Compression Wood Research Articles

Recent genome resequencing paraded COBRA-Like gene family roles in abiotic stress and wood formation in Poplar.

A cell wall determines the mechanical properties of a cell, serves as a barrier against plant stresses, and allows cell division and growth processes. The COBRA-Like (COBL) gene family encodes a putative glycosylphosphatidylinositol (GPI)-anchored protein that controls cellulose deposition and cell progression in plants by contributing to the microfibril orientation of a cell wall. Despite being studied in different plant species, there is a dearth of the comprehensive global analysis of COBL genes in poplar. Poplar is employed as a model woody plant to study abiotic stresses and biomass production in tree research. Improved genome resequencing has enabled the comprehensive exploration of the evolution and functional capacities of PtrCOBLs (Poplar COBRA-Like genes) in poplar. Phylogeny analysis has discerned and classified PtrCOBLs into two groups resembling the Arabidopsis COBL family, and group I genes possess longer proteins but have fewer exons than group II. Analysis of gene structure and motifs revealed PtrCOBLs maintained a rather stable motif and exon-intron pattern across members of the same group. Synteny and collinearity analyses exhibited that the evolution of the COBL gene family was heavily influenced by gene duplication events. PtrCOBL genes have undergone both segmental duplication and tandem duplication, followed by purifying selection. Promotor analysis flaunted various phytohormone-, growth- and stress-related cis-elements (e.g., MYB, ABA, MeJA, SA, AuxR, and ATBP1). Likewise, 29 Ptr-miRNAs of 20 families were found targeting 11 PtrCOBL genes. PtrCOBLs were found localized at the plasma membrane and extracellular matrix, while gene ontology analysis showed their involvement in plant development, plant growth, stress response, cellulose biosynthesis, and cell wall biogenesis. RNA-seq datasets depicted the bulk of PtrCOBL genes expression being found in plant stem tissues and leaves, rendering mechanical strength and rejoinders to environmental cues. PtrCOBL2, 3, 10, and 11 manifested the highest expression in vasculature and abiotic stress, and resemblant expression trends were upheld by qRT-PCR. Co-expression network analysis identified PtrCOBL2 and PtrCOBL3 as hub genes across all abiotic stresses and wood developing tissues. The current study reports regulating roles of PtrCOBLs in xylem differentiating tissues, tension wood formation, and abiotic stress latency that lay the groundwork for future functional studies of the PtrCOBL genes in poplar breeding.

Microscopic computed tomography aided finite element modelling as a methodology to estimate hygroexpansion coefficients of wood: a case study on opposite and compression wood in softwood branches

Abstract Microscopic X-ray computed tomography (XµCT) aided finite element (FE) modelling is a popular method in material science to relate material properties to heterogeneous microstructures. Recently, a methodology was developed for the XµCT aided FE modelling of wood, which characterises the process from specimen preparation to estimation of material properties. In the current research, this methodology is tested on branches of Norway spruce (Picea abies (L.) Karst.) to estimate the hygroexpansion coefficients of opposite (OW) and compression wood (CW). These properties are largely unknown and have engineering implications. The study is complemented by measurements of density, moisture content (MC) and elastic moduli. Results showed that the methodology assisted in the design of an integrated process and the identification of bottlenecks. It was seen that the level of detail of the numerical model had a strong influence on the obtained hygroexpansion properties. CW from branches showed higher density and longitudinal shrinkage coefficients, and elastic moduli less affected by MC. These differences are unlikely caused by MC, but more likely by the characteristics of the microstructure.

Modelling the growth stress in tree branches: eccentric growth vs. reaction wood

Modelling the growth stress in tree branches: eccentric growth vs. reaction wood

Seasonal Developing Xylem Transcriptome Analysis of Pinus densiflora Unveils Novel Insights for Compression Wood Formation.

Wood is the most important renewable resource not only for numerous practical utilizations but also for mitigating the global climate crisis by sequestering atmospheric carbon dioxide. The compressed wood (CW) of gymnosperms, such as conifers, plays a pivotal role in determining the structure of the tree through the reorientation of stems displaced by environmental forces and is characterized by a high content of lignin. Despite extensive studies on many genes involved in wood formation, the molecular mechanisms underlying seasonal and, particularly, CW formation remain unclear. This study examined the seasonal dynamics of two wood tissue types in Pinus densiflora: CW and opposite wood (OW). RNA sequencing of developing xylem for two consecutive years revealed comprehensive transcriptome changes and unique differences in CW and OW across seasons. During growth periods, such as spring and summer, we identified 2255 transcripts with differential expression in CW, with an upregulation in lignin biosynthesis genes and significant downregulation in stress response genes. Notably, among the laccases critical for monolignol polymerization, PdeLAC17 was found to be specifically expressed in CW, suggesting its vital role in CW formation. PdeERF4, an ERF transcription factor preferentially expressed in CW, seems to regulate PdeLAC17 activity. This research provides an initial insight into the transcriptional regulation of seasonal CW development in P. densiflora, forming a foundation for future studies to enhance our comprehension of wood formation in gymnosperms.

Is tree-ring anatomy the key? searching for an optimal dendrogeomorphic approach to distinguish shallow creep and true slide movements

Tree-ring based chronologies of slide movements can serve as an important basis for determining landslide activity, assessing risk and triggers, or potentially predicting future landslide behaviour. Unfortunately, however, the landslide areas studied are often affected by another type of slope movement, namely creep, which can introduce a considerable amount of noise into the resulting chronology. Thus, this study focused on testing different dendrogeomorphic approaches for the extraction of landslide signal from tree-ring series with respect to their ability to filter out the influence of creep movements and slide movements. For this purpose, 45 trees (Fagus sylvatica L. and Acer pseudoplatanus L.) growing on the steep slope of a sub-landslide block whose movement has been monitored for a long time and for which a single landslide event is known (second half of 2010) were sampled.The first tested approach was based on changing tree-ring eccentricity values and has shown its very high sensitivity to all slope movements and is therefore not applicable for creep filtering and slide movements unsuitable on steep slopes. The second approach, using macroscopic identification of the tension tree, proved to be the most suitable, accurately detecting the slide event and successfully filtering out all creep-induced noise. A final approach based on the detection of sudden anatomical changes in tree rings (independent of the presence of tension wood) also captured high amounts of noise and also slide event. Its use would be possible in the case of a suitably set event-response index threshold. The study also includes a discussion of the general effect of creep movements on tree growth, the limitations and advantages of each approach, and recommendations for future research.

Study of antioxidant enzymes’ activity in reaction wood of poplar tree (Populus alba L.)

Summary Studying biochemical variation in reaction wood is needed to understand the nature of reaction wood formation and the physiology of trees. The activity of antioxidant enzymes (superoxide dismutase (SOD), catalase (CAT), ascorbate peroxidase (APX) and guaiacol peroxidase (GPX)) was studied in normal wood (NW) and reaction wood (RW) (opposite wood (OW) and tension wood (TW)) in the bole of poplar trees (Populus alba L.). Four-year-old trees were induced to produce reaction wood by forced bending. The activity of antioxidant enzymes was studied with repeated sampling during a growing season. The results showed that total protein content and antioxidant enzymes are sensitive to mechanically induced stress. TW and OW showed higher activity of enzymes than NW as well as higher amounts of total protein. OW showed a higher concentration of CAT and SOD than TW. APX was more active than GPX in both TW and OW. In addition to mechanical stress, the effect of other climate factors was observed in the reactive oxygen species accumulation.

Selection history affects very early expression of wood properties in Pinus radiata

Trees in breeding programmes are often selected at 1/4–1/3 of rotation, called ‘early selection’, which is typically between 8 and 10 years in radiata pine. However, differences between populations and genotypes selected for either basic density or wood stiffness are already apparent at age 2. We report the application of very early screening techniques for wood properties in the New Zealand Radiata Pine Breeding Programme deployment populations. Approximately 3000 trees representing three populations with 92 families and 10 clones were grown in a common garden trial, leaning for 21 months to separate compression and opposite wood. The trees were harvested and analysis carried out separately by wood type. The trial showed the existence of large variability in wood properties at early age, in some cases similar to variability near rotation age, and moderate to high degree of genetic control (0.35≤h2≤0.71\\documentclass[12pt]{minimal} \\usepackage{amsmath} \\usepackage{wasysym} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{amsbsy} \\usepackage{mathrsfs} \\usepackage{upgreek} \\setlength{\\oddsidemargin}{-69pt} \\begin{document}$$0.35 \\le h^2 \\le 0.71$$\\end{document}). The genetic association between traits was strong, particularly between wood stiffness and longitudinal shrinkage (-0.69\\documentclass[12pt]{minimal} \\usepackage{amsmath} \\usepackage{wasysym} \\usepackage{amsfonts} \\usepackage{amssymb} \\usepackage{amsbsy} \\usepackage{mathrsfs} \\usepackage{upgreek} \\setlength{\\oddsidemargin}{-69pt} \\begin{document}$$-\\,0.69$$\\end{document}) and between longitudinal and volumetric shrinkage (0.83), suggesting that improving stiffness would also have a strong effect on improving dimensional stability. Basic density was also associated with stiffness and shrinkage, but with lower predictive capacity. These results can be used for roguing deployment populations-which already contain superior growing trees-and quickly upgrade the wood quality of seeds and clones currently available to New Zealand forest growers. We discuss necessary modifications to turn this research work into operation to screen any new material before commercial release.

Multiomics studies with co-transformation reveal microRNAs via miRNA-TF-mRNA network participating in wood formation in Hevea brasiliensis.

MicroRNAs (miRNAs) are small endogenous non-coding RNAs that play an important role in wood formation in plants. However, the significance of the link between miRNAs and their target transcripts in wood formation remains unclear in rubber tree (Hevea brasiliensis). In this study, we induced the formation of reaction wood by artificially bending rubber trees for 300 days and performed small RNA sequencing and transcriptome deep sequencing (RNA-seq) to describe the complement of miRNAs and their targets contributing to this process. We identified 5, 11, and 2 differentially abundant miRNAs in normal wood (NW) compared to tension wood (TW), in NW relative to opposite wood (OW), and between TW and OW, respectively. We also identified 12 novel miRNAs and 39 potential miRNA-mRNA pairs with different accumulation patterns in NW, TW, and OW. We noticed that many miRNAs targeted transcription factor genes, which were enriched in KEGG pathways associated with phenylpropanoid biosynthesis, phenylalanine metabolism, and pyruvate metabolism. Thus, miRNA-TF-mRNA network involved in wood formation via tension wood model were constructed. We validated the differential accumulation of miRNAs and their targets by RT-qPCR analysis and overexpressed miRNA in Nicotiana benthamiana with its potential target gene. These results will provide a reference for a deep exploration of growth and development in rubber tree.

Mesostructural changes in cellulose within wood cell wall upon hydrothermal treatment at 200 °C

Hydrothermal treatment between 150 °C and 230 °C is widely used in wood processing, from the steam treatment of timber for better dimensional stability and durability to the pretreatment for enzymatic saccharification and chemical pulping. Understanding the ultrastructural changes of wood cell walls through hydrothermal treatments is crucial for controlling and optimizing these hydrothermal treatment-based processes. Here, we studied the ultrastructure of wood cell walls of 24 hardwood species using simultaneous small- and wide-angle X-ray scattering measurements before and after the hydrothermal treatment at 200 °C. Most hardwoods show similar equatorial scattering features, representing the structure in the cross-section of the cell walls. In a water-saturated native state, there is a prominent correlation peak between 0.1 and 0.2 Å−1 and a second peak between 0.2 and 0.4 Å−1. The hydrothermal treatment above 160 ˚C drastically altered the structure at this nanometric scale: the two native correlation peaks disappeared, coincident with a buildup of a correlation peak in the 0.03–0.04 Å−1 range. The hydrothermal treatment likely removed the cell wall matrix component between the microfibrils through autohydrolysis and phase separation, leading to the collapse of microfibrils with each other in the normal wood. In cellulose-rich cell walls, such as the G-layer in tension wood, cellulose microfibrils are already collated in the native state.

Overload protection for forestry machines when using pressed wood friction material

The article proposes an original approach to the protection of forestry machines from overloads when using a friction material made of pressed wood. The use of compressed wood is an environmentally important aspect in the manufacture of friction materials for safety devices of forestry machines. This approach contributes to the development of green technologies in forestry engineering and ensures an increase in the environmental friendliness of logging operations. However, from an environmental point of view, self-ignition of pressed wood must be avoided. The definition of the transmitted and adjusting torque in relation to the frictional safety clutch is considered. It is noted that when the safety device slips, the friction material made of pressed wood will wear out unevenly, in the form of a wedge, if the working width of the contacting surfaces is large. A diagram of the calculated friction clutch is presented, as well as a diagram of the pressure fuse block, from which it can be seen how the rubber rings are not deformed in the same way due to their difference in diameters. It is noted that for forestry machines operating with frequent overloads, it is necessary to take into account the dynamic loads that occur during overload at the moment the fuse is triggered.

Evaluation of Physico-Mechanical Properties and Thermal Conduction to Energy-Saving Effects of Wood Compression Layered Structural Materials.

In this study, specimens were prepared from Japanese cedar (Cryptomeria japonica) with different thicknesses to determine the best hot-pressing conditions for wood compression layered structural materials (WCLS) through densification at various temperatures and compressing time conditions. However, residual stress-releasing after densification recovery can cause dimensional instability. To address this issue, the drying set method was combined with the compression-set recovery test to determine the best setting time. As a result, the bending strength and modulus of rupture (MOR) of WCLS increased by 9.98 ± 9.71 to 20.87 ± 13.09% and the modulus of elasticity (MOE) increased by 9.87 ± 11.92 to 22.40 ± 17.97%. The volumetric swelling coefficient (S), water absorption percent (WAP), and equivalent moisture content (EMC) decreased as the drying time increased. The anti-swelling efficiency (ASE) and moisture excluding efficiency (MEE) were found to be the highest at a drying time of 12 h, with values ranging from 13.20 ± 15.11 to 36.46 ± 6.83% and 15.18 ± 1.11 to 19.58 ± 8.31%, respectively. The drying method was found to be effective in increasing dimensional stability. The glass transition temperature (Tg) moved to a lower temperature as the compression-set increased, which was due to plasticization of wood caused by high temperature and pressure. The cell walls of WCLS presented viscous buckling, which provided effective dimensional stability. The thermal conductivity of Japanese cedar and each compression-set WCLS were 0.1863 ± 0.0071, 0.1520 ± 0.0147, 0.1817 ± 0.0106, and 0.1423 ± 0.0137 W/mK, respectively. The thermal conductivity of each WCLS increased with an increase in compression-set, but decreased by 10.67 to 22.52% compared to Japanese cedar. The total electricity energy consumption of each WCLS after 24 h of testing decreased with a trend of 30.50 ± 0.84, 29.83 ± 0.42, 29.57 ± 0.51, and 29.4 ± 0.36 kWH.

Maturation Stress and Wood Properties of Poplar (Populus × euramericana cv. ‘Zhonglin46’) Tension Wood

Understanding the maturation stress and wood properties of poplar tension wood is critical for improving lumber yields and utilization ratio. In this study, the released longitudinal maturation strains (RLMS), anatomical features, physical and mechanical properties, and nano-mechanical properties of the cell wall were analyzed at different peripheral positions and heights in nine artificially inclined, 12-year-old poplar (Populus × euramericana cv. ‘Zhonglin46’) trees. The correlations between the RLMS and the wood properties were determined. The results showed that there were mixed effects of inclination on wood quality and properties. The upper sides of inclined stems had higher RLMS, proportion of G-layer, bending modulus of elasticity, and indentation modulus of the cell wall but a lower microfibril angle than the lower sides. At heights between 0.7 m and 2.2 m, only the double-wall thickness increased with height; the RLMS and other wood properties such as fiber length and basic density fluctuated or changed little with height. The RLMS were good indicators of wood properties in the tension wood area and at heights between 0.7 m and 1.5 m. The results of this study present opportunities to better understand the interactions and effects of these two phenomena, which both occur quite frequently in poplar stands and can influence the wood quality of valuable assortments.

Anatomical characteristics of the leaning stem in Pinus merkusii seedling

Abstract. Safitri NM, Prastiwi FW, Salsabila S, Feriawan Y, Brilianto A, Nugroho WD. 2023. Anatomical characteristics of the leaning stem in Pinus merkusii seedling. Biodiversitas 24: 3796-3802. Pinus merkusii Jungh. & de Vriese is one of the tropical conifer species. The species formed reaction wood called compression wood (CW) when the stem grows in an abnormal position. CW in stem wood is frequently found to cause several problems for utilization in the use of its products. However, there are few studies on the anatomical characteristics of reaction wood in tropical conifer. This study aimed to observe the differences in anatomical characteristics between compression (CW) and opposite (OW) woods in leaning the stem of P. merkusii seedlings and their anatomical differences compared to normal wood (NW). Approx. 1-year-old P. merkusii seedlings from a nursery in Trenggalek, East Java, Indonesia, that were naturally bent or tilted were used in the study. Anatomical characteristics of wood, namely the dimension of tracheids, rays, resin canals, and the proportion of cells were observed in the study. The results showed that CW of P. merkusii seedlings formed a rounded tracheid shape and intercellular space. The differences between CW and OW in P. merkusii seedlings were significant in terms of height of ray cells, tracheid proportion, and resin canal proportion. However, tracheid length, tracheid diameter, tracheid wall thickness, resin canal diameter and frequency, rays frequency, and ray proportion were not significantly different compared to OW and NW.

The Impact of the Log-Sawing Patterns on the Quantitative and Qualitative Yield of Beech Timber (Fagus sylvatica L.)

The cost of raw materials has a major role in the wood industry. From this point of view, it is necessary to pay attention to the correct log-sawing pattern, which will optimize the sawing process and increase the quantity and quality of the material as well. For measurements, we used beech wood (Fagus sylvatica L.). Beech wood (Fagus sylvatica L.) is the most common wood species in the Slovak Republic; the total amount in forests is 34.8%. Beech wood has a high frequency of defects, such as red false heartwood, reaction wood (i.e., tension wood), dote, cracks, and so on. This research aimed to analyze the effect of the log-sawing patterns of through-and-through sawing, cant sawing, and quarter sawing on the resulting quantitative–qualitative yields of beech timber. The results showed that the highest quantitative yield was observed with the cant sawing pattern, at 84%, which was, on average, 17.3% higher compared to other types of cutting. It was found that quarter sawing resulted in a higher qualitative yield compared to through-and-through sawing or cant sawing (V5 = 62.69% and V6 = 47.86%).

Structure of cellulose in birch phloem fibres in tension wood: an X-ray nanodiffraction study

BackgroundTo gain a better understanding of bark layer structure and function, especially of the phloem fibres and their contribution to the posture control of trees, it is important to map the structural properties of these cells. The role of bark can also be linked to the reaction wood formation and properties which are essential when it comes to studying the questions related to tree growth. To offer new insights into the role of bark in the postural control of trees, we studied the micro- and nanoscale structures of the phloem and its nearest layers. This study is the first time, in which phloem fibres in trees have been extensively examined using X-ray diffraction (XRD).We determined the orientation of cellulose microfibrils in phloem fibres of Silver birch saplings by using scanning synchrotron nanodiffraction. The samples consisted of phloem fibres extracted from tension, opposite and normal wood (TW, OW, NW).ResultsUsing scanning XRD, we were able to obtain new information about the mean microfibril angle (MFA) in cellulose microfibrils in phloem fibres connected to reaction wood. A slight but consistent difference was detected in the average MFA values of phloem fibres between the TW and OW sides of the stem. Using scanning XRD, different contrast agents (intensity of the main cellulose reflection or calcium oxalate reflection, mean MFA value) were used to produce 2D images with 200 nm spatial resolution.ConclusionsBased on our results, the tension wood formation in the stem might be related to the structure and properties of phloem fibres. Thus, our results suggest that the nanostructure of phloem fibres is involved in the postural control of trees containing tension and opposite wood.

Combined X-ray diffraction tomography imaging of tension and opposite wood tissues in young hybrid aspen saplings

Combining spatially localized X-ray diffraction (XRD) with X-ray microtomography (XMT) enables the mapping of the micro- and nanoscale structures simultaneously. The combination of these methods results in a powerful tool when considering the structural studies of hierarchical materials, allowing one to couple the relationships and connections of the structures at various scales. In this study, XMT was used to map the anatomy and cellular structures in 3D in tension and opposite wood with 1.5 µm resolution, while XRD was used to determine the cellulose crystallite widths and microfibril orientations with 100 µm spatial resolution within the same tissues. Tension wood (TW) has an important biological function with clearly distinct properties to opposite (OW) and normal wood, e.g. differing cellular structures with a higher cellulose content. This is the first study of very young hybrid aspen saplings (1-month-old) using the combined diffraction tomography method. The TW tissues could be identified from the OW tissues based on both the XMT and XRD results: TW had a higher average size of the cellulose crystallites and smaller mean microfibril angles (mMFA) than those in OW. With the XRD data, we were able to reconstruct the images of the cross sections of the saplings using the structural parameters (cellulose crystallite width and mMFA) as contrast mechanisms. As far as the authors know, there are no previous studies with images on any TW samples using the XRD-based contrast. Home laboratory bench-top set-up offers its advantages for these studies, considering the number of samples characterized, time-dependent studies and larger field of views.

Three-dimensional distribution of mass movement signals in disturbed Picea abies (L.) Karst.

Dendrogeomorphologic approaches enable annual absolute dating of various potentially hazardous geomorphic processes. This study tested the dating precision of three commonly used dendrogeomorphic approaches used for the extraction of geomorphic signals from the tree ring series of disturbed trees. The first approach is based on the onset of reaction wood occurrence, and the other two approaches are based on the detection of abrupt changes in tree ring eccentricity. Nine 165 cm long stems of maximally 27 year old Norway spruces (Picea abies (L.) Karst.) growing on unstable slope affected by mass movement events of known age were analysed. Several sampling directions on stems were analysed to reveal the best sampling strategy for each tested approach. The relationships between stem curvature and the dating precision of the approaches were also analysed. The results of dating precision showed crucial differences between the tested approaches. Tree stem curvature did not influence dating precision. Overall, we concluded that the approaches based on eccentric increments are less precise than those based on reaction wood analysis when juvenile P. abies trees that grow on landslide slope are sampled. The importance of this study is in providing new insight into dating precision by analysing the whole tree stems of trees influenced by mass movements. This is unique since the approaches based on eccentricity increments have not been previously tested in such a detail.

An important contribution to the description of growth stresses in branches of adult trees based on a new model and an optimisation process with digitised branches.

A recommendation of: Arnoul VAN ROOIJ, Eric BADEL, Jean Francois BARCZI, Yves CARAGLIO, Tancrède ALMERAS, Joseph GRIL Modelling the growth stress in tree branches: eccentric growth vs. reaction wood https://hal.science/hal-03748026

Distribution of Plant Hormones and Their Precursors in Cambial Region Tissues of Quercus myrsinifolia and Castanopsis cuspidata var.sieboldii after Bending Stems or Applying Ethylene precursor

The role of plant hormones in tension wood (TW) formation has been studied but is still unclear. IAA, ABA, ACC, tZ, tZR, iP, and iPR in cambial region tissues were identified and quantified by liquid chromatography/mass spectrometry (LC/MS). We examined the distribution of plant hormones and their precursors in the stems of Quercus myrsinifolia Blume and Castanopsis cuspidata var.sieboldii Nakai after bending the stems or applying an ethylene precursor (ACC). After 3 weeks of bending, though not after 1 week of bending, the auxin (IAA) and abscisic acid (ABA) amounts were larger on the TW side than on the opposite wood (OW) side and in upright trees. After 2 weeks of bending, the peak concentrations of IAA in cambium on the TW side were obviously higher than those on the OW side. After 1 week of bending, the ACC amounts on both sides were larger than in upright trees, but after 3 weeks of bending, they were smaller than in upright trees. Applied ACC did not enhance TW formation but induced axical parenchyma and phloem formation in C. cuspidata var.sieboldii. These results indicated that the distribution patterns of IAA and ABA might have important roles in TW formation in these two species. The role of ACC might be limited in the early stages of TW formation.

A Review of Wood Compression along the Grain—After the 100th Anniversary of Pleating

This study focuses on the compression of wood along the grain (also known as pleating), a modification that improves the pliability of higher-density hardwoods with a moisture content above 20%. Pleated wood can be bent into small curves in any direction. The success of the industrial pleating process in some parts of the world is influenced by many factors, such as wood species, wood quality, moisture content, compression ratio, fixation time, etc. Pleating by 20% causes the modulus of elasticity to decrease to one-third for oak and beech, and the bending ratio can be increased above ½. Bending stress decreases and the absorbed energy increases multiple times. The impact bending strength also increases significantly. The walls of cells crinkle by pleating and the microfibrils of the fibres become distorted. Many patents, articles, and books on this subject have been published since 1917, and this review attempts to introduce and, where necessary, critically analyse them.