Engineering Small HOMO-LUMO Gaps in Polycyclic Aromatic Hydrocarbons with Topologically Protected States.

Topological phases in laterally confined low-dimensional nanographenes have emerged as versatile design tools that can imbue otherwise unremarkable materials with exotic band structures ranging from topological semiconductors and quantum dots to intrinsically metallic bands. The periodic boundary conditions that define the topology of a given lattice have thus far prevented the translation of this technology to the quasi-zero-dimensional (0D) domain of small molecular structures. Here, we describe the synthesis of a polycyclic aromatic hydrocarbon (PAH) featuring two localized zero modes (ZMs) formed by the topological junction interface between a trivial and nontrivial phase within a single molecule. First-principles density functional theory calculations predict a strong hybridization between adjacent ZMs that gives rise to an exceptionally small HOMO-LUMO gap. Scanning tunneling microscopy and spectroscopy corroborate the molecular structure of 9/7/9-double quantum dots and reveal an experimental quasiparticle gap of 0.16 eV, corresponding to a carbon-based small molecule long-wavelength infrared (LWIR) absorber.

Developmental transcriptomes predict adult social behaviours in the socially flexible sweat bee, Lasioglossum baleicum.

Natural variation can provide important insights into the genetic and environmental factors that shape social behaviour and its evolution. The sweat bee, Lasioglossum baleicum, is a socially flexible bee capable of producing both solitary and eusocial nests. We demonstrate that within a single nesting aggregation, soil temperatures are a strong predictor of the social structure of nests. Sites with warmer temperatures in the spring have a higher frequency of social nests than cooler sites, perhaps because warmer temperatures provide a longer reproductive window for those nests. To identify the molecular correlates of this behavioural variation, we generated a de novo genome assembly for L. baleicum, and we used transcriptomic profiling to compare adults and developing offspring from eusocial and solitary nests. We find that adult, reproductive females have similar expression profiles regardless of social structure in the nest, but that there are strong differences between reproductive females and workers from social nests. We also find substantial differences in the transcriptomic profiles of stage-matched pupae from warmer, social-biased sites compared to cooler, solitary-biased sites. These transcriptional differences are strongly predictive of adult reproductive state, suggesting that the developmental environment may set the stage for adult behaviours in L. baleicum. Together, our results help to characterize the molecular mechanisms shaping variation in social behaviour and highlight a potential role of environmental tuning during development as a factor shaping adult behaviour and physiology in this socially flexible bee.

Developmental transcriptomes predict adult social behaviors in the socially flexible sweat bee, Lasioglossum baleicum.

Natural variation can provide important insights into the genetic and environmental factors that shape social behavior and its evolution. The sweat bee, Lasioglossum baleicum , is a socially flexible bee capable of producing both solitary and eusocial nests. We demonstrate that within a single nesting aggregation, soil temperatures are a strong predictor of the social structure of nests. Sites with warmer temperatures in the spring have a higher frequency of social nests than cooler sites, perhaps because warmer temperatures provide a longer reproductive window for those nests. To identify the molecular correlates of this behavioral variation, we generated a de novo genome assembly for L. baleicum , and we used transcriptomic profiling to compare adults and developing offspring from eusocial and solitary nests. We find that adult, reproductive females have similar expression profiles regardless of social structure in the nest, but that there are strong differences between reproductive females and workers from social nests. We also find substantial differences in the transcriptomic profiles of stage-matched pupae from warmer, social-biased sites compared to cooler, solitary-biased sites. These transcriptional differences are strongly predictive of adult reproductive state, suggesting that the developmental environment may set the stage for adult behaviors in L. baleicum . Together, our results help to characterize the molecular mechanisms shaping variation in social behavior and highlight a potential role of environmental tuning during development as a factor shaping adult behavior and physiology in this socially flexible bee.

Group size rather than social status influences personal immune efficacy in a socially polymorphic bee.

The evolution of group living is associated with increased pressure from parasites and pathogens. This can be offset by greater investment in personal immune defences and/or the development of cooperative immune defences (social immunity). An enduring question in evolutionary biology is whether social-immune benefits arose in response to an increased need in more complex societies, or arose early in group living and helped facilitate the evolution of more complex societies. In this study, we shed light on this question through investigating how immunity varies intraspecifically in a socially polymorphic bee. Using a novel immune assay, we show that personal antibacterial efficacy in individuals from social nests is higher than that of solitary individuals, but that this can be explained by higher densities in social nests. We conclude that personal immune effects are likely to play a role in the social/solitary transition in this species. These patterns are consistent with the idea that social immunity evolved secondarily, following the evolution of group living. The flexibility of the individual immune system may have favoured a reliance on its use during the facultative phase early in social evolution.

Differential Gene Expression Correlates with Behavioural Polymorphism during Collective Behaviour in Cockroaches.

Consistent inter-individual variation in the propensity to perform different tasks (animal personality) can contribute significantly to the success of group-living organisms. The distribution of different personalities in a group influences collective actions and therefore how these organisms interact with their environment. However, we have little understanding of the proximate mechanisms underlying animal personality in animal groups, and research on this theme has often been biased towards organisms with advanced social systems. The goal of this study is to investigate the mechanistic basis for personality variation during collective behaviour in a species with rudimentary societies: the American cockroach. We thus use an approach which combines experimental classification of individuals into behavioural phenotypes ('bold' and 'shy' individuals) with comparative gene expression. Our analyses reveal differences in gene expression between behavioural phenotypes and suggest that social context may modulate gene expression related to bold/shy characteristics. We also discuss how cockroaches could be a valuable model for the study of genetic mechanisms underlying the early steps in the evolution of social behaviour and social complexity. This study provides a first step towards a better understanding of the molecular mechanisms associated with differences in boldness and behavioural plasticity in these organisms.

Trait Plasticity among Invasive Populations of the Ant Technomyrmex brunneus in Japan.

Characters in invasive populations often differ from those in the native range, and the ability to express different characters may enhance invasive potential. Ants are among the most pervasive and damaging invasive species, by virtue of their transportability and broad-ranging ecological interactions. Their success is often attributed to the ability to exhibit different characteristics in invasive populations, including the formation of large, unicolonial associations ('supercolonies'). It remains unclear, however, if such characteristics are a product or cause of the ecological dominance of invasive ants, and the advancement of our understanding has likely been restrained by the fact that studies to date have focused on a few globally important species with well-established invasions. In this study, we take advantage of an ongoing invasion of the tramp ant Technomyrmex brunneus in Japan to assess trait plasticity in the invasive range of this species. We find evidence for plasticity in social structure among island populations, with a supercolony evident on one of the three islands studied. Interestingly, we found no evidence of lower genetic diversity in this population, though natural isotope data indicate it was operating at a lower trophic level than other populations. These findings add weight to arguments that invasive species may benefit from the capacity to adaptively mould themselves to new ecological contexts.

Variation in personality can substitute for social feedback in coordinated animal movements.

Collective movements are essential for the effective function of animal societies, but are complicated by the need for consensus among group members. Consensus is typically assumed to arise via feedback mechanisms, but this ignores inter-individual variation in behavioural tendency ('personality'), which is known to underpin the successful function of many complex societies. In this study, we use a theoretical approach to examine the relative importance of personality and feedback in the emergence of collective movement decisions in animal groups. Our results show that variation in personality dramatically influences collective decisions and can partially or completely replace feedback depending on the directionality of relationships among individuals. The influence of personality increases with the exaggeration of differences among individuals. While it is likely that both feedback and personality interact in nature, our findings highlight the potential importance of personality in driving collective processes.

Geographic patterns in colonial reproductive strategy in Myrmecina nipponica: Links between biogeography and a key polymorphism in ants.

The ability to express different phenotypes can help define species distributions by allowing access to, and exploitation of, new environments. Social insects employ two markedly different reproductive strategies with contrasting cost/benefit characteristics: independent colony foundation (ICF), which is associated with high dispersal range and high risk, and dependent colony foundation (DCF), characterized by low risk but low dispersal. The ant Myrmecina nipponica employs both of these strategies, with the frequency of each apparently varying between populations. We combine molecular data with data on reproductive strategy from different populations of this species throughout Japan to explore how this polymorphism is linked to environmental factors and whether this relationship can help explain the current and historical biogeography of this species. Reproductive strategy exhibited a strong geographic pattern, with ICF predominant in southern populations and DCF more common in northern and southern highland populations. Molecular analyses clearly divided populations into broad geographic regions, with the southern lowland populations basal to (southern highland (+ northern)) populations. Intra-population polymorphism in colony-founding strategy was widespread, and polymorphism was reconstructed as the likely ancestral state. The frequency of different strategies was linked with climate, with DCF more common in colder areas. A recent inferred origin to the northern lineage suggests that colonization of northern Japan was a rapid event coincident with warming at the end of the Last Glacial Maxima, likely facilitated by the cold-adaptive advantages of DCF. We discuss how such polymorphisms could help explain the biogeography of this and other social insects.

Fussy groups thwart the collective burden of choice: A theoretical study of house-hunting ants.

Information is crucial to effective decision making, but too much information can be as detrimental as too little. Pooling information allows group-living organisms to benefit from the 'wisdom of the crowds', but could also increase the risk of information overload if not complemented by increased information processing capacity. In this study I use an agent-based model and empirical tests to explore the influence of group-size on information accumulation, and the impact of this on collective decision-making. Larger groups were more effective at sampling the environment in simulations and empirical trials. Simulations suggest this this could render them more susceptible to information overload in candidate-rich environments, in which larger groups suffered delays to decision making because of poorly focussed information sharing. However, the cost of excessive information could be ameliorated if individuals are 'fussy' and filter information by withholding information on poor candidates. This may explain observations from empirical trials, in which higher rates of information accumulation in larger groups did not have a clear impact on decision making. These results suggest groups face a potential collective burden of choice from excessive novel information in complex environments. However, collective information-filtering mechanisms analogous to those in the brains of unitary organisms could allow groups to exploit the wisdom of the crowds without suffering from information overload.

Supramolecular Assembly of Oriented Spherulitic Crystals of Conjugated Polymers Surrounding Carbon Nanotube Fibers.

The directed assembly of conjugated polymers into macroscopic organization with controlled orientation and placement is pivotal in improving device performance. Here, the supramolecular assembly of oriented spherulitic crystals of poly(3-butylthiophene) surrounding a single carbon nanotube fiber under controlled solvent evaporation of solution-cast films is reported. Oriented lamellar structures nucleate on the surface of the nanotube fiber in the form of a transcrystalline interphase. The factors influencing the formation of transcrystals are investigated in terms of chemical structure, crystallization temperature, and time. Dynamic process measurements exhibit the linear growth of transcrystals with time. Microstructural analysis of transcrystals reveals individual lamellar organization and crystal polymorphism. The form II modification occurs at low temperatures, while both form I and form II modifications coexist at high temperatures. A possible model is presented to interpret transcrystallization and polymorphism.

An agent-based model of nest-site selection in a mass-recruiting ant.

Complex systems are modular entities which can collectively generate sophisticated emergent solutions through interactions based on simple, local rules. In this study, I use an agent-based model to elucidate how numerous individual-level components contribute to the collective decision process during house-hunting in a mass-recruiting ant species. Myrmecina nipponica combines the use of pheromone trails with a quorum decision rule in collective decisions among nest sites when searching for a new home. The model employed only individual-level rules but accurately emulated group-level properties observed in empirical studies. Simulations suggest that in this system i) both social and private information are necessary for effective decision making, ii) decision making was effective even with very low numbers of 'discriminating' individuals, iii) individual acceptance thresholds were more influential than quorum thresholds in tuning decisions to emphasise speed or accuracy, and iv) acceptance thresholds could also help tune decisions to suit environmental complexity. Similar findings in species using one-to-one recruitment suggest that some individual parameters, such as acceptance thresholds, may hold key functions in collective decision making regardless of the form of recruitment.

Strategies of offspring investment and dispersal in a spatially structured environment: a theoretical study using ants.

BACKGROUND: Offspring investment strategies vary markedly between and within taxa, and much of this variation is thought to stem from the trade-off between offspring size and number. While producing larger offspring can increase their competitive ability, this often comes at a cost to their colonization ability. This competition-colonization trade-off (CCTO) is thought to be an important mechanism supporting coexistence of alternative strategies in a wide range of taxa. However, the relative importance of an alternative and possibly synergistic mechanism-spatial structuring of the environment-remains the topic of some debate. In this study, we explore the influence of these mechanisms on metacommunity structure using an agent-based model built around variable life-history traits. Our model combines explicit resource competition and spatial dynamics, allowing us to tease-apart the influence of, and explore the interaction between, the CCTO and the spatial structure of the environment. We test our model using two reproductive strategies which represent extremes of the CCTO and are common in ants. RESULTS: Our simulations show that colonisers outperform competitors in environments subject to higher temporal and spatial heterogeneity and are favoured when agents mature late and invest heavily in reproduction, whereas competitors dominate in low-disturbance, high resource environments and when maintenance costs are low. Varying life-history parameters has a marked influence on coexistence conditions and yields evolutionary stable strategies for both modes of reproduction. Nonetheless, we show that these strategies can coexist over a wide range of life-history and environmental parameter values, and that coexistence can in most cases be explained by a CCTO. By explicitly considering space, we are also able to demonstrate the importance of the interaction between dispersal and landscape structure. CONCLUSIONS: The CCTO permits species employing different reproductive strategies to coexist over a wide range of life-history and environmental parameters, and is likely to be an important factor in structuring ant communities. Our consideration of space highlights the importance of dispersal, which can limit the success of low-dispersers through kin competition, and enhance coexistence conditions for different strategies in spatially structured environments.

Ants work harder during consensus decision-making in small groups.

Individuals derive many benefits from being social, one of which is improved accuracy of decision-making, the so-called 'wisdom of the crowds' effect. This advantage arises because larger groups can pool information from more individuals. At present, limited empirical data indicate that larger groups outperform smaller ones during consensus decision-making in human and non-human animals. Inaccurate decisions can lead to significant costs, and we might therefore expect individuals in small groups to employ mechanisms to compensate for the lack of numbers. Small groups may be able to maintain decision accuracy if individuals are better informed than those in larger groups and/or by increasing the proportion of the group involved in collective decision-making relative to larger groups. In this study, we use interactive computer vision software to investigate individual contributions to consensus decision-making during house-hunting in different sized groups of the ant Myrmecina nipponica. We show that individuals in small colonies invest greater effort in the consensus decision process than those in large colonies and should be better informed as a result. This may act to ameliorate the limitations of group size, but could leave smaller groups more susceptible to additional stresses.

Ratio-dependent quantity discrimination in quorum sensing ants.

To optimise behaviour, organisms require information on the quantity of various components of their environment, and the ability of animals to discriminate quantity has been a subject of considerable recent interest. This body of research hints at generalised mechanisms of quantity discrimination in vertebrates, but data on invertebrates are still relatively scarce. In this study, I present data on the quantification abilities of an invertebrate in a novel context: quorum sensing. Quorum sensing generates a behavioural response in group-living animals once a threshold number of individuals, a 'quorum', is detected performing some key action. This process forms the basis for consensus decision-making in many species and allows group-living organisms to decide among mutually exclusive alternatives without compromising group integrity. To determine when a quorum is achieved, individuals must assess the number of group members performing the key action. Social insects employ quorum decisions to decide among potential nest sites when searching for a new home. In the Japanese ant, Myrmecina nipponica, quorum thresholds increase with colony size, providing an opportunity to assess the accuracy of quantity discrimination at different stimulus magnitudes. In this study, I demonstrate that the variation in individual quorum thresholds around the mean increases with increasing colony size. This indicates that the quantity discrimination ability of ants decreases with stimulus magnitude, and thus exhibits ratio dependence in the manner of Weber's Law. This may have implications for the accuracy of consensus decision-making and other collective actions in a range of group-living organisms.

Conditional use of social and private information guides house-hunting ants.

Social animals can use both social and private information to guide decision making. While social information can be relatively economical to acquire, it can lead to maladaptive information cascades if attention to environmental cues is supplanted by unconditional copying. Ants frequently employ pheromone trails, a form of social information, to guide collective processes, and this can include consensus decisions made when choosing a place to live. In this study, I examine how house-hunting ants balance social and private information when these information sources conflict to different degrees. Social information, in the form of pre-established pheromone trails, strongly influenced the decision process in choices between equivalent nests, and lead to a reduced relocation time. When trails lead to non-preferred types of nest, however, social information had less influence when this preference was weak and no influence when the preference was strong. These results suggest that social information is vetted against private information during the house-hunting process in this species. Private information is favoured in cases of conflict and this may help insure colonies against costly wrong decisions.

Recurrent evolution of dependent colony foundation across eusocial insects.

The spectacular success of eusocial insects can be attributed to their sophisticated cooperation, yet cooperation is conspicuously absent during colony foundation when queens are alone. Selection against this solitary stage has led to a dramatically different strategy in thousands of eusocial insect species in which colonies are started by groups of nestmates and the benefits of sociality are retained continuously. Dependent colony foundation (DCF) evolved recurrently multiple times across the ants, bees, and wasps, though its prevalence in termites remains unclear. We review adaptations at both the colony level (reproductive investment shifts from sexuals to workers) and the individual level (wingless queens evolve in ants), and other consequences for life history (invasiveness, parasite transmission). Although few studies have focused on DCF, the accumulated data from anecdotal reports, supported by indirect information including morphology, population genetics, and colony demographics, make it clear that this strategy is more diverse and widespread than is usually recognized.

The influence of intraspecific competition on resource allocation during dependent colony foundation in a social insect.

Organisms face a trade-off between investment in fewer, larger offspring, or more, smaller offspring. Most organisms can adjust investment through variation in the size and number of offspring in response to factors such as resource availability and competition. In some social animals, established colonies divide into groups of individuals that become autonomous, a process known as colony fission (also dependent colony foundation in social insects). Resource allocation under fission can be fine-tuned by adjusting the number of new groups (offspring number) and the number of individuals in each new group (offspring size). We assessed the influence of competition on resource allocation during fission in the ant Cataglyphis cursor, by allowing colonies to fission in experimental enclosures of high or low conspecific colony density. The pattern of colony fission was similar to that observed in the field: each fissioning colony produced a few new nests comprising a highly variable number of workers and a single queen, the old queen was often replaced, and new queens were produced in excess. The number of new nests produced depended on the available workforce in the parent colony but was not affected by differences in colony density. Comparison with data from fission under natural field conditions, however, indicates that colonies in enclosures produced fewer, larger new nests, suggesting that resource investment patterns during fission are indeed subject to extrinsic factors. The density of conspecific colonies in the immediate surroundings may be an unreliable estimate of competition intensity and other factors should be considered.

Unequal resource allocation among colonies produced by fission in the ant Cataglyphis cursor.

How organisms allocate limited resources to reproduction is critical to their fitness. The size and number of offspring produced have been the focus of many studies. Offspring size affects survival and growth and determines offspring number in the many species where there is a trade-off between size and number. Many social insects reproduce by colony fission, whereby young queens and accompanying workers split off from a colony to form new colonies. The size of a new colony (number of workers) is set at the time of the split, and this may allow fine tuning size to local conditions. Despite the prevalence of colony fission and the ecological importance of social insects, little is known of colony fission except in honey bees. We studied colony fission in the ant Cataglyphis cursor. For clarity, "colony" and "nest" refer to colonies before and after colony fission, respectively (i.e., each colony fissions into several nests). The reproductive effort of colonies was highly variable: Colonies that fissioned varied markedly in size, and many colonies that did not fission were as large as some of the fissioning colonies. The mother queen was replaced in half of the fissioning colonies, which produced 4.0 +/- 1.3 (mean +/- SD) nests of markedly varied size. Larger fissioning colonies produced larger nests but did not produce more nests, and resource allocation among nests was highly biased. When a colony produced several nests and the mother queen was not replaced, the nest containing the mother queen was larger than nests with a young queen. These results show that the pattern of resource allocation differs between C. cursor and honey bees. They also suggest that C. cursor may follow a bet-hedging strategy with regard to both the colony size at which fission occurs and the partitioning of resources among nests. In addition, colony fission may be influenced by the age and/or condition of the mother queen, and the fact that workers allocating resources among nests have incomplete knowledge of the size and number of nests produced. These results show that the process of colony fission is more diverse than currently acknowledged and that studies of additional species are needed.

Self-sacrifice in 'desperado' contests between relatives.

Intra-specific competition occurs in all animal species and can lead to escalated conflict. Overt fighting entails the risk of injury or death, and is usually avoided through the use of conventions or pre-fight assessments. However, overt fighting can be expected when value of the contest outweighs the value of the future, as contestants have little or nothing to lose. In these situations, respect for conventions and asymmetries between contestants can break down, and overt fighting becomes more likely (the desperado effect). Such conditions can arise in contests between queens over colony ownership in social insects, because the value of inheriting a colony of potentially thousands of helpers is huge and queens may have very limited alternative reproductive options. However, in social species the balance of possible outcomes may be influenced by inclusive fitness, as contestants are often relatives. Here we present a simple model based on social insects, which demonstrates that not fighting can be selectively advantageous when there is a risk posed by fighting to inclusive fitness, even when not fighting is likely to result in death. If contestants are related, a loser can still gain indirect fitness through the winner, whereas fighting introduces a risk that both queens will die and thereby obtain zero inclusive fitness. When relatedness is high and fighting poses a risk of all contestants dying, it can be advantageous to cede the contest and be killed, rather than risk everything by fighting.

Bourgeois queens and high stakes games in the ant Aphaenogaster senilis.

BACKGROUND: Many animals face some form of conflict over reproductive opportunities. Queen selection in social insect colonies represents a high-stakes conflict where competition occurs among multiple queens for a few or a single reproductive role(s). The outcome of the contest is critical to the fitness of all colony individuals as most are sterile, and thus represents a conflict at multiple levels. Aphaenogaster senilis is a monogynous, monandrous, fission performing ant, in which queen selection occurs during colony fission and when replacement queens are produced to overcome orphaning. First-born queens are usually behaviourally dominant over subsequent queens, and eventually inherit the colony. We investigated the importance of physical dominance in queen selection in orphaned groups by manipulating the fighting ability of first-born queens via mandibular ablation. RESULTS: First emerged queens were heavier than second emerged queens, performed almost all aggression, were behaviourally dominant 92% of the time, and prevailed in 76% of groups after co-existing for 16 days on average. Mandibular ablation had no effect on queen behaviour or contest outcome. CONCLUSION: Aggression is probably ritualised and contests are decided by workers based on relative queen fertility. First-born queens thus have an inherent advantage over second-born queens as they have more time to develop ovaries. Subordinates never retaliated against aggression from dominants and this lack of retaliation can be interpreted as a form of bourgeois strategy as dominants were almost always first-born. However, the lack of alternative reproductive options makes not-fighting effectively a form of suicide. High relatedness between full-sister queens means that subordinates may be better off sacrificing themselves than risking injury to both queens by fighting.