Comparative analysis of floral transition and floral organ formation in two contrasting species: Disocactus speciosus and D. eichlamii (Cactaceae).

KEY MESSAGE: Contrasting morphologies in Disocactus are the result of differential development of the vegetative and floral tissue where intercalary growth is involved, resulting in a complex structure, the floral axis. Species from the Cactaceae bear adaptations related with their growth in environments under hydric stress. These adaptations have translated into the reduction and modification of various structures such as leaves, stems, lateral branches, roots and the structuring of flowers in a so-called flower-shoot. While cacti flowers and fruits have a consistent structure with showy hermaphrodite or unisexual flowers that produce a fruit called cactidium, the developmental dynamics of vegetative and reproductive tissues comprising the reproductive unit have only been inferred through the analysis of pre-anthetic buds. Here we present a comparative analysis of two developmental series covering the early stages of flower formation and organ differentiation in Disocactus speciosus and Disocactus eichlamii, which have contrasting floral morphologies. We observe that within the areole, a shoot apical meristem commences to grow upward, producing lateral leaves with a spiral arrangement, rapidly transitioning to a floral meristem. The floral meristem produces tepal primordia and a staminal ring meristem from which numerous or few stamens develop in a centrifugal manner in D. speciosus and D. eichlamii, respectively. Also, the inferior ovary derives from the floral meristem flattening and an upward growth of the surrounding tissue of the underlying stem, producing the pericarpel. This structure is novel to cacti and lacks a clear anatomical delimitation with the carpel wall. Here, we present a first study that documents the early processes taking place during initial meristem determination related to pericarpel development and early floral organ formation in cacti until the establishment of mature floral organs.

Phylogenetic and morphological analyses of Pilosocereus leucocephalus group s.s. (Cactaceae) reveal new taxonomical implications.

Pilosocereus is one of the Cactaceae family's most relevant genera in terms of the number of species and its wide geographical range in the Americas. Within Pilosocereus, five informal taxonomic groups have been recognized, one of which is P. leucocephalus group s.s., whose phylogenetic relationships remain unresolved. Therefore, our objectives are to recognize the circumscriptions of the species in P. leucocephalus group s.s. and to corroborate the monophyly and phylogenetic relationships of this group through a set of morphological and molecular characters. This study is based on representative sampling along the broad distribution of this group in Mexico and Central America using multivariate and phylogenetic analyses. The morphological characters identified to contribute to species recognition and group formation are branch diameter, areole length, the areole length-width ratio, the distance between areoles, the length of the longest radial spine, and branch and spines colors. The chloroplast markers rpl16, trnL-trnF, and petL-psbE and the nuclear marker AT1G18270 support the monophyly of the P. leucocephalus group s.s., and two probable synapomorphies are suggested, including one transversion in rpl16 and another in petL-psbE. Together, our results demonstrate that sampled species of P. leucocephalus group s.s. encompass six species distributed in Mexico and Central America: P. alensis and P. purpusii in the western region, P. chrysacanthus and P. collinsii in the central region, and P. gaumeri and P. leucocephalus in the eastern region. A taxonomic key to recognized species is provided.

Pollen morphological investigations of family Cactaceae and its taxonomic implication by light microscopy and scanning electron microscopy.

The family Cactaceae is the diversified group of angiosperm plants whose pollen statistics has been used for taxonomic identification. In this article, we present the pollen morphology of eight species belong to seven taxonomically complex genera of Cactaceae including Astrophytum, Cylindropuntia, Echinocereus, Echinopsis, Mammillaria, Opuntia, and Thelocactus using light and scanning electron microscopy. The pollen grains were acetolyzed, measured, described, and electron photomicrographs were taken. Cactaceae can be characterized by presenting different palynomorphological features including pollen type, sculpturing, polar and equatorial diameter, aperture orientation, exine thickness, P/E ratio, and echini features. Four types of pollen shapes, that is, prolate spheroidal (three species), subprolate (two species), prolate (two species), and oblate spheroidal in Echinocereus reichenbachii were observed. The polar and equatorial diameter observed maximum in O. ficus indica 116.95 and 112.27 µm while minimum in M. compressa 38.42 and 21.05 µm. Pollen of two types, tricolpate in members of subfamily Cactioideae and pantoporate in the Opuntioideae were examined. The fertility percentage has been observed maximum in Opuntia macrocentra (83.84%) and minimum in Opuntia ficus-indica (57.89%). Exine sculpturing showing great variations such as granulate, reticulate, granulate perforate and micro-echinate foveolate ornamentation was examined only in Echinopsis eyriesii. A key to species, based on pollen micromorphological attributes, has been constructed for correct identification of complex cactus species.

Water droplet dynamics on bioinspired conical surfaces.

Cacti use the Laplace pressure gradient due to conical geometry as a mechanism for collecting water from fog. Bioinspired surfaces using conical geometry can be developed for water collection from fog for human consumption. A systematic study is presented which investigates the dynamics of water droplets on a bioinspired conical surface. A series of experiments was conducted where a known volume of droplets was deposited on the cone. This was followed by an investigation into droplet dynamics where the droplets are deposited from fog and the volume is unknown. This includes a study on the macroscopic level as well as the microscopic level. The main parameters that were varied for these tests were the tip angle and the cone orientation. The droplet movement observed was compared relatively. Based on captured videos of droplet movement, distance travelled and velocities were measured. The Laplace pressure gradient, gravity and droplet coalescence were found to be the mechanisms of droplet movement on a conical surface. The findings of this study should be of interest in designing bioinspired surfaces with high water collection. This article is part of the theme issue 'Bioinspired materials and surfaces for green science and technology (part 2)'.

The influence of cactus spine surface structure on puncture performance and anchoring ability is tuned for ecology.

Spines are common morphological features found in almost all major biological groups offering an opportunity to explore large-scale evolutionary convergence across disparate clades. As an example, opuntioid cacti have spines with barbed ornamentation that is remarkably similar in form and scale to that found on porcupine quills, suggesting specific biomechanical convergence across the animal and plant kingdoms. While the mechanics of porcupine quills as defensive mechanisms has been previously tested, the mechanics of cactus spines (which have evolved to fill a number of functions including defence, climbing and dispersal) has not been characterized. Here we study the puncturing and anchoring ability of six species of cactus, including both barbed and non-barbed spines. We found that barbed spines require less work to puncture a variety of targets than non-barbed spines. Barbed spines also require more work than non-barbed spines to withdraw from biological materials, owing to their barbs engaging with tissue fibres. These results closely match those found previously for barbed versus non-barbed porcupine quills, implying biomechanical convergence. The variation in performance of barbed versus non-barbed spines, as well as between barbed spines from different species, is probably tied to the diversity of ecological functions of cactus spines.

Gas exchange characteristics of giant cacti species varying in stem morphology and life history strategy.

PREMISE OF THE STUDY: Giant cacti species possess long cylindrical stems that store massive amounts of water and other resources to draw on for photosynthesis, growth, and reproduction during hot and dry conditions. Across all giant cacti taxa, stem photosynthetic surface area to volume ratio (S:V) varies by several fold. This broad morphological diversity leads to the hypothesis that giant cacti function along a predictable resource use continuum from a "safe" strategy reflected in low S:V, low relative growth rates (RGR), and low net assimilation rates (Anet ) to a high-risk strategy that is reflected in high S:V, RGR, and Anet . METHODS: To test this hypothesis, whole-plant gas exchange, chlorophyll fluorescence, and whole-spine-tissue carbon isotope ratios (δ13 C) were measured in four giant cacti species varying in stem morphology and RGR. Measurements were conducted on five well-watered, potted plants per species. KEY RESULTS: Under conditions of mild diel temperatures and low atmospheric vapor pressure deficit, Anet , transpiration (E), and stomatal conductance (Gs ) were significantly higher, and water-use efficiency (Anet : Gs ) was lower in fast-growing, multi-stemmed species compared to the slower growing, single-stemmed species. However, under warmer, less optimal conditions, gas exchange converged between stem types, and neither δ13 C nor chlorophyll fluorescence varied among species. CONCLUSIONS: The results add to a growing body of evidence that succulent-stemmed plants function along a similar economic spectrum as leaf-bearing plants such that functional traits including stem RGR, longevity, morphology, and gas exchange are correlated across species with varying life-history strategies.

Effect of flower orientation on the male and female traits of Myrtillocactus geometrizans (Cactaceae).

Intra-individual variation in the production and size of reproductive traits has been documented in columnar cacti, being higher in equator-facing flowers. Such variation is attributed to the high amount of PAR intercepted by stems oriented towards the equator. Most studies focused on this phenomenon have documented the existence of intra-individual variation on traits associated with the female function; however, its impact on traits associated with the male function has been neglected. We tested the hypothesis that equator-facing flowers of Myrtillocactus geometrizans exhibit higher values on traits associated with both male and female functions than flowers facing against it. Number and size of anthers and ovaries, pollen:ovule ratio and number and quality of pollen grains (diameter, germinability, viability and pollen tube length) were estimated from reproductive structures facing north and south, and compared with t-tests between orientations. Number of anthers per flower, number of pollen grains per anther and per floral bud; pollen size, viability and germinability; pollen tube length; ovary length and pollen:ovule ratio were significantly higher in reproductive structures oriented towards the south (i.e. equator). These findings suggest that intra-individual variation in floral traits of M. geometrizans might be associated with different availability of resources in branches with contrasting orientation. Our results provide new evidence of the existence of a response to an orientation-dependent extrinsic gradient. To our knowledge, this is the first study documenting the existence of intra-individual variation on pollen quality and P:O ratio in Cactaceae species.

Biomimetic optimisation of branched fibre-reinforced composites in engineering by detailed analyses of biological concept generators.

The aim of this study is the biomimetic optimisation of branched fibre-reinforced composites based on the detailed analysis of biological concept generators. The methods include analyses of the functional morphology and biomechanics of arborescent monocotyledons and columnar cacti as well as measurements and modelling of mechanical properties of biomimetic fibre-reinforced composites. The key results show evidence of notch stress reduction by optimised stem-branch-attachment morphology in monocotyledons and columnar cacti. It could be shown that some of these highly interesting properties can be transferred into biomimetic fibre-reinforced composites.

Occurrence and characterisation of calcium oxalate crystals in stems and fruits of Hylocereus costaricensis and Selenicereus megalanthus (Cactaceae: Hylocereeae).

Detailed description about occurrence of calcium oxalate (CaOx) crystals in the edible vine cactus species Hylocereus costaricensis and Selenicereus megalanthus is scarce. Therefore, we evaluated and characterized the presence, morphology and composition of CaOx crystals in both species. Crystals were isolated from greenhouse and in vitro vegetative stems, and from ripe fruit peels and pulp by enzymatic digestion and density centrifugation and quantified with a haemocytometer. Morphologies were studied using scanning electron microscopy, elemental composition with energy-dispersive X-ray spectroscopy and salt composition with X-ray powder diffraction. Analyses conducted confirmed that isolated crystals were exclusively composed by CaOx, both mono- and dihydrated. Highest crystal contents were measured in greenhouse stems, followed by the fruit peels. While very few crystals were quantified in in vitro plants, they were not detected in the fruit pulp at all, which is of advantage for its human consumption and could be linked to mechanisms of seed dispersal through animals. Different crystal morphologies were observed, sometimes varying between genotypes and tissues analysed. This is the first work known to the authors with a detailed characterization of CaOx crystals in vine cacti.

Hierarchical structures of cactus spines that aid in the directional movement of dew droplets.

Three species of cactus whose spines act as dew harvesters were chosen for this study: Copiapoa cinerea var. haseltoniana, Mammillaria columbiana subsp. yucatanensis and Parodia mammulosa and compared with Ferocactus wislizenii whose spines do not perform as dew harvesters. Time-lapse snapshots of C. cinerea showed movement of dew droplets from spine tips to their base, even against gravity. Spines emanating from one of the areoles of C. cinerea were submerged in water laced with fluorescent nanoparticles and this particular areole with its spines and a small area of stem was removed and imaged. These images clearly showed that fluorescent water had moved into the stem of the plant. Lines of vascular bundles radiating inwards from the surface areoles (from where the spines emanate) to the core of the stem were detected using magnetic resonance imaging, with the exception of F. wislizenii that does not harvest dew on its spines. Spine microstructures were examined using SEM images and surface roughness measurements (Ra and Rz) taken of the spines of C. cinerea It was found that a roughness gradient created by tapered microgrooves existed that could potentially direct surface water from a spine tip to its base.This article is part of the themed issue 'Bioinspired hierarchically structured surfaces for green science'.

Condensation on slippery asymmetric bumps.

Controlling dropwise condensation is fundamental to water-harvesting systems, desalination, thermal power generation, air conditioning, distillation towers, and numerous other applications. For any of these, it is essential to design surfaces that enable droplets to grow rapidly and to be shed as quickly as possible. However, approaches based on microscale, nanoscale or molecular-scale textures suffer from intrinsic trade-offs that make it difficult to optimize both growth and transport at once. Here we present a conceptually different design approach--based on principles derived from Namib desert beetles, cacti, and pitcher plants--that synergistically combines these aspects of condensation and substantially outperforms other synthetic surfaces. Inspired by an unconventional interpretation of the role of the beetle's bumpy surface geometry in promoting condensation, and using theoretical modelling, we show how to maximize vapour diffusion fluxat the apex of convex millimetric bumps by optimizing the radius of curvature and cross-sectional shape. Integrating this apex geometry with a widening slope, analogous to cactus spines, directly couples facilitated droplet growth with fast directional transport, by creating a free-energy profile that drives the droplet down the slope before its growth rate can decrease. This coupling is further enhanced by a slippery, pitcher-plant-inspired nanocoating that facilitates feedback between coalescence-driven growth and capillary-driven motion on the way down. Bumps that are rationally designed to integrate these mechanisms are able to grow and transport large droplets even against gravity and overcome the effect of an unfavourable temperature gradient. We further observe an unprecedented sixfold-higher exponent of growth rate, faster onset, higher steady-state turnover rate, and a greater volume of water collected compared to other surfaces. We envision that this fundamental understanding and rational design strategy can be applied to a wide range of water-harvesting and phase-change heat-transfer applications.

How specialised is bird pollination in the Cactaceae?

Many cactus species produce 'bird' flowers; however, the reproductive biology of the majority of these species has not been studied. Here, we report on a study of the pollination of two species from the Cleistocactus genus, cited as an ornithophilous genus, in the context of the different ways in which they are specialised to bird pollination. In addition, we re-evaluate the level of specialisation of previous studies of cacti with bird pollination and evaluate how common phenotypic specialisation to birds is in this family. Both Cleistocactus species exhibited ornithophilous floral traits. Cleistocactus baumannii was pollinated by hummingbirds, whereas Cleistocactus smaragdiflorus was pollinated by hummingbirds and bees. Pollination by birds has been recorded in 27 cactus species, many of which exhibit ornithophilous traits; however, they show generalised pollination systems with bees, bats or moths in addition to birds being their floral visitors. Of all cactus species, 27% have reddish flowers. This trait is associated with diurnal anthesis and a tubular shape. Phenotypic specialisation to bird pollination is recognised in many cactus species; however, it is not predictive of functional and ecological specialisation in this family.

Effective directional self-gathering of drops on spine of cactus with splayed capillary arrays.

We report that the fast droplet transport without additional energy expenditure can be achieved on the spine of cactus (Gymnocalycium baldianum) with the assistance of its special surface structure: the cactus spine exhibits a cone-like structure covered with tilted scales. A single scale and the spine surface under it cooperatively construct a splayed capillary tube. The arrays of capillary tube formed by the overlapping scales build up the out layer of the spine. The serial drops would be driven by the asymmetric structure resulted from tilt-up scales-by-scales on the cone-shaped spine, and move directionally toward the bottom from top of spine, by means of the Laplace pressure in differences. In addition, after the past of the first droplet, thin liquid film of drop is trapped in the splayed capillary micro-tube on the surface of spine, which greatly reduces the friction of subsequential droplet transport in efficiency. This finding provides a new biological model which could be used to transport droplet spontaneously and directionally. Also this work offers a way to reduce the surface adhesion by constructing liquid film on the surface, which has great significance in prompting droplet transport efficiency.

Physical and chemical characteristics of pitaya fruits at physiological maturity.

The aim of this study was to analyze the physical and chemical characteristics of the maturation process of pitaya fruit (Hylocereus undatus) to identify indicators that can be used to determine the point of physiological maturity and establish the optimal timing of physiological maturity for harvesting the fruit. A completely randomized experimental design was employed and four biological repeats were performed. Physiological maturity was assessed using various physical characteristics: longitudinal length (LL), equatorial diameter (ED), pericarp thickness (PeT), pulp thickness (PuT), fruit mass (FM), pulp mass (PuM), pericarp mass (PeM), pericarp percentage (%Pe), pulp percentage (%Pu), pulp/pericarp ratio (Pu/Pe), pericarp color index (CI), hue color angle (h°), lightness index (L*), chroma (C*), blue-yellow variation (b*), and green-red variation (a*). Additionally, chemical characteristics such as soluble solid content (SS), titratable acidity (TA), SS/TA ratio, and pH were screened. The data were statistically analyzed by fitting regression models and computing Pearson's correlation coefficients (P < 0.05). Physiological maturity in pitaya fruits occurred between the 30th and 32nd days after anthesis, and this proved to be the optimal period for harvest. At this time, the fruit was completely red with high SS, and had the recommended values of TA, pH, and SS/TA ratio. During this period, ED, PuT, FM, PuM, %Pu, and Pu/Pe increased while PeT, PeM, and %Pe fell; these changes are considered desirable by producers and/or consumers. PuM was the variable that displayed more strong's association with other variables in the analysis.

How and why does the areole meristem move in Echinocereus (Cactaceae)?

BACKGROUND AND AIMS: In Cactaceae, the areole is the organ that forms the leaves, spines and buds. Apparently, the genus Echinocereus develops enclosed buds that break through the epidermis of the stem adjacent to the areole; this trait most likely represents a synapomorphy of Echinocereus. The development of the areole is investigated here in order to understand the anatomical modifications that lead to internal bud development and to supplement anatomical knowledge of plants that do not behave according to classical shoot theory. METHODS: The external morphology of the areole was documented and the anatomy was studied using tissue clearing, scanning electron microscopy and light microscopy for 50 species that represent the recognized clades and sections of the traditional classification of the genus, including Morangaya pensilis (Echinocereus pensilis). KEY RESULTS: In Echinocereus, the areole is sealed by the periderm, and the areole meristem is moved and enclosed by the differential growth of the epidermis and surrounding cortex. The enclosed areole meristem is differentiated in a vegetative or floral bud, which develops internally and breaks through the epidermis of the stem. In Morangaya pensilis, the areole is not sealed by the periderm and the areole meristem is not enclosed. CONCLUSIONS: The enclosed areole meristem and internal bud development are understood to be an adaptation to protect the meristem and the bud from low temperatures. The anatomical evidence supports the hypothesis that the enclosed bud represents one synapomorphy for Echinocereus and also supports the exclusion of Morangaya from Echinocereus.

Comb-calibrated laser ranging for three-dimensional surface profiling with micrometer-level precision at a distance.

Non-contact surface mapping at a distance is interesting in diverse applications including industrial metrology, manufacturing, forensics, and artifact documentation and preservation. Frequency modulated continuous wave (FMCW) laser detection and ranging (LADAR) is a promising approach since it offers shot-noise limited precision/accuracy, high resolution and high sensitivity. We demonstrate a scanning imaging system based on a frequency-comb calibrated FMCW LADAR and real-time digital signal processing. This system can obtain three-dimensional images of a diffusely scattering surface at stand-off distances up to 10.5 m with sub-micrometer accuracy and with a precision below 10 µm, limited by fundamental speckle noise. Because of its shot-noise limited sensitivity, this comb-calibrated FMCW LADAR has a large dynamic range, which enables precise mapping of scenes with vastly differing reflectivities such as metal, dirt or vegetation. The current system is implemented with fiber-optic components, but the basic system architecture is compatible with future optically integrated, on-chip systems.

Ecophysiological and anatomical mechanisms behind the nurse effect: which are more important? A multivariate approach for cactus seedlings.

BACKGROUND: Cacti establish mostly occurs under the canopy of nurse plants which provide a less stressful micro-environment, although mechanisms underlying this process are unknown. The impact of the combination of light and watering treatments on Opuntia streptacantha (Cactaceae) seedlings was examined. METHODS/PRINCIPAL FINDINGS: Ecophysiological [titratable acidity, osmotic potential ('solute potential', Ψs ), relative growth rate (RGR) and their components (NAR, SLA, and LWR)], anatomical (chloroplast density, chloroplast frequency, and cell area), and environmental [photosynthetic photon flux density (PPFD) and air temperature] sets of variables were analyzed, assessing relationships between them and measuring the intensity of the relationships. Three harvests were carried out at days 15, 30, and 45. Ψs and acidity content were the most important responses for seedling establishment. The main anatomical and environmental variables were chloroplast density and water availability, respectively. Opuntia streptacantha seedlings establish better in the shade-watering treatment, due to higher Ψs and acidity, unaffected chloroplasts, and lower PPFD. In addition, the chloroplasts of cells under high-light and non-watering treatment were clumped closer to the center of the cytosol than those under shade-drought, to avoid photoinhibition and/or to better distribute or utilize the penetrating light in the green plant tissue. CONCLUSIONS: Opuntia seedlings grow better under the shade, although they can tolerate drought in open spaces by increasing and moving chloroplasts and avoiding drastic decreases in their Ψs . This tolerance could have important implications for predicting the impact of climate change on natural desert regeneration, as well as for planning reforestation-afforestation practices, and rural land uses.

Genetic diversity of spineless Cereus jamacaru accessions using morphological and molecular markers.

This is the first study to examine the genetic diversity of mandacaru cactus (Cereus jamacaru P. DC.). Plants of spineless mandacaru are commonly found in gardens and parks of urban areas in northeastern Brazil. In addition to exploring their ornamental potential, morphological, and genetic characterization may contribute to the development of plant materials that can be used as a source of macromolecules of potential economic interest. The goal of this study was to estimate the genetic variability of spineless mandacaru accessions using random amplified polymorphic DNA (RAPD) and inter-simple sequence repeat (ISSR) molecular markers, and to characterize their morphology. Ten samples of newly emitted shoots with differentiated areolas and ribs were collected from each accession from the Cactaceous Germplasm Collection of Embrapa Agroindústria Tropical, in Fortaleza, CE. Shoot shape and aspects of spine primordia (presence, location, grouping, and size of spines) were evaluated. The morphological analysis showed that the spineless mandacaru presented spine primordia. Twenty-six RAPD and 15 ISSR primers were polymorphic. A total of 262 markers were obtained, 129 of which were polymorphic. The average polymorphism of ISSR markers was higher than that of RAPD markers. The dendrograms for both analyses showed differentiation between accessions. Nevertheless, the molecular markers detected higher levels of diversity and a different pattern of diversity than those found using morphological markers. The molecular results revealed significant genetic variability both within and between groups.

Morphological, cytological and metabolic consequences of autopolyploidization in Hylocereus (Cactaceae) species.

BACKGROUND: Genome doubling may have multi-level effects on the morphology, viability and physiology of polyploids compared to diploids. We studied the changes associated with autopolyploidization in two systems of somatic newly induced polyploids, diploid-autotetraploid and triploid-autohexaploid, belonging to the genus Hylocereus (Cactaceae). Stomata, fruits, seeds, embryos, and pollen were studied. Fruit pulp and seeds were subjected to metabolite profiling using established gas chromatography-mass spectrometry (GC-MS) and ultra-performance liquid chromatography (UPLC) Q-TOF-MS/MS (time of flight)-protocols. RESULTS: Autopolyploid lines produced lower numbers of tetrads, larger pollen grains with lower viability, larger stomata with lower density, and smaller fruits with lower seed numbers and decreased seed viability. The abundance of sugars was lower in the fruits and seeds of the two duplicated lines than in their donor lines, accompanied by increased contents of amino acids, tricarboxylic acid (TCA) cycle intermediates, organic acids and flavonoids. Betacyanins, the major fruit pigments in diploid and triploid donors, decreased following genome doubling. Both autopolyploid Hylocereus lines thus exhibited unfavorable changes, with the outcome being more dramatic in the autohexaploid than in the autotetraploid line. CONCLUSION: Induced autotetraploid and autohexaploid lines exhibited morphological and cytological characteristics that differed from those of their donor plants and that were accompanied by significant metabolic alterations. It is suggested that a developmental arrest occurs in the fruits of the autohexaploid line, since their pericarp shows a greater abundance of acids and of reduced sugars. We conclude that genome doubling does not necessarily confer a fitness advantage and that the extent of alterations induced by autopolyploidization depends on the genetic background of the donor genotype.

Functional morphology and biomechanics of branch-stem junctions in columnar cacti.

Branching in columnar cacti features morphological and anatomical characteristics specific to the subfamily Cactoideae. The most conspicuous features are the pronounced constrictions at the branch-stem junctions, which are also present in the lignified vascular structures within the succulent cortex. Based on finite-element analyses of ramification models, we demonstrate that these indentations in the region of high flexural and torsional stresses are not regions of structural weakness (e.g. allowing vegetative propagation). On the contrary, they can be regarded as anatomical adaptations to increase the stability by fine-tuning the stress state and stress directions in the junction along prevalent fibre directions. Biomimetic adaptations improving the functionality of ramifications in technical components, inspired, in particular, by the fine-tuned geometrical shape and arrangement of lignified strengthening tissues of biological role models, might contribute to the development of alternative concepts for branched fibre-reinforced composite structures within a limited design space.