1 Graphical modeling using L-systems.- 1.1 Rewriting systems.- 1.2 DOL-systems.- 1.3 Turtle interpretation of strings.- 1.4 Synthesis of DOL-systems.- 1.4.1 Edge rewriting.- 1.4.2 Node rewriting.- 1.4.3 Relationship between edge and node rewriting.- 1.5 Modeling in three dimensions.- 1.6 Branching structures.- 1.6.1 Axial trees.- 1.6.2 Tree OL-systems.- 1.6.3 Bracketed OL-systems.- 1.7 Stochastic L-systems.- 1.8 Context-sensitive L-systems.- 1.9 Growth functions.- 1.10 Parametric L-systems.- 1.10.1 Parametric OL-systems.- 1.10.2 Parametric 2L-systems.- 1.10.3 Turtle interpretation of parametric words.- 2 Modeling of trees.- 3 Developmental models of herbaceous plants.- 3.1 Levels of model specification.- 3.1.1 Partial L-systems.- 3.1.2 Control mechanisms in plants.- 3.1.3 Complete models.- 3.2 Branching patterns.- 3.3 Models of inflorescences.- 3.3.1 Monopodial inflorescences.- 3.3.2 Sympodial inflorescences.- 3.3.3 Polypodial inflorescences.- 3.3.4 Modified racemes.- 4 Phyllotaxis.- 4.1 The planar model.- 4.2 The cylindrical model.- 5 Models of plant organs.- 5.1 Predefined surfaces.- 5.2 Developmental surface models.- 5.3 Models of compound leaves.- 6 Animation of plant development.- 6.1 Timed DOL-systems.- 6.2 Selection of growth functions.- 6.2.1 Development of nonbranching filaments.- 6.2.2 Development of branching structures.- 7 Modeling of cellular layers.- 7.1 Map L-systems.- 7.2 Graphical interpretation of maps.- 7.3 Microsorium linguaeforme.- 7.4 Dryopteris thelypteris.- 7.5 Modeling spherical cell layers.- 7.6 Modeling 3D cellular structures.- 8 Fractal properties of plants.- 8.1 Symmetry and self-similarity.- 8.2 Plant models and iterated function systems.- Epilogue.- Appendix A Software environment for plant modeling.- A.1 A virtual laboratory in botany.- A.2 List of laboratory programs.- Appendix B About the figures.- Turtle interpretation of symbols.

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The Algorithmic Beauty of Plants

Plant functional traits are the features (morphological, physiological, phenological) that represent ecological strategies and determine how plants respond to environmental factors, affect other trophic levels and influence ecosystem properties. Variation in plant functional traits, and trait syndromes, has proven useful for tackling many important ecological questions at a range of scales, giving rise to a demand for standardised ways to measure ecologically meaningful plant traits. This line of research has been among the most fruitful avenues for understanding ecological and evolutionary patterns and processes. It also has the potential both to build a predictive set of local, regional and global relationships between plants and environment and to quantify a wide range of natural and human-driven processes, including changes in biodiversity, the impacts of species invasions, alterations in biogeochemical processes and vegetation–atmosphere interactions. The importance of these topics dictates the urgent need for more and better data, and increases the value of standardised protocols for quantifying trait variation of different species, in particular for traits with power to predict plant- and ecosystem-level processes, and for traits that can be measured relatively easily. Updated and expanded from the widely used previous version, this handbook retains the focus on clearly presented, widely applicable, step-by-step recipes, with a minimum of text on theory, and not only includes updated methods for the traits previously covered, but also introduces many new protocols for further traits. This new handbook has a better balance between whole-plant traits, leaf traits, root and stem traits and regenerative traits, and puts particular emphasis on traits important for predicting species’ effects on key ecosystem properties. We hope this new handbook becomes a standard companion in local and global efforts to learn about the responses and impacts of different plant species with respect to environmental changes in the present, past and future.

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New handbook for standardised measurement of plant functional traits worldwide

Wood performs several essential functions in plants, including mechanically supporting aboveground tissue, storing water and other resources, and transporting sap. Woody tissues are likely to face physiological, structural and defensive trade-offs. How a plant optimizes among these competing functions can have major ecological implications, which have been under-appreciated by ecologists compared to the focus they have given to leaf function. To draw together our current understanding of wood function, we identify and collate data on the major wood functional traits, including the largest wood density database to date (8412 taxa), mechanical strength measures and anatomical features, as well as clade-specific features such as secondary chemistry. We then show how wood traits are related to one another, highlighting functional trade-offs, and to ecological and demographic plant features (growth form, growth rate, latitude, ecological setting). We suggest that, similar to the manifold that tree species leaf traits cluster around the 'leaf economics spectrum', a similar 'wood economics spectrum' may be defined. We then discuss the biogeography, evolution and biogeochemistry of the spectrum, and conclude by pointing out the major gaps in our current knowledge of wood functional traits.

Towards a worldwide wood economics spectrum

Summary 1. The leaf economics spectrum (LES) provides a useful framework for examining species strategies as shaped by their evolutionary history. However, that spectrum, as originally described, involved only two key resources (carbon and nutrients) and one of three economically important plant organs. Herein, I evaluate whether the economics spectrum idea can be broadly extended to water – the third key resource –stems, roots and entire plants and to individual, community and ecosystem scales. My overarching hypothesis is that strong selection along trait trade-off axes, in tandem with biophysical constraints, results in convergence for any taxon on a uniformly fast, medium or slow strategy (i.e. rates of resource acquisition and processing) for all organs and all resources. 2. Evidence for economic trait spectra exists for stems and roots as well as leaves, and for traits related to water as well as carbon and nutrients. These apply generally within and across scales (within and across communities, climate zones, biomes and lineages). 3. There are linkages across organs and coupling among resources, resulting in an integrated whole-plant economics spectrum. Species capable of moving water rapidly have low tissue density, short tissue life span and high rates of resource acquisition and flux at organ and individual scales. The reverse is true for species with the slow strategy. Different traits may be important in different conditions, but as being fast in one respect generally requires being fast in others, being fast or slow is a general feature of species. 4. Economic traits influence performance and fitness consistent with trait-based theory about underlying adaptive mechanisms. Traits help explain differences in growth and survival across resource gradients and thus help explain the distribution of species and the assembly of communities across light, water and nutrient gradients. Traits scale up – fast traits are associated with faster rates of ecosystem processes such as decomposition or primary productivity, and slow traits with slow process rates. 5. Synthesis. Traits matter. A single ‘fast–slow’ plant economics spectrum that integrates across leaves, stems and roots is a key feature of the plant universe and helps to explain individual ecological strategies, community assembly processes and the functioning of ecosystems.

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The world‐wide ‘fast–slow’ plant economics spectrum: a traits manifesto

Whole-plant energy capture depends not only on the photosynthetic response of individual leaves, but also on their integration into an effective canopy, and on the costs of producing and maintaining their photosynthetic capacity. This paper explores adaptation to irradiance level in this context, focusing on traits whose significance would be elusive if considered in terms of their impact at the leaf level alone. I review traditional approaches used to demonstrate or suggest adaptation to irradiance level, and outline three energetic tradeoffs likely to shape such adaptation, involving the economics of gas exchange, support, and biotic interactions. Recent models using these tradeoffs to account for trends in leaf nitrogen content, stornatal conductance, phyllotaxis, and defensive allocations in sun v. shade are evaluated. A re-evaluation of the classic study of acclimation of the photosynthetic light response in Atriplex, crucial to interpreting adaptation to irradiance in many traits, shows that it does not completely support the central dogma of adaptation to sun v. shade unless the results are analysed in terms of whole-plant energy capture. Calculations for Liriodendron show that the traditional light compensation point has little meaning for net carbon gain, and that the effective compensation point is profoundly influenced by the costs of night leaf respiration, leaf construction, and the construction of associated support and root tissue. The costs of support tissue are especially important, raising the effective compensation point by 140 pmol m- s - ' in trees 1 m tall, and by nearly 1350 pmol m - s - ' in trees 30 m tall. Effective compensation points give maximum tree heights as a function of irradiance, and shade tolerance as a function of tree height; calculations of maximum permissible height in Liriodendron correspond roughly with the height of the tallest known individual. Finally, new models for the evolution of canopy width/height ratio in response to irradiance and coverage within a tree stratum, and for the evolution of mottled leaves as a defensive measure in understory herbs, are outlined.

/pdf/adaptation-to-sun-and-shade-a-whole-plant-perspective-j0u9zbhbtr.pdf

Adaptation to Sun and Shade: a Whole-Plant Perspective

We investigated how water transport capacity, wood density and wood anatomy were related to leaf photosynthetic traits in two lowland forests in Panama. Leaf-specific hydraulic conductivity ( k(L)) of upper branches was positively correlated with maximum rates of net CO(2) assimilation per unit leaf area ( A(area)) and stomatal conductance ( g(s)) across 20 species of canopy trees. Maximum k(L) showed stronger correlation with A(area) than initial k(L) suggesting that allocation to photosynthetic potential is proportional to maximum water transport capacity. Terminal branch k(L) was negatively correlated with A(area)/ g(s) and positively correlated with photosynthesis per unit N, indicating a trade-off of efficient use of water against efficient use of N in photosynthesis as water transport efficiency varied. Specific hydraulic conductivity calculated from xylem anatomical characteristics ( k(theoretical)) was positively related to A(area) and k(L), consistent with relationships among physiological measurements. Branch wood density was negatively correlated with wood water storage at saturation, k(L), A(area), net CO(2) assimilation per unit leaf mass ( A(mass)), and minimum leaf water potential measured on covered leaves, suggesting that wood density constrains physiological function to specific operating ranges. Kinetic and static indices of branch water transport capacity thus exhibit considerable co-ordination with allocation to potential carbon gain. Our results indicate that understanding tree hydraulic architecture provides added insights to comparisons of leaf level measurements among species, and links photosynthetic allocation patterns with branch hydraulic processes.

https://www.fs.fed.us/pnw/pubs/journals/pnw_2004_santiago001.pdf

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

In a computer simulation of branching pattern and leaf cluster in Terminalia catappa, right and left branch angles were varied, and the effective leaf surface areas were calculated. Theoretical branch angles that result in maximum effective leaf area are close to the values observed in nature.

https://science.sciencemag.org/content/sci/199/4331/888.full.pdf

Tree Branch Angle: Maximizing Effective Leaf Area

Vessel dimensions (total diameter and length) were determined in tropical and subtropical plants of different growth forms with an emphasis upon lianas (woody vines). The paint infusion and compressed air methods were used on 38 species from 26 genera and 16 families in the most extensive survey of vessel length made to date. Within most stems there was a skewed frequency distribution of vessel lengths and diameter, with many short and narrow vessels and few long and wide ones. The longest vessel found (7.73 m) was in a stem of the liana (woody vine) Pithecoctenium crucigerum. Mean vessel length for 33 species of lianas was 0.38 m, average maximum length was 1.45 m. There was a statistically significant inter-species correlation between maximum vessel length and maximum vessel diameter. Among liana stems and among tree+shrub stems there were statistically significant correlations between stem xylem diameter and vessel dimensions. Lianas with different adaptations for climbing (tendril climbers, twiners, scramblers) were similar in their vessel dimensions except that scramblers tended to have shorter (but not narrower) vessels. Within one genus, Bauhinia, tendril climbing species had greater maximum vessel lengths and diameters than tree and shrub species. The few long and wide vessels of lianas are thought to hydraulically compensate for their narrow stem diameters. The many narrow and short vessels, which are present in the same liana stems, may provide a high resistance auxiliary transport system.

A survey of vessel dimensions in stems of tropical lianas and other growth forms

Results were compared between the latex paint and compressed air methods for determining total vessel lengths, and between the sectioning and maceration methods for determining vessel diameters. The minimum, mean, median, and maximum vessel diameters were less with the sectioning method than with the maceration technique. Vessel diameter distributions were always nonnormal and had roughly similar patterns with the two techniques, but were statistically different from one another. In all six species where the paint and air methods for determining vessel length were compared, both methods showed a similar skewed vessel length distribution, with many short vessels and few long ones. Although there was no consistent pattern to the difference in results with these two methods, the vessel length frequency distributions were statistically different from one another. With the paint method, many vessels, especially many of the narrowest ones, were not paint-filled at the paint infusion port. The air method utilized the paint method, in part, and, in addition, is based upon the incorrect assumption that all vessels in the stem are the same diameter. Both techniques tended to exclude vessel lengths of the narrowest vessels. However, the narrow vessels, although numerous, contributed an insignificant amount to the total theoretical hydraulic conductance in stems.

Techniques for measuring vessel lengths and diameters in stems of woody plants

Lateral branch and young leader axes of 122 species of dicotyledonous trees in 46 families, displaying a variety of architectural models, were surveyed for presence of reaction wood (RW) as defined by gelatinous wood fibers (G-fibers). Distinct G-fibers occurred in 56 species, mostly in the upper half of the axis. There was a positive correlation between changes in axis orientation and position of the RW in 22, or one-fourth, of the species showing changes in branch orientation. Many species show an increase in branch angle; i.e., branches become more horizontal with age. Among these, Hevea (Rauh's architectural model), Endospermum, and Terminalia (both Aubreville's model) first produce RW on the lower side near the pith and later on the upper side. The erection of a lateral branch, i.e., a decrease in branch angle, to form the new leader in Koriba's model is correlated with RW on the upper side of the branch in Hura, but not in other species with this model. Many species with Troll's model, in which the ...

Jack B. Fisher

Papers

Leaf photosynthetic traits scale with hydraulic conductivity and wood density in Panamanian forest canopy trees

Tree Branch Angle: Maximizing Effective Leaf Area

A survey of vessel dimensions in stems of tropical lianas and other growth forms

Techniques for measuring vessel lengths and diameters in stems of woody plants

Occurrence of Reaction Wood in Branches of Dicotyledons and Its Role in Tree Architecture