SCIENCE ROBOTICS | RESEARCH ARTICLE

C O L L E C T I V E B E H AV I O R Copyright © 2023 The

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in a swarm of sterically interacting robots American Association
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Matan Yah Ben Zion1,2,3*, Jeremy Fersula1,2, Nicolas Bredeche2, Olivier Dauchot1 to original U.S.
Government Works
Whereas naturally occurring swarms thrive when crowded, physical interactions in robotic swarms are either
avoided or carefully controlled, thus limiting their operational density. Here, we present a mechanical design
rule that allows robots to act in a collision-dominated environment. We introduce Morphobots, a robotic swarm
platform developed to implement embodied computation through a morpho-functional design. By engineering
a three-dimensional printed exoskeleton, we encode a reorientation response to an external body force (such as
gravity) or a surface force (such as a collision). We show that the force orientation response is generic and can
augment existing swarm robotic platforms (e.g., Kilobots) as well as custom robots even 10 times larger. At the

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individual level, the exoskeleton improves motility and stability and also allows encoding of two contrasting
dynamical behaviors in response to an external force or a collision (including collision with a wall or a
movable obstacle and on a dynamically tilting plane). This force orientation response adds a mechanical
layer to the robot’s sense-act cycle at the swarm level, leveraging steric interactions for collective phototaxis
when crowded. Enabling collisions also promotes information flow, facilitating online distributed learning.
Each robot runs an embedded algorithm that ultimately optimizes collective performance. We identify an effec-
tive parameter that controls the force orientation response and explore its implications in swarms that transition
from dilute to crowded. Experimenting with physical swarms (of up to 64 robots) and simulated swarms (of up to
8192 agents) shows that the effect of morphological computation increases with growing swarm size.

INTRODUCTION
A robotic swarm is a synergetic ensemble capable of a greater task
than its constituents (1). In a swarm, self-organization is an emer-
gent property, making the swarm resilient to the malfunction of an
individual. Swarm engineering poses an interdisciplinary challenge,
combining electrical and mechanical engineering, computer
science, and nonequilibrium statistical physics (2, 3). Discovering
robust design rules for a swarm offers an opportunity to simplify
the specifications of the individual robotic units, reducing manufac-
turing complexity and making production at-scale accessible (4–7).
Recent development in robotic swarms demonstrated potential ap-
plications in collective construction (8), coordinated motion for un-
crewed aerial vehicles (UAVs) (9–12), patrol in open water (13), or
underwater exploration (14, 15). These applications are character-
ized by swarms where robots explicitly avoid physical contact
(16). Another approach for swarm design is to keep a cohesive
swarm, where robots are constantly touching, keeping a physical in-
teraction that allows continuous transmission of information (17–
22). This approach showed success in self-assembly and morpho-
genesis (23–25) as well as coordination of a multicellular robotic
body (18, 26–28). To date, artificial swarms are designed to
operate exclusively in a dilute, collision-avoided setting or a cohe-
sive dense population. Resorting to in silico experiments to guide
swarm design in a collision-dominated swarm proves to be
limited by the ability to effectively account for the full mechanical
interaction. There is a need for physical design rules and robust
1
Gulliver UMR CNRS 7083, ESPCI, PSL Research University, 75005 Paris, France.
2
Institut des Systèmes Intelligents et de Robotique, Sorbonne Université, CNRS,
ISIR, F-75005 Paris, France. 3School of Physics and Astronomy and Center for
Physics and Chemistry of Living Systems, Tel Aviv University, Tel Aviv 6997801,
Israel.
*Corresponding author. Email: matanbz@gmail.com
algorithms where robots not only maintain a high degree of auton-
omy in solitude but also retain functionality in a crowded
environment.
Unlike artificial robotic swarms, naturally occurring swarms
show excellent flexibility in both dilute and highly dense environ-
ments throughout the animal kingdom. At the cellular scale, an in-
dividual bacterium not only can explore space by performing a
simple run-and-tumble (29) but can also form a dense, highly
active bacterial colony that displays internal turbulent flow (30),
thereby mixing in nutrients (31, 32). On the millimeter scale, an in-
dividual larva can search for food through chemotaxis but self-or-
ganize into a living fountain when a large localized food source is
present (33). Macroscopic animals such as fish and birds can indi-
vidually nest and mate but can come together into giant schools and
flocks to evade predation (34, 35). Even pedestrian flow can emerge
from simple body collisions (36). Swarm cooperation is found on a
range of length scales, suggesting that only simple primitive behav-
iors are required and physical interactions can be used for the
swarm’s advantage (37). Self-organization in dense assemblies is
not a prerogative of living beings. It has been shown that collective
motion can emerge from independent mindless robots physically
interacting with one another, considering a stripped-down flavor
of swarm robotics as a particular kind of active matter with no ex-
plicit decision computed at the level of the robots (28, 38–42).
Our motivation is to expand the roboticists’ toolbox by taking
advantage of a generic mechanical response: the tendency to reori-
ent in response to an external force. We then demonstrate the role of
this morphological response in different robotic tasks both for an
individual robot and a robotic swarm. We introduce two designs
to show that this force orientation response can come in different
flavors, aligning with or against an external force. We show that this
force orientation response can also be used in a robotic swarm

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SCIENCE ROBOTICS | RESEARCH ARTICLE
platform, giving control over the outcome of a robot-robot collision
and the resulting collective behavior.
To achieve this, we augment robots with a flexible exoskeleton
and show that the morphological difference is encoded in a single
effective parameter, labeled κ, that controls the outcome of colli-
sions. We show two different designs that have similar dynamics
for free individual robots (i.e., a single robot moving on a
smooth, flat surface, without experiencing any external force nor
touching any obstacles or robots). However, when experiencing
an external force or collision, the two designs have an opposite re-
sponse. We show that the passive dynamics of the design allow
tuning of the robot’s response to align or oppose an external
force. Upon collision, a robot can push against or slide along an ob-
stacle or another robot, based solely on its mechanical design. The
emerging collective dynamics add a morphology layer to a robot’s
computation, facilitating a combined physical and logical swarm ar-
chitecture (43, 44). We quantitatively analyzed the response to an
external force and found empirically that it can be described by a
single effective parameter, κ. We show experimentally, theoretically,
and numerically that a change in the sign of κ inverts the response to
an external force, making a robot go down or up a force field. We
find κ to be a key design parameter in the response of the robots
when encountering an obstacle (be it a wall or another robot).
Using positive and negative κ makes two primitives that qualitative-
ly alter the swarm’s dynamics in a collision-dominated, crowded en-
vironment. We also show that κ can be used to guide the design of
collective transport.
We then add a computational layer, where, in parallel to the im-
plicit morphological computation, robots execute an explicit sense-
act cycle. We experimentally studied phototaxis in a partially lit
arena, setting a swarm of N = 64 robots to search for the lit
region. We set up the illumination field to be unoriented (coming
directly from above) and without local light gradients, thus ensuring
that robots cannot use local information to guide their way to the lit
region. Moreover, the lit region was designed to be too small to host
the whole swarm. This restriction forced the robots to physically in-
teract, turning the phototaxis task into an exercise of collective ag-
gregation with sterical interactions, going from dilute (in the dark)
to crowded (in the light). Studying swarms with positive and nega-
tive κ revealed a qualitative and a quantitative different collective
phototaxis, directly attributed to the nature of this effective
parameter.
Last, we show that the swarm can collectively learn to execute the
phototactic task when operating in an environment where individ-
ual robots transition between dilute and crowded regions. We im-
plemented a decentralized reinforcement learning algorithm
inspired by social learning (45–48) and found that, when collisions
were allowed, the swarm fluidized and effectively converged to a
successful phototactic strategy despite the sparse and intermittent
communication network. We further support our experimental ob-
servations with agent-based model simulations, finding that mor-
phology is central to collective behavior and becomes increasingly
important with growing swarm size. The interplay of morphology
and computation is graphically illustrated in Fig. 1, summarizing
the findings of our work.
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
RESULTS
Individual dynamics
The Morphobots
We made Morphobots by augmenting Kilobots (23) with a three-
dimensional (3D) printed exoskeleton. The Kilobot is a 3.4-cm-
tall, 3.3-cm-diameter unit, standing on three rigid legs made of
thin metallic rods. Kilobots are capable of differential drive locomo-
tion using two vibrators. When either is activated, a Kilobot turns at
about 45°/s, and when both are activated, the Kilobot moves
forward. The power source is a rechargeable lithium-ion battery
that can keep the Kilobot running for a few hours. Kilobots are
equipped with an infrared transmitter and receiver so that they
can communicate with each other. The transmitter of a robot
sends light toward the surface that reflects up to the receiver of
another nearby robot. Obtaining straight motion over long distanc-
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es is difficult and requires lengthy calibration of the individual vi-
brators (49); moreover, the calibration itself drifts over time.
To both improve the robot’s nominal speed and eliminate the
need for individual calibration, we encapsulated Kilobots within
exoskeletons that changed how collision and locomotion are per-
formed. We made two tripod-based designs (a pair of flexible legs
and an opposing stiff leg), with both designs having a circular
chassis (diameter d = 4.8 cm). The two designs, which we call
fronter versus aligner, differ in the fore-versus-aft positioning of
their flexible legs (see Fig. 2, A to C, and Materials and Methods).
The natural vibration of the semiflexible legs is tuned near reso-
nance with the vibration motors to maximize the coupling with
their drive. Using an anisotropic flexible leg design (1 mm thick,
6 mm wide, and 20 mm long; see the Supplementary Materials), vi-
brations are coupled predominately to the forward stick-slip motion
of the robots. The exoskeleton did not interfere with the ability of
the robots to communicate and had multiple roles, including energy
storage of vibrations, near-resonance coupling to motors, and an-
isotropic design to translate motor vibrations into a forward
motion. The exoskeleton allowed a Morphobot to operate at an
order of magnitude higher speed than Kilobots (see Fig. 2D and
movie S1). It allowed control of the response to external forces
and collisions.
By tracking the motion of individual Morphobots, we found that
aligners and fronters show near-identical motility. Motility is
defined by the magnitude of the displacement of an individual
robot over different time scales and is measured by tracking the
time evolution of their 2D coordinates, ! r ðtÞ ¼ ðxðtÞ; yðtÞÞ, when
running in the experimental arena (50). Only a low concentration
of robots was used (less than 1% filling fraction) excluding the effect
of collisions, and robots were tracked only when away from the walls
of the arena. The mean square displacement, MSD = 〈Δr 2(Δt)〉N, of
an ensemble of noisy walkers presents two dynamical regimes (51).
On short time intervals Δt, the displacement of the robots was
aligned with the direction of their individual orientation, and dis-
placements were proportional to Δt. The motion is said to be bal-
listic when robots go roughly on a straight line at a constant speed:
hΔr2 ðΔtÞiN ¼ v20 Δt2 , where v0 is the instantaneous speed.
On long time intervals, the noisepffiffiffiffiffireoriented the robots several
times, and displacement grew as Δt as robots explored the space
randomly. The motion is now said to be diffusive 〈Δr 2(Δt)〉N =
4D0Δt, where D0 is called the diffusion constant. The noisy trajec-
tories of the robots originate from the stochastic nature of their
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SCIENCE ROBOTICS | RESEARCH ARTICLE
Fig. 1. Article at a glance. Swarm robotics design space spans both morphology
and computation. Experiments with 3D-printed flexible exoskeletons show that
they can direct the outcome of collisions between robots. The mechanical
design of the exoskeleton, fronter versus aligner, controls whether a robot
ascends or descends a hill, which translates to pushing against or sliding along
a wall. The exoskeletons’ morphological computation extends the ability of the
swarm to execute collective tasks, including transport, phototaxis, and decentral-
ized learning.
vibrational drive (52) as well as the roughness of theparena
ffiffiffiffiffi and in-
ternal electronic drift. The two regimes (∝Δt and / Δt ) are sepa-
rated by a crossover, which takes place on a time scale called the
persistence time, τp (see below), to which corresponds a persistence
1=2
length hΔr2 ðτp ÞiN . Figure 2D displays the MSD measured for a set
of Morphobots equipped with the two types of the exoskeleton, to-
gether with their ensemble average (the MSDs of eight robots of
each design were measured, shown as separate curves in Fig. 2D;
see also Table 1). One sees that the difference in motility between
the two designs is no larger than the robot-to-robot variation within
each design. The observed ballistic nature of the MSD (/v20 Δt2 ) in-
dicates that the persistence length of the motion is larger than the
size of the circular arena (diameter D = 150 cm). Fitting the ensem-
ble MSD in the ballistic regime, we found that the mean nominal
speeds are 5.2 ± 0.9 and 4.8 ± 0.1 cm/s for the aligners and fronters,
respectively. The measured effective parameters are summarized
in Table 1.
Together, the anisotropic leg design allows us to use higher
motor drive while keeping the robot’s trajectories straight, alleviat-
ing the need for individual robot calibration, and resulting in a
nominal speed of v0 ≃ 5 cm/s, a 10-fold increase relative to the
Table 1. Measured nominal speed and alignment parameter of aligner
and fronter designs.
v0(d/s) κ(1/d)
Aligners 1.1 ± 17% 0.3 ± 10%
Fronters 1.0 ± 2% −0.3 ± 20%
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
bare Kilobot design (see movie S1). The notion of using passive
dynamic machines (53) allowed us to achieve a similar feat in
bristle bots, using rows of soft legs (54), and in the commercially
available HEXBUG (27, 28, 55, 56). Using the differential drive,
robots can also pivot at 45°/s. Programming alternated straight
and pivoting motions of random duration and random direction,
we can set the Morphobots into a run-and-tumble motion (57),
where, on average, the robots run straight during a run time
τrun = 2 s, with a velocity to be specified, and tumble during a mean
tumble duration τtumble = 4 s. The total duration of a run-and-
tumble sequence sets τRT = τrun + τtumble, with the resulting persis-
tent time being τP ≈ 18 s (see the Supplementary Materials).
Fronters versus aligners response to an external force
We systematically measured the response of the two Morphobots’
designs to an external body force by placing the robots on a plane
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tilted by a small angle, ϕ. The two designs have a different polar re-
sponse: Aligners go downhill, whereas fronters climb against gravity
(Fig. 3, A and B, and movies S1 to S6). The effect is consistent, re-
gardless of the initial orientation of the robot being along the pos-
itive y direction (downhill to its right) or negative y direction
(downhill to its left), excluding the effect of an internal left/right
bias (see movies S3 to S6). We systematically increased the slope
(see Fig. 3C) and found that with increasing ϕ, the trajectories
turn increasingly curved until the robot orients downhill (aligner)
or uphill (fronter).
To quantify the different responses of the robots to an external
body force, we modified a model introduced in the context of active
matter to describe self-aligning particles (55). A Morphobot has two
translational !r ¼ ðx; yÞ and one orientational n ^ ¼ ðcosðθÞ; sinðθÞÞ
degree of freedom. In the overdamped limit, the deterministic dy-
namics of the velocity ! v and orientation n ^ of a Morphobot with a
!
nominal speed v0 subjected to a body force f (see Fig. 3D) obey the
following equations
d!r !
¼! ^þμf
v ¼ v0 n ð1Þ
dt
d^n
¼ κð^n�! vÞ�n ^ ð2Þ
dt
where μ is the particle mobility and κ is the alignment parameter, an
effective parameter describing the reorientation by force. The first
equation simply states that the velocity is the additive combination
^ and the speed induced by the external
of the propulsion velocity v0 n
!
force μ f . The second equation contains the key ingredient of the
model. It describes the reorientation of the Morphobot along the
direction of its motion. The coupling between an external force
and the orientation is captured through an effective alignment pa-
rameter, κ. This parameter measures to what extent an external
!
force, f , affects the orientation of a robot, n ^. In the most simple
case, κ = 0, and an external force does not affect the orientation
of a robot (Eq. 2 becomes redundant). However, generically, force
and orientation do couple, and κ is finite (κ ≠ 0). Moreover, κ is
signed and can be positive or negative. This means that, when sub-
jected to an external force, the robot does not only translate but also
rotates. The larger the magnitude of the alignment parameter, ∥κ∥,
the quicker a robot’s orientation will respond to an external force.
There are two important types: When positive (κ > 0), a robot aligns
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Fig. 2. Augmenting Kilobots with exoskeletons makes Morphobots. (A) A Kilobot with a light sensor, infrared (IR) communication, and two differential drive vibration
motors is fitted into one of two 3D-printed exoskeleton designs: (i) fronter or (ii) aligner. Both fronter (B) and aligner (C) exoskeletons are tripods, but the restitution vector
^r of the aligners is parallel to the orientation vector n ^ for the fronters. (D) MSD versus time shows the ballistic response for both designs (∝ t 2), with a
^ but antiparallel to n
typical speed of 5.2 cm/s for the aligners and 4.8 cm/s for the fronters. Also shown is the MSD of a bare Kilobot, which, after calibration, can also move ballistically but at a
speed one order of magnitude slower (0.53 cm/s). Inset: 12-s-long trajectories of fronters (green) and aligners (cyan) and a 36-s-long trajectory of bare Kilobots (gray)
inside the 150-cm-diameter arena.

parallel with an external force. Here, we call robots with positive κ
aligners. When negative (κ < 0), a robot aligns antiparallel to an ex-
ternal force. Here, we call robots with negative κ fronters. κ is an
effective parameter and results from the design details of the
robot (material properties, mass distribution, drive mechanism,
and so on), and yet κ is empirically well defined because it can be
experimentally measured and quantitatively compared between dif-
ferent robotic platforms. For a constant force in the ^x direction
! namics repeat, effectively tracing loops (see Fig. 4B and movie S2).
f ¼ f ^x, with f = mgsinϕ, where m is the mass of the Morphobot
and g is the acceleration of gravity, the dynamical evolution of the
orientation θ = θ(t), relative to the x axis, obeys a closed-form equa-
tion
dθ
¼ κμf sinθ ð3Þ
dt
Equation 3 is the equation for an overdamped simple pendulum,
which can be solved directly to give tan θ2 ¼ e κμft [for initial condi-
tion θ(t = 0) = 90°]. In the limit where v0 ≪ μf, when the orientation
is expressed as a function of the arclength θ(s), it is found to be in-
dependent of the external force and mobility (see the Supplemen-
tary Materials)
κs
θðsÞ ¼ 2atanðe Þ ð4Þ
To find the alignment parameter κ, we tracked the orientation of
individual robots as they moved across a fixed inclined plane (see
Fig. 3E, movies S3 and S4, and Materials and Methods). In the
absence of an external force (zero slope, ϕ = 0°), both fronters
and aligners move in roughly straight lines. For steeper inclines
(ϕ = 3° and ϕ = 6°), both designs show a similar rate for the conver-
gence of the orientation. Fitting the evolution of the orientation to
Eq. 4, we extract the alignment parameter κ. The magnitude of κ for
both designs is similar but with opposite signs: −κfronter ≈ κaligner ≈
0.06 cm−1. The measured effective parameters are summarized
in Table 1.
We find that the design rules controlling the alignment param-
eter apply also in a dynamically changing environment and to
robotic platforms of different sizes (see Fig. 4). Placing a Morphobot
of either type on a seesaw gives rise to a unique behavior (Fig. 4): An
aligner turns quickly downhill and falls off the seesaw, whereas a
fronter turns upward and climbs. Once a Morphobot passes the
center of the seesaw, the mass of the robot suffices to tip the
plane of the swinging table. The fronter is now facing downhill,
yet, again, as it moves, it turns around and recommences climbing
until passing the center of the seesaw once more, where these dy-
Moreover, the design rules for the Morphobot generalize to a
custom robotic platform independent of Kilobots (Fig. 4D) and
can be made 10 times larger (see movies S5 and S6).
Interaction with a wall
Using the response of the Morphobots to an external force, we can
predict their steric interaction with stationary walls, movable
objects, and with one another. Morphological interaction with a
boundary (stationary or movable) can be seen as a series of colli-
sions, at each of which the robot experiences an external force
(see Fig. 5A). With each collision, a Morphobot slightly reorients.
An aligner orientation approaches the normal force from the wall
(but leaves when parallel to the wall). A fronter turns against the
normal force (and toward the wall). This is confirmed by the
direct observation of Morphobots sent to move toward a wall of sta-
tionary robots (see Fig. 5B). Arranging turned-off robots into three
tight rows forms an effective wall with which a moving robot me-
chanically interacts. On average, both designs spend similar time
near the wall (∼3τp ≈ 1 min), but aligners travel a path along the
wall that is twice as long as that of the fronters (see Fig. 5C and
movie S1). Using a multiagent Brownian dynamics simulation of
active soft discs, we were able to predict the outcome of collisions
on the basis of alignment strength parameter κ. During a collision,
an agent with κ > 0 (aligner) turns toward the normal force and
away from the obstacle, whereas an agent with κ < 0 (fronter)
turns against the normal force and into the obstacle, thus pushing
it. In addition, simulating agents with κ < 0 near a passive particle
shows that the agents collectively move the passive object (see

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Fig. 3. The orientation response of aligners and fronters to an external force is equal and opposite. (A) Sketch of the experimental setup for measuring the dy-
namical response on an inclined plane (see movies S1 and S3 to S6). (B) Force diagram of a robot placed on an inclined plane. For small slopes (ϕ ≤ 6°), robots experience
an external body force. (C) Trajectories of aligners (cyan) and fronters (green) moving on different slopes (ϕ). Aligners and fronters have an opposite response to the
external force: Aligners’ orientation turns downward as they quickly descend, whereas fronters turn upward and slowly ascend. Scale bar, 20 cm. (D) Throughout its
!
motion, the orientation of the robot n ^ forms an angle θ with the external force f , together setting the net velocity !
v (see Eq. 1). (E) Tracking the orientations shows that
aligners turn downhill (θ → 0), aligning with the external force, whereas fronters turn uphill (θ → π), fronting the external force (see also movies S3 to S6). Solid curves are
fits to Eq. 4, from which κ can be extracted (see the Supplementary Materials). (F) Average fitted κ for each incline angle and robot design (error bars are the SE). For finite
tilt angle, the response of the two designs is about equal and opposite κaligner = −κfronter ≈ 0.3/d. At a slope near zero, the response is dominated by the robot’s internal
bias [left/right curving and internal noise; (52)] but averages to zero.

Fig. 5D). This has been confirmed experimentally using a group of
fronters that work together to push a circular object (see Fig. 5E and
movie S1). Previous work showed that collective transport can be
implemented using global control (58). Here, we extend this
notion and show that a single effective alignment parameter, κ, cap-
tures the embodied dynamics of sterically interacting agents, reduc-
ing the required computing resources for simulating a large swarm
in a crowded environment.
Collective dynamics
We turned to study the ability of a swarm of Morphobots to execute
a collective task, focusing on phototaxis, where robots explicitly use
their sensory and computational power in addition to the morpho-
logical interaction. Phototaxis is a common behavioral trait found
in nature where an organism changes its motion in response to
locally measured light. Phototaxis pervades the animal kingdom
from individual bacteria to groups of vertebrates. Phototaxis is clas-
sified as either positive, where the organism seeks to be present in
the light [such as moths; (59)], or negative, where the organism
prefers the dark [such as golden shiners; (60)]. Here, we tested
the ability of a robotic swarm to perform this widespread natural
behavior, with an emphasis on the collective ability to maximize
the time spent in the light (positive phototaxis). We programmed
the robots to change their speed in response to the locally measured
light intensity [also known as photokinesis; (61)] and show that the
robots’ morphology can be used to improve their collective photo-
taxis when crowded. Stimulus-induced locomotory movements (or
taxis) constitute an important building block in swarm robotics (2,
3, 19, 62–64), and several bioinspired algorithmic implementations
have been proposed for phototaxis using ambient light sensors (18,
65–68).
Collective phototaxis in the presence of steric interactions
The experimental setup for collective phototaxis was designed to
prevent an individual robot from accessing global information
that could guide its sense-act cycle. For this, we set a swarm
(N = 64 bots) of either aligner (κ > 0) or fronter (κ < 0) to explore a
circular arena (diameter D = 150 cm) with a lit spot that covers only
σ = 6% of the area of the arena (diameter b = 36 cm; see Fig. 6A and
the “Phototaxis arena” section in the Supplementary Materials). The
light is unoriented (coming from above), and there are no local light
intensity gradients to guide a robot to the lit region (the light field is
essentially flat). This excludes the use of traditional phototactic al-
gorithms such as steepest descent search (25, 69) or reorientation
(23). The lit region cannot accommodate more than 80% of the
robots (about 50 Morphobots), challenging the swarm to transition
between sparse and crowded.
As a first step, we consider ad hoc calibrated swarms, where the
Morphobots’ policies are defined as follows. Each Morphobot con-
tinuously monitors local light intensity: When in the dark (below a
predefined intensity threshold, Pth), the robot performs a run-and-
tumble motion, with a running velocity V0. When entering a bright
region (light intensity above Pth), the robot stops, V1 = 0. At early
times, when there are only a few robots in the lit region, the two
designs show similar performance as measured by the fraction of
the swarm that is in the light
N1
F; ð5Þ
N
where N1 is the number of bots taxied in the light. At the beginning
of every experiment, robots are placed in the arena with random

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SCIENCE ROBOTICS | RESEARCH ARTICLE
Fig. 4. Fronters and aligners respond differently to a dynamic environment.
(A) A Morphobot on a swinging table setup for testing morphological response to
a dynamically changing environment. (B) A fronter continually turns uphill,
moving in a circular motion while staying on the swinging table (see also movie
S2), whereas an aligner (C) quickly turns downhill and falls off. Top: Schematics.
Bottom: Measured trajectories. (D) The design rules apply also to custom-made
robots (not Kilobot-based), producing the expected force alignment response
with Morphobots as much as 10 times larger than a Kilobot (see movies S5 and
S6). Scale bars, 10 cm.
positions and orientations. At early times, ℱ is quantitatively cap-
tured as a diffusion-limited process (see Fig. 6F), with the kinetic
reaction constant found using Smoluchowski first passage compu-
tation for diffusive particles with an effective diffusion constant, Deff
≈ 4.22 cm/s (see the Supplementary Materials) (70). After about
10 min, both swarms deposit the first layer of robots at the perimeter
of the lit region (Fig. 6, B and D). From this point onward, the pho-
totaxis rate slows down and deviates from the simple diffusion-
limited reaction, because crowding of taxiing robots at the perime-
ter of the light spot physically prevents additional robots from en-
tering the lit region (Fig. 6, C and E). This steric effect is
quantitatively more pronounced for the aligner design, as can be
seen for the overall performance given by the fraction of the
swarm that has entered the lit region over long times (Fig. 6F).
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
The reason is that the collective success of each swarm depends
on the morphological interactions of a robot impinging from the
dark region on the first layer of robots at the perimeter of the lit
region. Individual Morphobots arriving at the wall of stationary
robots show qualitatively different dynamics for either design: An
aligner (κ > 0) that arrives at the wall tends to reorient along the
wall until it finally leaves, whereas a fronter Morphobot (κ < 0)
tends to “dig in,” pushing already taxied robots into the lit region
until being taxied itself (see Fig. 6, B and D). These individual
events add up: Fronting Morphobots progressively push the perim-
eter inward, constantly filling the bulk of the lit region with robots.
By contrast, the aligning Morphobots keep reorienting away from
the robots at the perimeter, leaving a hollow in the lit region (see
Fig. 6, C and E, and movie S1). This discrepancy exacerbates as
the swarm size increases. By simulating up to 8192 phototactic
Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
agents that follow Eqs. 1 and 2, we find that a swarm of aligners
(κ > 0) becomes considerably slower to phototaxis relative to an
equivalent swarm of fronters (κ < 0). The typical time, T1/2, to
have half of the swarm in the light (ℱ = 1/2) grows much faster
for aligners than for fronters (see Fig. 6, G and H). The above find-
ings demonstrate that the collective ability of the swarm to perform
a phototactic task at scale hinges on a subtle morpho-functional
difference.
Distributed learning of a phototactic strategy
The swarms considered thus far had the environmental information
hard-coded into individual robots before deployment—the sense-
act cycle of every robot had a preprogrammed ad hoc knowledge
of the correct light threshold for phototaxis (which is an unrealistic
assumption when robots are to be deployed in an unknown envi-
ronment). Concurrent challenges, however, require deployment of
robotic systems in unknown terrains (2, 3, 62, 63), including extra-
terrestrial exploration, underwater surveying, and search-and-
rescue missions (15, 71, 72). Under such circumstances, robots
will lack an a priori model of the environment that can be preprog-
rammed. This requires an added layer to the robots’ controller for
online tuning of the sense-act cycle based on continually acquired
knowledge. Under such conditions, a distributed robotic system
offers robustness with scale, improving both exploration and surviv-
al. A distributed system, however, also poses a challenge because the
swarm has to learn collectively.
In this section, we tested whether a swarm can collectively learn
even when robots collide. We show that, in a swarm of Morphobots,
both execution and learning can be made fully distributed (see
Fig. 7). We then proceeded to test the effect of the two designs on
the collective performance and found numerically that over a long
enough time, a swarm of fronters outperforms a swarm of aligners.
Because the robots are allowed to collide, the swarm is fluidized
(73). An individual collision event may be stochastic, but the
mean data flow within the swarm becomes predictable. Unlike a
solid or a jammed state (where individuals do not move), here,
the swarm acted as a fluid, meaning that it satisfied good mixing,
because robots frequently exchanged neighbors and visited
various parts of the arena, sustaining a flow of information (see
figs. S4 and S5). Each robot evaluated its performance and com-
pared it with other encountered robots. The selection algorithm
can be made distributed by implementing a decentralized online re-
inforcement learning scheme to make the swarm adaptive.
Previous work showed that a swarm can learn a phototactic strat-
egy with a policy defined by the weights of an artificial neural
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Fig. 5. Tuning interaction with a static and a moving obstacle using morphological computation. The morphological interaction of the Morphobots with walls can
be explained using their response to an external force. When a Morphobot collides with a wall, each impingement can be seen as a momentary normal force. (A) Aligners
turn to align their orientation (black arrow) with the normal force (red arrowheads). Once aligned along the wall, there is no further rotation because there are no more

Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
collisions, and the robot slides along. (B) When a fronter collides with a wall, its orientation (black) turns antiparallel with the normal force (red), and the robot continues to
push into the wall. When programmed to run-and-tumble, aligners and fronters interact differently with a wall of stationary robots, where aligners (A) cover a much larger
distance along the wall relative to fronters (B). (C) Multiple collision experiments show that aligners and fronters spend similar time near a wall (3τp ≈ 1 min), yet aligners
move twice as fast. (D) Brownian dynamics simulations predict that active particles with a negative alignment parameter, κ < 0 (fronters), push a movable object. (E) With
real robots, placing a round movable obstacle shows that a swarm of fronters is capable of transporting an object. Scale bar, 10 cm.

Fig. 6. The collective ability of a robotic swarm to phototaxis is constrained by steric interactions. (A) A swarm of phototactic robots is placed in an arena patterned
with an unoriented light field. Because there are no local gradients to guide a robot to the light source, robots perform a simple run-and-tumble in the dark and stand in
the light. (B) When a fronter arrives at the light spot, it collides with robots already taxied at the perimeter but progressively pushes its way in. Repeating this process
results in the complete coverage of the light spot (C). When an aligner arrives at the light spot (D), it slides along the wall of robots and leaves. This leads to only partial
filling of the light spot (E), leaving an empty void in the center. Scale bars, 20 cm. (F) The resulting fraction of swarm in the light as a function of time shows that at early
times (τ ≈ 30τp ≈ 10 min), both designs show equal performance, quantitatively consistent with a diffusion-limited reaction (dashed line). At later times, fronters out-
perform aligners (average of four realizations with error bars as SD). (G) Simulating swarms with more than two orders of magnitude more agents shows a sharp slow-
down in the phototaxis of aligners with swarm size. (H) The typical time for half of the swarm to enter the light T1/2 grows rapidly for an increasingly larger swarm of
aligners while having only a small effect on a swarm of fronters of similar size.

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SCIENCE ROBOTICS | RESEARCH ARTICLE

Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
Fig. 7. Distributed online reinforcement learning algorithm and deployment of a swarm of Morphobots. (A) The embedded algorithm for updating the phototactic
policy of each robot is composed of two main moduli: a sense-act cycle and a learning cycle. (B) The sense-act cycle is encoded into a perceptron: External stimuli are
weighted, summed, and evaluated through a sigmoid function, the output of which is binarized to select an action. The weights w define the policy π of the robot. (C) The
learning cycle is based on the conditional diffusion of the policies across the population, biased by a reward function ρ evaluated by each robot: When robot 2 receives a
message containing the policy control parameters {wi}1 and the reward ρ1 of robot 1, robot 2 will inherit the new policy if it is associated with a higher reward ρ1 > ρ2. (D)
Illustration of a centralized communication network where all agents (gray circles) are connected to a central operator (red circle). (E) An illustration of a sparsely con-
nected network where agents communicate only when in close proximity. (F) Illustrations of a sparse and intermittent communication where a robot (black curve) already
in the light source (F1) departs (F2) and broadcasts within communication range (yellow circle) its reward and policy to a robot in the dark (red curve) (F3), which inherits
the new policy (F4). Note that the broadcasting robot may have a wrong policy, effectively spreading false information, with neither robot knowing what is the best policy
for phototaxis. (G) A swarm of Morphobots with randomly initialized light thresholds collectively learns its environment, increasing the swarm fraction in the light. Results
are the average of four runs with error bars as the SD. Insets show that fronters (top) and aligners (bottom) develop a different spatial arrangement in the highlighted
region. (H) Evolution of the swarm fraction in the light for multiagent simulations of swarms of different sizes learning to phototaxis using the distributed online rein-
forcement learning. At earlier times, swarms of fronters and aligners develop the spatial arrangement found in the experiments. At later times (inaccessible experimen-
tally), the swarm of fronters compactly arranged inside the light spot, with a larger fraction of the swarm in the light compared with the arrangement of the swarm of
aligners (see also insets).

network as a controller, π = {wi} (74–76). There, the learning was
implemented using an evolutionary algorithm to act as a direct
policy search, using a reinforcement learning method, incremental-
ly refining behavioral policies by directly tuning the policies’ param-
eters (77, 78). The swarm is randomly initialized with a pool of
sense-act policies, and in an iterative selection process, only the
best-performing policies are carried over to the next generation.
In these algorithms, the execution is distributed over the swarm,
but it is a central computer performing selection and optimization
(see Fig. 7D). Such schemes are impossible to implement outside
the laboratory if the swarm has no access to an external operator
for guidance or a global communication channel to compare per-
formances. In what follows, we show that, despite having a sparse
and intermittent network, communication in a swarm of Morpho-
bots became effectively connected across time thanks to the swarm’s
fluidity, facilitating decentralized learning (see Fig. 7F).
For a successful reinforcement learning algorithm to converge, a
suitable reward function, ρ, must be defined (79). As an input, the
reward function receives the robot’s sensory information ρ = ρ({ii})
and outputs a scalar score (see Fig. 7A and algorithm S1 in the Sup-
plementary Materials). The combination of an individual robot
sense-act policy, π = π({wi}), and the environment that it sampled
results in a robot’s reward at a given time. Robots can also commu-
nicate when within communication range and exchange their pol-
icies and rewards (Fig. 7, C and F). After such an exchange, a robot
can choose to adopt the received policy based on its associated
reward, operating under the assumption that a higher reward is
the result of a superior policy. Unlike classical evolutionary algo-
rithms that use a centralized computer to compare performance,
the scheme presented here allows embedding of both the evaluation
and the selection within the robot. In other words, robots collective-
ly performed a kind of social learning, where successful individual
innovations diffuse over the population (45, 46, 48). Note that the
assumption that a higher reward is a result of a superior policy may
not be always true. From time to time, robots with an incidental fa-
vorable history can have a disproportionally higher reward. Such
bots will propagate a poor policy globally to the swarm, effectively
spreading “fake news.” This local-to-global gap challenges the

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SCIENCE ROBOTICS | RESEARCH ARTICLE
chosen reward function to be robust to such fluctuations. The fluid
nature of the swarm of Morphobots allows us to choose ρ that over-
comes this culprit. Most swarm robotic systems use a collision-
avoidance algorithm that slows down their collective dynamics,
jamming the information flow and creating deadlocks (8). By con-
trast, the Morphobots are allowed to collide, sustaining good mixing
and offering a quantitative link between global and local informa-
tion (see figs. S4 and S5).
To formulate a proper reward function, we note that the collec-
tive phototaxis, ℱ, as defined in Eq. 5 is an ensemble average, 〈〉N,
counting the fraction of the swarm that is in the light at a given time.
ℱ cannot be directly optimized by an individual robot because it
stores global information. However, in a system with good
mixing, a time average can approximate an ensemble average
ϕ ≡ 〈〉T ≈ 〈〉N. A time average equivalent to the ensemble average
given in Eq. 5 is the mean time that an individual robot spent in
the light, Φ ≡ Tin light/T ≈ ℱ. In a distributed robotic system with
good mixing, a reward function that estimates the mean time that
the robot spent in the light can serve as a proxy for the global per-
formance of a swarm of its kind, ρ ∝ Φ ≈ ℱ. Robot i can evaluate the
mean time that it spent in the light through a temporal average of its
light sensor signal
1MX1
ρi ¼ Pi ð6Þ
M k¼0 k
where M is the number of averaged measurements and Pik is the kth
measurement taken at constant time intervals. Because the time that
a robot spends in the light is inversely proportional to its speed in
the light, Φ ∝ ρ ∝ 1/V1, the above choice of reward function is an-
alytically linked to a given policy. This link allows us to show that,
when formulated as a dynamical process, collective learning will
converge on average (see the Supplementary Materials for detailed
derivation).
To test the validity of the proposed decentralized learning
scheme, we deployed a swarm of Morphobots in the same light-pat-
terned arena but this time with randomly initialized policies (see
Fig. 7 and movie S1). A “correct” policy corresponds to a set of
weights {wi} of the perception where the robot slows down in the
light. A “wrong” policy corresponds to a set of weights where the
robot does not slow down in the light. When randomly distributed
in the large arena used (150 cm), robots are on average more than 16
cm apart (see, for example, 01:41 in movie S1). This is more than
twice the maximum communication distance supported by the
Kilobot platform (7 cm). At such a separation, many of the robots
are out of communication range from one another, making the
network sparse. Not only is the network sparse, but the network
is also intermittent. Even when two robots do come within commu-
nication range, they quickly move away. On average, the subgroup
of the swarm that interacts at a given time makes up only a small
fraction of the swarm, and this subgroup is constantly changing, to-
gether making a sparse and intermittent communication network.
We also programmed the robots to start with random initial policies
by drawing their weights at equal probability from values ranging
between 0 and 255 (see Materials and Methods and the Supplemen-
tary Materials for details). At very early times, only a few robots
slowed down in the light—most robots either stood in the dark or
moved quickly through the lit region, yet the swarm fraction in the
light progressively increased with time. We tested swarms of both
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
aligners and fronters and found a consistent increase in the fraction
of the swarm in the light reaching about ℱ ≈ 0.4. At these swarm
fractions in the light, the difference between fronters and aligners is
not expected to be substantial as is evident here (compare with
Fig. 6F at early times). The learning swarms had eight times more
robots in the light than the case of random placement that is the
fraction of the lit region, σ = 0.06. Given that Kilobots are 10
times slower and that diffusion scales as the speed square (/v20 ;
see the Supplementary Materials), a swarm of bare Kilobots
would have taken 100 times longer to converge, making collective
learning impossible. The fact that the adaptive swarm performed
almost but not quite as well as the precalibrated swarm is not unex-
pected and may be attributed to the fact that dynamics of about half
of the randomly initialized population are very slow (because they
stand in the dark), and given enough time, they will also eventually
Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
learn. This may be resolved by expanding the learning space of the
sense-act cycle to have the robots also learn the speed in the
light, V1.
We found that the difference between learning swarms of align-
ers and fronters increased with time. By extending the multiagent
numerical engine, we tested the learning process over a longer du-
ration (inaccessible experimentally) and for larger swarms (up to
eight times larger; see Fig. 7H and fig. S3). Implementing algorithm
S1 numerically, we found that, after about τ ≈ 300 to 400 from their
initialization, swarms of either aligners or fronters reached the same
fraction of robots in the light (ℱ ≈ 0.4; Fig. 7G), consistent with
Morphobots studied in the laboratory. In both experiments and
simulations, we found that, at these intermediate times, the
swarm of fronters tended to occupy the lit region, whereas aligners
remained on the perimeter (see Fig. 7, G and H). Simulations show
that, at later times (τ > 400), this qualitative difference grew: The
fraction of aligners in the light remained the same, whereas the
swarm of fronters continued to crowd the lit region, reaching
ℱ ≈ 0.55. This effect is consistent with growing swarm size,
showing the significance of morphological interaction at scale in
adaptive swarms.
CONCLUSION
Here, we introduced a robotic swarm that can operate in both dilute
and dense settings, where collisions between robots occur frequent-
ly. We show that the robots’ physical morphology played an impor-
tant role in terms of collective behaviors, where a seemingly small
change in the shape led to very different outcomes. Applying tools
from statistical physics of active matter to swarm robotics, we show
that behavior found in natural swarms [such as ant colonies; (80)],
including collective self-aggregation and transport, can be attained
with limited use of software-controlled behaviors by capitalizing on
the morphological properties of robots, demonstrating morpholog-
ical computation at work within the swarm.
We also showed that distributed online evolutionary reinforce-
ment learning can be implemented in a dense robot swarm to
exploit or mitigate physical contingencies after deployment in the
open. We found that this kind of artificial social learning, when
robots locally exchanged information about their behavioral strate-
gies, can be modeled as a conditional diffusion process.
Future collective learning implementations in swarm robotics
can go beyond the diffusion of behavioral strategies. An extreme
case is of sparse swarms, where individual robots or groups may
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be temporarily disconnected from the whole (81). Although dense
and sparse robot swarms may be considered separately, a robot
swarm in the open will likely have to switch from one configuration
to another, requiring versatile social learning capabilities. A natural
extension can be motivated by cultural evolution found in nature
(82), which is built on reformulation and combining previously
learned behaviors to continuously discover behaviors of growing
complexity.
Future physical models of a learning swarm should go beyond
the mean-field approximation used here and account for the
spatial density fluctuations [as is done for describing the motility-
induced phase separation observed in active fluids; (57, 83)]. From
that point of view, our work shows that, building upon the recent
theoretical progress made in describing the population of active
particles, one can envision a rapid development of a statistical me-
chanics description of smart active matter for future swarm
engineering.
MATERIALS AND METHODS
Exoskeleton design and manufacturing
Exoskeleton manufacturing
Exoskeletons were 3D-printed using one of the following 3D print-
ers and materials to give similar results: Stratasys Objet350 Connex3
printer using VeroClear (modulus of elasticity E = 1 to 2 GPa and
density ρ = 1.2 g/cm3), 3D Systems ProJet 2500 plus printer using
VisiJet M2R-CL (MJP) (density ρ = 1.14 g/cm3 and elastic modulus
E ≈ 1 GPa), and Prusa i3 printer using polylactic acid (PLA)
(density ρ = 1.2 g/cm3 and elastic modulus E = 5 GPa).
Data acquisition and experimental setups
Phototaxis experiments
Phototaxis experiments were performed on a 5-mm translucent
plexiglass sheet placed on the floor (PMMA 27000 Diffusant
Laiteux 40%, WEBER METAUX). A circular region (6% of the
arena) was lit using a projector (EPSON EB-1795F), at red green
blue (RGB) = (10, 255, 255), at about the spectral sensitivity peak
(570 nm) of the Kilobot’s light sensor (TEPT5700, Vishay Semicon-
ductors). To reduce interference of ambient light with the light
sensor, the arena was lit using four desk lamps directed at the sur-
rounding walls (for a homogeneous diffusive illumination) and
covered with red cellophane (where the robot’s light sensor’s sensi-
tivity is at a minimum). Red cellophane also covered the camera
used for acquisition (PIXELINK.PL-D734MU) to reduce the satu-
ration from the lit region and imaging of the robots. Image acqui-
sition was carried out using μicromanager (84), at 2 frames/s.
Robots were counted when fully in the light, and results shown in
Figs. 6F and 7G are the average of four realizations with error bars as
the SD.
Code used
Robots were programmed to phototaxis using variations of the fol-
lowing two codes: (i) phototaxisAdhoc.c: Robots stand when their
photoperceptron is activated given a predefined threshold; (ii) pho-
totaxisLearning.c: Robots stand when photoperceptron is activated,
but light thresholds are randomly initialized and evolve through the
learning algorithm.
Inclined plane experiments
Coupling between external force and robot’s orientation was mea-
sured by letting robots run on a plane made of 5-mm-thick
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
plexiglass placed at an incline and imaged using a SonyAlphaS7
camera, with video-rate image acquisition at 30 frames/s. See
movies S3 to S6.
Data analysis
Raw images were preprocessed using ImageJ (85) or FFmpeg, fol-
lowed by particle locating using trackpy package (50) or a custom
code, and trajectories were linked using trackpy (50).
Simulating Morphobots
Computer simulations were carried out by adding a stochastic term
to Eqs. 1 and 2 and using fifth-order Runge-Kutta for the time prop-
agation of Brownian dynamics of active soft discs by adopting code
from previous publications (86–88).
Custom Morphobots
Design files and bill of materials for custom Morphobots (Fig. 4) are
found in the extended data. At their core, custom Morphobots are
Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
made from a 3D-printed flat chassis onto which a battery (LIR
2477), an electric switch, and two vibration motors (BestTong
14kRPM 3 V) are mounted. The chassis has two holes for mounting
flexible curved legs (which can be 3D-printed separately).
Supplementary Materials
This PDF file includes:
Supplementary Text
Figs. S1 to S5
Algorithm S1
References (89–95)
Other Supplementary Material for this
manuscript includes the following:
Movies S1 to S6
Exoskeletons CAD files
REFERENCES AND NOTES
1. G. Beni, J. Wang, Swarm intelligence in cellular robotics systems, in Robots and Biological
Systems: Towards a New Bionics? NATO ASI Series, P. Dario, G. Sandini, P. Aebischer, Eds.
(Springer, 1993), vol. 102, pp. 703–712.
2. M. Brambilla, E. Ferrante, M. Birattari, M. Dorigo, Swarm robotics: A review from the swarm
engineering perspective. Swarm Intell. 7, 1–41 (2013).
3. H. Hamann, Swarm Robotics: A Formal Approach (Springer International Publishing, 2018).
4. R. A. Brooks, A. M. Flynn, “Fast, cheap and out of control” (Massachusetts Inst of Tech
Cambridge Artificial Intelligence Lab, 1989).
5. M. Dorigo, G. Théraulaz, V. Trianni, Reflections on the future of swarm robotics. Sci. Rob. 5,
abe4385 (2020).
6. D. Floreano, H. Lipson, From individual robots to robot societies. Sci. Rob. 6,
eabk2787 (2021).
7. M. Dorigo, G. Theraulaz, V. Trianni, Swarm robotics: Past, present, and future. Proc. IEEE
109, 1152–1165 (2021).
8. J. Werfel, K. Petersen, R. Nagpal, Designing collective behavior in a termite-inspired robot
construction team. Science 343, 754–758 (2014).
9. S. Hauert, J.-C. Zufferey, D. Floreano, Evolved swarming without positioning information:
An application in aerial communication relay. Auton. Robots 26, 21–32 (2008).
10. C. Virágh, G. Vásárhelyi, N. Tarcai, T. Szörényi, G.⍰. Somorjai, T. Nepusz, T. Vicsek, Flocking
algorithm for autonomous flying robots. Bioinspir. Biomim. 9, 025012 (2014).
11. G. Vásárhelyi, C. Virágh, G. Somorjai, T. Nepusz, A. E. Eiben, T. Vicsek, Optimized flocking of
autonomous drones in confined environments. Sci. Robot. 3, eaat3536 (2018).
12. K. McGuire, C. De Wagter, K. Tuyls, H. Kappen, G. C. de Croon, Minimal navigation solution
for a swarm of tiny flying robots to explore an unknown environment. Sci. Robot. 4,
eaaw9710 (2019).
13. J. Gomes, M. Duarte, P. Mariano, A. L. Christensen, in International Conference on Parallel
Problem Solving from Nature (Springer, 2016), pp. 591–601.
14. P. Zahadat, T. Schmickl, Division of labor in a swarm of autonomous underwater robots by
improved partitioning social inhibition. Adaptive Behavior 24, 87–101 (2016).
10 of 12
SCIENCE ROBOTICS | RESEARCH ARTICLE
15. F. Berlinger, M. Gauci, R. Nagpal, Implicit coordination for 3D underwater collective be-
haviors in a fish-inspired robot swarm. Sci. Robot. 6, eabd8668 (2021).
16. M. Schranz, M. Umlauft, M. Sende, W. Elmenreich, Swarm robotic behaviors and current
applications. Front. Robot. AI. 7, 36 (2020).
17. D. Saldaña, B. Gabrich, M. Whitzer, A. Prook, M. F. M. Campos, M. Yim, V. Kumar, A decen-
tralized algorithm for assembling structures with modular robots, in 2017 IEEE/RSJ Inter-
national Conference on Intelligent Robots and Systems (IROS), Vancouver, BC, Canada,
September 2017, pp. 2736–2743.
18. S. Li, R. Batra, D. Brown, H.-D. Chang, N. Ranganathan, C. Hoberman, D. Rus, H. Lipson,
Particle robotics based on statistical mechanics of loosely coupled components. Nature
567, 361–365 (2019).
19. W. Savoie, T. A. Berrueta, Z. Jackson, A. Pervan, R. Warkentin, S. Li, T. D. Murphey,
K. Wiesenfeld, D. I. Goldman, A robot made of robots: Emergent transport and control of a
smarticle ensemble. Sci. Robot. 4, eaax4316 (2019).
20. X. Du, H. Cui, T. Xu, C. Huang, Y. Wang, Q. Zhao, Y. Xu, X. Wu, Reconfiguration, camouflage,
and color-shifting for bioinspired adaptive hydrogel-based millirobots, camouflage, and
color-shifting for bioinspired adaptive hydrogel-based millirobots. Adv. Funct. Mater. 30,
1909202 (2020).
21. S. Li, B. Dutta, S. Cannon, J. J. Daymude, R. Avinery, E. Aydin, A. W. Richa, D. I. Goldman,
D. Randal, Programming active cohesive granular matter with mechanically induced phase
changes. Sci. Adv. 7, eabe8494 (2021).
22. Y. Ozkan-Aydin, D. I. Goldman, M. Saad Bhamla, Collective dynamics in entangled worm
and robot blobs. Proc. Nat. Acad. Sci. U.S.A. 118, e2010542118 (2021).
23. M. Rubenstein, A. Cornejo, R. Nagpal, Robotics. Programmable self-assembly in a thou-
sand-robot swarm. Science 345, 795–799 (2014).
24. I. Slavkov, D. Carrillo-Zapata, N. Carranza, X. Diego, F. Jansson, J. Kaandorp, S. Hauert,
J. Sharpe, Morphogenesis in robot swarms. Sci. Robot. 3, eaau9178 (2018).
25. G. Wang, T. V. Phan, S. Li, M. Wombacher, J. Qu, Y. Peng, G. Chen, D. I. Goldman, S. A. Levin,
R. H. Austin, L. Liu, Emergent field-driven robot swarm states. Phys. Rev. Lett. 126,
108002 (2021).
26. G. Oliveri, L. C. van Laake, C. Carissimo, C. Miette, J. T. B. Overvelde, Continuous learning of
emergent behavior in robotic matter. Proc. Natl. Acad. Sci. U.S.A. 118, e2017015118 (2021).
27. A. Deblais, T. Barois, T. Guerin, P. H. Delville, R. Vaudaine, J. S. Lintuvuori, J. F. Boudet,
J. C. Baret, H. Kellay, Boundaries control collective dynamics of inertial self-propelled
robots. Phys. Rev. Lett. 120, 188002 (2018).
28. J. F. Boudet, J. Lintuvuori, C. Lacouture, T. Barois, A. Deblais, K. Xie, S. Cassagnere, B. Tregon,
D. B. Brückner, J. C. Baret, H. Kellay, From collections of independent, mindless robots to
flexible, mobile, and directional superstructures. Sci. Robot. 6, eabd0272 (2021).
29. H. C. Berg, Motile behavior of bacteria. Phys. Today 53, 24–29 (2000).
30. H. H. Wensink, J. Dunkel, S. Heidenreich, K. Drescher, R. E. Goldstein, H. Löwen,
J. M. Yeomans, Meso-scale turbulence in living fluids. Proc. Natl. Acad. Sci. U.S.A. 109,
14308–14313 (2012).
31. H. Xu, J. Dauparas, D. Das, E. Lauga, Y. Wu, Self-organization of swimmers drives long-range
fluid transport in bacterial colonies. Nat. Commun. 10, 1792 (2019).
32. J. W. Costerton, P. S. Stewart, E. P. Greenberg, Bacterial biofilms: A common cause of
persistent infections. Science 284, 1318–1322 (1999).
33. O. Shishkov, M. Hu, C. Johnson, D. L. Hu, Black soldier fly larvae feed by forming a fountain
around food. J. R. Soc. Interface. 16, 20180735 (2019).
34. A. Procaccini, A. Orlandi, A. Cavagna, I. Giardina, F. Zoratto, D. Santucci, F. Chiarotti,
C. K. Hemelrijk, E. Alleva, G. Parisi, C. Carere, Propagating waves in starling, Sturnus vulgaris,
flocks under predation. Anim. Behav. 82, 759–765 (2011).
35. BBC, Feeding frenzy, from the hunt: Hunger at sea (oceans) (2015).
36. M. Moussaïd, D. Helbing, G. Theraulaz, How simple rules determine pedestrian behavior
and crowd disasters. Proc. Natl. Acad. Sci. U.S.A. 108, 6884–6888 (2011).
37. C. Bechinger, R. di Leonardo, H. Löwen, C. Reichhardt, G. Volpe, G. Volpe, Active particles in
complex and crowded environments. Rev. Mod. Phys. 88, 045006 (2016).
38. F. Hara, R. Pfeifer, Eds., Morpho-Functional Machines: The New Species (Springer, 2003).
39. J. Deseigne, O. Dauchot, H. Chaté, Collective motion of vibrated polar disks. Phys. Rev. Lett.
105, 098001 (2010).
40. T. Vicsek, A. Zafeiris, Collective motion. Phys. Rep. 517, 71–140 (2012).
41. K.-D. N. T. Lam, M. Schindler, O. Dauchot, Self-propelled hard disks: Implicit alignment and
transition to collective motion. New J. Phys. 17, 113056 (2015).
42. A. Bricard, J.-B. Caussin, N. Desreumaux, O. Dauchot, D. Bartolo, Emergence of macroscopic
directed motion in populations of motile colloids. Nature 503, 95–98 (2013).
43. R. Pfeifer, J. Bongard, in How the Body Shapes the Way We Think: A New View of Intelligence
(MIT Press, 2006), pp. 418.
44. R. Pfeifer, G. Gómez, in Creating Brain-Like Intelligence (Springer, 2009), pp. 66–83.
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
45. R. A. Watson, S. G. Ficici, J. B. Pollack, Embodied evolution: Distributing an evolutionary
algorithm in a population of robots. Rob. Auton. Syst. 39, 1–18 (2002).
46. N. Bredeche, E. Haasdijk, A. Prieto, Embodied evolution in collective robotics: A review.
Front. Robot. AI 5, 12 (2018).
47. N. Fontbonne, O. Dauchot, N. Bredeche, in 2020 IEEE Congress on Evolutionary Computation
(CEC) (2020), pp. 1–8.
48. N. Bredeche, N. Fontbonne, Social learning in swarm robotics. Philos. Trans. R. Soc. B 377,
20200309 (2022).
49. M. Rubenstein, C. Ahler, N. Hoff, A. Cabrera, R. Nagpal, Kilobot: A low cost robot with
scalable operations designed for collective behaviors. Rob. Auton. Syst. 62,
966–975 (2014).
50. D. B. Allan, T. Caswell, N. C. Keim, C. M. van de Wel, Soft-matter/trackpy: Trackpy v0.4.2
(v0.4.2). Zenodo, 10.5281/zenodo.3492186 (2019).
51. É. Fodor, M. Cristina Marchetti, The statistical physics of active matter: From self-catalytic
colloids to living cells. Phys. A. Stat. Mech. Appl. 504, 106–120 (2018).
52. C. Scholz, S. D’Silva, T. Pöschel, Ratcheting and tumbling motion of Vibrots. New J. Phys. 18,
123001 (2016).
Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
53. S. Collins, A. Ruina, R. Tedrake, M. Wisse, Efficient bipedal robots based on passive-dynamic
walkers. Science 307, 1082–1085 (2005).
54. L. Giomi, N. Hawley-Weld, L. Mahadevan, Swarming, swirling and stasis in sequestered
bristle-bots. Proc. R. Soc. A. Math. Phys. Eng. Sci. 469, (2013).
55. O. Dauchot, V. Démery, Dynamics of a self-propelled particle in a harmonic trap. Phys. Rev.
Lett. 122, 068002 (2019).
56. P. Baconnier, D. Shohat, C. H. López, C. Coulais, V. Démery, G. Düring, O. Dauchot, Selective
and collective actuation in active solids. Nat. Phys. 18, 1234–1239 (2022).
57. J. Tailleur, M. E. Cates, Statistical mechanics of interacting run-and-tumble bacteria. Phys.
Rev. Lett. 100, 218103 (2008).
58. S. Shahrokhi, A. T. Becker, Object manipulation and position control using a swarm with
global inputs, in IEEE International Conference on Automation Science and Engineering
(CASE), Fort Worth, TX, USA, November 2016, pp. 561–566.
59. D. Lees, A. Zilli, Moths: A Complete Guide to Biology and Behavior (Smithsonian Books, 2019).
60. A. Berdahl, C. J. Torney, C. C. Ioannou, J. J. Faria, I. D. Couzin, Emergent sensing of complex
environments by mobile animal groups. Science 339, 574–576 (2013).
61. W. NULTSCH, Phototactic and photokinetic action spectra of the diatom nitzschia com-
munis. Photochem. Photobiol. 14, 705–712 (1971).
62. E. Şahin, in International Workshop on Swarm Robotics (Springer, 2004), pp. 10–20.
63. L. Bayindir, A review of swarm robotics tasks. Neurocomputing 172, 292–321 (2016).
64. W. Savoie, S. Cannon, J. J. Daymude, R. Warkentin, S. Li, A. W. Richa, D. Randall,
D. I. Goldman, Phototactic supersmarticles. Artif. Life Robot. 23, 459–468 (2018).
65. H. Hamann, H. Worn, K. Crailsheim, T. Schmickl, in 2008 IEEE/RSJ International Conference on
Intelligent Robots and Systems (IEEE, 2008), pp. 1415–1420.
66. T. Schmickl, R. Thenius, C. Moeslinger, G. Radspieler, S. Kernbach, M. Szymanski,
K. Crailsheim, Get in touch: Cooperative decision making based on robot-to-robot colli-
sions. Auton. Agents Multi-Agent Syst. 18, 133–155 (2009).
67. S. Kernbach, R. Thenius, O. Kernbach, T. Schmickl, Re-embodiment of honeybee aggre-
gation behavior in an artificial micro-robotic system. Adapt. Behav. 17, 237–259 (2009).
68. F. Arvin, K. Samsudin, A. R. Ramli, M. Bekravi, Imitation of honeybee aggregation with
collective behavior of swarm robots. Int. J. Comput. Intell. Syst. 4, 739–748 (2011).
69. T. Schmickl, H. Hamann, BEECLUST: A swarm algorithm derived from honeybees, in Bio-
Inspired Computing and Communication Networks (CRC Press, ed. 1, 2011), pp. 95–137.
70. M. V. Smoluchowski, Drei Vortrage uber Diffusion, Brownsche Bewegung und Koagulation
von Kolloidteilchen. Physik. Zeit. 17, 557–585 (1916).
71. K. Uckert, A. Parness, N. Chanover, E. J. Eshelman, N. Abcouwer, J. Nash, R. Detry, C. Fuller,
D. Voelz, R. Hull, D. Flannery, R. Bhartia, K. S. Manatt, W. J. Abbey, P. Boston, Investigating
habitability with an integrated rock-climbing robot and astrobiology instrument suite.
Astrobiology 20, 1427–1449 (2020).
72. I. Fiorello, E. Del Dottore, F. Tramacere, B. Mazzolai, Taking inspiration from climbing plants:
Methodologies and benchmarks—A review. Bioinspir. Biomim. 15, 031001 (2020).
73. R. Solé, M. Moses, S. Forrest, Liquid brains, solid brains. Philos. Trans. R. Soc. B: Biol. Sci. 374,
20190040 (2019).
74. V. Trianni, R. Groß, T. H. Labella, E. Şahin, M. Dorigo, Evolving aggregation behaviors in a
swarm of robots, in Advances in Artificial Life ECAL 2003, W. Banzhaf, J. Ziegler, T. Christaller,
P. Dittrich, J. T. Kim, Eds. (Springer, Berlin, Heidelberg, 2003), pp. 865–874.
75. M. Dorigo, V. Trianni, E. Şahin, R. Groß, T. H. Labella, G. Baldassarre, S. Nolfi,
J. L. Deneubourg, F. Mondada, D. Floreano, L. M. Gambardella, Evolving self-organizing
behaviors for a swarm-bot. Auton. Robots 17, 223–245 (2004).
76. O. Soysal, E. Bahçeci, E. Şahin, Aggregation in swarm robotic systems: Evolution and
probabilistic control. Turk. J. Elec. Engin. 15, 199–225 (2007).
11 of 12
SCIENCE ROBOTICS | RESEARCH ARTICLE
77. S. Nolfi, D. Floreano, Evolutionary robotics: The biology, intelligence, and technology of
self-organizing machines, in Intelligent Robotics and Autonomous Agents Series (The MIT
Press, 2000), p. 332.
78. S. Doncieux, N. Bredeche, J.-B. Mouret, A. Eiben, Evolutionary robotics: What, why, and
where to. Front. Robot. AI 2, 4 (2015).
79. L. P. Kaelbling, M. L. Littman, A. W. Moore, Reinforcement learning: A survey. J. Artif. Intell.
Res. 4, 237–285 (1996).
80. O. Feinerman, I. Pinkoviezky, A. Gelblum, E. Fonio, N. S. Gov, The physics of cooperative
transport in groups of ants. Nat. Phys. 14, 683–693 (2018).
81. D. Tarapore, R. Groß, K.-P. Zauner, Sparse robot swarms: Moving swarms to real-world
applications. Front. Robot. AI 7, 83 (2020).
82. A. Whiten, The burgeoning reach of animal culture. Science 372, eabe6514 (2021).
83. M. Y. Ben Zion, Y. Caba, A. Modin, P. M. Chaikin, Cooperation in a fluid swarm of fuel-free
micro-swimmers. Nat. Commun. 13, 184 (2022).
84. A. D. Edelstein, M. A. Tsuchida, N. Amodaj, H. Pinkard, R. D. Vale, N. Stuurman, Advanced
methods of microscope control using μManager software. J. Biol. Methods 1, e10 (2014).
85. C. T. Rueden, J. Schindelin, M. C. Hiner, B. E. DeZonia, A. E. Walter, E. T. Arena, K. W. Eliceiri,
ImageJ2: ImageJ for the next generation of scientific image data. BMC Bioinformatics 18,
529 (2017).
86. N. Oppenheimer, D. B. Stein, M. J. Shelley, Rotating membrane inclusions crystallize
through hydrodynamic and steric interactions. Phys. Rev. Lett. 123, 148101 (2019).
87. N. Oppenheimer, D. B. Stein, M. Y. Ben Zion, M. J. Shelley, Hyperuniformity and phase
enrichment in vortex and rotor assemblies. Nat. Commun. 13, 804 (2022).
88. M. Y. Ben Zion, A. Modin, P. M. Chaikin, Hydrodynamic spin-orbit coupling in asynchronous
optically driven micro-rotors. arXiv:2203.11051 [cond-mat.soft] (2022).
89. M. Y. Ben Zion, X. He, C. C. Maass, R. Sha, N. C. Seeman, P. M. Chaikin, Self-assembled three-
dimensional chiral colloidal architecture. Science 358, 633–636 (2017).
90. G. Zhu, M. Hannel, R. Sha, F. Zhou, M. Y. Ben Zion, Y. Zhang, K. Bishop, D. Grier, N. Seeman,
P. Chaikin, Microchemomechanical devices using DNA hybridization. Proc. Natl. Acad. Sci.
U.S.A. 118, e2023508118 (2021).
Ben Zion et al., Sci. Robot. 8, eabo6140 (2023) 22 February 2023
91. J. R. Howse, R. A. L. Jones, A. J. Ryan, T. Gough, R. Vafabakhsh, R. Golestanian, Self-motile
colloidal particles: From directed propulsion to random walk. Phys. Rev. Lett. 99,
048102 (2007).
92. M. Y. Ben Zion, Y. Caba, R. Sha, N. C. Seeman, P. M. Chaikin, Mix and match—A versatile
equilibrium approach for hybrid colloidal synthesis. Soft Matter 16, 4358–4365 (2020).
93. M. Yoones, M. Y. Ben Zion, O. Dauchot, N. Bredeche, in GECCO 2022 - Proceedings of the 2022
Genetic and Evolutionary Computation Conference (Association for Computing Machinery,
2022), pp. 104–112.
94. M. Y. Ben Zion, J. Fersula, N. Bredeche, O. Dauchot, Distributed on-line reinforcement
learning in a swarm of sterically interacting robots. https://doi.org/10.48550/arXiv.2111.
06953 (2021).
95. M. J. Schnitzer, Theory of continuum random walks and application to chemotaxis. Phys.
Rev. E 48, 2553–2568 (1993).
Acknowledgments: We thank Y. Lahini and N. Oppenheimer. Funding: This work was
supported by the MSR project funded by the Agence Nationale pour la Recherche under grant
no. ANR-18-CE33-0006. M.Y.B.Z. acknowledges support from the Ministry of Aliyah and
Immigrant Absorption of Israel. Author contributions: O.D., N.B., and M.Y.B.Z. conceived the
Downloaded from https://www.science.org at Chinese University of Hong Kong on March 29, 2023
research. M.Y.B.Z. designed the experiments, theoretical models, and analysis. M.Y.B.Z. and J.F.
performed the experiments and data analysis. M.Y.B.Z., O.D., and N.B. wrote the manuscript.
Competing interests: The authors declare that they have no competing interests. Data and
materials availability: All data needed to support the conclusions of this manuscript are
included in the main text or Supplementary Materials. Code is available at the following DOI: 10.
5281/zenodo.7573761.
Submitted 18 March 2022
Accepted 30 January 2023
Published 22 February 2023
10.1126/scirobotics.abo6140
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 Morphological computation and decentralized learning in a swarm of sterically
interacting robots
Matan Yah Ben Zion, Jeremy Fersula, Nicolas Bredeche, and Olivier Dauchot

Sci. Robot., 8 (75), eabo6140.

DOI: 10.1126/scirobotics.abo6140

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