1
Created by Peter Downing – Educational Media Access and Production © 2011 Latency (Left - Right) (s) -10 -5 0 5 10 15 20 M97 Latency M127 Latency M99 Latency TOC Time (s) -6 -5 -4 -3 -2 -1 0 1 EMG Spike frequency (Spikes/s) 10 12 14 16 18 20 22 24 LM97 RM97 LM127 RM127 LM99 RM99 TOC Time (s) -0.6 -0.4 -0.2 0.0 0.2 X (mm) 420 440 460 480 500 520 540 Z (mm) 260 280 300 320 340 360 X Z TOC X (mm) 420 440 460 480 500 520 Z (mm) 260 280 300 320 340 Locusts (Locusta migratoria) are ideal model systems to study complex behaviours, such as flight responses to objects approaching on a collision course. Thus far, flight muscle activity, wing kinematics and aerodynamic forces of locusts have been recorded during collision avoidance behaviour and measured from rigidly-tethered locusts flying in open-loop conditions. However, loosely-tethered flying locusts are capable of changing orientation in response to looming stimuli within a single wing beat, and generate avoidance responses within a single downstroke. To better understand neural control of flight steering, we placed a loosely-tethered flying locust inside an existing flight simulator, and presented visual stimuli of objects moving along complex trajectories. INTRODUCTION METHODS RESULTS Muscle activity and corresponding physiological changes Acknowledgements Funding provided by the Natural Science and Engineering Research Council of Canada, the Canada Foundation for Innovation and the University of Saskatchewan. References [1] McMillan, G. A., Loessin, V., & Gray, J. R. (2013). The Journal of experimental biology, 216(17), 3369-3380. [2] Silva, A. C., McMillan, G. A., Santos, C. P., & Gray, J. R. (2015). Journal of neurophysiology, 113(1), 218- 231. [3] Santer, R. D., Simmons, P. J., & Rind, F. C. (2005). Journal of Comparative Physiology A, 191(1), 61-73. [4] McMillan, G. A., & Gray, J. R. (2012). Journal of neurophysiology, 108(4), 1052-1068. SUMMARY ACKNOWLEDGEMENTS & REFERENCES Locust flight muscle activity and body orientation in response to objects moving along complex trajectories Sinan Zhang and John R. Gray, Dept. Biology, University of Saskatchewan Stimuli-independent EMG frequency change Behavioural responses during collision avoidance Locusts generate steering behaviour during collision avoidance, without preference in directions Changes in flight direction can be reflected by the latency between bilateral flight muscle spikes Within a flow field background, wing beat frequency decreased during object approach. High speed camera recordings (1028 x 1028 pixels at 225fps) revealed the trajectory of locust flight during collision avoidance. Images from two cameras, allowed 3D tracking of the locust position. A) 3D tracking results showing the locust generating a maintained right turn while avoiding the disc coming from left at 45°. B) Video capture from one camera, showing positions of the locust before and during avoidance behaviour; C) Movement in x and z planes showing the overall right turn (changes in the y plane were minor). Time aligned with TOC (Time of collision). Time A) Experimental setup, including visual stimuli projection, high speed cameras and EMG recordings C) Muscles recorded in this experiment: M97 (forewing 1st basalar), M99 (forewing subalar), and M127 (hindwing 1st basalar) B) Visual stimuli with flow field background and tether Loosely tethered locusts (n=3) were placed in the center of a rear projection dome screen. Head-on air flow (v=3m/s) was provided to maintain straight flight. Visual stimuli (7 cm disc travelling at 3 m/s) were projected from different directions (left 45°, right 45°, 0°), with a flow field background (B). Four markers on the tether monitored 3D body position. (C) We recorded from 6 flight muscles known to be associated with steering. Activity of 3 bilaterally paired steering muscles were also recorded during collision avoidance. Previous studies have shown that the timing of these muscles are associated with attempted steering manoeuvres. A) Sample EMG recordings (Left 45° looming); B) Bilateral latency increases indicating a steering behaviour; C) EMG spike frequency (reflecting wing beat frequency) decreased as the disc approached. A) B) C) B) Stimulus generation Stimulus frame pulses LCD projector Rear projection dome screen Data collection and analysis EMG Recordings Time (s) -4 -3 -2 -1 0 EMG Frequency (Spikes/s) 5 10 15 20 25 TOC EMG Frequency (Spikes/s) 5 10 15 20 25 EMG Frequency (Spikes/s) 5 10 15 20 25 B) C) A) C) Locust Locust Looming stimuli Looming stimuli Before During A) 3D motion tracking of a locust avoiding a disc looming from the left at 45°. The coordinate system is right-handed: positive x to the right, positive y down, positive z to the front (from the locust’s perspective of view) Locust Tether EMG spike frequency across all three types of looming stimuli, displaying a stimuli-independent decreasing trend over time. A)Left 45°; B)Right 45°; C)0°. Front (Z) Down (Y) Right (X)

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Page 1: Locust flight muscle activity and body orientation in

Created by Peter Downing – Educational Media Access and Production © 2011

Late

ncy (

Left

- R

igh

t) (

s)

-10

-5

0

5

10

15

20

M97 Latency

M127 Latency

M99 Latency

TOC

Time (s)

-6 -5 -4 -3 -2 -1 0 1

EM

G S

pik

e fr

equ

ency

(S

pik

es/s

)

10

12

14

16

18

20

22

24

LM97

RM97

LM127

RM127

LM99

RM99

TOC

Time (s)

-0.6 -0.4 -0.2 0.0 0.2

X (

mm

)

420

440

460

480

500

520

540

Z (

mm

)

260

280

300

320

340

360

X

Z

TOC

X (mm)

420 440 460 480 500 520

Z (

mm

)

260

280

300

320

340

Locusts (Locusta migratoria) are ideal model

systems to study complex behaviours, such as flight

responses to objects approaching on a collision

course. Thus far, flight muscle activity, wing

kinematics and aerodynamic forces of locusts have

been recorded during collision avoidance

behaviour and measured from rigidly-tethered

locusts flying in open-loop conditions. However,

loosely-tethered flying locusts are capable of

changing orientation in response to looming stimuli

within a single wing beat, and generate avoidance

responses within a single downstroke. To better

understand neural control of flight steering, we

placed a loosely-tethered flying locust inside an

existing flight simulator, and presented visual

stimuli of objects moving along complex

trajectories.

INTRODUCTION

METHODS

RESULTS

Muscle activity and corresponding physiological changes

Acknowledgements

Funding provided by the Natural Science and Engineering Research Council

of Canada, the Canada Foundation for Innovation and the University of

Saskatchewan.

References

[1] McMillan, G. A., Loessin, V., & Gray, J. R. (2013). The Journal of

experimental biology, 216(17), 3369-3380. [2] Silva, A. C., McMillan, G. A.,

Santos, C. P., & Gray, J. R. (2015). Journal of neurophysiology, 113(1), 218-

231. [3] Santer, R. D., Simmons, P. J., & Rind, F. C. (2005). Journal of

Comparative Physiology A, 191(1), 61-73. [4] McMillan, G. A., & Gray, J. R.

(2012). Journal of neurophysiology, 108(4), 1052-1068.

SUMMARY

ACKNOWLEDGEMENTS &

REFERENCES

Locust flight muscle activity and body orientation in response to

objects moving along complex trajectoriesSinan Zhang and John R. Gray, Dept. Biology, University of Saskatchewan

Stimuli-independent EMG frequency change

Behavioural responses during collision avoidance

• Locusts generate steering behaviour

during collision avoidance, without

preference in directions

• Changes in flight direction can be

reflected by the latency between

bilateral flight muscle spikes

• Within a flow field background, wing

beat frequency decreased during object

approach.

High speed camera recordings (1028 x 1028 pixels at 225fps) revealed the trajectory of locust flight during

collision avoidance. Images from two cameras, allowed 3D tracking of the locust position. A) 3D tracking

results showing the locust generating a maintained right turn while avoiding the disc coming from left at 45°.

B) Video capture from one camera, showing positions of the locust before and during avoidance behaviour;

C) Movement in x and z planes showing the overall right turn (changes in the y plane were minor). Time

aligned with TOC (Time of collision).

Time

A) Experimental setup, including visual stimuli projection, high

speed cameras and EMG recordings

C) Muscles recorded in this

experiment: M97 (forewing 1st

basalar), M99 (forewing subalar),

and M127 (hindwing 1st basalar)

B) Visual stimuli with flow

field background and tether

Loosely tethered locusts (n=3) were placed in the

center of a rear projection dome screen. Head-on air

flow (v=3m/s) was provided to maintain straight

flight. Visual stimuli (7 cm disc travelling at 3 m/s)

were projected from different directions (left 45°,

right 45°, 0°), with a flow field background (B).

Four markers on the tether monitored 3D body

position. (C) We recorded from 6 flight muscles

known to be associated with steering.Activity of 3 bilaterally paired steering muscles were also recorded during collision avoidance. Previous

studies have shown that the timing of these muscles are associated with attempted steering manoeuvres. A)

Sample EMG recordings (Left 45° looming); B) Bilateral latency increases indicating a steering behaviour;

C) EMG spike frequency (reflecting wing beat frequency) decreased as the disc approached.

A)

B)

C)

B)

Stimulus

generation

Stimulus frame

pulses

LCD

projector

Rear projection

dome screen

Data collection

and analysis

EMG Recordings

Time (s)

-4 -3 -2 -1 0

EM

G F

req

uen

cy (

Sp

ikes/

s)

5

10

15

20

25

TOC

EM

G F

req

uen

cy (

Sp

ikes/

s)

5

10

15

20

25

EM

G F

req

uen

cy (

Sp

ikes/

s)

5

10

15

20

25

B)

C)

A)

C)

LocustLocust

Looming

stimuliLooming

stimuli

Before During

A) 3D motion tracking of a locust avoiding a disc looming

from the left at 45°. The coordinate system is right-handed:

positive x to the right, positive y down, positive z to the front

(from the locust’s perspective of view)

Locust Tether

EMG spike frequency across all three types of

looming stimuli, displaying a stimuli-independent

decreasing trend over time.

A)Left 45°; B)Right 45°; C)0°.

Front (Z)

Down (Y)

Right (X)