下降导致树木高生长减慢.营养限制假说认为土壤中营养元素(特别是氮素)在植物活体或枯落物中积累使土壤中可利用的养分含量降低,细根生物量增加和叶片光合能力下降导致了地上部分生长的减缓.呼吸假说认为边材呼吸消耗随个体发育的增加使投入到高生长的碳减少.水力限制假说认为水分运输阻力随高度增加而增加导致了叶片总光合碳同化下降,分配到高生长的生物量减少.树木发展假说认为植物用多种调节机制克服随个体发育增加的水力阻力,包括叶片结构和生理特征的变化,叶/边材面积比降低,边材渗透性和树干储水能力的增加等.水力限制假说得到了较多的关注,对不同高度树木的叶比导率、光合特征和树干生长量等测定结果支持这一假说.但对这一假说也存在很多的争议,主要表现在:水力阻力是否确实随高度的增加而增加,水力阻力的分布,补偿机制的作用和生物量分配转变等.本文综述了树木高生长限制的4个假说以及争论的焦点,并总结了目前研究的热点问题和今后的研究方向.     

Bending of apricot tree branches under the weight of axillary growth: test of a mechanical model with experimental data

Stem orientation is an important factor for fruit tree growth and branching habit since it influences fruit production as well as training practices. A mechanical model of the bending of a stem under axillary load was written and evaluated using experimental data on apricot trees (Prunus armeniaca L.). A set of 15 1-year-old stems of various shapes was observed during the early stage of the growing season when radial growth is still negligible and the loading of the stem increases considerably. The structural modulus of elasticity (MOE) of the stems was estimated through in situ bending tests assuming homogeneous material behaviour. The effect of viscoelasticity was observed through creep tests performed on similar stems during winter. Inputs of the model are initial shape, initial diameter, and final load, defined at various positions along the stem. The final shape was simulated based on different mechanical assumptions, and compared to observations. Assuming small deflections resulted in an underestimate of the mean slope variation of 48%, accounting for large displacements reduced this underestimate to 29% and accounting for viscoelasticity reduced it further to 14%. An adjustment of the structural MOE to fit the final shape led to an excellent fit of the data in most cases, the residual errors for some axes being attributed to material heterogeneity. The use of biomechanical models to predict the shape of fruit trees based on growth parameters, provided adequate assumptions are made, is discussed.     

Modeling decay rates of dead wood in a neotropical forest

Variation of dead wood decay rates among tropical trees remains one source of uncertainty in global models of the carbon cycle. Taking advantage of a broad forest plot network surveyed for tree mortality over a 23-year period, we measured the remaining fraction of boles from 367 dead trees from 26 neotropical species widely varying in wood density (0.23–1.24 g cm⁻³) and tree circumference at death time (31.5–272.0 cm). We modeled decay rates within a Bayesian framework assuming a first order differential equation to model the decomposition process and tested for the effects of forest management (selective logging vs. unexploited), of mode of death (standing vs. downed) and of topographical levels (bottomlands vs. hillsides vs. hilltops) on wood decay rates. The general decay model predicts the observed remaining fraction of dead wood (R ² = 60%) with only two biological predictors: tree circumference at death time and wood specific density. Neither selective logging nor local topography had a differential effect on wood decay rates. Including the mode of death into the model revealed that standing dead trees decomposed faster than downed dead trees, but the gain of model accuracy remains rather marginal. Overall, these results suggest that the release of carbon from tropical dead trees to the atmosphere can be simply estimated using tree circumference at death time and wood density.     

Physical strain-mediated microtubule reorientation in the epidermis of gravitropically or phototropically stimulated maize coleoptiles

During gravitropic and phototropic curvature of the maize coleoptile, the cortical microtubules (MTs) adjacent to the outer epidermal cell wall assume opposite orientations at the two sides of the organ. Starting from a uniformly random pattern during straight growth in darkness, the MTs reorientate perpendicularly to the organ axis at the outer (faster growing) side and parallel to the organ axis at the inner (slower growing) side. As similar reorientations can be induced during straight growth by increasing or decreasing the effective auxin concentration, it has been proposed that these reorientations may be used as a diagnostic test for assessing the auxin status of the epidermal cells during tropic curvature. This idea was tested by determining the MT orientations in the coleoptile of intact maize seedlings in which the gravitropic or phototropic curvature was prevented or inversed by an appropriate mechanical counterforce. Forces that just prevented the coleoptile from curving in a gravity or light field prevented reorientations of the MTs. Forces strong enough to overcompensate the tropic stimuli by enforcing curvature in the opposite direction induced reorientations of the MTs opposite to those produced by tropic stimulation. These results show that the MTs at the outer surface of the coleoptile respond to changes in mechanical tissue strain rather than to gravitropic or phototropic stimuli and associated changes at the level of auxin or any other element in the signal transduction chain between perception of tropic stimuli and asymmetric growth response. It is proposed that cortical MTs can act as strain gauges in a positive feed-back regulatory circle utilized for amplification and stabilization of environmentally induced changes in the direction of elongation growth.     

Using an evapo-transpiration model (ETpN) to predict the risk and expression of symptoms of pine wilt disease (PWD) across Europe

The pine wood nematode (PWN) Bursaphelenchus xylophilus is the causal agent of pine wilt disease (PWD), a xylem restricting disease of pine trees. PWN, a native of North America where it very rarely kills native pine trees, has spread internationally killing host trees in China, Japan, Korea, Taiwan and Portugal, with isolated incursions into Spain. Based on the locations where tree mortality has been recorded, it appears that pine trees growing in hot, dry conditions are more susceptible to pine wilt disease. This paper describes the ETpN model, an evapo-transpiration model (previously developed by Forest Research), which has been modified to incorporate the presence of PWN inside a tree and which predicts the regions of Europe that are likely to succumb to PWD. ETpN acts independently of the vector beetle (Monochamus spp.), predicting the likelihood of PWD on the assumption that a tree in a particular region has already been infested by the pine wood nematode. Different regions across Europe are included to investigate and demonstrate how different climates affect PWD incidence significantly. Simplified, “lite” and latency models have been developed to allow a non-specialist user to determine respectively the risk of PWD at a particular location and the likelihood of delays (latency) in expression of wilt symptoms.