
approach to applying a separation principle: control inputs are specifically designed to excite sensory
receptors, presumably in service to the state estimator. This may be, at least in part, because biological
sensory systems often stop responding to persistent (i.e. “DC”) stimuli, via sensory “adaptation” [22–26]
or “perceptual fading” [27,28].
In this paper, we formalize a class of nonlinear systems that have a simple high-pass sensory output that
mimics sensory adaptation or perceptual fading. Under some simplified modeling assumptions, reviewed
below, this implies that linear observability is lost, which means the usual LQG-style framework does not
apply. However, under some interesting modeling conditions, nonlinear observability persists. Critically,
nonlinear observability does not necessarily afford a separation principle: the control signal may need
to contain ancillary energy that is expressly for the purpose of state estimation, and may be in conflict
with task goals. Indeed, the energy expended for active sensing movements do not necessarily directly
serve a motor control goal, and are instead believed to improve sensory feedback and prevent perceptual
fading [12, 27, 28].
The organization of the paper is as follows. Section 3 motivates the model structure from prior work
and Section 4 generalizes the model and presents the main theorem. Section 5 has the proof of the main
theorem. The Appendix provides some background concepts for the ease of understanding the tools used
in Section 3 and 5.
3 Biological motivation and simplified system
Station keeping behavior in weakly electric fish, Eigenmannia virescens, provides an ideal system for
investigating the interplay between active sensing and task-level control [11, 12, 29, 30]. These fish rou-
tinely maintain their position relative to a moving refuge and uses both vision and electrosense to collect
the necessary sensory information from its environment [31–34]. While tracking the refuge position (i.e.,
task-level control), the fish additionally produce rapid “whisking-like” forward and backward swimming
movements (i.e., active sensing). When vision is limited (for example, in darkness), the fish increase
their active sensing movements [12,30]. This increased motion compensates the lack of visual cues [11].
Suppose xis the position of an animal and z= ˙xis its velocity as it moves in one degree of freedom.
We assume that a sensory receptor measures only the local rate of change of a stimulus, s(x) as the
animal moves relative to the sensory scene, i.e. y=d
dts(x). Defining γ(x) := d
dxs(x), we arrive at a
2-dimensional, single-input, single-output normalized mass-damper system of the following form [35,36]:
˙x=z, x ∈R
˙z=−z+u, z, u ∈R
y=d
dts(x) = γ(x)z, y ∈R
(1)
where the mass and the damping constant both are assumed to be unity. Linearization of the above
system (1) around any equilibrium, (x∗,0), is given by (A, B, C) as follows:
A=0 1
0−1, B =0
1, C =0γ∗,
where γ∗=γ(x∗). Clearly (A,C) is not observable irrespective of γ∗[37]. Indeed, the output introduces
a zero at the origin that cancels a pole at the origin, rendering xunobservable. Assuming no input u,
we can write the system (1) as, ˙
ξ=f(ξ), y =h(ξ),(2)
where ξ= (x, z)>, f = (z, −z)>and h(ξ) = γ(x)z. We can construct the observation space, O(set of
all infinitesimal observables) by taking y=γ(x)zwith all repeated time derivatives
y(k)=L(k)
f(γ(x)z)
2