More than a half century ago, Shinozaki et al. (Shinozaki K, Yoda K, Hozumi K, Kira T. 1964a. A quantitative analysis of plant form – the pipe model theory. I. Basic analyses. Japanese Journal of Ecology B: 97–105) proposed an elegant conceptual framework, the pipe model theory (PMT), to interpret the observed linear relationship between the amount of stem tissue and corresponding supported leaves. The PMT brought a satisfactory answer to two vividly debated problems that were unresolved at the moment of its publication: (1) What determines tree form and which rules drive biomass allocation to the foliar versus stem compartments in plants? (2) How can foliar area or mass in an individual plant, in a stand or at even larger scales be estimated? Since its initial formulation, the PMT has been reinterpreted and used in applications, and has undoubtedly become an important milestone in the mathematical interpretation of plant form and functioning. This article aims to review the PMT by going back to its initial formulation, stating its explicit and implicit properties and discussing them in the light of current biological knowledge and experimental evidence in order to identify the validity and range of applicability of the theory. We also discuss the use of the theory in tree biomechanics and hydraulics as well as in functional–structural plant modelling. Scrutinizing the PMT in the light of modern biological knowledge revealed that most of its properties are not valid as a general rule. The hydraulic framework derived from the PMT has attracted much more attention than its mechanical counterpart and implies that only the conductive portion of a stem cross-section should be proportional to the supported foliage amount rather than the whole of it. The facts that this conductive portion is experimentally difficult to measure and varies with environmental conditions and tree ontogeny might cause the commonly reported non-linear relationships between foliage and stem metrics. Nevertheless, the PMT can still be considered as a portfolio of properties providing a unified framework to integrate and analyse functional–structural relationships.

Increasing or decreasing wood density (WD) from pith to bark is commonly observed in tropical tree species. The different types of WD radial variations, long been considered to depict the diversity of growth and mechanical strategies among forest guilds (heliophilic vs. shade-tolerant), were never analyzed in the light of heartwood (HW) formation. Yet, the additional mass of chemical extractives associated to HW formation increases WD and might affect both WD radial gradient (i.e., the slope of the relation between WD and radial distance) and pattern (i.e., linear or nonlinear variation). We studied 16 legumes species from French Guiana representing a wide diversity of growth strategies and positions on the shade-tolerance continuum. Using WD measurements and available HW extractives content values, we computed WD corrected by the extractive content and analyzed the effect of HW on WD radial gradients and patterns. We also related WD variations to demographic variables, such as sapling growth and mortality rates. Regardless of the position along the shade-tolerance continuum, correcting WD gradients reveals only increasing gradients. We determined three types of corrected WD patterns: (1) the upward curvilinear pattern is a specific feature of heliophilic species, whereas (2) the linear and (3) the downward curvilinear patterns are observed in both mid- and late-successional species. In addition, we found that saplings growth and mortality rates are better correlated with the corrected WD at stem center than with the uncorrected value: taking into account the effect of HW extractives on WD radial variations provides unbiased interpretation of biomass accumulation and tree mechanical strategies. Rather than a specific feature of heliophilic species, the increasing WD gradient is a shared strategy regardless of the shade tolerance habit. Finally, our study stresses to consider the occurrence of HW when using WD.

Peach palm (Bactris gasipaes Kunth) is a Neotropical palm domesticated by Native Americans. Its domestication resulted in a set of landraces (var. gasipaes), some with very starchy fruit used for fermentation, others with an equilibrium of starch and oil used as snacks. Which of the three wild types (var. chichagui) was involved and where the domestication process began are unclear, with three hypotheses under discussion: an origin in southwestern Amazonia; or in northwestern South America; or multiple origins. We reevaluate one of the wild types, defining it as the incipient domesticate, and then evaluate these hypotheses using the Brazilian peach palm Core Collection and selected herbaria samples to: (1) model the potential distributions of wild and domesticated populations; (2) identify the probable origin of domestication with a phylogeographic analysis of chloroplast DNA sequences; and (3) determine the dispersal routes after domestication using spatial analysis of genetic diversity based on 17 nuclear microsatellite loci. The two very small-fruited wild types have distinct distributions in the northern Andes region and across southern Amazonia, both under moderately humid climates, while the incipient domesticate, partly sympatric with the southern wild type, is also found along the Equatorial Andes, in a more humid climatic envelope, more similar to that of the domesticated landraces. Two distribution models for Last Glacial Maximum conditions (CCSM4, MIROC) also suggest distinct distributions for the two wild populations. The chloroplast DNA phylogeographic network confirms the area of sympatry of the incipient domesticate and the southern wild type in southwestern Amazonia as the origin of domestication. The spatial patterns of genetic diversity confirm the proposal of two dispersals, one along the Ucayali River, into western Amazonia, northwestern South America and finally Central America; the other along the Madeira River into central and then eastern Amazonia. The first dispersal resulted in very starchy fruit for fermentation, while the second may have been later and resulted in snack fruits. Further explorations of southwestern Amazonia are essential for more precise identification of the earliest events, both with new archeological methods and genetic analyses with larger samples.

Wood density (WD) relates to important tree functions such as stem mechanics and resistance against pathogens. This functional trait can exhibit high intraindividual variability both radially and vertically. With the rise of LiDAR-based methodologies allowing nondestructive tree volume estimations, failing to account for WD variations related to tree function and biomass investment strategies may lead to large systematic bias in AGB estimations. Here, we use a unique destructive dataset from 822 trees belonging to 51 phylogenetically dispersed tree species harvested across forest types in Central Africa to determine vertical gradients in WD from the stump to the branch tips, how these gradients relate to regeneration guilds and their implications for AGB estimations. We find that decreasing WD from the tree base to the branch tips is characteristic of shade-tolerant species, while light-demanding and pioneer species exhibit stationary or increasing vertical trends. Across all species, the WD range is narrower in tree crowns than at the tree base, reflecting more similar physiological and mechanical constraints in the canopy. Vertical gradients in WD induce significant bias (10%) in AGB estimates when using database-derived species-average WD data. However, the correlation between the vertical gradients and basal WD allows the derivation of general correction models. With the ongoing development of remote sensing products providing 3D information for entire trees and forest stands, our findings indicate promising ways to improve greenhouse gas accounting in tropical countries and advance our understanding of adaptive strategies allowing trees to grow and survive in dense rainforests.

Understanding tree growth and carbon sequestration are of crucial interest to forecast the feedback of forests to climate change. To have a global understanding of the wood formation, it is necessary to develop new methodologies for xylogenesis measurements, valid across diverse wood structures and applicable to both angiosperms and gymnosperms. In this study, the authors present a new workflow to study xylogenesis using high-resolution X-ray computed tomography (HRXCT), which is generic and offers high potential for automatization. The HXRCT-based approach was benchmarked with the current classical approach (microtomy) on three tree species with contrasted wood anatomy (Pinus nigra, Fagus sylvatica, and Quercus robur). HRXCT proved to estimate the relevant xylogenesis parameters (timing, duration, and growth rates) across species with high accuracy. HRXCT showed to be an efficient avenue to investigate tree xylogenesis for a wide range of wood anatomies, structures, and species. HRXCT also showed its potential to provide quantification of intra-annual dynamics of biomass production through high-resolution 3D mapping of wood biomass within the forming growth ring.