So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

Genetics and analysis of quantitative traits

Trees, and their derivative products, have been used by societies around the world for thousands of years. Contemporary construction of tall buildings from timber, in whole or in part, suggests a growing interest in the potential for building with wood at a scale not previously attainable. As wood is the only significant building material that is grown, we have a natural inclination that building in wood is good for the environment. But under what conditions is this really the case? The environmental benefits of using timber are not straightforward; although it is a natural product, a large amount of energy is used to dry and process it. Much of this can come from the biomass of the tree itself, but that requires investment in plant, which is not always possible in an industry that is widely distributed among many small producers. And what should we build with wood? Are skyscrapers in timber a good use of this natural resource, or are there other aspects of civil and structural engineering, or large-scale infrastructure, that would be a better use of wood? Here, we consider a holistic picture ranging in scale from the science of the cell wall to the engineering and global policies that could maximise forestry and timber construction as a boon to both people and the planet.

The wood from the trees: The use of timber in construction

Bordered pits are cavities in the lignified cell walls of xylem conduits (vessels and tracheids) that are essential components in the water-transport system of higher plants. The pit membrane, which lies in the center of each pit, allows water to pass between xylem conduits but limits the spread of embolism and vascular pathogens in the xylem. Averaged across a wide range of species, pits account for > 50% of total xylem hydraulic resistance, indicating that they are an important factor in the overall hydraulic efficiency of plants. The structure of pits varies dramatically across species, with large differences evident in the porosity and thickness of pit membranes. Because greater porosity reduces hydraulic resistance but increases vulnerability to embolism, differences in pit structure are expected to correlate with trade-offs between efficiency and safety of water transport. However, trade-offs in hydraulic function are influenced both by pit-level differences in structure (e.g. average porosity of pit membranes) and by tissue-level changes in conduit allometry (average length, diameter) and the total surface area of pit membranes that connects vessels. In this review we address the impact of variation in pit structure on water transport in plants from the level of individual pits to the whole plant.

Structure and function of bordered pits: new discoveries and impacts on whole-plant hydraulic function

Solar cell substrates require high optical transparency but also prefer high optical haze to increase the light scattering and consequently the absorption in the active materials. Unfortunately, there is a trade-off between these optical properties, which is exemplified by common trans- parent paper substrates exhibiting a transparency of about 90% yet a low optical haze (<20%). In this work, we introduce a novel transparent paper made of wood fibers that displays both ultrahigh optical transparency (∼96%) and ultrahigh haze (∼60%), thus delivering an optimal substrate design for solar cell devices. Compared to previously demonstrated nanopaper composed of wood-based cellulose nanofibers, our novel transparent paper has better dual performance in transmittance and haze but also is fabricated at a much lower cost. This high-performance, low-cost transparent paper is a potentially revolutionary material that may influence a new generation of environmentally friendly printed electronics.

https://terpconnect.umd.edu/~lit/publications/TengLi-Pub58-NL-2014.pdf

Novel Nanostructured Paper with Ultrahigh Transparency and Ultrahigh Haze for Solar Cells

The cell wall of wood tracheids is made up of various layers, distinguished from one other by the alignment of the innumerable, fine crystalline cellulose microfibrils within each layer that helically wind about the cell lumen. Microfibrils themselves are embedded in a more compliant, water-reactive matrix of amorphous lignin and hemicelluloses. The average inclination of microfibrils relative to the axis of the cell affects axial rigidity and dimensional stability of wood which are the two most important properties of wood. High and variable microfibril angles can be found in juvenile and compression wood, thus resulting in variations in product performance of forest products. For instance, seemingly identical trees in a plantation can have moduli of elasticity that differ by a factor of two or more. This is why the future is often seen in engineered wood products, where wood may be chipped, fiberised and blended before being glued together again: the average property values are little changed, but the range—the variability—is greatly reduced. There is the opportunity for better wood allocation and processing of timber, if averaged values for individual log characteristics, such as average microfibril angle, can be identified before the processing. In parallel there is genetic potential to select trees with low average microfibril angles. Unfortunately, determination of the average microfibril angle is a time-consuming, laboratory-based task. Preferably, a non-destructive, simple, field-hardened method should be employed that reflects the average microfibril angle in a given piece of wood. For this reason, acoustic methods have been developed to measure the velocity of sound propagation directly related to the stiffness of wood and in turn is dependent on the ultrastructure of the tracheid cell wall. In the fundamental equation, Edynamic=ρV2, the acoustic modulus is derived from two components, density, ρ, and velocity of sound, V. The latter relates to the intrinsic wood quality and ultrastructure of the tracheid wall. It is shown that acoustic methods can sort and grade trees and logs according to their suitability for structural lumber and for a range of fiber properties of interest to papermakers. Thus, acoustic methods have applications in tree breeding, harvesting, and wood processing.

Cell wall structure and wood properties determined by acoustics—a selective review

clones with high modulus of elasticity (MOE). Various ways of measuring MOE are examined: traditional static bending, axial compression of bolts, existing sonic methods, as well as a tool specifically configured for resonance on short bolts. Wood characteristics and microfibril angle are measured on discs taken from each tree. Results show a good correlation between acoustic and static measurements of modulus of elasticity. Moreover, the selected seven radiata clones differ in stiffness by a factor of two, and much of this variation seems to relate to differences in microfibril angle. The results validate the assumption that there is potential to improve wood stiffness of radiata pine genetically. This work does not offer definitive solutions but explores a number of approaches that could be utilised as a selection tool in tree breeding for better product performance of radiata lumber. Here, development of methods based on acoustics is shown to assist in the necessary mass screening of clones for stiffness properties.

Methods for measuring stiffness of young trees

Tracheid length, microfibril angle, spiral grain, wood density, MOE, MOR, compressive strength, modulus of rigidity and their inter-relationships were investigated for fifty Siberian larch trees (Larix sibirica) collected from five natural stands. No inter-stand variation in tracheid length, microfibril angle and spiral grain was observed. No effect of tracheid length and microfibril angle on the mechanical properties was observed. Inter-stand variation in wood density was significant, especially for mature wood. The wood from the Baikal site in the eastern range of the natural distribution had a very high density. The wood from the Altai site in the mountain range had a low density, especially within the latewood. Minimum density within a ring seemed to be the major source of variation for average density within the heartwood, whereas the effect of maximum density on average density was greater within the sapwood. Wood density, which was much higher for a given ring width than in plantation-grown Japanese larch (Larix kaempferi), had a major effect on the mechanical properties.

https://brill.com/downloadpdf/journals/iawa/24/4/article-p355_2.pdf

Anatomical characteristics and mechanical properties of larix sibirica grown in south-central siberia

Heartwood in tree stems provides natural durability and ornamental values to wood products as well as antimicrobial properties to protect living stems from decay. Control of the amount and quality of heartwood is required because of its importance in wood utilization. Despite the importance, the mechanism of heartwood formation has been poorly understood. To obtain further knowledge for understanding this mechanism, we studied the seasonal variation of various components of heartwood formation in Larix kaempferi (Lamb.) Carriere. Dehydration of tracheids at the boundary between sapwood and intermediate wood occurred continuously during the study period. The death of ray parenchyma cells at intermediate wood occurred from spring to summer. The deposition of heartwood substance took place in autumn to winter. Thus, different components of heartwood formation were active in different seasons. Since heartwood formation is a system consisting of several components within a series of continuing processes, understanding the seasonal change of heartwood formation requires monitoring these components simultaneously.

Seasonal variation of heartwood formation in Larix kaempferi

Bordered pit membranes of Cryptomeria japonica were examined successively from the outermost sapwood to the heartwood by scanning electron microscopy and by ultraviolet microspectrophotometry in an attempt to evaluate the time course of the secondary deposition of incrusting materials and to gain clues to their chemical composition. Scanning electron microscopy revealed that the bordered pit membranes were covered by incrusting materials from the middle layer of the sapwood to the heartwood. Both the amount and the appearance of the deposited incrusting materials differed among four regions of the wood, namely, the middle to inner layer of the sapwood, the innermost layer of the sapwood, the intermediate wood and the heartwood. From our results it appears that, in C. japonica, incrusting materials are deposited on bordered pit membranes by stages over several years. Apparent absorption of ultraviolet light by the bordered pit membranes was detected in the analysis of the innermost layer of the sapwood, the intermediate wood and the heartwood. The incrusting materials deposited in these zones were probably phenolic compounds. However, differences in the manner and extent of the absorption of ultraviolet light were found between these three regions of the wood. The results of microspectrophotometric analysis also suggested the phased deposition of incrusting materials at the bordered pit membranes of C. japonica.

/pdf/time-course-of-the-secondary-deposition-of-incrusting-159bka7781.pdf

Ryogo Nakada

Papers

Cell wall structure and wood properties determined by acoustics—a selective review

Methods for measuring stiffness of young trees

Anatomical characteristics and mechanical properties of larix sibirica grown in south-central siberia

Seasonal variation of heartwood formation in Larix kaempferi

Time Course of the Secondary Deposition of Incrusting Materials on Bordered Pit Membranes in Cryptomeria Japonica