Use of Magnetic Map Information by a Short-Distance Migrant
Newts deprived of directional visual, olfactory, magnetic and inertial
cues during displacement from their home ponds ('route deprivation')
have been shown to exhibit accurate homeward orientation when given
access to both magnetic and non-magnetic cues at the testing site,
indicating that they can use map information to determine the home
direction (Phillips
et al. 1995). More recent experiments investigated the possibility
that magnetic inclination is used to derive one coordinate of a
unicoordinate or bicoordinate map (Fischer
et al. 2001, Phillips
et al. 2002). Newts were displaced from their home ponds 42
km north-northeast to a testing site (home direction = 207 degrees)
where the inclination was slightly steeper than the inclination
at their home ponds. When exposed to a +2 degree change in inclination
resulting in a value that should occur in the same general direction
as our testing site, but at a greater distance from the home pond,
the newts were homeward oriented (Fig 1A) and indistinguishable
from controls (Fig 1B). In contrast, when newts were exposed to
a -2 degree change in inclination resulting in a value that should
occur on the other side of the home ponds, they oriented in the
opposite direction (Fig 1E). Identical changes in inclination had
no effect on shoreward magnetic compass orientation, which does
not involve the use of map information (see Fig 4 in Fischer
et al. 2001), suggesting that the change in orientation in Fig
1E was due to an effect on the map rather than the compass component
of homing.
In subsequent experiments (Phillips
et al. 2002), newts were exposed to smaller changes in magnetic
inclination to behaviorally "titrate" the home value (see
Phillips 1996). As predicted, the transition from homeward to
anti-homeward orientation coincided with the value of inclination
measured in the vicinity of the newts' home ponds (Fig 1B-D). Moreover,
newts exposed to a -0.15 + 0.03 degree decrease in inclination,
which coincided with the home value of -0.17 + 0.04 degree , failed
to exhibit a consistent direction of orientation (Fig 1C). Disorientation
is consistent with the newts having a unicoordinate map that would
require other information (e.g., familiar landmarks) to specify
their east-west position relative to home, or a bicoordinate map
with the value of the second coordinate also coinciding with the
home value.
| Figure 1. Effects of changes in magnetic
inclination on the homing orientation of newts (Data from
Fischer et al., 2001; Phillips et al. 2002). A) Newts exposed
to a 2 + 0.2 degree increase in inclination oriented in
the home direction. B) Newts exposed to the ambient magnetic
field at the testing site (controls) also oriented in the
home direction. C) Newts exposed to a decrease in inclination
of -0.15 + 0.03 degree , which approximated the home value,
failed to exhibit a consistent direction of orientation.
D) Newts exposed to a decrease in inclination of -0.48 +
0.04 degree were significantly oriented opposite the home
direction. E) Newts exposed to a -2.0 + 0.2 degree decrease
also oriented ~opposite the home direction. Methods: Data
points are magnetic bearings of newts tested in one of four
alignments of an Earth-strength magnetic field (magnetic
north at North, East, South or West). Prior to testing,
newts were held in outdoor tanks with an artificial shore
to the North (diamond-shaped symbols) or West (round symbols).
Arrows inside the circles indicate mean vectors that were
significant by the Rayleigh test (p < 0.05). Mean vector
length, r, is proportional to the strength of orientation
(radius of circle corresponds to r = 1). Dashed lines show
the 95% confidence interval for the mean vector bearings.
The black triangle outside each circle indicates the home
direction. Values on the left are the inclination changes
relative to the ambient value at the testing site (B). |
|
In subsequent experiments (Phillips
et al. 2002), newts were exposed to smaller changes in magnetic
inclination to behaviorally "titrate" the home value (see Phillips
1996). As predicted, the transition from homeward to anti-homeward
orientation coincided with the value of inclination measured in
the vicinity of the newts' home ponds (Fig 1B-D). Moreover, newts
exposed to a -0.15 + 0.03 degree decrease in inclination, which
coincided with the home value of -0.17 + 0.04 degree , failed to
exhibit a consistent direction of orientation (Fig 1C). Disorientation
is consistent with the newts having a unicoordinate map that would
require other information (e.g., familiar landmarks) to specify
their east-west position relative to home, or a bicoordinate map
with the value of the second coordinate also coinciding with the
home value.
Use of Magnetic Map Information by a Long-Distance Migrant
Behavioral experiments with experienced adult migratory birds have
also provided evidence for the geomagnetic field's involvement in
deriving map information (Fischer
et al. 2002). Adult Tasmanian silvereyes were tested in Armidale,
NSW near the mid point of their fall migration up the southeastern
coast of Australia. In the ambient field of Armidale, the birds
exhibited seasonally-appropriate north-northeasterly (NNE) orientation.
The birds were then divided into two groups and exposed to magnetic
intensity and inclination values that would normally be encountered
near the beginning (simulated south displacement, "SimS") or end
(simulated north displacement, "SimN") of their fall migration.
Consistent with the predictions of the magnetic map hypothesis,
adult migrants exposed to the SimS condition continued to exhibit
NNE orientation, while those exposed to the SimN condition failed
to exhibit a consistent direction of orientation (Fig 2).
| |
Figure 2. Effects of changes in magnetic intensity
and inclination on the migratory orientation of adult Tasmanian
silvereyes. A) SimS birds tested in a magnetic field with
values of approximately 60,200 nT (total intensity) and
-65 degree (inclination) oriented to the NNE (38 degrees,
r = 0.75, p < 0.01, one-sample Hotelling's test); and
did not differ significantly from the responses of these
birds in the control condition (p > 0.10, two-sample
Hotelling's test; control data, data included in B). B)
Controls tested in the ambient magnetic field with values
of approximately 55,000 nT (total intensity) and -62 degree
(inclination) exhibited seasonally appropriate orientation
to the NNE (22 degrees, r = 0.52, p < 0.001, one-sample
Hotelling's test). C) Responses of SimN birds tested in
a magnetic field with values of approximately 49,950 nT
(total intensity) and -58.5 degree (inclination) were randomly
distributed (p > 0.10; one-sample Hotelling's test) and
differed significantly from their responses in the control
condition (p < 0.01, two-sample Hotelling's test; data
included in B). Responses of silvereyes in SimN and SimS
(A vs C) also differed significantly (p < 0.01, Hotelling's
two-sample test). Data points outside circles indicate nightly
mean vector bearings of individual birds relative to magnetic
north. Vectors originating at centers of circles are mean
vector bearings for each of the birds, calculated by vector
addition from each bird's nightly mean vector bearings (Batschelet
1981). Radii of circles correspond to a mean vector length
of 1. Ellipses indicate 95% confidence intervals for second
order mean vectors. |
To determine whether the change in orientation in the SimN condition
was due to an effect on the map or compass, more recent experiments
were carried out to compare the effects of somewhat larger magnetic
field changes on the fall migratory orientation of young, inexperienced
birds captured in Tasmania shortly after fledging ("Juveniles")
with the effects on experienced adult birds that had completed at
least one round trip migration ("Adults"). Birds on their first
migration are believed to utilize only compass information combined
with a temporal program that specifies distance, while experienced
adults exhibit true navigation that utilizes both map and compass
information. Consistent with the magnetic map hypothesis, the SimS
treatment did not affect either group of birds, while the SimN treatment
affected only the older birds. While consistent with a "map effect",
the failure of experienced adult birds in the SimN condition to
reorient back to the south remains to be explained (Deutschlander
et al. 2003).
Evidence for a Specialized "Map Detector"
Behavioral studies of newts (Phillips
1986b, Phillips
& Borland 1994, Phillips et al. submitted), and silvereyes
(Munro
et al. 1997a, Munro
et al. 1997b) have provided evidence for the involvement of
a second magnetoreception mechanism in deriving map information.
In contrast to the light-dependent magnetic compass, the putative
"map detector" appears to be non-light-dependent, and to involve
permanent magnetic material, possibly magnetite (e.g.,
Brassart et al. 1999, Phillips
et al. 2002).
This material is based upon work supported by the National
Science Foundation under Grant Nos. IBN95-07826 and IBN98-08420
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