First published online January 8, 2007
Journal of Experimental Biology 210, 198-207 (2007)
Published by The Company of Biologists 2007
doi: 10.1242/jeb.02657
The desert ant odometer: a stride integrator that accounts for stride length and walking speed
Matthias Wittlinger1,*,
Rüdiger Wehner2 and
Harald Wolf1
1 Institute of Neurobiology, University of Ulm, Albert-Einstein-Allee 11,
D-89069 Ulm, Germany
2 Institute of Zoology, University of Zürich, Winterthurerstrasse 190,
CH-8057 Zürich, Switzerland

View larger version (5K):
[in this window]
[in a new window]
|
Fig. 1. Experimental situation. Schematic diagram of channel layout; training
channel, top; test channel, bottom (broken lines indicate extended length of
test channel to 24 m). The search behaviour exhibited by Cataglyphis
foragers after having run off their home vector is illustrated schematically
below the test channel. Not drawn to scale. N, north.
|
|

View larger version (8K):
[in this window]
[in a new window]
|
Fig. 2. Manipulation of leg length. The right hind leg of a Cataglyphis
fortis worker is shown from the anterior [adapted from
fig. 1 in Wehner
(Wehner, 1983 )], and the
manipulations performed in the present study are indicated: stumps II and I,
the removal of the distal part of the leg from the mid tibia, and removal of
the tarsi, respectively; a normal leg; and stilts, the addition of pig
bristles to lengthen the leg.
|
|

View larger version (64K):
[in this window]
[in a new window]
|
Fig. 3. Tripod gait of a Cataglyphis individual before (normal) and after
leg shortening (stumps II). The six complete strides captured by the
high-speed video are shown, and indicated as the tripods performed by the
animal in its normal tripod gate. Tripods formed by the left front and hind
(L1, L3) and right middle (R2) legs are drawn in blue; those formed by the
right front and hind (R1, R3) and left middle (L2) legs are drawn in red.
Stride length (s2) was determined as the distance between two
successive middle leg footfalls. (A) Untreated normal situation, showing the
typical tripod gait (s2=13.3 mm; v=0.28 m s-1).
(B) After leg shortening the ant still shows the typical tripod gait but with
obviously decreased stride length (s2=10.5 mm; v=0.27 m
s-1). Single video frames of the ant, taken during the 1st and 6th
captured strides, are pasted into the tripod analysis.
|
|

View larger version (32K):
[in this window]
[in a new window]
|
Fig. 4. Homing distances of experimental ants. The top panels show search density
plots (abscissae, homing distance; ordinates, cumulated relative search
densities between the first and sixth turning points; see Materials and
methods), the bottom panels show box-and-whisker plots (medians of the initial
six turning points; compare Fig.
1,bottom), derived from the same data sets (N=25 ants for
each experimental situation). (A) The ants were tested immediately after the
lengths of their legs had been modified at the feeding site, that is, leg
lengths were normal during the outbound journey but manipulated during the
homebound run (Test 1). (B) The ants were tested after re-emerging from the
nest after previous manipulation. In this situation leg lengths were equal,
though manipulated, during outbound and homebound runs (Test 2). The hatched
box plots in the lower panel of A illustrate the search centres as predicted
from the high-speed video analyses of stride lengths in normal and manipulated
animals; for details see text. Colour code: red, stilts; blue, normal; yellow,
stumps I; green, stumps II.
|
|

View larger version (8K):
[in this window]
[in a new window]
|
Fig. 5. Widths of search density distributions change with altered leg length. The
half widths (width of graph at half-maximum height) of the search density
distributions in Fig. 4 were
determined. They are shown for the different experimental situations, Test 1,
top, Test 2, bottom. Colour code: red, stilts; blue, normal; yellow, stumps I;
green, stumps II.
|
|

View larger version (14K):
[in this window]
[in a new window]
|
Fig. 6. Relationship between (relative) stride length and stride frequency in
normal ants and after manipulation of leg length. Stride length was normalised
with regard to body size (alitrunk length, see Materials and methods), and is
shown on the ordinate; the abscissa gives stride frequency. Each data point
represents one individual. Best fit regression lines are indicated; the
respective equations are, from top to bottom: stilts, stride
length=0.124xstride frequency+2.14; normal, stride
length=0.084xstride frequency+1.89; stumps I, stride
length=0.061xstride frequency+1.788; stumps II, stride
length=0.057xstride frequency+1.42 (stride frequency = Hz). Colour code:
red, stilts; blue, normal; yellow, stumps I; green, stumps II.
|
|

View larger version (24K):
[in this window]
[in a new window]
|
Fig. 7. The relationship between walking speed and (A) stride frequency, (B)
(relative) stride length. The different experimental situations are
colour-coded as in the previous figures: red, stilts; blue, normal; yellow,
stumps I; green, stumps II. Linear regression lines are indicated. The
regression lines for the normal ants: stride frequency=0.054xwalking
speed+7.72 (R2=0.87) in A; stride
length=0.018xwalking speed+6.80 (R2=0.81) in B
(stride frequency = Hz, stride length = mm and walking speed = mm
s-1).
|
|
© The Company of Biologists Ltd 2007