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Contributions From Parallel Strategies For Spatial Orientation in C. Elegans
Contributions From Parallel Strategies For Spatial Orientation in C. Elegans
elegans
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diffuse over the course of the simulated assays. We also
replicate the parameters of chemotaxis assays, such as spot
locations and amounts, model worm starting position, and
duration of the assay.
We have embraced a computational modeling approach
for several reasons. Most importantly, we are able to se-
lectively enable and disable each strategy. With a compu-
tational approach we are also afforded precise control over
our environments, and even able to make manipulations that
would not be possible in the real world. We are also able
to collect more fine-grained measurements of the worm’s
chemotaxis than we would be able to in an empirical set-
ting, as well as run an otherwise impossible number of (sim-
ulated) assays.
Model
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equation for two-dimensional diffusion without a border:
2
C = No e−r 400Dt
/4πdDt (2)
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0.65
Frequency of solutions
Chemotaxis Index 0.60 1.5
0.55
0.50 1.0
0.45
0.40 0.5
0.35
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Figure 5: Strategies on the radial assay. The four combinations of strategies we examined are shown in the columns: (A) Dual
strategy; (B) Klinotaxis-only strategy; (C) Klinokinesis-only strategy; and (D) Null strategy. Top row depicts a bird’s eye view
of the traces of the worms over time, for five representative examples. Middle row depicts the density plots for the locations
visited using 10,000 worms. Bottom row depicts distance to the peak of the gradient over time for the five model worms in the
top row and the median (black) and mean (dashed) of the population from the middle row.
ness of the gradient has on the contribution of each strategy. Effect of noise on the effectiveness
To address this we designed a steepness experiment, where
the concentration gradient was subject to a multiplicative All the work we have shared so far involves concentration
scale parameter, which increased or decreased the gradient’s gradients that are perfectly smooth and lacking noise. How-
steepness but otherwise left the gradient undistorted. A scale ever, this is not the case in the real world - the gradient itself
parameter of 1.00 reflects the original gradient. We analyzed may be noisy and imperfect, or the worm’s sensory appa-
the effect of gradient steepness on the contributions of the ratus may be imperfect. Therefore, we wish to study how
two strategies in the original grid assay using the Chemo- noise will affect the model worm’s performance. To probe
taxis Index (Fig. 7) and in the radial assay using a wider this we applied additive noise to the model worm’s sensed
range of metrics (Fig. 8). In both assays, we can see that concentration, which was sampled from a normal distribu-
the effectiveness of the overall spatial orientation behavior tion. Additionally, a modification was made to how klinoki-
improves with the steepness of the gradient (blue traces). In nesis senses the environment. Previously the sensorimo-
both assays, we can also see that klinotaxis (orange) con- tor transformation was acting on the instantaneous change
tributes more than klinokinesis (green) consistently across in the gradient’s concentration, but in this experiment the
the full range of the scale considered. Crucially, the over- model was parameterized to accommodate time windows of
all contribution from klinotaxis becomes relatively a larger greater length. This implementation takes the average of the
portion of the full behavior with steepness. In other words, differences between the sensed concentrations in the time
klinotaxis plays a larger role in steep gradients than in shal- window. This is consistent with C. elegans neuronal trace
low gradients. The opposite is true for klinokinesis, the rela- observations that suggest klinokinesis integrates the differ-
tive contribution to the overall behavior is larger in the shal- ence in concentration sensed over a time window of several
low gradients than it is in the steeper gradients. seconds (Kato et al., 2014).
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Figure 6: Across all four measures the dual strategy is most effective, followed by klinotaxis-only, then by klinokinesis-only
(N=10,000). First row: histograms of dual strategy. Second row: klinotaxis-only. Third row: klinokinesis-only.
N-spot experiment
We have investigated environments that are steep, shallow,
noisy, or smooth, but they still all have one thing in com-
mon: they all contain one NaCl spot. How will the contri-
bution of each strategy change with many NaCl spots? To
test this, we selectively disabled spots in the grid assay, cre-
ating symmetrical environments of 12, 8, 4, 2, and 1 spot(s).
Figure 7: Effect of gradient steepness on the contribu- The amount of total NaCl was held constant, such that the
tions of the two strategies in the grid assay. Depicted spot in the 1-spot environment contained 12 times as much
are the dual strategy (blue), klinotaxis-only strategy (or- NaCl as the spots in the 12-spot environment. These spots
ange), klinokinesis-only strategy (green), and the null strat- also tend to be steeper than those in the radial assay, as they
egy (dashed black). are allowed to diffuse for only 1 hour before the experiment
(versus 4-19 hours in the radial assay).
In the 1-spot condition, all three strategies are at near-
chance performance (Fig. 10) probably because the gradient
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Figure 10: In environments with few spots all three strate-
gies exhibit similar effectiveness, but as the number of spots
increases, klinotaxis becomes more important. Depicted
are the dual strategy (blue), weathervane strategy (orange),
pirouette strategy (green), and no strategy (dashed black).
Discussion
We set out to examine the contributions of the klinokinesis
and klinotaxis strategy to the overall spatial orientation be-
havior in the worm across different environments. Our anal-
ysis suggests klinotaxis is the primary contributor to chemo-
taxis in most environments, although klinokinesis always
contributes. We found that klinotaxis contributes more in
steep gradients and klinokinesis contributes more in shallow
gradients.
The two strategies are complementary in terms of
Figure 9: Longer pirouette time windows are more robust
timescales and in terms of the bearing between the worm’s
in the face of noise, with diminishing returns for windows
heading and the source of the gradient. In terms of
longer than about 12 seconds. Each cell represents one pop-
timescales, klinotaxis operates on a faster timescale than kli-
ulation of 2,000 model worms.
nokinesis. The time scale of klinotaxis is determined by the
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length of lateral head sweeps, whereas klinokinesis operates Foundation under Grant No. 1845322.
over the range of the worm’s translational movement for-
ward. In terms of spatial scales, klinotaxis contributes the References
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This work suggests several future experiments and model
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