Chemical Trails Research Articles

Trail using ants follow idiosyncratic routes in complex landscapes

A large volume of research on individually navigating ants has shown how path integration and visually guided navigation form a major part of the ant navigation toolkit for many species and are sufficient mechanisms for successful navigation. One of the behavioural markers of the interaction of these mechanisms is that experienced foragers develop idiosyncratic routes that require that individual ants have personal and unique visual memories that they use to guide habitual routes between the nest and feeding sites. The majority of ants, however, inhabit complex cluttered environments and social pheromone trails are often part of the collective recruitment, organisation and navigation of these foragers. We do not know how individual navigation interacts with collective behaviour along shared trails in complex natural environments. We thus asked here if wood ants that forage through densely cluttered woodlands where they travel along shared trails repeatedly follow the same routes or if they choose a spread of paths within the shared trail. We recorded three long homing trajectories of 20 individual wood ants in their natural woodland habitat. We found that wood ants follow idiosyncratic routes when navigating along shared trails through highly complex visual landscapes. This shows that ants rely on individual memories for habitual route guidance even in cluttered environments when chemical trail information is available. We argue that visual cues are likely to be the dominant sensory modality for the idiosyncratic routes. These experiments shed new light on how ants, or insects in general, navigate through complex multimodal environments.

Dynamics, statistics, and task allocation of foraging ants.

Ant foraging is one of the most fascinating examples of cooperative behavior observed in nature. It is well studied from an entomology viewpoint, but there is currently a lack of mathematical synthesis of this phenomenon. We address this by constructing an ant foraging model that incorporates simple behavioral rules within three task groups of the ant colony during foraging (foragers, transporters, and followers), pheromone trails, and memory effects. The motion of an ant is modeled as a discrete correlated random walk, with a characteristic zigzag path that is congruent with experimental data. We simulate the foraging cycle, which consists of ants searching for food, transporting food, and depositing chemical trails to recruit and orient more ants (en masse) to the food source. This allows us to gain insights into the basic mechanism of the cooperative interactions between ants and the dynamical division of labor within an ant colony during foraging to achieve optimal efficiency. We observe a disorder-order phase transition from the start to the end of a foraging process, signaling collective motion at the population level. Finally, we present a set of time delay ODEs that corroborates with numerical simulations.

Self-Solidifying Active Droplets Showing Memory-Induced Chirality.

Most synthetic microswimmers do not reach the autonomy of their biological counterparts in terms of energy supply and diversity of motions. Here, this work reports the first all-aqueous droplet swimmer powered by self-generated polyelectrolyte gradients, which shows memory-induced chirality while self-solidifying. An aqueous solution of surface tension-lowering polyelectrolytes self-solidifies on the surface of acidic water, during which polyelectrolytes are gradually emitted into the surrounding water and induce linear self-propulsion via spontaneous symmetry breaking. The low diffusion coefficient of the polyelectrolytes leads to long-lived chemical trails which cause memory effects that drive a transition from linear to chiral motion without requiring any imposed symmetry breaking. The droplet swimmer is capable of highly efficient removal (up to 85%) of uranium from aqueous solutions within 90min, benefiting from self-propulsion and flow-induced mixing. These results provide a route to fueling self-propelled agents which can autonomously perform chiral motion and collect toxins.

Chemical Activity of Low Altitude (50 km) Sprite Streamers

AbstractA three‐stage simulation is used to explore the chemical influence of low altitude (50 km) sprite streamers on the atmosphere, including the chemical trail after the streamer has faded away. In the first stage (streamer phase) a 2D electrodynamical streamer model quantifies the generation of NOx and N2O, and the removal of ozone (O3) by a downward propagating streamer during Δtstreamer = 80 μs. This streamer propagation leads to a distinctive region in the streamer channel, the glow, where the electric field is enhanced. In the second stage (glow phase), the computed densities of the first stage are used as initial conditions for a 0D model to study the chemical evolution of the streamer channel, where we assume a remanent field of 100 Td for the glow and 0 Td elsewhere. This stage lasts Δtglow = 85 μs, the typical glow lifetime at 50 km. Finally, in the third stage (post‐streamer phase), we use the same 0D model, switch off the field in the glow region and let the whole streamer wake evolve roughly 100 s (100 s − Δtglow). Results show a key species such as O3 is mainly depleted during the streamer phase while NOx and N2O are predominantly produced during the same phase. We also compute the local increase of NO2 by sprite streamers at ∼50 km and find out that it could account for the measurable NO2 anomaly over thunderstorms reported from satellite‐based measurements.

Extension and dynamical phases in random walkers depositing and following chemical trails

Active walker models have proved to be extremely effective in understanding the evolution of a large class of systems in biology like ant trail formation and pedestrian trails. We propose a simple model of a random walker which modifies its local environment that in turn influences the motion of the walker at a later time. We perform direct numerical simulations of the walker in a discrete lattice with the walker actively depositing a chemical which attracts the walker trajectory and also evaporates in time. We propose a method to look at the structural transitions of the trajectory using radius of gyration for finite time walks. The extension over a definite time window shows a non-monotonic change with the deposition rate characteristic of a coil-globule transition. At certain regions of the parameter space of the chemical deposition and evaporation rates, the extensions of the walker shows a re-entrant behavior. The dynamics, characterised by the mean-squared displacement, shows deviation from diffusive scaling at intermediate time scales, returning to diffusive behavior asymptotically. A mean-field theory captures the variation of the asymptotic diffusivity.

Biofilm matrix cloaks bacterial quorum sensing chemoattractants from predator detection.

Microbes often secrete high levels of quorum sensing (QS) autoinducers into the environment to coordinate gene expression and biofilm formation, but risk detection and subsequent predation by bacterivorous predators. With such prominent signaling molecules acting as chemoattractants that diffuse into the environment at alarmingly high concentrations, it is unclear if bacterial cells can mask their chemical trails from predator detection. Here, we describe a microbial-based anti-detection adaptation, termed as "biofilm cloak", where the biofilm prey produced biofilm matrix exopolysaccharides that "locked" and reduced the leaching of autoinducers into the milieu, thereby concealing their trails to the detection by the bacterivorous Caenorhabditis elegans nematode. The exopolysaccharides act as common good for the non-producers to hide their autoinducers from predator detection. Deficiency in chemosensory gene odr-10 in mutant animals abrogated their ability to detect autoinducers and migrate toward their prey in a directed manner, which led to lower population growth rate of animals. Hence, restriction of bacterial communication activities to the confinements of biofilms is a novel approach for predator evasion, which plays a fundamental role in shaping ecological dynamics of microbial communities and predator-prey interactions.

Flying ant robot for chemical trail detection and localization

This paper presents first advances in the area of aerial chemical trail following. For that purpose, we equipped a palm-size aerial robot, based on the Crazyflie 2.0 quadrocopter, with a small lightweight metal oxide gas sensor for measuring evaporated ethanol from chemical trails. To detect and localize a 5 cm wide chemical trail, a detection criterion was developed that uses only relative changes in the transient phase of the sensor response. The reduction in signal strength dependence improves the robustness of its application. We tested our setup in first crossing-trail experiments showing that our flying ant robot can correlate an odor hit with the chemical trail within 0.14 m. Principally, this could enable aerial chemical trail following in the future.

Do predators prefer toxic animals? A case of chemical discrimination by an Asian snake that sequesters firefly toxins.

Several Asian natricine snakes of the genus Rhabdophis feed on toads and sequester steroidal cardiac toxins known as bufadienolides (BDs) from them. A recent study revealed that species of the Rhabdophis nuchalis Group ingest lampyrine fireflies to sequester BDs. Although several species of fireflies are distributed in the habitat of the R. nuchalis Group, only lampyrine fireflies, which have BDs, are included in the diet of these snakes. Thus, we hypothesized that the R. nuchalis Group chemically distinguishes fireflies that have BDs from those that do not have BDs. We also predicted that the R. nuchalis Group detects BDs as the chemical cue of toxin source. To test these predictions, we conducted 3 behavioral experiments using Rhabdophis chiwen, which belongs to the R. nuchalis Group. In the first experiment, R. chiwen showed a moderate tongue flicking response to cinobufagin, a compound of BDs. On the other hand, the snake showed a higher response to the chemical stimuli of lampyrine fireflies (BD fireflies) than those of lucioline fireflies (non-BD fireflies). In the second experiment, in which we provided live BD and non-BD fireflies, the snake voluntarily consumed only the former. In the third, a Y-maze experiment, the snake tended to select the chemical trail of BD fireflies more frequently than that of non-BD fireflies. These results demonstrated that R. chiwen discriminates BD fireflies from non-BD fireflies, but the prediction that BDs are involved in this discrimination was not fully supported. To identify the proximate mechanisms of the recognition of novel toxic prey in the R. nuchalis Group, further investigation is necessary.

Cuticular Hydrocarbon Trails Released by Host Larvae Lose their Kairomonal Activity for Parasitoids by Solidification

Successful host search by parasitic wasps is often mediated by host-associated chemical cues. The ectoparasitoid Holepyris sylvanidis is known to follow chemical trails released by host larvae of the confused flour beetle, Tribolium confusum, for short-range host location. Although the hexane-extractable trails consist of stable, long-chain cuticular hydrocarbons (CHCs) with low volatility, the kairomonal activity of a trail is lost two days after release. Here, we studied whether this loss of kairomonal activity is due to changes in the chemical trail composition induced by microbial activity. We chemically analyzed trails consisting of hexane extracts of T. confusum larvae after different time intervals past deposition under sterile and non-sterile conditions. GC-MS analyses revealed that the qualitative and quantitative pattern of the long-chain CHCs of larval trails did not significantly change over time, neither under non-sterile nor sterile conditions. Hence, our results show that the loss of kairomonal activity of host trails is not due to microbially induced changes of the CHC pattern of a trail. Interestingly, the kairomonal activity of trails consisting of host larval CHC extracts was recoverable after two days by applying hexane to them. After hexane evaporation, the parasitoids followed the reactivated host trails as they followed freshly laid ones. Cryo-scanning electron microscopy showed that the trails gradually formed filament-shaped microstructures within two days. This self-assemblage of CHCs was reversible by hexane application. Our study suggests that the long-chain CHCs of a host trail slowly undergo solidification by a self-assembling process, which reduces the accessibility of CHCs to the parasitoid’s receptors as such that the trail is no longer eliciting trail-following behavior.

ALMA observations of doubly deuterated water: inheritance of water from the prestellar environment

Context. Establishing the origin of the water D/H ratio in the Solar System is central to our understanding of the chemical trail of water during the star and planet formation process. Recent modeling suggests that comparisons of the D2O/HDO and HDO/H2O ratios are a powerful way to trace the chemical evolution of water and, in particular, determine whether the D/H ratio is inherited from the molecular cloud or established locally. Aims. We seek to determine the D2O column density and derive the D2O/HDO ratios in the warm region toward the low-mass Class 0 sources B335 and L483. The results are compared with astrochemical models and previous observations to determine their implications for the chemical evolution of water. Methods. We present ALMA observations of the D2O 11,0–10,1 transition at 316.8 GHz toward B335 and L483 at ≲0.′′5 (≲100 au) resolution, probing the inner warm envelope gas. The column densities of D2O, HDO, and H218O are determined by synthetic spectrum modeling and direct Gaussian fitting, under the assumption of a single excitation temperature and similar spatial extent for the three water isotopologs. Results. D2O is detected toward both sources in the inner warm envelope. The derived D2O/HDO ratio is (1.0 ± 0.2) × 10−2 for L483 and (1.4 ± 0.1) × 10−2 for B335. These values indicate that the D2O/HDO ratio is higher than the HDO/H2O ratios by a factor of ≳2 toward both sources. Conclusions. The high D2O/HDO ratios are a strong indication of chemical inheritance of water from the prestellar phase down to the inner warm envelope. This implies that the local cloud conditions in the prestellar phase, such as temperatures and timescales, determine the water chemistry at later stages and could provide a source of chemical differentiation in young systems. In addition, the observed D2O/H2O ratios support an observed dichotomy in the deuterium fractionation of water toward isolated and clustered protostars, namely, a higher D/H ratio toward isolated sources.

Collective decision‐making, caste roles and sequential transport during colony emigration in the small carpenter ant, Camponotus yamaokai

AbstractEffective coordination of group actions underlies the success of group‐living organisms. Making decisions collectively is a crucial step in executing behaviours affecting the whole group. In social insects, decision‐making is typically achieved in a self‐organised manner and incorporates a quorum process, which can improve decision accuracy and be tailored to different conditions. Studies of the nest‐selection process in different social insect species have revealed remarkable consistency in the mode of decision‐making despite marked differences in life history and colony size. However, an often‐overlooked aspect of this decision process is how it might be influenced by diversity in behavioural propensity among colony members. Higher diversity can improve collective processes, but could also represent a cost if certain classes of individual do not contribute to the cooperative activity. Many ant species exhibit caste polymorphism, with different worker castes specialised to different functions. These castes could thus enhance or erode the collective decision process depending on whether they contribute to this process or not. In this study, we explore the process of colony emigration and nest‐site choice in the small carpenter ant Camponotus yamaokai, a species with two distinct worker castes. We show that this species is likely to use a quorum decision process facilitated by navigation with chemical trails. Both major and minor castes contributed to the collective decision and emigration process. However, although major workers comprised 24% of the colony on average, they were 5.5 times less likely than expected to be involved in assessing new nests and 4.6 times less likely than expected to contribute to brood transport. This may represent a previously unrecognised cost to the colony in the emigration process. In addition, we document the first observations of sequential brood transfer during emigration behaviour and speculate upon the function of this behaviour.

Conspecific chemical cues facilitate mate trailing by invasive Argentine black and white tegus.

Squamate reptiles (snakes and lizards) rely on chemical cues from conspecifics to search the environment for potential mates. How such cues are used by invasive species to facilitate reproduction, especially seasonally, is a key question that can inform management practices. The Argentine black and white tegu (Salvator merianae) is an invasive reptile species in south Florida threatening native fauna in biodiverse regions such as Everglades National Park. While some information exists on the reproductive ecology of this species in its native range in South America, the chemical ecology of S. merianae is unclear especially in its invasive range. By testing both male (n = 7) and female (n = 7) tegus in a Y-maze apparatus, we assessed if either sex follows chemical trails left by conspecifics and if behaviors were sex- or season-specific. We conducted three types of trials where conspecifics created odor trails: Male-only (male scent only in base and one arm of Y), Female-only, and Male vs. female. Males did not preferentially follow scent trails from either sex, but they did differentially investigate conspecific scent from both sexes. Seasonally, males showed increased rates of chemosensory sampling (rates of tongue-flicking) during the spring (breeding season; March-May) compared to fall (non-breeding season; September-November). Males also had reduced turning and pausing behavior while trailing in the spring. Female tegus exhibited stronger conspecific trailing abilities than males, following both male and female scent trails, and they explored the maze less before making an arm choice. Females also investigated the scent trails intensely compared to males (more passes in scented arms, more time with scent trails). Our results demonstrate for the first time that females of an invasive reptile species can follow conspecific scent trails. Given the strong female responses to odor, sex-specific targeting of tegus via application of a conspecific chemical cue in traps could enhance removal rates of females during the breeding season.

Male Burmese pythons follow female scent trails and show sex-specific behaviors.

Animals communicate with potential mates using species-specific signals, and pheromones are powerful sexual signals that modify conspecific behavior to facilitate mate location. Among the vertebrates, snakes are especially adept in mate searching via chemical trailing, which is particularly relevant given that many snake species are invasive outside their native ranges. Chemical signals used in mate choice are, thus, potentially valuable tools for management of invasive snake species. The Burmese python (Python bivittatus) is an invasive snake in the Florida Everglades where it is negatively impacting native fauna. In this study, we sought to: (i) determine if males can follow conspecific chemical trails in a Y-maze; and (ii) describe the mate searching behaviors exhibited by males while trailing. All males consistently followed a single female scent trail in the maze, but when only a male scent trail was present they did not discriminate between the male and blank arms. Rate of tongue-flicking, a proxy for chemosensory sampling, was also marginally higher when males were following female versus male scent trails. However, when both female and male scent trails were simultaneously present in the Y-maze, males did not show a preference for the female arm, although the tongue-flick rate was higher in the female-only trial compared to female versus male. Analyses of multiple male behaviors individually and using an ethogram revealed that behaviors were more frequent and complex in the female-only trials compared to male-only trials. Additional behavioral trials are needed to determine if an effective pheromonal approach to Burmese python management is possible.

Moth and Ant Inspired Routing in Hierarchical Airborne Networks with Multi-Beam Antennas

A set of novel routing protocols is proposed for directional hierarchical airborne networks. These networks use a two-level architecture, where the higher-level network is sparse with directional long-distance links that support the high data-rate communication. The lower-level network consists of high-density nodes with short-distance and low data-rate links. We assume that the higher-level nodes are equipped with multi-beam antennas, whereas the lower-level nodes have omni-directional antennas. We use the bio-inspired algorithms (based on moth and ant behaviors) to design the routing schemes for both levels. Specifically, we use the male moth’s light source pursuing pattern for routing the data from an event node to the highly mobile sink in lower-level network, and the ant’s chemical trail maintenance principle to trace the trajectory of commander node to deliver the data from lower-level node to the commander node in the higher-level network, with minimum delay. In addition, we construct a weighted fence routing topology among higher-level nodes with multi-beam antennas, in order to achieve high throughput. Our simulation results demonstrate that significant performance improvement is achieved by the bio-inspired routing schemes, compared with conventional ad hoc routing schemes.

Haptically assisted chemotaxis for odor source localization

Detection of chemical trails has multiple applications in industry and exploration, where prompt localization of dangerous substances encourages development of robust sensors, systems, and algorithms. Current chemical sensing robotics has focused on autonomous navigation, while transduction of chemical information to visual, tactile, or other types of sensory cues have not been thoroughly addressed. This work proposes the inclusion of a human operator in order to solve the robot navigation problem and proposes visual and haptic feedback arrangements that carry the information transmitted by the chemical sensors. A chemical source is placed in a simulated environment, which is navigated by a tracked robot controlled by human drivers. A multipoint haptic interface operates as the robot controller and provides force feedback according to the direction of the chemical gradient detected by the robot. Seven experiments test the tracking performance of three combinations of haptic and visual feedback under different air current configurations, which consistently prove the feasibility of a chemotaxis-based navigation system. Moreover, this work demonstrates with statistical significance that gradient information transmitted using haptic feedback minimizes the required time for reaching the chemical source. This enables the operator's visual capability in other important tasks during navigation (i.e., obstacle avoidance).

Synthetic Chemotaxis and Collective Behavior in Active Matter.

The ability to navigate in chemical gradients, called chemotaxis, is crucial for the survival of microorganisms. It allows them to find food and to escape from toxins. Many microorganisms can produce the chemicals to which they respond themselves and use chemotaxis for signaling, which can be seen as a basic form of communication, allowing ensembles of microorganisms to coordinate their behavior, for example, during embryogenesis, biofilm formation, or cellular aggregation. For example, Dictyostelium cells use signaling as a survival strategy: when starving, they produce certain chemicals toward which other cells show taxis. This leads to aggregation of the cells resulting in a multicellular aggregate that can sustain long starvation periods. Remarkably, the past decade has led to the development of synthetic microswimmers, which can self-propel through a solvent, analogously to bacteria and other microorganisms. The mechanism underlying the self-propulsion of synthetic microswimmers like camphor boats, droplet swimmers, and in particular autophoretic Janus colloids involves the production of certain chemicals. As we will discuss in this Account, the same chemicals (phoretic fields) involved in the self-propulsion of a (Janus) microswimmer also act on other ones and bias their swimming direction toward (or away from) the producing microswimmer. Synthetic microswimmers therefore provide a synthetic analogue to motile microorganisms interacting by taxis toward (or away from) self-produced chemical fields. In this Account, we review recent progress in the theoretical description of synthetic chemotaxis mainly based on simulations and field theoretical descriptions. We will begin with single motile particles leaving chemical trails behind with which they interact themselves, leading to effects like self-trapping or self-avoidance. Besides these self-interactions, in ensembles of synthetic motile particles each particle also responds to the chemicals produced by other particles, inducing chemical (or phoretic) cross-interactions. When these interactions are attractive, they commonly lead to clusters, even at low particle density. These clusters may either proceed toward macrophase separation, resembling Dictyostelium aggregation, or, as shown very recently, lead to dynamic clusters of self-limited size (dynamic clustering) as seen in experiments in autophoretic Janus colloids. Besides the classical case where chemical interactions are attractive, this Account discusses, as its main focus, repulsive chemical interactions, which can create a new and less known avenue to pattern formation in active systems leading to a variety of pattern, including clusters which are surrounded by shells of chemicals, traveling waves and more complex continuously reshaping patterns. In all these cases "synthetic signalling" can crucially determine the collective behavior of synthetic microswimmer ensembles and can be used as a design principle to create patterns in motile active particles.

Behavioral and mechanistic characteristics of the predator-prey interaction between the dinoflagellate Dinophysis acuminata and the ciliate Mesodinium rubrum

Predator-prey interactions of planktonic protists are fundamental to plankton dynamics and include prey selection, detection, and capture as well as predator detection and avoidance. Propulsive, morphology-specific behaviors modulate these interactions and therefore bloom dynamics. Here, interactions between the mixotrophic, harmful algal bloom (HAB) dinoflagellate Dinophysis acuminata and its ciliate prey Mesodinium rubrum were investigated through quantitative microvideography using a high-speed microscale imaging system (HSMIS). The dinoflagellate D. acuminata is shown to detect its M. rubrum prey via chemoreception while M. rubrum is alerted to D. acuminata via mechanoreception at much shorter distances (89 ± 39 μm versus 41 ± 32 μm). On detection, D. acuminata approaches M. rubrum with reduced speed. The ciliate M. rubrum responds through escape jumps that are long enough to detach its chemical trail from its surface, thereby disorienting the predator. To prevail, D. acuminata uses capture filaments and/or releases mucus to slow and eventually immobilize M. rubrum cells for easier capture. Mechanistically, results support the notion that the desmokont flagellar arrangement of D. acuminata lends itself to phagotrophy. In particular, the longitudinal flagellum plays a dominant role in generating thrust for the cell to swim forward, while at other times, it beats to supply a tethering or anchoring force to aid the generation of a posteriorly-directed, cone-shaped scanning current by the transverse flagellum. The latter is strategically positioned to generate flow for enhanced chemoreception and hydrodynamic camouflage, such that D. acuminata can detect and stealthily approach resting M. rubrum cells in the water column.

Increase in scout trips due to forager removal in Atta sexdens (Hymenoptera, Formicidae) (Forel, 1908)

Social information exchange through physical contacts and chemical trail deposition forms the basis of food recruitment in leaf-cutting ants. The scout initiates the process and passes the information to nestmates that recruit more foragers, thus amplifying the stimulus and ensuring the success of foraging. An interruption of the contact between workers and a reduction in trail laying can diminish the effectiveness of mass recruitment and alter scouting activity and forager flow. This study verified an increase in scout trips as a consequence of inbound workers (with or without a plant load) removal during Atta sexdens foraging, sustaining the outbound flow of foragers, and consequently foraging activity, either through direct contact or chemical trail deposition. Data indicate as one of the roles of unladen workers along the foraging trail must be to stimulate other workers to go out and so speed up the recruitment process The remarkable ability to organize themselves without central control is a major strength of social insects and the increase in scouting activity observed here is an example of this behavioral flexibility in leaf-cutting ants. Although foraging performance is enhanced through communication between workers, the simple adjustment in scouting activity can maintain the outbound flow of foragers which is an essential activity of the colony.

Airplane clouds: From chemtrail pseudoscience to the science of contrails

Les afirmacions i els arguments mes frequents trobats en forums i pagines web de pseudociencia en favor de l’existencia dels deixants dels avions, que anomenen chemtrails , es refuten a partir de l’explicacio cientifica del proces de formacio dels deixants de condensacio o sublimacio dels avions, els coneguts contrails . Es descarta la hipotesi de l’existencia dels chemtrails , ja que aquests no difereixen en res dels contrails , els quals son de natura hidrica i resultat d’un proces fisic referenciat en nombrosos estudis desenvolupats des de comencament de l’era de l’aviacio, a comencament del segle XX, fins a l’actualitat. L’article permet concloure que els chemtrails de la pseudociencia no son res mes que els contrails que defineix la ciencia.

Detection and tracking of chemical trails in bio-inspired sensory systems

Many aquatic organisms exhibit remarkable abilities to detect and track chemical signals when foraging, mating and escaping. For example, the male copepod T. longicornis identifies the female in th...