Ant tunneling—a granular media perspective

Laboratory and field data show that the digging habits of ants and the resulting nest architecture vary with soil conditions, yet, the geomechanical understanding of ant tunneling is lacking. We study the excavation strategies used by harvester ants in clay, silt, sand, and gravel at water contents that range from dry to saturated. The study focuses on the conditions at the tunnel face that determine particle removal methods, digging rate, the development of branches, and tunneling patterns. Analytical and numerical models provide particle-level insight into the experimental observations and help identify the causal links that relate ants digging performance and nest geometric patterns with the properties of the granular medium such as grain size, moisture, and packing density. Results highlight ants’ exceptional ability to sense the prevailing geomechanical conditions in tunnels, and to adapt excavation strategies, transport methods and tunneling patterns to those conditions, within their inherent size and strength limitations. The resulting tunnel structure emerges as a mechanically-convenient and energy-efficient topology based on local information gather by ants along the tunnel and at the tunnel face.     

Effects of worker size on the dynamics of fire ant tunnel construction

Social insects work together to complete tasks. However, different individuals within a colony may vary in task proficiency. We investigated if fire ant (Solenopsis invicta) worker body size influenced the ability to construct tunnels—a key component of subterranean nests. We monitored excavation by worker groups in a substrate of small wetted glass particles in quasi-two-dimensional arenas. Morphological and network features of the tunnel system were measured. Total tunnel area did not differ significantly between groups of large and small workers, although the tunnel area of control sized workers was significantly larger than that of large workers. Moreover, large workers created wider but shorter tunnels, with slower growth rate of tunnel number. However, edge–vertex scaling and degree distribution of the tunnel network were similar across all treatments. In all cases, the amount of excavated material was correlated with the number of active workers. Our study reveals that morphological features of excavated tunnels show modest variation when constructed by workers of varying sizes, but topological features associated with the tunnel network are conserved. These results suggest that important behavioural aspects of tunnel construction—and thus nest building—are similar among morphologically distinct members of fire ant societies.     

Local cost minimization in ant transport networks: from small-scale data to large-scale trade-offs

Transport networks distribute resources and information in many human and biological systems. Their construction requires optimization and balance of conflicting criteria such as robustness against disruptions, transport efficiency and building cost. The colonies of the polydomous Australian meat ant Iridomyrmex purpureus are a striking example of such a decentralized network, consisting of trails that connect spatially separated nests. Here we study the rules that underlie network construction in these ants. We find that a simple model of network growth, which we call the minimum linking model (MLM), is sufficient to explain the growth of real ant colonies. For larger networks, the MLM shows a qualitative similarity with a Euclidean minimum spanning tree, prioritizing cost and efficiency over robustness. We introduce a variant of our model to show that a balance between cost, efficiency and robustness can be also reproduced at larger scales than ant colonies. Remarkably, such a balance is influenced by a parameter reflecting the specific features of the modelled transport system. The extended MLM could thus be a suitable source of inspiration for the construction of cheap and efficient transport networks with non-zero robustness, suggesting possible applications in the design of human-made networks.     

The effect of individual variation on the structure and function of interaction networks in harvester ants

Social insects exhibit coordinated behaviour without central control. Local interactions among individuals determine their behaviour and regulate the activity of the colony. Harvester ants are recruited for outside work, using networks of brief antennal contacts, in the nest chamber closest to the nest exit: the entrance chamber. Here, we combine empirical observations, image analysis and computer simulations to investigate the structure and function of the interaction network in the entrance chamber. Ant interactions were distributed heterogeneously in the chamber, with an interaction hot-spot at the entrance leading further into the nest. The distribution of the total interactions per ant followed a right-skewed distribution, indicating the presence of highly connected individuals. Numbers of ant encounters observed positively correlated with the duration of observation. Individuals varied in interaction frequency, even after accounting for the duration of observation. An ant''s interaction frequency was explained by its path shape and location within the entrance chamber. Computer simulations demonstrate that variation among individuals in connectivity accelerates information flow to an extent equivalent to an increase in the total number of interactions. Individual variation in connectivity, arising from variation among ants in location and spatial behaviour, creates interaction centres, which may expedite information flow.     

Treadmilling and dynamic protrusions in fire ant rafts

Fire ants (Solenopsis invicta) are exemplary for their formation of cohered, buoyant and dynamic structures composed entirely of their own bodies when exposed to flooded environments. Here, we observe tether-like protrusions that emerge from aggregated fire ant rafts when docked to stationary, vertical rods. Ant rafts comprise a floating, structural network of interconnected ants on which a layer of freely active ants walk. We show here that sustained shape evolution is permitted by the competing mechanisms of perpetual raft contraction aided by the transition of bulk structural ants to the free active layer and outward raft expansion owing to the deposition of free ants into the structural network at the edges, culminating in global treadmilling. Furthermore, we see that protrusions emerge as a result of asymmetries in the edge deposition rate of free ants. Employing both experimental characterization and a model for self-propelled particles in strong confinement, we interpret that these asymmetries are likely to occur stochastically owing to wall accumulation effects and directional motion of active ants when strongly confined by the protrusions'' relatively narrow boundaries. Together, these effects may realize the cooperative, yet spontaneous formation of protrusions that fire ants sometimes use for functional exploration and to escape flooded environments.     

The importance of accurate muscle modelling for biomechanical analyses: a case study with a lizard skull

Computer-based simulation techniques such as multi-body dynamics analysis are becoming increasingly popular in the field of skull mechanics. Multi-body models can be used for studying the relationships between skull architecture, muscle morphology and feeding performance. However, to be confident in the modelling results, models need to be validated against experimental data, and the effects of uncertainties or inaccuracies in the chosen model attributes need to be assessed with sensitivity analyses. Here, we compare the bite forces predicted by a multi-body model of a lizard (Tupinambis merianae) with in vivo measurements, using anatomical data collected from the same specimen. This subject-specific model predicts bite forces that are very close to the in vivo measurements and also shows a consistent increase in bite force as the bite position is moved posteriorly on the jaw. However, the model is very sensitive to changes in muscle attributes such as fibre length, intrinsic muscle strength and force orientation, with bite force predictions varying considerably when these three variables are altered. We conclude that accurate muscle measurements are crucial to building realistic multi-body models and that subject-specific data should be used whenever possible.     

Defect-Mediated Growth of Crystallographic Shear Plane

As representative extended planar defects, crystallographic shear (CS) planes, namely Wadsley defects, play an important role in modifying the physical and chemical properties of metal oxides. Although these special structures have been intensively investigated for high-rate anode materials and catalysts, it is still experimentally unclear how the CS planes form and propagate at the atomic scale. Here, the CS plane evolution in monoclinic WO₃ is directly imaged via in situ scanning transmission electron microscope. It is found that the CS planes nucleate preferentially at the edge step defects and proceed by the cooperative migration of WO₆ octahedrons along particular crystallographic orientations, passing through a series of intermediate states. The local reconstruction of atomic columns tends to form (102) CS planes featured with four edge-sharing octahedrons in preference to the (103) planes, which matches well with the theoretical calculations. Associated with the structure evolution, the sample undergoes a semiconductor-to-metal transition. In addition, the controlled growth of CS planes and V-shaped CS structures can be achieved by artificial defects for the first time. These findings enable an atomic-scale understanding of CS structure evolution dynamics.     

Group dynamics and record signals in the ant Temnothorax albipennis

Many purely physical complex systems, in which there are both stochasticity and local interactions between the components, exhibit record dynamics. The temporal statistics of record dynamics is a Poisson process operating on a logarithmic rather than a linear time scale (i.e. a log-Poisson process). Record dynamics often drive substantial changes in complex systems when new high water marks in partially stochastic processes trigger new events. Social insect colonies are exemplary complex biological systems in which many of the local interactions of the components have been moulded by natural selection for the common good. Here, we combine experimental manipulation of ant colony demography with modelling to test the hypothesis that social interactions are the mechanism underlying the record dynamics. We found that compared with the control, log-Poisson statistics were disrupted in colonies in which the pattern of interactions was modified by the removal of the brood, and disappeared completely in ‘callow’ colonies composed entirely of very young workers from the same age cohort. We conclude that a subtle interplay between the demography of the society and the pattern of the interactions between the ants is crucial for the emergence of record dynamics. This could help identify what makes an ant colony a cohesive society.     

The interplay between a self-organized process and an environmental template: corpse clustering under the influence of air currents in ants

Many spatial patterns observed in nature emerge from local processes and their interactions with the local environment. The clustering of objects by social insects represents such a pattern formation process that can be observed at both the individual and the collective level. In this paper, we study the interaction between air currents and clustering behaviour in order to address the coordinating mechanisms at the individual level that underlie the spatial pattern formation process in a heterogeneous environment. We choose the corpse clustering behaviour of the ant Messor sanctus as an experimental paradigm. In a specifically designed experimental set-up with a well-controlled laminar air flow (approx. 1 cm s-1), we first quantify the modulation of the individual corpse aggregation behaviour as a function of corpse density, air flow intensity and the ant's position with respect to corpse piles and air flow direction. We then explore by numerical simulation how the forming corpse piles modify the laminar air flow around them and link this result with the individual behaviour modulation. Finally, we demonstrate on the collective level that this laminar air flow leads to an elongation and a slow displacement of the formed corpse piles in the direction of the air current. Both the individual behaviour modulated by air flow and the air flow modulated by the forming corpse piles can explain the pile patterns observed on the collective level as a stigmergic process. We discuss the generality of this coordinating mechanism to explain the clustering phenomena in heterogeneous environments reported in the literature.     

Reorientation patterns in central-place foraging: internal clocks and klinokinesis

We study central-place foraging patterns of Aphaenogaster senilis ants at a population level by video framing individual ant trajectories in a circular arena with a nest connected to its centre. The ants naturally leave and enter the nest and forage generating non-trivial movement patterns around the nest. Our data analysis indicated that the trajectories observed can be classified into two strategies: the risk-averse strategy, which involves wandering around the nest without departing far from it and the risk-prone strategy, which involves long exploration paths with periodic returns to the central region, nearby the nest. We found that both risk-prone and risk-averse strategies exhibit qualitatively the same reorientation patterns, with the time between consecutive reorientations covering a wide range of scales, and fitting a stretched exponential function. Nevertheless, differences in the temporal scales and the time variability of such reorientation events differ, together with other aspects of motion, such as average speed and turns. Our results give experimental evidence that the internal mechanisms driving reorientations in ants tend to favour frequently long relocations, as theory predicts for efficient exploration in patchy landscapes, but ants engaged in central-place foraging can modulate such behaviour to control distances from the nest. Previous works on the species support the idea that risk-prone and risk-averse strategies may reflect actual differences between individuals age and experience; these factors (age and experience) should be then relevant in modulating the internal reorientation clocks. To support the validity of our findings, we develop a random-walk model combining stretched exponential reorientation clocks with klinokinesis that fits the time length and the travelled distance distributions of the observed trajectories.     

Atomic structure of hardening precipitates in an Al–Mg–Zn–Cu alloy determined by HAADF-STEM and first-principles calculations: relation to η-MgZn2

The structures of two nanoscale plate precipitates prevalent at maximum strength and over-aged conditions in a 7449 Al–Mg–Zn–Cu alloy were investigated. Models derived from images of high angle annular dark field scanning transmission electron microscopy were supported by first-principles calculations. Both structures are closely linked to the η-MgZn₂ Laves phase through similar layers of a rhombohedral atomic subunit. The finest plate contains one such layer together with a layer of an orthorhombic unit. The second plate contains rhombohedral layers only, normally four, but rotated relatively to form different stacking variants, one of which may be likened to η. For both structures, the same atomic planes describe the main interface with Al. Both plates could be described in space group P3. The unit cells comprise interface and arbitrary numbers of {111}Al (habit) planes. Eight Al-planes were included in the first-principles calculations. The enthalpy indicates high layer/unit stability. The plate thickness can be understood by a simple mismatch formulation.     

Coordination-induced formation of nanobelts of organic-inorganic hybrid materials at room temperature

The unexpected finding that the direct mix of o-phenylenediamine and H₂PdCl₄ aqueous solution at room temperature leads to supramolecular nanobelts of coordination polymers was reported. The morphology of the structures was characterized by scanning electron microscopy (SEM) and the chemical composition was examined by energy-dispersed spectrum (EDS) and X-ray maps analysis. The possible formation process of such supramolecular nanobelts is given and the influence of the molar ratio of reactants on the formation of these nanobelts is also examined. It is also found that the molar ratio of reactants has strong influence on the formation of the supramolecular structures.     

Housing tubes from the marine worm Chaetopterus sp.: biomaterials with exceptionally broad thermomechanical properties

The housing tube material of the marine worm Chaetopterus sp. exhibits thermal stability up to 250°C, similar to other biological materials such as mulberry silkworm cocoons. Interestingly, however, dynamic mechanical thermal analysis conducted in both air and water elucidated the lack of a glass transition in the organic tube wall material. In fact, the viscoelastic properties of the anhydrous and undried tube were remarkably stable (i.e. constant and reversible) between –75°C and 200°C in air, and 5°C and 75°C in water, respectively. Moreover, it was found that hydration and associated-water plasticization were key to the rubber-like flexible properties of the tube; dehydration transformed the material behaviour to glass-like. The tube is made of bionanocomposite fibrils in highly oriented arrangement, which we argue favours the biomaterial to be highly crystalline or cross-linked, with extensive hydrogen and/or covalent bonds. Mechanical property characterization in the longitudinal and transverse directions ascertained that the tubes were not quasi-isotropic structures. In general, the higher stiffness and strength in the transverse direction implied that there were more nanofibrils orientated at ±45° and ±65° than at 0° to the tube axis. The order of the mechanical properties of the soft–tough tubes was similar to synthetic rubber-like elastomers and even some viscid silks. The complex structure–property relations observed indicated that the worm has evolved to produce a tubular housing structure which can (i) function stably over a broad range of temperatures, (ii) endure mechanical stresses from specific planes/axes, and (iii) facilitate rapid growth or repair.     

Computer-assisted design for scaling up systems based on DNA reaction networks

In the past few years, there have been many exciting advances in the field of molecular programming, reaching a point where implementation of non-trivial systems, such as neural networks or switchable bistable networks, is a reality. Such systems require nonlinearity, be it through signal amplification, digitalization or the generation of autonomous dynamics such as oscillations. The biochemistry of DNA systems provides such mechanisms, but assembling them in a constructive manner is still a difficult and sometimes counterintuitive process. Moreover, realistic prediction of the actual evolution of concentrations over time requires a number of side reactions, such as leaks, cross-talks or competitive interactions, to be taken into account. In this case, the design of a system targeting a given function takes much trial and error before the correct architecture can be found. To speed up this process, we have created DNA Artificial Circuits Computer-Assisted Design (DACCAD), a computer-assisted design software that supports the construction of systems for the DNA toolbox. DACCAD is ultimately aimed to design actual in vitro implementations, which is made possible by building on the experimental knowledge available on the DNA toolbox. We illustrate its effectiveness by designing various systems, from Montagne et al.''s Oligator or Padirac et al.''s bistable system to new and complex networks, including a two-bit counter or a frequency divider as well as an example of very large system encoding the game Mastermind. In the process, we highlight a variety of behaviours, such as enzymatic saturation and load effect, which would be hard to handle or even predict with a simpler model. We also show that those mechanisms, while generally seen as detrimental, can be used in a positive way, as functional part of a design. Additionally, the number of parameters included in these simulations can be large, especially in the case of complex systems. For this reason, we included the possibility to use CMA-ES, a state-of-the-art optimization algorithm that will automatically evolve parameters chosen by the user to try to match a specified behaviour. Finally, because all possible functionality cannot be captured by a single software, DACCAD includes the possibility to export a system in the synthetic biology markup language, a widely used language for describing biological reaction systems. DACCAD can be downloaded online at http://www.yannick-rondelez.com/downloads/.     

Structure and formation of ant transportation networks

Many biological systems use extensive networks for the transport of resources and information. Ants are no exception. How do biological systems achieve efficient transportation networks in the absence of centralized control and without global knowledge of the environment? Here, we address this question by studying the formation and properties of inter-nest transportation networks in the Argentine ant (Linepithema humile). We find that the formation of inter-nest networks depends on the number of ants involved in the construction process. When the number of ants is sufficient and networks do form, they tend to have short total length but a low level of robustness. These networks are topologically similar to either minimum spanning trees or Steiner networks. The process of network formation involves an initial construction of multiple links followed by a pruning process that reduces the number of trails. Our study thus illuminates the conditions under and the process by which minimal biological transport networks can be constructed.     

Characterisation of 3D woven composite internal architecture and effect of compaction

Yarn waviness and resin rich regions play a major role in 3D woven composite mechanical properties and failure. A detailed study of two angle interlock 3D woven carbon fabrics has been carried out to characterise these architectural features and how they change under increasing levels of compaction. Computerised tomography (CT) X-ray scanning was used to capture the internal architecture of the fabrics. The study focussed on the out-of-plane crimp of the yarns and the size and shape of resin rich regions in consolidated panels. Results showed areas of high local crimp at weft/weaver interlace points. This was exacerbated at low levels of compaction but was eventually lowered under higher compaction levels. The appearance of resin channels was found to be heavily dependent on weave style, with large resin pockets appearing in weaver yarn planes which decreased significantly in size under compaction with distinctive changes in profile.     

The influence of sensory delay on the yaw dynamics of a flapping insect

In closed-loop systems, sensor feedback delays may have disastrous implications for performance and stability. Flies have evolved multiple specializations to reduce this latency, but the fastest feedback during flight involves a delay that is still significant on the timescale of body dynamics. We explored the effect of sensor delay on flight stability and performance for yaw turns using a dynamically scaled robotic model of the fruitfly, Drosophila. The robot was equipped with a real-time feedback system that performed active turns in response to measured torque about the functional yaw axis. We performed system response experiments for a proportional controller in yaw velocity for a range of feedback delays, similar in dimensionless timescale to those experienced by a fly. The results show a fundamental trade-off between sensor delay and permissible feedback gain, and suggest that fast mechanosensory feedback in flies, and most probably in other insects, provide a source of active damping which compliments that contributed by passive effects. Presented in the context of these findings, a control architecture whereby a haltere-mediated inner-loop proportional controller provides damping for slower visually mediated feedback is consistent with tethered-flight measurements, free-flight observations and engineering design principles.     

Fabrication of a novel micro-nano fibrous nonwoven scaffold with Antheraea assama silk fibroin for use in tissue engineering

The success of developing artificial organs by tissue engineering depends on scaffold properties and architecture. Here, we describe the fabrication of an Antheraea assama fibroin based novel micro-nano fibrous nonwoven scaffold. The morphological and chemical characterization was done by scanning electron microscope (SEM) and Fourier transform infrared spectroscopy (FTIR) respectively, which demonstrated the formation of scaffold with micro-nano architecture. The biocompatibility was assessed in vitro by haemolysis and cytotoxicity assays, whereby the scaffold was found to be nontoxic and efficient in supporting cell adhesion and growth.     

Synchrotron-based dynamic computed tomography of tissue motion for regional lung function measurement

During breathing, lung inflation is a dynamic process involving a balance of mechanical factors, including trans-pulmonary pressure gradients, tissue compliance and airway resistance. Current techniques lack the capacity for dynamic measurement of ventilation in vivo at sufficient spatial and temporal resolution to allow the spatio-temporal patterns of ventilation to be precisely defined. As a result, little is known of the regional dynamics of lung inflation, in either health or disease. Using fast synchrotron-based imaging (up to 60 frames s⁻¹), we have combined dynamic computed tomography (CT) with cross-correlation velocimetry to measure regional time constants and expansion within the mammalian lung in vivo. Additionally, our new technique provides estimation of the airflow distribution throughout the bronchial tree during the ventilation cycle. Measurements of lung expansion and airflow in mice and rabbit pups are shown to agree with independent measures. The ability to measure lung function at a regional level will provide invaluable information for studies into normal and pathological lung dynamics, and may provide new pathways for diagnosis of regional lung diseases. Although proof-of-concept data were acquired on a synchrotron, the methodology developed potentially lends itself to clinical CT scanning and therefore offers translational research opportunities.     

The microscopic network structure of mussel (Mytilus) adhesive plaques

Marine mussels of the genus Mytilus live in the hostile intertidal zone, attached to rocks, bio-fouled surfaces and each other via collagen-rich threads ending in adhesive pads, the plaques. Plaques adhere in salty, alkaline seawater, withstanding waves and tidal currents. Each plaque requires a force of several newtons to detach. Although the molecular composition of the plaques has been well studied, a complete understanding of supra-molecular plaque architecture and its role in maintaining adhesive strength remains elusive. Here, electron microscopy and neutron scattering studies of plaques harvested from Mytilus californianus and Mytilus galloprovincialis reveal a complex network structure reminiscent of structural foams. Two characteristic length scales are observed characterizing a dense meshwork (approx. 100 nm) with large interpenetrating pores (approx. 1 µm). The network withstands chemical denaturation, indicating significant cross-linking. Plaques formed at lower temperatures have finer network struts, from which we hypothesize a kinetically controlled formation mechanism. When mussels are induced to create plaques, the resulting structure lacks a well-defined network architecture, showcasing the importance of processing over self-assembly. Together, these new data provide essential insight into plaque structure and formation and set the foundation to understand the role of plaque structure in stress distribution and toughening in natural and biomimetic materials.