The legacy and evolvability of Pere Alberch's ideas
Pere Alberch played a pivotal role in shaping the field of evolutionary developmental biology during the 1980s and 1990s. Whereas initially his contributions were sidelined by the empirical advancements of the molecular revolution in developmental and evolutionary biology, his theoretical insights have left a lasting impact on the discipline. This article provides a comprehensive review of the legacy and evolvability of Alberch's ideas in contemporary evo-devo, which included the study of morphogenesis as the proper level of developmental causation, the interplay between developmental constraints and natural selection, the epistemic role of teratologies, the origin of evolutionary novelties and the concept of evolvability.
The logic of monsters: development and morphological diversity in stem-cell-based embryo models
Organoids and stem-cell-based embryo models (SEMs) are imperfect organ or embryo representations that explore a much larger space of possible forms, or morphospace, compared to their counterparts. Here, we discuss SEM biology in light of seminal work by Pere Alberch, a leading figure in early evo-devo, interpreting SEMs as developmental 'monstrosities' in the Alberchian sense. Alberch suggested that ordered patterns in aberrant development-i.e. 'the logic of monsters'-reveal developmental constraints on possible morphologies. In the same vein, we detail how SEMs have begun to shed light on structural features of normal development, such as developmental variability, the relative importance of internal versus external constraints, boundary conditions and design principles governing robustness and canalization. We argue that SEMs represent a powerful experimental tool to explore and expand developmental morphospace and propose that the 'monstrosity' of SEMs can be leveraged to uncover the 'hidden' rules and developmental constraints that robustly shape and pattern the embryo.
Fundamental constraints to the logic of living systems
It has been argued that the historical nature of evolution makes it a highly path-dependent process. Under this view, the outcome of evolutionary dynamics could have resulted in organisms with different forms and functions. At the same time, there is ample evidence that convergence and constraints strongly limit the domain of the potential design principles that evolution can achieve. Are these limitations relevant in shaping the fabric of the possible? Here, we argue that fundamental constraints are associated with the logic of living matter. We illustrate this idea by considering the thermodynamic properties of living systems, the linear nature of molecular information, the cellular nature of the building blocks of life, multicellularity and development, the threshold nature of computations in cognitive systems and the discrete nature of the architecture of ecosystems. In all these examples, we present available evidence and suggest potential avenues towards a well-defined theoretical formulation.
Capacity building in porous materials research for sustainable energy applications
The project aimed to develop porous materials for sustainable energy applications, namely, hydrogen storage, and valorization of biomass to renewable fuels. At the core of the project was a training programme for Africa-based researchers in (i) the exploitation of renewable locally available raw materials; (ii) the use of advanced state-of-the-art techniques for the design and synthesis of porous materials (zeolites and metal-organic frameworks (MOFs)) for energy storage; and (iii) the valorization of sustainable low-value feedstock to renewable fuels. We found that compaction of the UiO-66 MOF at high pressure improves volumetric hydrogen storage capacity without any loss in gravimetric uptake, and experimentally demonstrated the temperature-dependent dynamic behaviour of UiO-66, which allowed us to propose an activation temperature of ≤ 150°C for UiO-66. Co-pelletization was used to fabricate UiO-66/nanofibre monoliths as hierarchical porous materials with enhanced usable (i.e. deliverable) hydrogen storage capacity. We clarified the use of naturally occurring kaolin as a source of silica and alumina species for zeolite synthesis. The kaolin-derived zeolite X was successfully used as a catalyst for the transesterification of oil (from non-edible biomass) to biodiesel. We also prepared porous composites (i.e. carbon/UiO-66, organoclay/UiO-66 and zeolite/carbon) that were successfully applied in electrochemical sensing.
RS-DFID Africa capacity-building initiative programme grant: harnessing unsteady phase-change heat exchange in high-performance concentrated solar power systems
The Royal Society and UK Department for International Development supported a consortium of three universities across sub-Saharan Africa and Imperial College London with the aim of developing new knowledge on direct-steam-generation concentrated solar power (CSP) plants and supporting relevant capacity building across the Universities of Lagos, Mauritius and Pretoria. Key research findings from the programme include an improved flow-classification scheme for two-phase, liquid-liquid flows; testing of advanced surfaces with much-improved steady-state heat transfer performance-the commercial nanoFLUX surface showed up to 200% higher heat-transfer coefficients (HTCs) in pool boiling compared with other surfaces with R-134a/R-245fa; first-of-a-kind measurements of transient flow boiling HTCs, which were up to 30% lower in step perturbations than quasi-steady-state expectations in horizontal pipes with R-245fa; error estimation and corrections for laser-induced fluorescence (LIF) measurements, leading to the development of an adapted planar LIF technique with uncertainty <10% for local, instantaneous film thickness measurements in annular flows, and the application of such diagnostic methods to pool, falling-film and flow boiling in pipes; and predictions of an ~80% increase in the net present value of a case-study CSP plant when integrated with solid storage media.
Exploring the West African forest island phenomenon: scientific insights gained, successes achieved and capacities strengthened
Anthropogenic activities around local villages in mesic savanna landscapes of West Africa have resulted in soil improvement and forest establishment outside their climatic zones. Such unique 'forest islands' have been reported to provide ecosystem services including biodiversity conservation. However, the science underpinning their formations is limitedly studied. In 2015 and with funding support from the Royal Society-DFID (now FCDO), we set out to investigate the biogeochemistry of the forest islands in comparison with adjacent natural savanna and farmlands across 11 locations in Burkina Faso, Ghana and Nigeria. Our results showed that the forest islands do not differ significantly from the adjoining ecosystems in soil mineralogy implying that their formation was anthropogenically driven. We observed greater soil organic carbon and nutrient distributions in the forest islands, which also had more stable macro (>500 μm) and meso-aggregates (500-250 μm) than the adjoining agricultural lands. We found that soil micro-aggregate (250-53 μm) stability was climate (precipitation) driven in the West African ecosystems while meso- and macro-aggregate stability was land-use driven. In one of the unique forest islands we studied in the Mole National Park of Ghana, we found its mineral-associated organic carbon over 40% greater than the adjoining natural savanna with potential implications for the achievement of the global initiative of the '4p1000' in West Africa. We conclude that the North-South-South research collaboration has established clearly, the science underlying the age-long West African forest island phenomenon and has, among many successes, led to capacity building of young scientists driving cutting-edge research in climate change adaptation and food systems transformation in the sub-region.
Creating sustainable capacity for river science in the Congo basin through the CRuHM project
In this article, we examine the scientific and sustainable research capacity outcomes of the 'Congo River: user Hydraulics and Morphology' or CRuHM project, a six-year effort supported by the Royal Society's Africa Capacity Building Initiative. This project brought together a consortium of African and UK universities to undertake the first large-scale scientific expeditions to the Congo basin of the modern era in order to better understand the hydraulics and geomorphology of this understudied but globally important river. The river is essential for navigation, irrigation, drinking water and hydroelectric power generation for the 10 basin countries and is critically important for biodiversity and global nutrient, carbon and climatological cycles. This article summarizes the new scientific understanding contributed by the project and the steps taken to ensure a meaningful legacy that would continue long beyond the finite lifetime of available funding. Actions taken to achieve this include establishing a new hydrology research centre at the University of Kinshasa as well as steps to build a wider international community of Congo basin researchers. In this way, we hope to build momentum for future funding initiatives and collaboration.
Doctoral training to support sustainable soil geochemistry research in Africa
Africa's potential for scientific research is not yet being realized, for various reasons including a lack of researchers in many fields and insufficient funding. Strengthened research capacity through doctoral training programmes in higher education institutes (HEIs) in Africa, to include collaboration with national, regional and international research institutions, can facilitate self-reliant and sustainable research to support socio-economic development. In 2012, the Royal Society and the UK's Department for International Development (now the Foreign, Commonwealth and Development Office) launched the Africa Capacity Building Initiative (ACBI) Doctoral Training Network which aimed to strengthen research capacity and training across sub-Saharan Africa. The ACBI supported 30 core PhD scholarships, all registered/supervised within African HEIs with advisory support from the UK-based institutes. Our 'Soil geochemistry to inform agriculture and health policies' consortium project, which was part of the ACBI doctoral training programme network, was implemented in Malawi, Zambia and Zimbabwe between 2014 and 2020. The aims of our consortium were to explore linkages between soil geochemistry, agriculture and public health for increased crop productivity, nutrition and safety of food systems and support wider training and research activities in soil science. Highlights from our consortium included: (i) the generation of new scientific evidence on linkages between soils, crops and human nutrition; (ii) securing new projects to translate science into policy and practice; and (iii) maintaining sustainable collaborative learning across the consortium. Our consortium delivered high-quality science outputs and secured new research and doctoral training funding from a variety of sources to ensure the continuation of research and training activities. For example, follow-on Global Challenges Research Funded Translation Award provided a strong evidence base on the prevalence of deficiencies in children under 5 years of age and women of reproductive age in Zimbabwe. This new evidence will contribute towards the design and implementation of a nationally representative micronutrient survey as an integral part of the Zimbabwe Demographic and Health Surveys conducted by the Ministry of Health and Child Care. The award also generated new evidence and a road map for creating quality innovative doctorates through a doctoral training landscape activity led by the Zimbabwe Council for Higher Education. Although our project and the wider ACBI has contributed to increasing the self-reliance and sustainability of research within the region, many challenges remain and ongoing investment is required.
Materials modelling in the University of Limpopo
This article provides insights into building research capacity in computational modelling of materials at the University of Limpopo (UL), formerly University of the North, in South Africa, through a collaboration with a consortium of universities in the United Kingdom (UK) through the support of the National Research Foundation (NRF), formerly the Foundation for Research and Development, and the Royal Society (RS). A background that led to the choice of building research capacity at historically disadvantaged universities in South Africa, including the UL, is given. The of the collaboration between the UL and several UK universities on computational modelling of materials is outlined, together with the scientific highlights that were achieved in themes of minerals, energy storage and alloy development. The capacity built in terms of human capital and institutions set up is shared, which is followed by a discussion of the continuing research activities after the formal NRF-RS collaboration ceased with more alignment to industrial applications with national and international support. We conclude by highlighting the success of the project in capacity-building and consolidating the Materials Modelling Centre with developments of high-performance computing in South Africa and the African continent. We comment on the lessons learned regarding successful capacity-building programmes.
Portrait of a UK-Africa Capacity Building Initiative Consortium 2015-2022: the Cameroon, Ghana, South Africa and United Kingdom Materials Initiative (CaGSUMI) for developing materials for solar cells
The CaGSUMI consortium was funded by the Royal Society-Department for International Development (later the Foreign, Commonwealth & Development Office) on the Africa Capacity Building Initiative programme between the years 2015 and 2022 and involved three Sub-Saharan African universities: Kwame Nkrumah University of Science and Technology, Kumasi, Ghana, University of Yaoundé I, Cameroon, and the University of Zululand, South Africa; and the University of Manchester in the United Kingdom. The project was used to cement an emergent UK-Africa network in the areas of materials chemistry related to renewable energy generation with both thin films and nanomaterials. The consortium's outputs led to numerous publications of African science in international journals, a number of graduated PhDs who went on to permanent academic positions and prestigious fellowships, the establishment of a capacity-building plan relevant to the chemistry departments in each of the African countries, and the installation of a number of first-in-kind pieces of kit for African laboratories that will keep them on a competitive footing at an international level for the next decade and more.
Chem4Energy: a consortium of the Royal Society Africa Capacity-Building Initiative
The Africa Capacity-Building Initiative is a Royal Society programme funded by the former UK Department for International Development to develop collaborative research between scientists in sub-Saharan Africa and the UK. Initially, four institutions were involved in the Chem4Energy consortium: Cardiff University in the UK and three African partners, the Kwame Nkrumah University of Science and Technology, Ghana, the University of Namibia and the University of Botswana, soon also including the Botswana International University of Science and Technology. The Chem4Energy research programme focused on 'New materials for a sustainable energy future: linking computation with experiment', aiming to deploy the synergy between state-of-the-art computational and experimental techniques to design and optimize new catalysts and semiconductor materials for renewable energy applications, based on materials that are abundant and readily available in African countries. The Chem4Energy consortium has achieved ambitious research goals, graduated seven PhD students and delivered a high-quality cross-disciplinary training programme in materials science and simulation techniques relevant to renewable energy applications. Since 2021, the extended consortium, including North-West University and the Centre for High-Performance Computing in South Africa, has remained active through an annual Chem4Energy conference series, with the sixth meeting taking place in Namibia in April 2025.
Interaction of a non-axisymmetric artificial single spatula with rough surfaces
Hair-like attachment structures are frequently used by animals to create stable contact with rough surfaces. Previous studies focused primarily on axisymmetric biomimetic models of artificial spatulas, such as those with a mushroom-shaped and cylinder-shaped geometry, in order to simulate the so-called gecko effect. Here, two geometric prototypes of artificial adhesive structures with non-axisymmetric properties were designed. The investigation of the prototype's interactions with rough surfaces was carried out using the finite element software ABAQUS. Under increasing vertical displacement, the effect of asperity size on the contact pressure evolution of the spatula was investigated. It has been demonstrated that the contact behaviour is greatly affected by the flexibility of the spatula, which is caused by its variable thickness. The thinner spatula shows a higher nominal contact area and attaches more strongly to various rough surfaces. Although a thicker spatula is more susceptible to the 'leverage' phenomenon, which occurs when excessively applied displacements prematurely reduce the nominal contact area, it obtains the ability to regulate attachment during unidirectional loading. Two non-axisymmetric prototypes provide different design concepts for the artificial adhesives. It is hoped that this study will provide fresh viewpoints and innovations that contribute to the development of biologically inspired adhesives.
Prey size reflected in tooth wear: a comparison of two wolf populations from Sweden and Alaska
Ingesta leaves distinct patterns on mammalian teeth during mastication. However, an unresolved challenge is how to include intraspecific variability into dietary reconstruction and the biomechanical aspects of chewing. Two extant populations of the grey wolf (), one from Alaska and one from Sweden, were analysed with consideration to intraspecific dietary variability related to prey size depending on geographical origin, sex and individual age as well as tooth function. Occlusal enamel facets of the upper fourth premolars, first molars and the second lower molar were analysed via three-dimensional surface texture analysis. The Swedish wolves displayed facets characterized by higher peaks and deeper, more voluminous dales, featuring an overall rougher surface than the wolves from Alaska. Compared to females, the Swedish male wolves had a slightly larger dale area and hill volume on their facets. Upper fourth premolars are smoother and had higher values in texture direction compared to upper first molars. The upper first molars were rougher than the occluding lower second molars and were characterized by larger and deeper dales. We find evidence supporting intraspecific dietary segregation, and antagonistic asymmetry in occlusal wear signatures. The data offer new insights into the roles of apex predators like the grey wolf.
Elasmoid fish scales as a natural fibre composite: microscopic heterogeneities in structure, mineral distribution, and mechanical properties
The elasmoid scales in teleost fish serve as exemplary models for natural fibre composites with integrated flexibility and protection. Yet, limited research has been focused on the potential structural, chemical, and mechanical heterogeneity within individual scales. This study presents systematic characterizations of the elasmoid scales from black drum fish () at different zones within individual scales as a natural fibre composite, focusing on the microscopic structural heterogeneities and corresponding mechanical effects. The focus field at the centre of the scales exhibits a classical tri-layered collagen-based composite design, consisting of the mineralized outermost limiting layer, external elasmodine layer in the middle, and the unmineralized internal elasmodine layer. In comparison, the rostral field at the anterior end of the scales exhibits a two-layered design: the mineralized outermost limiting layer exhibits radii sections on the outer surface, and the inner elasmodine layer consists of collagen fibre-based sublayers with alternating mineralization levels. Chemical and nanoindentation analysis suggests a close correlation between the mineralization levels and the local nanomechanical properties. Comparative finite element modelling shows that the rostral-field scales achieve increased flexibility under both concave and convex bending. Moreover, the evolving geometries of isolated Mandle's corpuscles in the internal elasmodine layer, transitioning from irregular shapes to faceted octahedrons, suggest the mechanisms of mineral growth and space-filling to thicken the mineralized layers in scales during growth, which enhances the bonding strength between the adjacent collagen fibre layers. This work offers new insights into the structural variations in individual elasmoid scales, providing strategies for bioinspired fibre composite designs with local-adapted functional requirements.
The biomechanics of splitting hairs
Splitting of hair, creating 'split ends', is a very common problem which has been extensively documented. However, the mechanics underlying the splitting phenomenon are poorly understood. This is partly owing to the lack of a test in which splitting can be generated and quantified under laboratory conditions. We developed three new tests, known as 'loop tensile', 'moving loop' and 'moving loop fatigue', aiming to simulate the mechanical environment of tangles of hair strands during combing. We tested straight strands of human hair, comparing low-quality hair (from a subject who experienced split ends) with hair from a control (non-splitting) subject. Significant differences were found, especially in the moving loop fatigue test where the low-quality hair failed in fewer cycles. Splitting occurred in both types of hair, but with the crucial difference that in the low-quality hair, splits originated inside the hair strand and propagated longitudinally over considerable distances, while in the control hair, splits originated at the strand surface and remained short. Bleaching of the control hair changed its behaviour, making it similar to that of the low-quality hair. Some simple calculations emphasized the role of longitudinal shear stress and shear stress intensity in generating microcracks which could then propagate within the moving loop, paving the way for a future theoretical model of the splitting mechanism.
Driving macro-scale transformations in three-dimensional-printed biopolymers through controlled induction of molecular anisotropy at the nanoscale
Motivated by the need to harness the properties of renewable and biodegradable polymers for the design and manufacturing of multi-scale structures with complex geometries, we have employed our additive manufacturing platform that leverages molecular self-assembly for the production of metre-scale structures characterized by complex geometries and heterogeneous material composition. As a precursor material, we used chitosan, a chemically modified form of chitin, an abundant and sustainable structural polysaccharide. We demonstrate the ability to control concentration-dependent crystallization as well as the induction of the preferred orientation of the polymer chains through the combination of extrusion-based robotic fabrication and directional toolpathing. Anisotropy is demonstrated and assessed through high-resolution micro-X-ray diffraction in conjunction with finite element simulations. Using this approach, we can leverage controlled and user-defined small-scale propagation of residual stresses to induce large-scale folding of the resulting structures.
The thermal 'Buddha Board'-application of microstructured polyolefin films for variable thermal infrared transparency materials
In this study, we explore the innovative application of biological principles of scattering foams and structural colouration of white materials to manipulate the transmission properties of thermal infrared (IR) radiation, particularly within the 8-14 μm wavelength range in polyolefin materials. Inspired by the complex skin of organisms such as chameleons, which can dynamically change colour through structural alterations, as well as more mundane technologies such as Buddha Boards and magic water colouring books, we are developing methods to control thermal IR transmission using common thermoplastic materials that are semi-transparent to thermal IR radiation. Polyethylene and polypropylene, known for their versatility and cost-effectiveness, can be engineered into microstructured sheets with feature sizes spanning from 5 to 100 μm. By integrating these precisely moulded microstructures with index-matching fluids, specifically IR transparent oils, we achieve a reversible modification of the thermal transmission properties. This novel approach not only mimics the adaptive functionality of natural systems but also offers a practical and scalable solution for dynamic thermal management. Our results indicate a promising pathway for the development of new materials that can adapt their IR properties in real time, paving the way for smarter thermal management solutions via radiative emission/absorption.
Analysis of micropolar elastic multi-laminated composite and its application to bioceramic materials for bone reconstruction
The asymptotic homogenization method is applied to characterize the effective behaviour of periodic multi-laminated micropolar elastic heterogeneous composites under perfect contact conditions. The local problem formulations and the analytical expressions for the effective stiffness and torque coefficients are derived for the centrosymmetric case. One of the main findings in this work is the analysis of the rotations effect of the layers' constitutive properties on the mechanical response of bi-laminated composites. The effects of microstructure and interfacial interactions on the composite's mechanical behaviour are captured through the independent effective moduli. Comparisons with the classical elastic case show the approach validation. Some numerical examples are shown. Furthermore, considering the micropolar media's prevalence in bio-inspired systems, the model's applicability is evaluated for reconstructing bone fractures using multi-laminated biocomposites. An important finding in this bio-inspired simulation is related to the analysis of a periodic bi-laminated micropolar composite whose isotropic constituents are a bioceramic material and a compact bone. This artificial bio-inspired material should integrate with host tissue to support cell growth and be stable and compatible. These characteristics are crucial in the enhancement of the fractured bone.
Effects of mineralization on the hierarchical organization of collagen-a synchrotron X-ray scattering and polarized second harmonic generation study
The process of mineralization fundamentally alters collagenous tissue biomechanics. While the structure and organization of mineral particles have been widely studied, the impact of mineralization on collagen matrix structure, particularly at the molecular scale, requires further investigation. In this study, synchrotron X-ray scattering (XRD) and polarization-resolved second harmonic generation microscopy (pSHG) were used to study normally mineralizing turkey leg tendon in tissue zones representing different stages of mineralization. XRD data demonstrated statistically significant differences in collagen D-period, intermolecular spacing, fibril and molecular dispersion and relative supramolecular twists between non-mineralizing, early mineralizing and late mineralizing zones. pSHG analysis of the same tendon zones showed the degree of collagen fibril organization was significantly greater in early and late mineralizing zones compared to non-mineralizing zones. The combination of XRD and pSHG data provide new insights into hierarchical collagen-mineral interactions, notably concerning possible cleavage of intra- or interfibrillar bonds, occlusion and reorganization of collagen by mineral with time. The complementary application of XRD and fast, label-free and non-destructive pSHG optical measurements presents a pathway for future investigations into the dynamics of molecular scale changes in collagen in the presence of increasing mineral deposition.
Hierarchical looping results in extreme extensibility of silk fibre composites produced by Southern house spiders ()
Spider silk is a tough and versatile biological material combining high tensile strength and extensibility through nanocomposite structure and its nonlinear elastic behaviour. Notably, spiders rarely use single silk fibres in isolation, but instead process them into more complex composites, such as silk fibre bundles, sheets and anchorages, involving a combination of spinneret, leg and body movements. While the material properties of single silk fibres have been extensively studied, the mechanical properties of silk composites and meta-structures are poorly understood and exhibit a hereto largely untapped potential for the bio-inspired design of novel fabrics with outstanding mechanical properties. In this study, we report on the tensile mechanics of the adhesive capture threads of the Southern house spider (), which exhibit extreme extensibility, surpassing that of the viscid capture threads of orb weavers by up to tenfold. By combining high-resolution mechanical testing, microscopy and experiments based on a hierarchical modified version of the Fibre Bundle Model, we demonstrate that extreme extensibility is based on a hierarchical loops-on-loops structure combining linear and coiled elements. The stepwise unravelling of the loops leads to the repeated fracture of the connected linear fibres, delaying terminal failure and enhancing energy absorption. This principle could be used to achieve tailored fabrics and materials that are able to sustain high deformation without failure.
Hierarchical nested honeycomb-based energy absorbers: design factors and tailorable mechanical properties
This study presents a novel hierarchical nested honeycomb drawing inspiration from the hierarchical structures found in energy-absorbing citrus peels. Our investigation reveals that integrating secondary hierarchical units into primary honeycomb cells results in energy absorption profiles featuring two distinct plateaus. Notably, we found that these profiles can be finely tuned by adjusting the thickness of primary and secondary cell walls. Additionally, our study demonstrates a strategic removal of cell walls at key positions, reducing material consumption without compromising specific energy absorption. By establishing comprehensive structure-property relationships, we offer valuable insights into the design and optimization of hierarchical cellular materials. Compared with traditional honeycomb structures, the nested honeycomb structure shows a twofold increase in compressive strength and a fivefold increase in specific energy absorption, positioning them as promising candidates for applications requiring two-step impact protection and tunable performance, ranging from packaging to high-speed automobiles.