spacer gif spacer gif spacer gif spacer gif spacer gif
 QUICK SEARCH:   [advanced]


spacer gif
     Home     Help     Feedback     Subscriptions     Archive     Search     Table of Contents    

First published online March 27, 2009
Journal of Experimental Biology 212, 1120-1130 (2009)
Published by The Company of Biologists 2009
doi: 10.1242/jeb.020768
This Article
Right arrow Summary Freely available
Right arrow Full Text
Right arrow Full Text (PDF)
Right arrow Supplementary Material
Right arrow Alert me when this article is cited
Right arrow Alert me if a correction is posted
Services
Right arrow Email this article to a friend
Right arrow Similar articles in this journal
Right arrow Alert me to new issues of the journal
Right arrow Download to citation manager
Right arrow reprints & permissions
Citing Articles
Right arrow Citing Articles via Google Scholar
Google Scholar
Right arrow Articles by Fry, S. N.
Right arrow Articles by Dickinson, M. H.
Right arrow Search for Related Content
PubMed
Right arrow Articles by Fry, S. N.
Right arrow Articles by Dickinson, M. H.
Social Bookmarking
 Add to CiteULike   Add to Complore   Add to Connotea   Add to Del.icio.us   Add to Digg   Add to Reddit   Add to Technorati   Add to Twitter  
What's this?

Visual control of flight speed in Drosophila melanogaster

Steven N. Fry1,2,*, Nicola Rohrseitz1, Andrew D. Straw3 and Michael H. Dickinson3

1 Institute of Neuroinformatics, University of Zürich and ETH Zürich, Switzerland
2 Institute of Robotics and Intelligent Systems, ETH Zürich, Switzerland
3 Bioengineering, California Institute of Technology, MC 138-78, Pasadena, CA 91125, USA


Figure 1
View larger version (26K):
[in this window]
[in a new window]

 
Fig. 1. Experimental analysis of optomotor turning and visual flight speed responses. (A) Optomotor turning response paradigm. An insect is tethered within a patterned rotating drum and its steady-state turning responses are measured. Because the body is fixed, the behavioral reactions have no effect on the perceived stimulus (open-loop condition). (B) Idealized spatio–temporal tuning of optomotor turning reactions. The response surface represents the steady-state output of a basic correlation-type motion detector scheme, calculated using the equations applicable for sine grating stimuli provided by Borst and Bahde (Borst and Bahde, 1986Go). Normalized response strength (green and dark red correspond to values between 0 and 1, respectively) is plotted as a function of the temporal frequency (tf, in s–1) and angular spatial frequency (sf, in deg.–1) of sine grating patterns. Optomotor turning responses are maximally elicited by a particular optimal temporal frequency (tfopt) and optimal spatial frequency (sfopt). (C) Standard flight speed paradigm. An insect is induced to fly along a patterned channel and its flight speed is measured under steady-state conditions. The insect adjusts its flight speed according to the perceived visual feedback from the pattern (closed-loop condition). (D) Flight speed measurements in the presence of sine grating patterns with varying linear spatial frequency (SF, in m–1). Insects adjust their flight speed as to hold constant the ratio of temporal and linear spatial frequency, such that measurements (red circles) fall on a diagonal line in the TF–SF parameter space, which corresponds to the insect's preferred velocity (Vpref=TF/SF) in m s–1. (E) One-parameter open-loop paradigm, which allows arbitrary patterns to be defined with respect to the fly's body coordinates, irrespective of its flight speed. Open-loop stimulation requires measuring the fly's position (symbolized by the two cameras) and controlling the pattern in real-time (black arrow). (F) Open-loop stimulation is required to characterize the transfer properties of the visual speed response in the two-dimensional TF–SF parameter space.

 

Figure 2
View larger version (30K):
[in this window]
[in a new window]

 
Fig. 2. Experimental setup and measurement procedure. (A) TrackFly. The free-flight experimental setup consisted of a wind tunnel (only the working section is shown) equipped with a real-time 3-D position tracking system (Trackit 3D) (Fry et al., 2000Go; Fry et al., 2004Go) and custom-programmed graphical rendering software (based on the VisionEgg) (Straw and O'Carroll, 2003Go; Straw et al., 2006Go). Flies were induced to fly upwind (red dotted arrow shows the flight direction of fly; blue arrows indicate wind direction), while their position was tracked in real time (green arrows pointing from cameras). Visual stimuli were projected onto the sidewalls via pairs of mirrors (yellow arrows show the light path from the projector to the screen for one side of the wind tunnel). The virtual reality features of TrackFly were used to implement a one-parameter visual open-loop paradigm [see text and Fry et al. (Fry et al., 2008Go) for further details]. (B) Definition of linear and angular coordinate systems. A plan view of the screens (sine gratings are represented by stripes for clarity) is shown to scale. The linear spatial frequency (SF, in m–1) of the pattern displayed on the screen corresponds to the inverse of the linear spatial pattern period ({lambda}, in m). In the example, SF=10 m–1 and thus {Lambda}=0.1 m. The angular period ({lambda}, unit: deg.) of the displayed patterns depends on the azimuth, with {lambda} decreasing toward frontal and caudal positions (note red arrows). Angular spatial frequency (sf, in deg.–1), the inverse of {lambda}, therefore increases toward frontal and caudal positions. The linear wavelength ({lambda}1, {lambda}2) and linear spatial frequency SF remain constant. (C) Sample acceleration responses and parameter extraction. The time course of body position along the wind tunnel (red traces) is shown for 11 measurements performed under identical stimulus conditions. Flies were first held near the middle of the wind tunnel by controlling the pattern speed (t<0). At t≥0, the flies were stimulated in open-loop (TF=4 s–1; SF=12.5 m–1; stimulus condition is marked with an asterisk in Fig. 4A). Flies reacted to the back-to-front image motion by accelerating forward, as indicated by the exponential increase of the position function (red traces, right part of the plot). Mean acceleration of each sample was measured from the fitting parameters of a parabola (t>0.1 s, black traces). These values were then averaged over the trials to obtain the response strength for a single TF–SF combination (each marked with a dot in Fig. 3 and Fig. 4). Figure modified from Fry et al. (Fry et al., 2008Go).

 

Figure 3
View larger version (46K):
[in this window]
[in a new window]

 
Fig. 3. Spatio–temporal tuning of speed responses. Body acceleration is shown color coded as a function of the open-loop temporal frequency (TF) and linear spatial frequency (SF) of presented sine grating stimuli. Contour lines show iso-response curves at intervals of 0.5 ms–2. The response surface was obtained by linearly interpolating over 435 parameter conditions, each indicated by a dot. These were calculated from a total of 12,711 trials.

 

Figure 4
View larger version (54K):
[in this window]
[in a new window]

 
Fig. 4. Velocity dependence and locomotor limits. (A) Response surface relevant to forward acceleration responses. The surface and iso-response curves are re-plotted from Fig. 3 (subset of 166 parameter conditions and 6245 trials); the zero iso-response line shown in bold and indicated with a black arrow. Labeled diagonal lines represent pattern iso-velocity lines (pattern velocity, V=TF/SF). Note the close correspondence between the iso-response curves and the pattern iso-velocity lines for pattern speeds up to 0.5 m s–1. Colored arcs follow the gradient of pattern velocity for varying radii in the spatio–temporal parameter space. (B) Pattern-invariant response properties. The response surface was evaluated along the four colored arcs shown in A to obtain response strength as a function of pattern slip speed, which is plotted in the respective colors together with the standard deviation. The responses depend on pattern velocity over a large range of the TF–SF parameter space. (C) Velocity-dependent responses to sine gratings and a naturalistic image. The mean responses to sine gratings moving at a particular velocity were obtained by averaging the response surface along the respective pattern iso-velocity line (within the range of the red and blue arcs). These are plotted against pattern velocity in black together with the standard deviation (gray shaded area). The red line shows a linear regression for pattern velocities up to V<0.6 m s–1 (R2=0.985; see text for further details). Mean responses to a photographic image moving at various velocities are plotted in green together with the standard deviation (shaded). (D) Air speed saturation. Median and 25 percentile values of maximal forward accelerations induced with strong motion stimuli (V>=0.6 m s–1; SF<25 m–1) in presence of low (left: –0.29 m s–1; N=941) and high (right: –0.73 m s–1; N=45) wind speeds. The measured accelerations reveal a significant dependence on the wind speed, suggesting an air-speed-dependent response saturation (Wilcoxon test, P<0.001).

 

Figure 5
View larger version (5K):
[in this window]
[in a new window]

 
Fig. 5. Flight speed control scheme. Optic flow was delivered in open-loop using TrackFly, such as to deliver a constant pattern slip speed during the short trials. The visual system (`V') extracts a speed signal, which is converted to a flight speed command. The actual measured flight speed is subject to air speed saturation. A quantitative control model and experimental verification under open- and closed-loop conditions is to be presented elsewhere (N.R. and S.N.F., in preparation).

 

Add to CiteULike CiteULike   Add to Complore Complore   Add to Connotea Connotea   Add to Del.icio.us Del.icio.us   Add to Digg Digg   Add to Reddit Reddit   Add to Technorati Technorati   Add to Twitter Twitter    What's this?




© The Company of Biologists Ltd 2009