Traveling through light clutter: Path integration and panorama guided navigation in the Sonoran Desert ant, Novomessor cockerelli.

Foraging ants use multiple navigational strategies, including path integration and visual panorama cues, which are used simultaneously and weighted based upon context, the environment and the species' sensory ecology. In particular, the amount of visual clutter in the habitat predicts the weighting given to the forager's path integrator and surrounding panorama cues. Here, we characterize the individual cue use and cue weighting of the Sonoran Desert ant, Novomessor cockerelli, by testing foragers after local and distant displacement. Foragers attend to both a path-integration-based vector and the surrounding panorama to navigate, on and off foraging routes. When both cues were present, foragers initially oriented to their path integrator alone, yet weighting was dynamic, with foragers abandoning the vector and switching to panorama-based navigation after a few meters. If displaced to unfamiliar locations, experienced foragers travelled almost their full homeward vector (∼85 %) before the onset of search. Through panorama analysis, we show views acquired on-route provide sufficient information for orientation over only short distances, with rapid parallel decreases in panorama similarity and navigational performance after even small local displacements. These findings are consistent with heavy path integrator weighting over the panorama when the local habitat contains few prominent terrestrial cues.

Pheromone cue triggers switch between vectors in the desert harvest ant, Veromessor pergandei.

The desert harvester ant (Veromessor pergandei) employs a mixture of social and individual navigational strategies at separate stages of their foraging trip. Individuals leave the nest along a pheromone-based column, travelling 3-40 m before spreading out to forage individually in a fan. Foragers use path integration while in this fan, accumulating a direction and distance estimate (vector) to return to the end of the column (column head), yet foragers' potential use of path integration in the pheromone-based column is less understood. Here we show foragers rely on path integration both in the foraging fan and while in the column to return to the nest, using separate vectors depending on their current foraging stage in the fan or column. Returning foragers displaced while in the fan oriented and travelled to the column head location while those displaced after reaching the column travel in the nest direction, signifying the maintenance of a two-vector system with separate fan and column vectors directing a forager to two separate spatial locations. Interestingly, the trail pheromone and not the surrounding terrestrial cues mediate use of these distinct vectors, as fan foragers briefly exposed to the pheromone cues of the column in isolation altered their paths to a combination of the fan and column vectors. The pheromone acts as a contextual cue triggering both the retrieval of the column-vector memory and its integration with the forager's current fan-vector.

Same but different: Socially foraging ants backtrack like individually foraging ants but use different mechanisms.

Diverse species may adopt behaviourally identical solutions to similar environmental challenges. However, the underlying mechanisms dictating these responses may be quite different and are often associated with the specific ecology or habitat of these species. Foraging desert ants use multiple strategies in order to successfully navigate. In individually foraging ants, these strategies are largely visually-based; this includes path integration and learned panorama cues, with systematic search and backtracking acting as backup mechanisms. Backtracking is believed to be controlled, at least in solitary foraging species, by three criteria: 1) foragers must have recent exposure to the nest panorama, 2) the path integrator must be near zero, and 3) the ant must be displaced to an unfamiliar location. Instead of searching for the nest, under these conditions, foragers head in the opposite compass direction of the one in which they were recently travelling. Here, we explore backtracking in the socially foraging desert harvester ant (Veromessor pergandei), which exhibits a foraging ecology consisting of a combination of social and individual cues in a column and fan structure. We find that backtracking in V. pergandei, similar to solitary foraging species, is dependent on celestial cues, and in particular on the sun's position. However, unlike solitary foraging species, backtracking in V. pergandei is not mediated by the same criteria. Instead the expression of this behaviour is dependent on the presence of the social cues of the column and the proportion of the column that foragers have completed prior to displacement.

Not just going with the flow: foraging ants attend to polarised light even while on the pheromone trail.

The polarisation pattern of skylight serves as an orientation cue for many invertebrates. Solitary foraging ants, in particular, rely on polarised light to orient along with a number of other visual cues. Yet it is unknown, if this cue is actively used in socially foraging species that use pheromone trails to navigate. Here, we explore the use of polarised light in the presence of the pheromone cues of the foraging trail. The desert harvester ant, Veromessor pergandei, relies on pheromone cues and path integration in separate stages of their foraging ecology (column and fan, respectively). Here, we show that foragers actively orient to an altered overhead polarisation pattern, both while navigating individually in the fan and while on the pheromone-based column. These heading changes occurred during twilight, as well as in the early morning and late afternoon before sunset. Differences in shift size indicate that foragers attend to both the polarisation pattern and the sun's position when available, yet during twilight, headings are dominated by the polarisation pattern. Finally, when the sun's position was experimentally blocked before sunset, shift sizes increased similar to twilight testing. These findings show that celestial cues provide directional information on the pheromone trail.

Self-organizing conflicts: Group assessment and the spatio-temporal dynamics of ant territory battles.

Territorial battles among ants exhibit temporal and spatial patterns that self-organize, arising spontaneously from distributed decisions by large numbers of individuals. We describe agent-based models of inter-group fights in ants and show that two behavioral mechanisms that are rarely quantified have large effects on the dynamics of intraspecific battles; specifically, the pattern of search by unengaged ants, and assessment of relative numbers. In the absence of assessment, recruitment by both colonies rises to steady averages. Alternatively, if ants tend to lay trails only when they detect that their nestmates outnumber opponents, fights can be rapidly resolved as one colony ceases recruiting. If ants tend to lay trails when their nestmates are locally outnumbered, the position of the battle may oscillate. We show that the collective ability of fighting ants to accurately compare group sizes may be high even if each ant has limited perception and memory. However, amplification of small initial numerical advantages can lead to priority effects favoring the first colony to recruit even if it is the smaller colony.

New exocrine glands in ants: the hypostomal gland and basitarsal gland in the genus Melissotarsus (Hymenoptera: Formicidae).

Fisher and Robertson (Insect Soc 46: 78-83, 1999) discovered the production of silk-like secretions emerging from slit-shaped openings along the anterior margin of the ventral hypostoma of Melissotarsus ant workers. The current histological study describes a hitherto unknown hypostomal gland from which this silk-like substance originates. In addition, this study describes a new basitarsal gland in the three pairs of legs of Melissotarsus workers.

Chemical communication during foraging in the harvesting ants Messor pergandei and Messor andrei.

We combined behavioral analyses in the laboratory and field to investigate chemical communication in the formation of foraging columns in two Nearctic seed harvesting ants, Messor pergandei and Messor andrei. We demonstrate that both species use poison gland secretions to lay recruitment trails. In M. pergandei, the recruitment effect of the poison gland is enhanced by adding pygidial gland secretions. The poison glands of both species contain 1-phenyl ethanol. Minute quantities (3 µl of a 0.1 ppm solution) of 1-phenyl ethanol drawn out along a 40 cm long trail released trail following behavior in M. pergandei, while M. andrei required higher concentrations (0.5-1 ppm). Messor pergandei workers showed weak trail following to 5 ppm trails of the pyrazines 2,5-dimethylpyrazine and 2,3,5-trimethylpyrazine, whereas M. andrei workers showed no behavioral response. Minute quantities of pyrazines were detected in M. pergandei but not in M. andrei poison glands using single ion monitoring gas chromatography-mass spectrometry.

An empirically based simulation of group foraging in the harvesting ant, Messor pergandei.

We present an empirically based group model of foraging interactions in Messor pergandei, the Sonoran desert harvesting ant. M. pergandei colonies send out daily foraging columns consisting of tens of thousands of individual ants. Each day, the directions of the columns may change depending on the resource availability and the neighbor interactions. If neighboring columns meet, ants fight, and subsequent foraging is suppressed. M. pergandei colonies face a general problem which is present in many systems: dynamic spatial partitioning in a constantly changing environment, while simultaneously minimizing negative competitive interactions with multiple neighbors. Our simulation model of a population of column foragers is spatially explicit and includes neighbor interactions. We study how different behavioral strategies influence resource exploitation and space use for different nest distributions and densities. Column foraging in M. pergandei is adapted to the spatial and temporal properties of their natural habitat. Resource and space use is maximized both at the colony and the population level by a model with a behavioral strategy including learning and fast forgetting rates.

Pygidial gland chemistry and potential alarm-recruitment function in column foraging, but not solitary, Nearctic Messor harvesting ants (Hymenoptera: Formicidae: Myrmicinae).

We investigated the role of the pygidial gland on foraging behavior in two ecologically dominant column foraging Nearctic harvesting ants (Messor pergandei and Messor andrei). Using chemical analyses and behavioral tests, we show that n-tridecane is the major biologically active compound of pygidial gland secretions in both species, and that this chemical functions as a powerful alarm-recruitment pheromone. Another major compound of pygidial gland contents is benzaldehyde; this substance does not release behavioral reactions in M. pergandei workers but might function as a defensive secretion. Six solitary foraging Nearctic Messor and two column foraging Palearctic Messor species, did not have large pygidial gland reservoirs.

Metalloproteinase-dependent cleavage of neuregulin and autocrine stimulation of vascular endothelial cells.

Inflammation is often accompanied by robust angiogenesis. Vascular endothelial cells (ECs) express erbB receptors and their ligand, neuregulin-1, and can respond to neuregulin by proliferation and angiogenesis. We hypothesized that some growth factor-like responses of ECs to inflammatory cytokines can be explained by cleavage of transmembrane neuregulin with subsequent release of its extracellular epidermal growth factor-like-containing domain and autocrine activation. Using a model of cultured human ECs, we found that interleukin-6 or interferon-gamma causes rapid cleavage and release of transmembrane neuregulin. Inhibitors of metalloproteinases abolish this effect. The addition of an inhibitor of tumor necrosis factor-alpha converting enzyme (TACE) blocks cytokine-induced neuregulin release. Silencing of TACE expression increases the amount of basal proneuregulin present in ECs but does not block neuregulin release in response to phorbol myristate acetate (PMA), suggesting that other proteinases are responsible for mediating protein kinase C-dependent cleavage. Cytokines capable of inducing neuregulin cleavage stimulated ERK activation and in vitro angiogenesis (Matrigel cord formation). This effect is blocked by inhibitors that block neuregulin cleavage, erbB protein tyrosine kinase inhibitors, or antineuregulin-neutralizing antibodies. Cytokine-activated metalloproteinase cleavage of neuregulin may play an important role in autocrine activation of EC signaling pathways, contributing to key biological effects, perhaps including inflammation-associated angiogenesis.

An empirical test of Lanchester's square law: mortality during battles of the fire ant Solenopsis invicta.

Lanchester's models of attrition describe casualty rates during battles between groups as functions of the numbers of individuals and their fighting abilities. Originally developed to describe human warfare, Lanchester's square law has been hypothesized to apply broadly to social animals as well, with important consequences for their aggressive behaviour and social structure. According to the square law, the fighting ability of a group is proportional to the square of the number of individuals, but rises only linearly with fighting ability of individuals within the group. By analyzing mortality rates of fire ants (Solenopsis invicta) fighting in different numerical ratios, we provide the first quantitative test of Lanchester's model for a non-human animal. Casualty rates of fire ants were not consistent with the square law; instead, group fighting ability was an approximately linear function of group size. This implies that the relative numbers of casualties incurred by two fighting groups are not strongly affected by relative group sizes and that battles do not disproportionately favour group size over individual prowess.