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Paul Graham

Publications and source records attributed to Paul Graham.

9 recordsLinked to original sources
PubMed2006

Bi-directional route learning in wood ants.

Some ants and bees readily learn visually guided routes between their nests and feeding sites. They can learn the appearance of visual landmarks for the food-bound or homeward segment of the route when these landmarks are only present during that particular segment of their round trip. We show here that wood ants can also acquire landmark information for guiding their homeward path while running their food-bound path, and that this information may be picked up, when ants briefly reverse direction and retrace their steps for a short distance. These short periods of looking back tend to occur early in route acquisition and are more frequent on homeward than on food-bound segments.

Animals↗
PubMed2005

Ant navigation: priming of visual route memories.

Ants travelling to and fro between their nest and a foraging area may follow stereotyped foodward and homeward routes that are guided by different visual and directional memory sequences. Honeybees are known to fly a feeder-to-hive or hive-to-feeder vector according to whether or not they have recently fed--their feeding state controls which compass direction they select. We show here that the feeding state of the wood ant Formica rufa also determines the choice between an outward or inward journey, but by priming the selective retrieval of visual landmark memories.

Animal Feed↗
PubMed2004

The binding and recall of snapshot memories in wood ants (Formica rufa L.).

Insects can locate spatial goals by means of 2-D retinotopic views of the surrounding landmarks, which they memorise from the vantage point of the goal. Wood ants acquire such snapshot memories while fixating conspicuous landmarks with frontal retina, and their snapshots extend horizontally at least 120 degrees into the periphery. Are spatially separate items within such an extended snapshot bound together so that a snapshot is recalled as a whole, or are its components recognised individually? We approached this question by training ants to find food midway between two upright black cylinders of different sizes and then examined where they searched when they were given two cylinders of the same size. If the ants know which cylinder replaces the small cylinder and which the large, they should search at a position where the two equal-sized cylinders subtend the same angles as do the training cylinders when viewed from the feeder. Ants conformed to this prediction under one condition, searching at a shorter distance from the substitute for the large cylinder than from the substitute for the small cylinder. But, under another condition, ants were unable to distinguish between the two equal-sized cylinders. Ants failed when white curtains completely surrounded the platform on which the cylinders were placed. They succeeded when one side of the platform had a patterned curtain. We suggest that ants take two snapshots at the feeding site, one when facing the small cylinder and one when facing the large cylinder, and that each snapshot includes the patterned curtain, if it is there. Ants will view the patterned curtain with the lateral retina of one eye when facing the small cylinder and with the lateral retina of the other eye when facing the large cylinder. Our data suggest that there may be associative links between these spatially separate components of the snapshot, which cause the memory of the small cylinder or the large cylinder to be recalled according to which eye sees the curtain. It seems that an extended snapshot not only enhances the accuracy of localisation but can also increase the reliability of snapshot recall, provided that the components of a snapshot are bound together.

Animals↗
PubMed2004

Switching destinations: memory change in wood ants.

We have studied the changing use of spatial memories in wood ants by charting how the ants' paths transform when ants are first trained to feed at one site and must then switch to another site. Because ants, which are trained to approach a single feeding site from a single starting point, are attracted directly to that goal when started from unfamiliar positions, we describe the ants' paths in terms of the use of two stored snapshots. Each snapshot consists of retinotopic views of the ants' surroundings acquired at one of the two feeding sites. When a snapshot is activated, it draws an ant to the related site from a wide range of directions. Here, we focus on routes that occur before ants have learnt to go directly from the start to the second site. The initial direction of the ant's path is then mostly aimed either at the first site or between the two sites. On 62.2% of all recorded paths, this segment is followed by an abrupt turn, after which the ant often aims directly at the second feeding site. The details of this behaviour suggest that, after the turn, control of the path switches from the snapshot recorded at the first feeding site (or some combination of the two snapshots) to the snapshot recorded at the second feeding site. We discuss different ways in which control might be transferred from one snapshot to the other.

Animals↗
PubMed2003

Snapshot memories and landmark guidance in wood ants.

Insects are thought to pinpoint a place by using memorized "snapshots," i.e., two-dimensional retinotopic views of the surrounding landmarks recorded when at the place (reviewed in ). Insects then reach the place by moving until their current view matches their snapshot. To determine when snapshots are recalled, and how differences between view and snapshot are translated into appropriate movements, we analyzed the approaches of wood ants to a feeding site that was located in the center of an array of two or three cylinders. In ants, contrary to flying hymenopterans, body orientation and direction of travel are collinear, so that an ant approaching an object always looks at it with frontal visual field. On their way to a food site, ants fixated and approached a cylinder predominantly when its angular size was smaller than when viewed from the food site. This finding implies that ants store snapshots at this place while fixating landmarks with frontal retina, so simplifying the later alignment of snapshots with their current view. It also means that ants recall snapshots well in advance of reaching the place. Although snapshots are centered on a landmark, we show that they extend at least 120 degrees into the periphery.

Animals↗
PubMed2003

Route learning by insects.

Ants and other insects often follow fixed routes from their nest to a foraging site. The shape of an ant's route is set, initially, by navigational strategies, such as path integration and the ant's innate responses to landmarks, which depend minimally on memory. With increasing experience, these early routes are stabilised through the learning of views of landmarks and of associated actions. The substitution of memory-based strategies makes an insect's route more robust and precise. The ability to select between different learnt routes might incur additional memory requirements to those needed for performing a route, and lead to the associative grouping of those memories that relate to a particular route.

Animals↗
PubMed2003

The influence of beacon-aiming on the routes of wood ants.

Many insects have an innate propensity to approach conspicuous objects. We explore how such beacon aiming determines the shape of a wood ant's habitual route. We find that a single large black cylinder within an arena biases the route taken by ants as they run from a start position at one end of the arena to reach a feeder at the other. Ants learn a stable route with the first segment of their trajectory aimed at the cylinder, which becomes an intermediate goal on the way to the feeder. When in occasional tests the cylinder is removed or displaced, ants head for the usual site of the cylinder. They also aim for the same site when the cylinder is removed and the ant's normal start position is changed. This behaviour suggests that visual features of the arena are learnt from the vantage point of the cylinder and that this stored snapshot guides the ant to that site. Ants thus reinforce their ability to reach the cylinder by learning other visual features in their surroundings that can also steer them to its location. The use of beacon aiming in fixing routes has several benefits. Because the same path will be traversed on every trial, beacon aiming facilitates the acquisition of routes. Beacon aiming also increases the robustness of learnt routes: ants straying from the route will be attracted to the closest beacon and so regain their habitual paths.

Animals↗
PubMed2002

View-based navigation in insects: how wood ants (Formica rufa L.) look at and are guided by extended landmarks.

Bees, wasps and ants learn landmarks as views from particular vantage points, storing the retinal positions of landmark edges. By moving so as to minimise the difference between their stored and current view, they can return to the vantage point from which a view was taken. We have examined what wood ants learn about a laterally placed, extended landmark, a wall, while walking parallel to it to reach a feeder and how they use this stored information to guide their path. Manipulation of the height of the wall and the ant's starting distance from it reveals that ants maintain a desired distance from the wall by keeping the image of the top of the wall at a particular retinal elevation. Ants can thus employ image matching both for returning to a place and for following a fixed route. Unlike many flying insects, an ant's direction of motion while walking is always along its longitudinal body axis and, perhaps for this reason, it favours its frontal retina for viewing discrete landmarks. We find that ants also use their frontal retina for viewing a laterally placed wall. On a coarse scale, the ant's path along the wall is straight, but on a finer scale it is roughly sinusoidal, allowing the ant to scan the surrounding landscape with its frontal retina. The ant's side-to-side scanning means that the wall is viewed with its frontal retina for phases of the scanning cycle throughout its trajectory. Details of the scanning pattern depend on the scene. Ants scan further to the side that is empty of the wall than to the side containing the wall, and they scan further into the wall side when the wall is of a lower apparent height. We conclude that frontal retina is employed for image storage and for path control.

Animals↗