How Navigational Guidance Systems Are Combined in a Desert Ant

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How Navigational Guidance Systems Are Combined in a Desert Ant Matthew Collett  Current Biology  Volume 22, Issue 10, Pages 927-932 (May 2012) DOI: 10.1016/j.cub.2012.03.049 Copyright © 2012 Elsevier Ltd Terms and Conditions

Figure 1 Manipulating PI along a Habitual Route (A) Schematic of the training route. An enclosure (circle) surrounded the nest, opening into a moveable tray (rectangle) that sat in the channel (parallel lines). The feeder (square) lay 6 m from the ramp at the end of the channel. The dashed arrow represents the local vector route memory. The PI output vector calculated at the end of the channel is represented by the solid arrow and by the coordinates in square brackets. The coordinates in curved brackets indicate an ant's global PI coordinates at significant positions. The PI-based goal is indicated by the cross (in this condition it coincides with the feeder). (B) Schematic of the PI manipulation on the training route. An ant is carried 4 m in the tray (curved arrow) before being released to run along the channel. The ant's global PI coordinates and the position of its PI-based goal (cross) are thus shifted 4 m with respect to training. The local vector (dashed arrow) and PI output vector (solid arrow) therefore no longer coincide. (C) Photo of experimental area. The channel and feeder were dug into the ground to make them inconspicuous. The wheel tracks, although striking from above, were not followed by the ants. Note the distant trees that are visible on the landward side of the panorama. More photos in Figure S1. (D) Trajectories along the habitual route from the training channel. (i) Reference trajectories. (ii) Test trajectories after PI manipulation. The coordinate system reflects an ant's PI global coordinates (in meters). Thus for the test trajectories, the feeder is at (6,6), indicated by the square, although the PI output vector is directed toward (10,6), indicated by the cross. Final convergence to the feeder (squares) can be guided by visual, and in some cases odor (see Figure S1), cues. (E) Relationship between the initial directions (over first 3 m) of an ant's reference trajectory (–0.7 ± 9.8°) and its test trajectory (8.1 ± 10.2°). Throughout, data are given as mean ± SD. Angles are measured clockwise from the direct path. Each point is a single ant (n = 21). The diagonal line indicates where the two would be equal. (F) Histogram of the residual directions (corresponds to vertical distance from diagonal line of points in E). The triangle beneath the scale shows the mean, and the diamond indicates the PI-based prediction—an accurate PI output vector minus the mean reference direction. See also Figure S1. Current Biology 2012 22, 927-932DOI: (10.1016/j.cub.2012.03.049) Copyright © 2012 Elsevier Ltd Terms and Conditions

Figure 2 Manipulations of PI State on Test Ground (A) Trajectories from the test channel. Training is with configuration B (see Figure S1). Coordinates reflect the ants' global PI coordinates. The abknicht on each trajectory is indicated by a point, and the beginning of the subsequent search is shown in light gray. (i) 2 m manipulation; (ii) 6 m manipulation; (iii) 10 m manipulation; (iv) 14 m manipulation. (B) Histograms of residual directions between test and reference trajectories in (A). (i) 2 m manipulation. Residual directions = 22 ± 14.3° (mean ± SD), n = 17. (ii) 6 m manipulation. Residual directions = 13.2 ± 6.7°, n = 14; (iii) 10 m manipulation. Residual directions = 5.7 ± 12.4°, n = 14; (iv) 14 m manipulation. Residual directions = −0.8 ± 7.8°, n = 14. (C) Manipulation with an oblique channel during training with Landmark configuration A. Residual directions = 4.0 ± 7.1°. Note that the trajectories do not rotate with the direction of the preceding channel. (D) Histogram of lengths of trajectories before abknicht. (i) 2 m manipulation. Lengths = 5.8 ± 2.9 m, n = 18; (ii) 6 m manipulation. Lengths = 6.3 ± 1.5 m, n = 15; (iii) 10 m manipulation. Lengths = 5.9 ± 1.7 m, n = 15; (iv) 14 m manipulation. Lengths = 6.3 ± 2.2 m, n = 15; (v) Oblique manipulation. Lengths = 3.9 ± 0.7 m, n = 12. See also Figure S2. Current Biology 2012 22, 927-932DOI: (10.1016/j.cub.2012.03.049) Copyright © 2012 Elsevier Ltd Terms and Conditions

Figure 3 Convergence of Independent Navigational Guidance Systems into a Common Population Encoding of Heading Direction (A) Schematic of navigational architecture showing the processing stage at which information may be combined. (B) Activation patterns in a common population encoding. The resultant (solid line with peak indicated by a triangle) arises from a simple superposition of the heading directions from two guidance systems (dashed lines with peaks indicated by open circles). (i) The compromise is biased when two activation patterns have equal variance. (ii) The compromise is biased when two activation patterns have unequal variance. (iii) There are multiple peaks when activation patterns are widely divergent. Current Biology 2012 22, 927-932DOI: (10.1016/j.cub.2012.03.049) Copyright © 2012 Elsevier Ltd Terms and Conditions

Figure 4 Influence of Landmarks on Habitual Route During training, landmarks were in configuration A. (A) Reference trajectories; n = 27. Abknichts are shown in the four trajectories that do not reach the feeder directly (15%). (B) Feeder-landmark removed. Note that the convergence is not simply beaconing, because removing the feeder-landmark alone does not abolish the convergence; n = 23. Abknichts shown in nine trajectories (39%). (C) All three artificial landmarks removed; n = 17. Abknichts shown in 15 trajectories (88%). (D) Correlations between reference and test trajectories when only the feeder landmark is removed (filled circles, n = 20. Residual directions = 5.0 ± 9.2°) and when all three artificial landmarks are removed (open circles, n = 12. Residual directions = 7.3 ± 10.0°). Means of reference and test trajectories shown as triangles (filled and open) on the axes. (E) Superposition of three guidance systems. Further details of the model are in Supplemental Experimental Procedures. (i) Training with a rich landmark configuration. GDIM is centered at 0° with SD = 25°. The PI output vector is centered at 10° (reflecting systematic under-turning) with SD = 25°. Both patterns are shown as pale dashed lines with peak directions indicated by pale circles. The local vector (dark dashed line) is derived from the superposition of GDIM and the PI output vector. Peak direction (dark circle) lies under the peak direction of resultant (triangle). (ii) Superposition when artificial landmarks are removed. GDIM is given SD = 45° but all other parameter remain constant. (iii) Extended training with landmarks removed. The local vector reflects the superposition of the PI output vector and new GDIM. See also Figure S3. Current Biology 2012 22, 927-932DOI: (10.1016/j.cub.2012.03.049) Copyright © 2012 Elsevier Ltd Terms and Conditions