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Collective Motion
Collective Motion
Collective Motion
1
Department of Biological Physics, E
otvos University - P
azmany Peter stny. 1A, Budapest, Hungary
H-1117
2
Statistical and Biological Physics Research Group of HAS - P
azmany Peter stny. 1A, Budapest, Hungary
H-1117
3
Department of Mathematics, National University of Athens - Panepistimioupolis 15784 Athens, Greece
Abstract
We review the observations and the basic laws describing the essential aspects of collective motion being
one of the most common and spectacular manifestation of coordinated behavior. Our aim is to provide
a balanced discussion of the various facets of this highly multidisciplinary field, including experiments,
mathematical methods and models for simulations, so that readers with a variety of background could get
both the basics and a broader, more detailed picture of the field. The observations we report on include
systems consisting of units ranging from macromolecules through metallic rods and robots to groups of
animals and people. Some emphasis is put on models that are simple and realistic enough to reproduce
the numerous related observations and are useful for developing concepts for a better understanding of
the complexity of systems consisting of many simultaneously moving entities. As such, these models allow
the establishing of a few fundamental principles of flocking. In particular, it is demonstrated, that in
spite of considerable differences, a number of deep analogies exist between equilibrium statistical physics
systems and those made of self-propelled (in most cases living) units. In both cases only a few well
defined macroscopic/collective states occur and the transitions between these states follow a similar scenario,
involving discontinuity and algebraic divergences.
2 1 INTRODUCTION
1.4 Goals to be achieved give a somewhat more detailed list of features char-
acterizing flocking in general. Thus we assume that a
The approach of treating flocks, or even crowds, as system exhibiting collective motion is made of units
systems of particles naturally leads to the idea of ap-
plying the successful methods of statistical physics, that are rather similar
such as computer simulations or theories on scaling,
to the detailed description of the collective behavior moving with a nearly constant absolute velocity
of organisms. Naturally, for better progress, obser- and are capable of changing their direction
vations/experiments and modeling have to be inti-
interacting within a specific interaction range by
mately related. Indeed, over the past few decades, an
changing their direction of motion, in a way in-
increasingly growing number of significant attempts
volving an effective alignment
have been made to both observe and describe flocking
as well as modeling (simulation) the most conspicu- which are subject to a noise of a varying magni-
ous features of the observed natural systems ranging tude
from molecules to groups of mammals.
It would be quite an achievement if we could es-
tablish a systematic chart of the types of collective
2.1 Principles and concepts
motion, since many times understanding is achieved In a general sense, phase transition is a process, dur-
through classification. There are reasons and argu- ing which a system, consisting of a huge number of
ments for thinking that the same patterns of collec- interacting particles, undergoes a transition from one
tive motion apply to the collection of molecules up phase to another, as a function of one or more exter-
to groups of humans. There must be some still to nal parameters. The most familiar phases in which
be discovered laws of such systems from which the a physical system can be, are the solid, liquid and
above observation follows. gaseous phases, and the best known example for a
phase transition is the freezing of a fluid when the
temperature drops. In this case the temperature is
2 Basics of the statistical me- the external or control parameter.
chanics of flocking Phase transitions are defined by the change of one
or more specific system variables, called order param-
Throughout this overview the notion of flocking is eters. This name, order parameter, comes from the
used as a synonym of any kind of coherent motion observation that phase transitions usually involve an
of individual units. However, the notion of coher- abrupt change in a symmetry property of the system.
ent motion needs some further elaboration since, as For example, in the solid state of matter, the atoms
it turns out, it can be manifested in a number of have a well-defined average position on the sites of an
specific ways. In any case, coherent or ordered mo- ordered crystal lattice, whereas positions in the liq-
tion is assumed to be a counterpart of disordered, uid and gaseous phases are disordered and random.
random motion. In the various models of flocking it Accordingly, the order parameter refers to the degree
emerges through a kind of transition (from disorder of symmetry that characterizes a phase. Mathemat-
to order) as a function of the relevant parameter(s) of ically, this value is usually zero in one phase (in the
the models. To demonstrate more clearly this aspect disordered phase) and non-zero in the other (which is
of collective motion the best approach is to adopt a the ordered phase). In the case of collective motion
few related definitions motivated mainly by statisti- the most naturally (but not necessarily) chosen order
cal mechanics or statistical physics (the physics of parameter is the average normalized velocity ,
many interacting molecules).
N
However, before we turn to the discussion of the 1 X
= ~vi , (1)
N v
statistical mechanics aspects of collective motion we 0 i=1
6 2 BASICS OF THE STATISTICAL MECHANICS OF FLOCKING
where N is the total number of the units and v0 is that the investigations on the various universal be-
the average absolute velocity of the units in the sys- haviors that can be associated with non-equilibrium
tem. If the motion is disordered, the velocities of the systems have by now grown into a sub-discipline with
individual units point in random directions and av- its own language and formalism including specific
erage out to give a small magnitude vector, whereas processes the related research concentrates on.
for ordered motion the velocities all add up to a vec- In particular, most of the representative reviews
tor of absolute velocity close to N v0 (thus the order
on non-equilibrium phase transitions (see, e.g., Odor
parameter for large N can vary from about zero to [2004] or Henkel et al. [2009], Non-Equilibrium Phase
about 1). Transitions: Volume 1: Absorbing Phase Transi-
If the order parameter changes discontinuously tions) are centered on such features as i) absorbing
during the phase transition, we talk about a first or- states of reaction-diffusion-type systems, ii) mapping
der transition. For example, waters volume changes to the universal behavior of interface growth mod-
like this (discontinuously) when it freezes to ice. In els, iii) dynamical scaling in far-from-equilibrium re-
contrast, during second order (or continuous) phase laxation behavior and ageing, iv) extension to non-
transition the order parameter changes continuously, equilibrium systems by using directed percolation as
while its derivative, with respect to the control pa- the main paradigm of absorbing phase transitions.
rameter, is discontinuous. Second order phase tran- On the other hand, there are some specific features
sitions are always accompanied by large fluctuations of collective motion such as giant number fluctuations
of some of the relevant quantities. (GNF, see below) or various unusual transitions (e.g.,
During a phase transition a spontaneous symme- to jamming) which obviously do not occur in systems
try breaking takes place, i.e., the symmetry of the at equilibrium, but we still have to consider them in
system changes as we pass the critical value of the this review. On balance, we find that the language
control parameter (e.g., temperature, pressure, etc.). and the spectrum of the various aspects of equilib-
In the case of one of the simplest representations of rium phase transitions is surprisingly suitable for in-
a continuous phase transition (the Ising model), the terpreting most of the observed phenomena in the
spins are placed on a lattice, interact with their clos- context of collective motion; and this is an important
est neighbors and can point up or down. For high point we intend to make.
temperatures the spins point in random directions A further important aspect of the phenomena tak-
(with a probability 1/2 either up or down), while ing place during collective motion is that in contrast
for low temperatures (way below the critical point) with the standard assumptions of statistical mechan-
most of them point in the same direction (which is ics, the number of units involved in the collective be-
either up or down), selected spontaneously. This is havior typically ranges from a few dozens to a few
an example for the up and down symmetry (high thousands (in rare cases tens of thousands). On the
temperatures) breaking during the transition (one of other hand, for example, phase transitions in the
the directions becomes dominant). If there are in- framework of statistical mechanics are truly mean-
finitely many possible directions (continuous symme- ingful only for very large system sizes (consisting a
try), during a transition a single preferred direction number of units approaching infinity). Quantities like
can still emerge spontaneously. critical exponents cannot be properly interpreted for
Phase transitions can occur in both equilibrium flocks of even moderate sizes. However, a simple tran-
and non-equilibrium systems. In the context of col- sition from a disordered to an ordered state can take
lective motion although it is a truly non-equilibrium place even in cases when the number of units is in
phenomenon there are reasons for the preference the range of a few dozens. Most of the real-life ob-
of the possible analogies with the equilibrium phase servations and the experiments involve this so called
transitions rather than with those studied in the mesoscopic scale. The states (e.g. rotation) and
framework common in the interpretation of non- the transitions (e.g., from random to ordered motion)
equilibrium phenomena. One important reason is we later describe can be associated with the phenom-
2.2 Definitions and expressions 7
We end this Section with a brief discussion of two of the main points in this review is that the kinds of
characteristic phenomena which occur in systems of systems and the types of collective motion patterns
self-propelled particles but have no direct analogy in have a greater variety than originally thought of. Be-
processes occurring in equilibrium. Firstly, we point low we give a naturally incomplete list of systems
out that in a confined geometry any group of persis- in which collective motion has been observed (with
tently moving units having a finite size have a chance only some of the representative references included):
to be trapped or jammed. The presence of walls
or even a too narrow exit would inevitably lead Non-living systems: nematic fluids, shaken
to this phenomenon (see, e.g., [Kudrolli et al., 2008, metallic rods, nano swimmers, simple robots,
boats, etc. [Kudrolli, 2010, Ibele et al., 2009,
Helbing et al., 2000]).
Suematsu et al., 2010, Narayan et al., 2007]
Another interesting feature is a very specific form
of density changes called as giant number fluctua- Macromolecules [Schaller et al., 2010,
tions or GNF.This expression stands for the follow- Butt et al., 2010]
ing property of a system of self-propelled units: The
fluctuation of the number of units in an increasing Bacteria colonies [Czirok et al., 1996, 2001,
area of the system scale with the number of units (N) Sokolov et al., 2007, Cisneros et al., 2007]
in this area linearly. In addition, these fluctuations
Amoeba [Kessler and Levine, 1993, Nagano,
relax anomalously slowly. This is in sharp contrast
1998, Rappel et al., 1999]
with a theoretical result valid for equilibrium systems
according to which (along with the law of large num- Cells [Szab
o et al., 2006, Friedl and Gilmour,
bers) the fluctuations in the number of units grow as 2009, Arboleda-Estudillo et al., 2010,
a square root of N . This interesting feature was first Belmonte et al., 2008]
pointed out in Narayan et al. [2007] for a system of
shaken elongated rods and subsequently commented Insects [Buhl et al., 2006, Couzin and Franks,
upon by Aranson et al. [2008] showing that inelastic 2003]
collisions can lead to GNF even for spherical parti- Fish [Hemelrijk and Kunz, 2005, Parrish et al.,
cles. 2002, Becco et al., 2006, Ward et al., 2008]
Since shaking introduces an average velocity and
nematic or inelastic collisions involve a tendency for Birds [Heppner, 1997, Ballerini et al., 2008,
the particles to align, all these findings are expected Hayakawa, 2010, Bajec and Heppner, 2009]
to occur (and will be later discussed) in a variety of
Mammals [Fischhoff et al., 2007,
systems with collective motion.
Sueur and Petit, 2008, King et al., 2008]
Humans [Faria et al., 2010b, Helbing et al.,
3 Observations and experi- 1997, 2000, Moussad et al., 2011]
ments Although throughout the present review we pri-
It seems that collective motion (or flocking: these two marily classified the observations and experiments
notions will be used synonymously in this review al- based on the organizational complexity of the units
though in principle there are some subtle differences constituting the systems, there are various other as-
in their meanings) is displayed by almost every living pects as well, by which valid and meaningful catego-
system consisting of at least dozens of units. 2 One rizations can be made, such as:
2 In such small systems, we associate with flocking a state The patterns the units form (coherently moving
of the group in which the units assume an approximately com- clusters, mills, stripes, etc)
mon direction (or orientation) developing through local com-
munications among the entities. From an energetic viewpoint:
3.1 Data collection techniques 11
The way it is introduced to the system (uni- particles. These particles are assumed to follow
formly, at the boundaries, at special points, the flow dynamics accurately, and it is their motion
etc.) that is then used to calculate velocity information.
if the particles can preserve it (animals) Cisneros et al. [2007] applied this method in order to
or they dissipate it almost immediately evaluate the velocity filed of thousands of swimming
(shaken rods) bacteria. The task of tracing cells share many similar-
ities with the challenge of tracing the motion of bac-
The way the units interact with each other. This teria. Czirok et al. [1998] used computer controlled
can be phase contrast video microscope system in order to
physical, chemical (chemotaxis), visual or follow the collective motion of cells. The trajecto-
medium-mediated ries were recorded for several days to determine the
velocities of each cell of the types.
isotropic or anisotropic
The movement of vertebrate flocks has been
polar or apolar tracked mainly by camera-based techniques. Here,
short or long-range (regarding its temporal the observed animals are bigger, but the space in
characteristics) which they move is often unconfined. The simpler
through metric or topological distance case is when the group to be observed moves only
in two dimensions. In the 70-es Sinclair [1977] used
aerial photos to investigate the individuals spatial
3.1 Data collection techniques
positions within grazing African buffalo herds. Ex-
The main challenge during the implementation of ob- actly because of the difficulties of analyzing three
servations and/or experiments on collective motion, dimensional group motions, Becco et al. [2006] con-
is to keep a record on the individual trajectories of fined the motion of fish to two dimensions by putting
the group-members. This can easily turn out to be a them into a container which was basically two di-
difficult task, since (i) most of the colonies or groups mensional (in the sense that it was very shallow):
being investigated consist of many members which 40cm X 30cm X 2cm. This arrangement was con-
(ii) usually look very much alike (iii) and are often venient to track fish with a single video recorder. A
moving fast just think of a flock of starlings. The homogeneous light source was placed above the con-
applied technology is chosen by considering two fac- tainer and a CCD camera recorded the fish from be-
tors: (i) the size of the moving units, and (ii) the size low. (Obviously, the usage of the container confined
and dimension of the space in which the group can the area of motion as well.)
move. Both parameters range through many scales: In order to reconstruct the three-dimensional posi-
bacteria and cells are subjects of such experiments as tions and orientations of the fish, Cullen et al. [1965]
well as African buffalos or whales. Also, the area in used the so called shadow method. With this tech-
which the observation is carried out can range from nology, Partridge et al. [1980] investigated position-
a Petri dish [Keller and Segel, 1971] to the Georges ing behavior in fish groups of up to 30 individuals.
Bank (a region separating the Gulf of Maine from the Parrish and Turchin [1997] recorded the trajectories
Atlantic Ocean, [Makris et al., 2009]). Accordingly, of fish in three dimensions with three orthogonally
there is a variety of technologies that have been ap- positioned video cameras.
plied during the last decades. Major and Dill. [1978] were the first to apply the
The method called Particle Image Velocimetry, stereo photography technique in order to record the
(PIV) [Raffel et al., 2002] is used to visualize the three dimensional positions of birds within flocks
motion of small particles moving in a well-confined of European starlings and dunlins. By using the
area. Originally it is an optical technique used to pro- same technique, recently Ballerini et al. [2008] recon-
duce the two dimensional instantaneous velocity vec- structed the three-dimensional positions of hundreds
tor field of fluids, by seeding the media with tracer of starlings in airborne flocks with high precision.
12 3 OBSERVATIONS AND EXPERIMENTS
The stereo photography method allowed the detailed lective motion in deionized water under UV illumi-
and accurate analysis of nearest neighbor distances nation. The autonomous motion these particles ex-
in large flocks, but still did not make the trajectory hibit under the above circumstances (deionized wa-
reconstruction of the individual flock members possi- ter and UV light) is due to their asymmetric photo-
ble. decomposition, and the spatial self-organization is
As technology advances, newer and newer methods due to the ions which are secreted by the AgCl par-
and ideas show up with the purpose of studying col- ticles as they move.
lective motion. Here we mention two of these: The re- Various experiments on non-living self propelled
cently developed OWARIS (Acoustic Waveguide Re- particles (SPPs) possessing diverse features advo-
mote Sensing) exploits the wave propagation prop- cate that the shape and symmetry of the SPPs
erties of the ocean environment [Makris et al., 2006], play an important role in their collective dynam-
and makes the instantaneous imaging and continu- ics, and that large-scale inhomogeneity and coher-
ous monitoring of fish populations possible, covering ent motion can appear in a system in which parti-
thousands of square kilometers, that is, an area tens cles do not communicate except by contact. Sym-
of thousands to millions of times greater than that metric (or apolar) rods on vibrating surfaces have
of conventional methods. The other new method is been observed to form nematic order and under cer-
based on the usage of the Global Positioning Sys- tain conditions found to exhibit persistent swirling
tem, commonly known as GPS. The idea is to put as well [Narayan et al., 2006, Galanis et al., 2006].
small GPS devices on moving animals by which the Narayan et al. [2007] have also investigated symmet-
problem of trajectory-recording is basically solved. ric macroscopic rods and have found giant number
With this method the trajectory of flock members fluctuations lasting long, decaying only as a logarith-
can be collected with high temporal resolution in mic function of time (see Fig. 6). This finding is
their natural environment. The limits of this tech- in obtrusive contrast with the expected behaviour,
nique at this moment are on the one hand the grow- since the the central limit theorem predicts number
ing cost of the research with the growing number of fluctuations proportional to the square root of the
tracked flock members, and on the other hand the particle number, in the homogeneous ordered phase
limited accuracy of the devices. Biro et al. [2006] of equilibrium systems, away from the transition. By
and Nagy et al. [2010] analyzed GPS logged flight conducting complementary experiments with spheri-
tracks of homing pigeon pairs in order to investigate cal particles, Aranson et al. [2008] demonstrated that
hierarchical leadership relations inside the group and the giant number fluctuation phenomenon can arise
DellAriccia et al. [2008] used this method to study either from dynamic inelastic clustering or from per-
the homing efficiency of a pigeon group consist of 6 sistent density inhomogenity as well.
birds. Periodically, vertically vibrated granular rods form
vortex patterns [Blair et al., 2003]. Above a critical
3.2 Physical, chemical and biomolec- packing fraction, the ordered domains consisting of
nearly vertical rods spontaneously form and coex-
ular systems
ist with horizontal rods (see Fig. 4). The vortices
Along with the accumulating observations and exper- nucleate and grow as a function of time. Exper-
iments clarified the recognition, that flocking collec- iments performed in an annulus with a single row
tive motion emerges not only in systems consisting of rods revealed that the rod motion was generated
of living beings, but also among interacting physical when these objects were inclined from the vertical,
objects, based on mere physical interactions without and was always in the direction of the inclination (see
communication. A very simple system has been de- Fig. 5). The relationship between the covered area
scribed by Ibele et al. [2009] who reported about sim- fraction and the diffusion properties in the case of
ple autonomous micromotors, which are micrometer- self-propelled rods was also studied [Kudrolli, 2010].
sized silver chloride (AgCl) particles exhibiting col- Kudrolli et al. [2008] have made experiments with
3.2 Physical, chemical and biomolecular systems 13
ever, under certain hostile environmental conditions ticity field. By simultaneously measuring the posi-
(like limited nutrient source or hard agar surface) the tions, velocities and orientations of around one thou-
complexity of the colony as a whole increases, charac- sand bacteria, Zhang et al. [2010] demonstrated that
terized by the appearance of cell-differentiation and under specific conditions, colonies of wild-type Bacil-
long-range information transmission [Shapiro, 1988]. lus subtilis exhibit giant number fluctuations. More
For describing the observed hydrodynamics (vortex- specifically, they found that the number of bacteria
formation, migration of clusters) of the bacteria in per unit area shows fluctuations far larger than those
an intermediate level, Czirok et al. [1996] proposed for populations in thermal equilibrium.
a simpler model of self-propelled particles, and more The collective behavior of motile aerobic bacte-
complex ones taking into account further biologi- ria (aerobic are those bacteria which need the
cal details to capture the more elaborated collective presence of oxygen for their survival), primarily in
behaviors, like the vortex and colony formation (see high cell-concentration, is governed by the inter-
Sec. 5.1.3). Figure 10 depicts a typical bacterium play between buoyancy, oxygen consumption, mix-
colony formed by the vortex morphotype. ing and hydrodynamic interactions. The pattern-
This kind of bacterium, Bacillus subtilis, when formation of these bacteria is often governed by an-
the cells are very concentrated (nearly close-packed), other physical mechanism as well, called bioconvec-
forms a special kind of collective phase called Zoom- tion [Pedley and Kessler, 1992], which appears on
ing BioNematics (ZBN) [Cisneros et al., 2007]. This fluid medium having a surface open to the air. The
phase is characterized by large scale orientational co- authors argued that the patterns appear because bac-
herence, analogous to the molecular alignment of ne- teria, which are denser than the fluid they swim
matic liquid crystals, in which the cells assemble to- in, gather at the surface creating a heavy layer on
gether into co-directionally swimming clusters, which the top of a lighter medium. When the density of
often move at speeds larger then the average speed the bacteria-cells exceed a certain threshold, this ar-
of single bacteria. Figure 11 shows a snapshot of rangement becomes unstable resulting in a large-scale
swimming Bacillus subtilis cells exhibiting collective cell circulation (or convection).
dynamics, and Fig. 12 depicts the corresponding vor- Sokolov et al. [2009] have investigated the onset of
3.3 Bacterial colonies 17
gating the collective motion of the cells Dictyostelium neurons to the olfactory bulb (RMS) and neural crest
discoideum (commonly referred to as slime mold ). (NC) cells migrating from the developing neural tube
In order to describe the dynamics of these cells, to many distant locations in the embryos of mam-
Rappel et al. [1999] took into account their shape mals; sperm cells. Although the collective motion
and plasticity. Based on the observations regard- of these cells are often guided by chemical signals, in
ing how these amoeba cells aggregate into rotat- some cases they can also form patterns based on mere
ing pancake-form structures, the authors built a hydrodynamic effects [Riedel et al., 2005].
model of the dynamics of self-propelled deformable ii) Other migrating groups are more tightly asso-
objects (see also in Sec. 5.1.2). According to ciated and the cells normally never dissociate. Ex-
the experiments, these cells tended to form round amples are the fish lateral line, structures perform-
structures which rotated around the center clock- ing branching and sprouting morphogenesis such as
wise or counterclockwise (depending on some ini- trachea or the vasculature and finally moving sheets
tial conditions) often persisting for tens of hours. of cells in morphogenesis or wound healing. These
Using the same genus, Dictyostelium discoideum, groups have an additional feature, in that the mov-
McCann et al. [2010] conducted experiments in order ing structure has an inherent polarity, a free front
to elucidate the role of signal relay a process during and an attached back.
which the individual cells amplify chemotactic signals iii) Drosophila border cells are a group or clus-
by secreting additional attractants upon stimulation ter of cells performing a directional movement dur-
in the collective motion of these cells. They found ing oogenesis. These migrating cells are associ-
that this process enhances the recruitment range, but ated tightly but the cluster is free, without an in-
does not effect the speed or directionality of the cells. herent back. A particularly nice visualization of
Next we discuss the collective motion of cells in the collectively moving cells during the develop-
highly structured, multi-cellular organisms, in which ment of zebra fish (by three dimensional tracing
cell migration plays a major role in both embry- of live-stained cell nuclei) very well demonstrates
onic development (e.g. gastrulation, neural crest the relevance of collective motion during morpho-
migration) and the normal physiological or patho- genesis [Schoetz, 2008]. Lecaudey et al. [2008] in-
physiological responses of adults (e.g. wound heal- vestigate the mechanism that organize cells behind
ing, immune response or cancer metastasis). In these the leading edge. In particular, they study the
mechanisms cells have to be both motile and adhere role of the fibroblast growth factors (Fgfs) a sig-
to one other. In order to describe these features, naling method known to regulate many types of
Szab o et al. [2010] expanded the cellular Potts model developmental processes [Affolter and Weijer, 2005,
by including active cell motility, and studied a cor- Ciruna and Rossant, 2001] during the formation of
responding computer simulations as well, which was sensory organs in zebrafish. The role of chemokine
compatible with the experimental findings. signaling in regulating the self-organizing migration
In living organisms different strategies exist for of tissues during the morphogenesis of zebrafish is
cell movement, including both individual cell mi- investigated by Haas and Gilmour [2006].
gration and the coordinated movement of groups of By using an interdisciplinary approach,
cells [Rorth, 2007]. Figure 16 summarizes the basic Diz-Munoz et al. [2010] identified MCA (membrane-
types of collective cell migrations with respect to the to-cortex attachement) as a key component in
strength of the contact among the cells moving to- controlling directed cell migration during zebrafish
gether. gastrulation. They showed that reducing MCA in
i) Groups can be associated loosely with occasional mesendoderm progenitor cells during gastrulation
contact and much of the apparent cohesion might reduces the directionality of the cell migration as
come from essentially solitary cells following the same well. By investigating the same process, zebrafish
tracks and cues. Examples are germ cells in many gastrulation, Arboleda-Estudillo et al. [2010] found
organisms; the rostral migratory stream supplying that individually moving mesendoderm cells are
3.4 Cells 21
scribed in tumors in vivo. The first results from readers can find abundant literature on the topic, for
protruding sheets and strands that maintain contact example [Holldobler and Wilson, 2008].
with the primary site, yet generate local invasion. Many species of butterflies (e.g. Red Admiral,
The second shows detached cell clusters or cell files, Painted Lady) and moths (Humming-bird Hawk-
histologically seen as nests, which detach from their moth, Silver-Y moth) migrate twice a year between
origin and frequently extend along interstitial tissue the two hemispheres: when it is autumn on the
gaps and paths of least resistance, as seen in epithe- Northern hemisphere they form huge clouds and
lial cancer and melanoma [Friedl, 2004]. Collective fly to south and come back only when spring arrives.
migration represents the predominant mode of tissue Other insects being famous for exhibiting collective
invasion in most epithelial cancers. motion are ants. Many of them create tracks between
Furthermore, recently it has been argued that the nest and the food sources very efficiently, using
malignant tumor cells may be capable of devel- pheromone trails. For example New World army ant
oping collective patterns that resemble to evolved Eciton burchelii whose colonies may consist of mil-
adaptive behaviors like collective decision-making lion or more workers stage huge swarm raids with
or collective sensing of environmental conditions. up to 200 thousand individuals forming trail systems
Deisboeck and Couzin [2009] presented a concept as that are in length up to 100 m or even more, and 20
to how these abilities could arise in tumors and why m wide [Gotwald, 1995, Franks et al., 1991]. Based
the emergence of such sophisticated swarm-like be- on the observations Couzin and Franks [2003] have
havior would endow advantageous properties to the investigated the formation of these elaborated traffic
spatio-temporal expansion of tumors. lanes, and created a corresponding model exploring
In a recently published review article the influences of turning rates and local perception on
Friedl and Gilmour [2009] draw attention on traffic flow. Furthermore, Beekman et al. [2001] have
the presumed common mechanistic themes under- investigated another fundamental question regarding
lying the different collective cell migration types by these formations, namely what is the minimum num-
comparing them at the molecular and cellular level. ber of workers that are required for this kind of self
This review paper summarizes the topic from a more organization to occur. They have observed Pharaoh
biological point of view. ants and they actually discovered that small groups
forage in a disorganized way while larger ones are
organized. Thus for the first time they have
3.5 Insects
provided experimental evidence on a behavioral first-
Insects are one of the most diverse animal groups order phase-transition exhibiting hysteresis between
on Earth including more than a million described organized and disorganized states.
species which makes them represent more than half of Traditionally, an aggregate is considered to be an
all known living organisms [Chapman, 2009]. They evolutionarily advantageous state for its members: it
can move about by walking, flying or occasionally provides protection, information and choice of mates
swimming. Some of their species (like water strid- on the cost of limited resources and increased prob-
ers) are even able to walk on the surface of water. ability for various infections [Wilson, 1975a]. How-
Most of them live a solitary life, but some insects ever, according to some recent studies, in the case
(such as certain ants, bees or termites) are social of some insect-species the depletion of nutritional re-
and are famous for their large and well-organized sources may easily lead to cannibalism among group-
colonies. Some of these so called eusocial insects members. Simpson et al. [2006] reported how the lo-
have evolved sophisticated communication system, cal availability of protein and salt influenced the ex-
such as the round dance and waggle dance of tent to which Mormon cricket bands marched, both
Western honey bee, Apis mellifera. However, motion- through the direct effect of nutrient state on loco-
patterns based on such highly developed communica- motion and indirectly through the threat of canni-
tion are out of the scope of our review, but interested balism by resource-deprived specimens. Similarly,
24 3 OBSERVATIONS AND EXPERIMENTS
spect to the effect of the animal-density on the tran- other in a looser manner than in a school, and they
sition between disordered and ordered states. The might include fish of various species as well [Pitcher,
experimental results are depicted on Fig. 20, which 1983]. Shoals are more vulnerable to predator at-
shows the alignment of the motion in the func- tack. In contrast, in a school fish swim in a more
tion of locust density, for three different cases: (a) tightly organized way considering their speed and di-
low density ( = 0.26 0.86 103 expressed in rection, thus a school can be considered as a spe-
terms of normalized density), (b) intermediate den- cial case of shoal [Helfman et al., 1997]. At the same
= 1.23 3.1 103 ) and (c) high density
sity ( time, from one second to the other a shoal can or-
= 3.7 103 ). Normalized density is calculated
( ganize itself into a disciplined school and vice versa,
as usually, = /max , where max is the maximal according to the changes in the momentary activ-
hopper density estimated to be 20000 hopper/m2 af- ity: avoiding a predator, resting, feeding or traveling
ter [Murphy, 1967, Symmons and Cressman, 2001]. [Moyle and Cech, 2003, Hoare et al., 2004].
As it can be seen, coordinated marching behavior Schooling is a very basic feature of aquatic species
strongly depends on the animal-density. With these and may have appeared in a very early stage of verte-
experiments they also confirmed that the transition brate evolution [Shaw, 1978]. Over 50 percent of bony
followed the theoretical predictions of the SPP-model fish species school and the same behavior has been
[Vicsek et al., 1995], and identified the critical den- reported in a number of cartilaginous fish species as
sity for the onset of coordinated marching as well. well [Shaw, 1978, Benoit-Bird and Au, 2003]. Since
Although zooplankton are not insects, here we the large-scale coherent motion of fish is also very
briefly mention an interesting study in which important from a practical point of view (fishing in-
Daphnia-swarms were artificially induced to carry dustry), the observational and simulational aspects of
out vortex motion by using optical stimulus. When fish schools have played a central role in the studies
the density of these tiny creatures is small, they ex- of coherent motion.
hibit circular motion around a vertical shaft of light, Becco et al. [2006] recorded the trajectories of
to which they are attracted. Ordemann et al. [2003] young fish in a school (see Fig. 22). Both individ-
found that above a density-threshold a swarm-like ual and collective behavior were studied as a func-
motion emerges in which all Daphnia circle in the tion of fish-density, and a transition from disor-
same randomly chosen direction. In order to dered to correlated motion was found. Also by tra-
reproduce the observed behavior, the authors devel- jectory analysis, Katz et al. [2011] inferred the struc-
oped a self-propelled agent based model based on ran- ture of the interactions among schooling golden shin-
dom walks. They found that with two ingredients of ers, Notemigonus crysoleucas. They found that it is
the model the observed circular motion can be repro- not an alignment rule but a speed regulation which
duced: (i) a short-range temporal correlation of the is the key aspect during interaction, i.e., changes
velocities (which is in the experiment the short-range in speed effecting conspecifics both behind and in
alignment resulting from the water drag), and (ii) front of the fish are essential. They argue that align-
an attraction to a central point proportional to the ment only modulates the strength of speed regulation,
agents distance from it (which is in the experiment rather than being an explicit force itself. Another
the attraction generated by the light beam.) important claim of the study is that the observed
interactions can not be decomposed into the sum of
two-body interactions, but rather three-body interac-
3.6 Fish schools and shoals
tions are necessary in order to explain the observed
The largest groups of vertebrates exhibiting a rich set dynamics.
of collective motion patterns are certainly fish shoals Using a novel technique called Ocean Acoustic
and schools. Although these two terms cover very Waveguide Remote Sensing, OAWRS [Makris et al.,
similar behaviors and thus are often mixed their 2006], which enables instantaneous imaging and con-
meaning slightly differs: in a shoal fish relate to each tinuous monitoring of oceanic fish shoals over tens of
26 3 OBSERVATIONS AND EXPERIMENTS
Akos et al., 2008]. not all, [Nagy et al., 2010]). Such groups might reach
Applying GPS data-loggers in six highly pre- a consensus either without leader (by quorum re-
trained pigeons, the efficiency of a flock was investi- sponse, mean value, etc.) or with a leader. However,
gated by DellAriccia et al. [2008]. They found that even in this latter case, leadership is temporal since
the homing performance of the birds flying as a flock it is based on temporal differences, such as pertinent
was significantly better than that of the birds released information of food location or differences in some
30 3 OBSERVATIONS AND EXPERIMENTS
bras, Equus burchellii. Identity covers both dom- members are needed to lead the group. Dyer et al.
inance and kinship, while the inner state was inter- [2008] tested these predictions on human groups in
preted as the reproductive state (if the individual is which the experimental subjects were nave and they
in its lactation period or not). Zebra harems consist did not use verbal communication or any other ac-
of tightly knit individuals in which females were ob- tive signaling. The experiments indeed supported the
served to have habitual roles (personal differences) predictions. Other experiments investigated the re-
in the initiation of group movements. The authors lationship between the spatial position of informed
also found that lactating females initiate movements individuals and the speed and accuracy of the group
more often than non-lactating ones, thus lactation, motion [Dyer et al., 2009]. The results proved valid
as inner state, plays an important role in leader- in larger crowds as well (100 and 200 people) which
ship. Others find more direct relationship to hier- can have important implications on plans aiming to
archy. S arova et al. [2010] recorded the motion of a guide human groups for example in case of emer-
herd of 15 beef cows, Bos taurus, for a three-week gency.
period using GPS devices. They found that short- Faria et al. [2010b] studied the effect of the knowl-
distance travels and foraging movements are not lead edge regarding the presence and identity of a leader in
by particular individual, instead, they are rather in- small human groups, and also investigated those in-
fluenced in a graded manner, i.e., the higher an in- advertent social cues by which group members might
dividual was in the group hierarchy, the bigger influ- identify leaders. With this object they conducted 3
ence it exerted on the motion of the herd. Accord- treatments: in the first, participants did not know
ing to the observations, Rhesus macaques (Macaca that there was a leader, in the second treatment they
mulatta) preferred to join related or high-ranking in- were instructed to follow the leader but they did
dividuals too, whereas Tonkean macaques (Macaca not know who it was, while in the third they knew
tonkeana) exhibited no specific order at departure who the leader was. The experiments took place in
[Sueur and Petit, 2008]. In a recent review article a circular area with 10m diameter labeled by num-
Petit and Bon [2010] interpreted the process of col- bers from 1 to 16. These marks were spaced equally
lective decision making (regarding group movements) around the perimeter, as shown in Fig. 30. In all the
as a combination of two kinds of rules: individual- trials, participants were instructed (i) not to talk or
based and self-organized. The first one covers the to make any gesture, (ii) to walk continuously, and
differences of the [mostly inner] states of the an- (iii) to remain together as a group. Further instruc-
imals, that is, differences in social status, physi- tions were provided on a piece of paper: a (randomly
ology, energetic state, etc. The second one, self- chosen) person was asked to move to a (randomly
organization, corresponds to the interactions, simple chosen) target but stay with the group. She/he was
responses among individuals. the informed individual, the leader. The rest of
Regarding the case when leadership emerges solely the group was uninformed whose instructions differed
from differences in the inner states of the group mem- from treatment to treatment: In the first one, they
bers (those ones lead who have pertinent informa- were only asked to stay with the group. In the sec-
tion), Couzin et al. [2005] suggested a simple model ond treatment they were told to follow the leader, but
to show how a few informed individuals can lead a they did not know who it was. In the third one, they
whole group. In this model (which is detailed in Sec. were asked to follow the leader whose identity was
5.4) group members do not signal and do not know provided (by the color if his/her sash). Although the
which of them (if any) has information regarding the accuracy of the group movement significantly differed
desired direction. This model predicts that even if from treatment to treatment, the leader always suc-
the portion of the informed individuals within the ceeded to guide the group to the target. The least ac-
group is very small, the group as a whole can achieve curate group motions were measured during the first
great accuracy in its movement. In fact, the larger treatment, while the second and third ones resulted
the group size, the smaller the portion of informed group motions whose accuracy were close to the pos-
32 3 OBSERVATIONS AND EXPERIMENTS
hv~j (t)iR
v~i (t + 1) = v0 + perturbation (13)
hv~j (t)iR
tion patterns, such as marching groups, mills, rotat- favor of their picture that L diverges in various lim-
ing chains, bands, etc. (see Sec. 4.2 and 4.3). In some its: both the low and high density limits, as well as
cases (i.e. by applying some certain parameter and in the small velocity limit. In particular, an extrap-
initial condition settings) these ordered phases can olation of their estimates towards the small velocity
exhibit some remarkable features as well, such as gi- regimes considered in prior works gives values of L
ant number fluctuations (GNF, see for example page so large that do not make the corresponding simula-
40, 52) or band formation. According to the studies, tions feasible.
noise, density, the type of interaction (attractive or Studies aiming to reveal the nature of the above
repulsive, polar or apolar, the range of the interac- phase transition (whether it is first or second or-
tion) and the boundary conditions (in case of finite der) find that the noise (more precisely, the way it
size models) all proved to play an important role in is introduced into the system), and the velocity with
the formation of certain patterns. which the particles move, play a key role. Accord-
ingly, while simulations show that for relatively large
4.1.1 The order of the phase transition velocities (v0 > 0.5) the transition is discontinuous,
Baglietto and Albano [2009b] demonstrated that for
In the paper introducing the original variant (OVM) smaller velocities, even in the limit when the veloc-
of the SVM [Vicsek et al., 1995], a second order phase ity goes to zero (except when it is exactly equal to
transition from disordered to ordered motion was zero), the transition to ordering is continuous (is in-
shown to exist. In particular, in the thermodynamic dependent of the actual value of the velocity, as it
limit, the model was argued to exhibit a kinetic phase can be seen in Fig. 32 b). Very recently Ihle [2011]
transition analogous to the continuous ones in equi- and Mishra et al. [2010] were able to see band-like
librium systems, that is, structures in their solutions obtained from a contin-
uum theory approach. Bands usually signal first or-
[c () ] der phase transition, however, Ihle [2011] found them
[ c ()] (16) for a large velocity case, while Mishra et al. [2010]
assumed throughout their calculation that the tran-
which defines the behavior of the order parameter sition from disorder to order was continuous.
at criticality, in the case of a standard second order Aldana et al. [2007] demonstrated that the type of
transition. and are critical exponents, is the the phase transition depends on the way in which the
noise (in the form of random perturbations), is the noise is introduced into the system. They analyzed
particle density, and c () and c () are the critical two network models that capture some of the main
noise and critical density, respectively, for L . aspects characterizing the interactions in systems of
(L is the linear size of the system.) self-propelled particles. In the so called vectorial
However, the continuous nature of this transition noise model the perturbation (in the form of a ran-
has been questioned [Gregoire and Chate, 2004] re- dom vector) is first added to the average of the veloc-
sulting in a number of studies investigating this fun- ities and the final direction is determined only after
damental aspect of collective motion. Chate and this [Gregoire and Chate, 2004]. When the average
coauthors [Chate et al., 2008b], in their extensive velocity is small (disordered motion) this seemingly
follow-up study, presented numerical results indicat- subtle difference in the definition of the final direction
ing that there exists a crossover system size, which leads to a qualitatively different ordering mechanism
they call L , beyond which the discontinuous char- (sudden first order-type transition to the ordered
acter of the transition appears independent of the state).
magnitude of the velocity. They demonstrate that Correspondingly, Aldana et al. [2009] analyzed the
this discontinuous character is the true asymp- order-disorder phase transitions driven by two dif-
totic behavior in the infinite-size limit. Importantly, ferent kinds of noises: intrinsic (the original form,
Chate et al. [2008b] showed and presented results in perturbing the final angle) and extrinsic (the vec-
36 4 BASIC MODELS
torial one, perturbing the direction of the individ- Similarly, the fluctuation of the order parameter,
ual particles before averaging). Intrinsic is related = 2 L2 , takes the form
to the decision mechanism through which the par-
ticles update their positions, while extrinsic affects (, L) = L/ ((
c )L1/ ), (18)
the signal that the particles receive from the environ-
ment. The first one calls continuous phase transitions where is a suitable scaling function,
is
the sus-
2
forth, whereas the second type produces discontinu- ceptibility critical exponent, and 2 2 hi is
ous phase transitions [Pimentel et al., 2008]. Finally, the variance of the order parameter. In the thermo-
Nagy et al. [2007] showed that vectorial noise results dynamic limit, obeys ( c ) . (See also Eq.
in a behavior which can be associated with an insta- (4))
bility. Equations (17) and (18) are convenient to deter-
mine the critical exponents within the framework of
finite size scaling theory. As a crucial result, the au-
4.1.2 Finite size scaling thors found that the exponents they calculated satisfy
So far, the most complete study regarding the scaling the so-called hyperscaling relationship
behavior of systems of self-propelled particles exhibit-
d 2 = (19)
ing simple alignment plus perturbation, has been car-
ried out by Baglietto and Albano [2008]. They per- which is, in general, valid for standard (equilibrium)
formed extensive simulations of the SVM, and ana- critical phenomena. d denotes the dimension, d = 2.
lyzed them both by a finite-size scaling method (a The nature of intermittency intermittent
method used to determine the values of the criti- bursts during which the order is temporarily lost in
cal exponents and of the critical point by observing such systems has also been a subject of investiga-
how the measured quantities vary for different lat- tions recently [Huepe and Aldana, 2004].
tice sizes), and by a dynamic scaling approach. They
observed the transition to be continuous. In addi-
tion they demonstrated the existence of a complete 4.2 Variants of the original SPP
set of critical exponents for the two dimensional case model
3
(including those corresponding to finite size scaling ) Several variants of the above-introduced, simplest
and numerically determined their values as well. In SPP model have been proposed over the years. One
particular, within the framework of finite-size scaling of the main directions comprises those studies that
theory, the scaling ansatz for the order parameter investigate systems in which the particles (units) do
of the SVM has been rewritten as not follow any kind of explicit alignment rule, only
collisions occur between them in the presence of some
(, L) = L/ (
c )L1/ , (17) kind of interaction potential. We shall overview this
where L is the finite size of the system, is a suitable approach in Sec. 4.2.1. Models assuming some kind
scaling function, and finally, and are two of the of alignment rule for the units, will be dealt with in
critical exponents in question: is the one belonging Sec. 4.2.2.
to the order parameter, and is the correlation length
critical exponent. 4.2.1 Models without explicit alignment rule
3 Numerical simulations carried out on systems having fi- As mentioned in Sec. 4.1, in the most simple SPP
nite size L in at least one space dimension exhibit so called models, an alignment term is assumed. However,
finite size effects, most importantly rounding and shifting ef- according to very recent studies (see Sec. 3.2), the
fects during second-order phase transitions. These artifacts are
particularly emphasized near the critical points, but they can
motion of particles may become ordered even if no
be accounted for by means of the so called finite-size scaling. explicit alignment rule is applied, but alignment is
See more on this topic in [Cardy, 1996, Brankov et al., 2000]. introduced into the collision in an indirect way by
4.2 Variants of the original SPP model 37
tems, however, at the same time, also to interpret the In Eq. (24), the terms , > 0 give v a nonzero
so called active matter systems associated mainly magnitude, DL,1,2 are diffusion constants and ~ is an
with applications from physics, such as active nemat- uncorrelated Gaussian random noise. The terms on
ics and active suspensions. the left hand side of the equation are the analogs of
The first theory describing the full nonlinear the usual convective derivative of the coarse-grained
higher dimensional dynamics was presented in velocity field ~v in the Navier-Stokes equation. Here
[Toner and Tu, 1995, 1998]. Tu and Toner followed the absence of Galilean invariance allows all three
the historical precedent [Forster et al., 1977] of the combinations of one spatial gradient and two veloci-
Navier-Stokes equation by deriving the continuum, ties that transform like vectors; if Galilean invariance
long-wavelength description not by explicitly coarse did hold, it would force 2 = 3 = 0 and 1 = 1.
graining the microscopic dynamics, but, rather, by However, Galilean invariance does not hold, and so
writing down the most general continuum equations all three coefficients can be non-zero phenomenolog-
of motion for the velocity field ~v and density con- ical parameters whose values are determined by the
sistent with the symmetries and conservation laws of microscopic rules. Eq. (25) reflects the conservation
the problem. This approach allows to introduce a of mass (birds). The pressure P depends on the local
few phenomenological parameters (like the viscosity density only, as given by the expansion
in the Navier-Stokes equation), whose numerical val-
ues will depend on the detailed microscopic behavior X
P = P () = n ( 0 )n (26)
of the particles. The terms in the equations describ- n=1
ing the large-scale behavior, however, should depend
only on symmetries and conservation laws, and not where 0 is the mean of the local number density and
on the microscopic rules. n is a coefficient in the pressure expansion.
The only symmetry of the system is rotation invari- It is possible to treat the whole problem analyt-
ance: since the particles lack a compass, all direction ically using dynamical renormalization group and
of space are equivalent to other directions. Thus, show the existence of an ordered phase in 2D,
the hydrodynamic equation of motion cannot have and extract exponents characterizing the velocity-
built into it any special direction picked a priori; all velocity and density-density correlation functions
directions must be spontaneously selected. Note that [Toner and Tu, 1998]. The most dramatic result is
the model does not have Galilean invariance: chang- that an intrinsically non-equilibrium and nonlinear
ing the velocities of all the particles by some constant feature, namely, convection, suppresses fluctuations
boost ~vb does not leave the model invariant. of the velocity ~v at long wavelengths, making them
To reduce the complexity of the equations of mo- much smaller than the analogous fluctuations found
tion still further, a spatial-temporal gradient expan- in ferromagnets, for all spatial dimensions d < 4.
sion can be performed keeping only the lowest order In other words, the existence of the convective term
terms in gradients and time derivatives of ~v and . makes the dynamics non-potential and further sta-
This is motivated and justified by the aim to con- bilizes the ordered phase. Heuristically, this term
sider only the long distance, long time properties of accounts for the stabilization effect resulting from
the system. The resulting equations are the feature that the actual neighbours of each unit
continually change due to the local differences in
2 the direction. Thus, particles (birds) which initially
t~v + 1 (~v )~v + 2 (~v )~v + 3 (|~v | ) =
were not neighbours, and thus did not interact with
~v |~v |2~v P +DL (~v )+D1 2~v +D2 (~v )2~v +~ each other, at a later time-step might be within each
(24) others interaction range.
Further predictions of the above model were tested
and by numerically studying a discrete model very sim-
t + (~v ) = 0. (25) ilar to the SVM. Compared to the original model,
4.3 Continuous media and mean-field approaches 43
an extra interaction term was introduced, in order to for all transport coefficients as a function of the three
prevent cluster formation: main parameters, noise, density and velocity.
3 2 ! Over the last 4 years, the hydrodynamic equations
l0 l0 became increasingly precise by including higher order
~gij = g0 (~
ri r~j ) (27)
rij rij terms and more precise coefficients. Very recently
Mishra et al. [2010] solved the equations describing
This expression sets the average distance between the collective motion of self-propelled polar rods mov-
boids in the flock to be l0 by creating an attraction ing on an inert substrate. From their theoretical con-
force between particles i and j if the rij distance be- siderations and numerical analysis, the authors ob-
tween them is bigger than than l0 , and making them tained a remarkable phase diagram for this system
repel each other if rij < l0 . g0 is a model param- (Fig. 41). They showed that the same physics that
eter, defining the strength of the above mentioned leads to global ordering destabilizes the homogeneous
attraction-repulsive force. The linear size L of the ordered state above a critical value of self-propulsion
simulated system was L = 400 with N = 320, 000 speed and allows the nonlinear equations to admit
boids moving in it. The range of interaction was set a propagating front solution that yields the striped
to be unit (1), g0 = 0.6, the velocity v0 = 1.0 and phase identified numerically. The two phases they
l0 = 0.707. In finite size systems, the average direc- observed, namely, the striped phase and the fluctu-
tion of the flock < ~v > slowly changes in time, due ating flocking phase, have been identified earlier in
to the noise. In contrast, analytic results assume in- the context of numerical studies of the SVM model,
finite systems size in which the direction < ~v > is thus, the approach of Mishra et al. [2010] identified
constant. In order to handle this disagreement, the the origin of these phenomena in the model indepen-
boundary condition in one direction (say the x) was dent framework of the dynamics of conserved quan-
set to be periodic, while in the other (the y) a re- tities and broken symmetry variable.
flective boundary condition was applied. Hence, the It is important to point out that the above
symmetry broken velocity was forced to lie along the mentioned equations [Bertin et al., 2009, 2006,
x direction. As an interesting phenomenon, in the Mishra et al., 2010] obtained as a result of detailed
direction y (the one perpendicular to < ~v >) the in- derivations based on microscopic dynamics have an
dividual boids exhibit an anomalous diffusion. Fur- analogous structure and contain the same major
thermore, as an even more surprising result, in the terms as the ones (Eqs. 24 and 25) proposed by Toner
flocks moving direction (x), the fluctuations of the and Tu inspired by general considerations.
velocity and the that of the density were propagating A further important approach involving contin-
with different velocity. uum mechanics is based on considering the hydro-
Bertin et al. [2006, 2009] made a very important dynamic properties of systems consisting of micro-
step towards a fundamental theory of collective mo- scopic swimmers (see also Sec. 5.1.1). By develop-
tion by deriving the hydrodynamic equations for the ing a kinetic theory, Saintillan and Shelley [2008b]a
density and velocity fields of a gas of self-propelled and Saintillan and Shelley [2008a]b studied the col-
particles with binary interactions from the corre- lective dynamics and pattern formation in suspen-
sponding microscopic rules. They gave explicit ex- sions of self-propelled particles. They investigated
pressions for the transport coefficients as a function the stability both of aligned and isotropic sus-
of the microscopic parameters. Comparison with pensions, and by generalizing the predictions of
numerical simulations on a standard model of self- Simha and Ramaswamy [2002] they showed that
propelled particles (SVM, see Sec. 4.1) resulted in aligned suspensions of self-propelled particles are al-
an agreement as well as in a demonstration of the ways unstable to fluctuations. Furthermore, they
robustness of the phase diagram they obtained. Ihle showed that in the case of initially isotropic suspen-
[2011] showed how to explicitly coarse grain the mi- sions an instability for the particle stress takes place
croscopic dynamics of the SVM to obtain expressions for pushers particles propelled from the rear but
44 4 BASIC MODELS
t~rL = ~u(~rL ),
t~rS = ~u(~rS ), (28)
studying the transition from normal state (with non- common direction which is in this case the topic of
zero resistance) to the zero-resistance phase, the au- consensus. The position of the ith bird is given by
thors find analogy with flocking systems. In partic- xi ( 3 ). (Of course, xi = xi (t).) Let us define the
ular, the two frames become identical in the limit of adjacency matrix A = (aij ), where the element aij
zero magnetic field and short range electron-electron measures the ability of birds i and j to communicate
interactions. with each other, or one could say, the influence they
The equations of motion for the current and den- exert on each other. The elements of A should take
sity fluctuations are constructed based on symmetries values from the interval (0...1], and the closer unit
and conservation laws. Two cases were analyzed: a i is to unit j, the bigger aij should be (since they
model valid for small length scales (on which the den- influence each other stronger). is a tuning pa-
sity does not vary noticeably) characterized by an im- rameter, effecting the strength of the influence. An
posed symmetry under a global uniform shift of the appropriate expression for the above requirements is
density, and a description argued to be appropriate
1
for describing the system on long length scales. In aij = , (30)
2
the first case, the type of the phase transition is pre- (1 + kxi xj k )
dicted to be continuous in case of short-range inter-
actions and first order otherwise (long-range), while where 0 (not to be confused with the critical ex-
the second model predicts first order phase transition ponent introduced in Sec. 2.1). The main advantage
in both cases. of this form of the distance dependence of the interac-
tion is that it is a smooth function allowing analyt-
This is a quickly growing field of its own has re-
ical treatment. Importantly, this adjacency matrix
cently been reviewed by Lauga and Powers [2009].
A changes with time, since the positions of the birds
change with time.
4.4 Exact results For the more manifest usage of graphs the authors
introduce the Laplacian matrix of A as well, L =
By exact results here we mean results obtained
D A, where D is a k k diagonal matrix
Pk whose ith
with a minimum or completely missing amount of
diagonal element is defined as di = j=1 aij . The
any kind of assumptions or approximations concern-
Laplacian matrix a form by which a graph can be
ing the behavior of the moving units (beyond the
represented in matrix-form is often used to find
rules/definitions they obey). Originally most results
various properties of a graph. In particular, as we will
in this area were obtained only for systems in which
see, the eigenvalues of L bear important information.
the noise (an otherwise essential aspect of flocking)
Denoting the velocity of bird i at time t by vi (t)(
was completely neglected. Thus, one could consider
3 ),
the related systems as fully deterministic. However,
it has recently been shown (see later) that the the- k
X
orems we review below are in most cases valid for vi (t + h) vi (t) = h aij (vj vi ) (31)
systems with a low level of perturbations as well. j=1
on (3 )k (3 )k , where L gives the local averaging. of the conditions needed to be satisfied for the emer-
(Note that the matrices A and L are acting on (3 )k gence of flocking.
by mapping (v1 , . . . , vk ) to (ai1 v1 + . . . + aik vk )ik .) Importantly, in the case of flocking, F = F (t) (it
is a function of time), because the elements of G de-
After the above preparations, one can ask that pend on the xi positions of the individual birds. By
under what conditions does a system (described by using the Fiedler-number, we can say that one ob-
the above equations) exhibit flocking behavior? Or in tains flocking, if and only if
other words, when do the solutions of vi (t) converge
to a common v ( 3 )? 0 < const F = F (x(t)). (35)
One of the most important results of
Cucker and Smale [2007a]a is that the emer- Otherwise the flock disperses.
gence of the flocking behavior depends on ; namely The above definitions can be extended to weighted,
if is small enough ( < 1/2) then flocking always general matrices as well.
emerges. Formally,
Theorem: For the equations of flocking (Eqs. 32) In addition, Cucker and Dong [2010] extended the
there exists a unique solution for all t . model by adding to it a repelling force between par-
If < 1/2 then the velocities vi (t) tend to a com- ticles. They showed that, for this modified model,
mon limit v ( 3 ) as t , where v is indepen- convergence to flocking is established along the same
dent of i, and the vectors xi xj tend to a limit-vector lines while, in addition, avoidance of collisions (i.e.,
ij for all i, j k, as t , that is, the relative po- the respect of a minimal distance between particles)
x
sitions remain bounded. is ensured.
If 1/2 dispersal, the split-up of the flock is The main differences between the systems de-
possible. But, provided that some certain initial scribed by Cucker and Smale and by the SVM are,
conditions are satisfied, flocking will occur. from the one hand, the definition of the range of in-
teraction, and from the other hand, the existence (or
To obtain more general results for the conditions absence) of noise. The SVM comprises noise, while
of flocking, one can investigate the eigenvalues of the the original Cucker-Smale model does not. Regard-
corresponding Laplacian matrix L. Let G denote a ing the range of interaction, in the present model it is
graph and A be the corresponding adjacency matrix a long-range effect decaying with the distance accord-
defined as usually, that is, ing to (see Eq. (30)), while in the SVM it has the
same intensity for all the neighboring units around a
1 if i and j are connected,
aij = (33) given particle, but only within a well-defined range
0 if not (see Eq. (13)).
Very recently perturbations have also been consid-
Let D be a diagonal matrix P with the same dimensions ered in the CS model. This has been done with var-
as A, defined by d(i, i) = j a(i, j). Then the general
ious forms of noise by Cucker and Mordecki [2008],
form of the Laplacian matrix of G, is given as L =
Shang [2009], modeled with stochastic differential
L(G) = DA. The eigenvalues of L can be expressed
equations by Ha et al. [2009], and taking into ac-
by
count random failures between agents connections
0 = 1 2 3 . . . (34)
by Dalmao and Mordecki [2009].
1 , the first eigenvalue is always zero. The second Other works developing the Cucker-Smale (CS)
one in ascending order, 2 , is the so-called Fiedler- model in several directions include an extension to
number, F , which is zero if the graph G is separated fluid-like swarms [Ha and Liu, 2009, Ha and Tadmor,
(in this case the flock disintegrates to two or more 2008, Carrillo et al., 2010], collision avoiding flocking
smaller flocks), and non-zero if and only if G is con- [Cucker and Dong, 2010], the inclusion of agents with
nected. This number is a crucial descriptive measure a preferred velocity direction [Cucker and Huepe,
48 4 BASIC MODELS
2008], and its proposal as a control law for the space- spite the absence of centralized coordination and de-
crafts of the Darwin mission of the European Space spite the fact that each agents set of nearest neigh-
Agency [Perea et al., 2009]. bors changes in time. By addressing the question of
global ordering in models analogous to Eqs. (13) and
4.4.2 Network and control theoretical as- (15) they presented some rigorous conditions for the
pects graph of interactions needed for arriving at a consen-
sus.
Networks have recently been proposed to repre- Several further control theory inspired papers dis-
sent a useful approach to the interpretation of cussed both the question of convergence of the
the intricate underlying structure of connections simplest SPP models as well as the close re-
among the elements of complex systems. A lation of flocking to such alternative problems
number of important features of such networks as consensus finding, synchronization and gossip
have been uncovered [Albert and Barab asi, 2002, algorithms[Blondel et al., 2005, Boyd et al., 2005].
Watts and Strogatz, 1998]. It has been shown that Ren and Beard [2005] considered the problem of
in many complex systems ranging from the set of consensus finding under the conditions of limited
protein interactions to the collaboration of scientists and unreliable information exchange for both discrete
the distribution of the number of connections is de- and continuous update schemes. They found that
scribed by a power law as opposed to a previously in systems with dynamic interaction-topologies con-
supposed Poissonian. Most of the networks in life and sensus can be reached asymptotically, if the union
technology are dynamically changing and are highly of the directed communication network across some
structured. In particular, such networks are typically time intervals has spanning trees frequently enough,
made of modules that are relatively more densely con- as the system evolves. Similarly, Xiao and Wang
nected parts within the entire network (e.g., interact- [2006] found the existence of spanning trees to be
ing flocks) [Newman, 2004, 2006, Palla et al., 2005, crucial in the directed graphs representing the inter-
Scott, 2000]. The evolution of these modules plays a action topologies, in systems in which the topology,
central role in the behavior of the system as a whole weighting factors and time delays are time-invariant.
[Palla et al., 2007]. They studied the consensus-problem for dynamic net-
In these terms, a dynamically changing network works with bounded time-varying communication de-
can be associated with a flock of collectively moving lays under discrete-time updating scheme, based on
organisms (or robots, agents, units, dynamic systems, the properties of non-negative matrices.
etc.). In such a network two units are connected if An efficient algorithm controlling a flock of un-
they interact. Obviously, if two units are closer in manned aerial vehicles (UAVs) is considered by
space have a better chance to influence the motion of Ben-Asher et al. [2008]. The units are organized into
each other, but their interaction can also be disabled a minimal set of rooted spanning trees (preserving
by environment or internal disturbances. Since the the geographical distances) which can be used for
units are moving and the environment is also chang- both distributed computing and for communication
ing, the network of momentarily interacting units is as well, in addition to computation and propagation
evolving in time in a complex way. Using the con- of the task assignment commands. The proposed
ventional terminology of control theory, this kind of protocol continually attempts to keep the number of
topology (that is, when certain number of edges are trees minimal by fusing separate adjacent trees into
added or removed from the graph from time to time), single ones: as soon as radio connection between two
is called switching topology. nodes belonging to separate trees occurs, the cor-
Jadbabaie et al. [2003] investigated a theoretical responding networks fuse. This arrangement over-
explanation for a fundamental aspect of the SVM, comes the typical deficiencies of a centralized solu-
namely, that by applying the nearest neighbor rule, tion. The motion of the certain UAVs is coordinated
all particles tend to align into the same direction de- by Reynoldss algorithm [Reynolds, 1987] (see also
4.5 Relation to collective robotics 49
moving there upon switching on the light. Jij is introduced to account for the observation
In a more theoretical work, Sugawara et al. [2007] that animals tend to align with each other [Hunter,
investigated the formations of motile elements 1966] through an interaction which is supposed to
(robots) as a function of various control parameters. decrease in the linear function of distance between
Their kinetic model inspired by living creatures, individuals i and j:
such as birds, fishes, etc. is defined by Eqs. (39) 1
and (40). |~rj ~ri |
Jij = k , (42)
rc
d~vi
ij f~ij + ~gi
X
m = ~vi + a~ni + (39) where k is the control parameter and rc is the pre-
dt ferred distance between neighbors. The term gi is a
j6=i
force directed towards the center of the group, and fi-
nally, fij denotes a mutual attractive/repulsive force
di X between elements i and j, in analogy with the inter-
= sin(i i ) + Jij sin(j i ), (40)
dt molecular forces, as suggested by Breder [1954].
j6=i
a general mechanism that builds on preferential at- neighboring segments i and i + 1. Kb and K are the
traction/attachment to elongated structures. The stretching and the bending coefficients, respectively,
proposed interactive particle model exhibits robust defining the extent to which the length of the seg-
sprouting dynamics and results in patterns that are ments and the angles between them can vary. Both
similar to native primordal vascular plexuses with- of them are dimensionless values, and are the same
out any assumptions involving mechano-chemical sig- for all the segments and angles.
naling or chemotaxis. Regarding the active motion of the certain cells,
Wang and Wolynes [2011] proposed a model for first the head-node moves in a particular direction,
structures like the cytoskeleton, that is, for systems followed by the other nodes which take positions so
consisting of many interacting bio-macromolecules that the Hamiltonian function belonging to the new
driven by energy-consuming motors. Readers inter- configuration is minimal. Since according to the ob-
ested in the models of active polar gels will find more servations, Myxococcus xanthus cells do not have any
details in [J
ulicher et al., 2007]. kind of long-range communicating systems [Kaiser,
2003], the model takes the interactions only among
5.1.3 Swarming bacteria neighboring cells into account.
The experimentally-observed reversals (sudden
By using models and simulations, experimentally ob- changes in the direction with 180o) are most proba-
served behaviors which are seemingly unintelligible bly regulated by an internal biochemical clock, which
might also be elucidated. Recently, as described in is independent of the actual interactions of the given
Sec. 3.3, bacteria belonging to Myxococcus xanthus cell. Therefore, the model takes into account these
swarms were observed to reverse their gliding di- reversals by simply switching the roles of the head-
rections regularly, while the colony itself expanded nodes and the tail-nodes, according to an internal
[Wu et al., 2009]. To compass this seemingly energy- clock.
wasting behavior, the authors simulated the observed Simulations based on the above model did not re-
phenomena using a cell-based model, taking into ac- sult in swarming of the non-reversing cells in contrast
count only the contact-mediated, local interactions to the simpler models by Peruani et al. [2006] and
[Wu et al., 2007]. The individual cells are represented Ginelli et al. [2010]. On the other hand, it was found
by a flexible string of N nodes, consisting of N 1 that the expansion rate of the colony depends on the
segments, as depicted on Fig. 49 (basically a bend- length of the reversal period. Notably, the biggest
able rod, bended in N 2 points, being able to move expansion is obtained within the same time-period
in 2-D space). Each segment has the same length that was experimentally observed, that is 8 min.
r. In the simulations N was chosen to be 3, thus The cellular motion and the emerged patterns deep
each cell had two segments, as the rod was blended inside the colony was also modeled. As it can bee
in one point, in the middle. The orientations of the seen on Fig. 50, the considered social interactions
cells are defined by the vectors directed from the tail result an enhanced order regarding the collective cel-
nodes to the head nodes. In order to keep the shapes lular motion. It should be noted, that in a very re-
of the cells within an interval that agrees with the cent preprint Peruani et al. [2010] found signs of both
observations, a Hamiltonian function was defined to ordering and clustering in experiments with a non-
characterize the certain node-configurations, as given reversing, genetically modified mutant of a myxobac-
by Eq. (47). teria strain.
One of the earliest works on the collective motion of
N 1 N 2
X X bacteria pointing out the reason why such models are
H= Kb (ri r0 )2 + K i2 , (47) important, is done by Czirok et al. [1996]. The au-
i=0 i=0
thors emphasized that the study of bacterial colonies
where ri is the length of the ith segment, r0 is its can lead to interesting insights into the functioning of
target length and i is the angle enclosed by the self-organized biological systems which rest on com-
5.1 Systems involving physical and chemical interactions 57
if rnm < rc ,
rnm
fnm = 1 re if rc < rnm < r0 , (53)
0 if rnm > r0 ,
Figure 59: The global collective motion emerging 5.3.2 Moving in three dimensions fish and
from escape and pursuit behaviors. s is the rescaled birds
density, which is defined as s = N ls2 /L2 , where N
The main goal of the first system-specific models aim-
is the total number of the individuals, ls is their
ing to simulate the motion of animals moving in 3
interaction range and L is the size of the simula-
dimensions (primarily birds and fish) was to pro-
tion field. The simulations were carried out by us-
duce realistically looking collective motion [Reynolds,
ing periodic boundary conditions. The column de-
1987], to give system-specific models taking into
noted by p shows the typical spatial configurations
account many parameters [McFarland and Okubo,
for the pursuit-only case, e for the escape-only case,
1997], or to create systems in which some charac-
and p + e when both interactions are present. The
teristics (for example nearest neighbor distance or
large arrows indicate the direction and speed of the
density) resembles to an actual biological system
mean migration. As it can be seen on the top row, at
[Huth and Wissel, 1994]. Later various other aspects
low rescaled densities the emergent patterns strongly
and features were also studied, such as the func-
vary according to the strength of the escape and pur-
tion and mechanism of line versus cluster forma-
suit interactions. From Romanczuk et al. [2009].
tion in bird flocks [Bajec and Heppner, 2009], how
the fish size and kinship correlates with the spatial
characteristics (e.g., animal density) of fish schools
to collective motion, but with an opposite effect on [Hemelrijk and Kunz, 2005], the effect of social con-
the density distribution. Whereas pursuit leads to nections (social network) on the collective motion
density-inhomogeneities (that is, to the appearance of the group [Bode et al., 2011b], the cohort depar-
of clusters, as it can be seen on the first column on ture of bird flocks [Heppner, 1997], the collective be-
Fig. 59), escape calls to forth homogenization. Thus, havior in an ecological context (in which not only the
the collective dynamic in which both behavior-types external stimuli, but the internal state of the individ-
are present, is a competition between the two oppo- uals are also taken into account) [Vab and Skaret,
site effects. 2008], or the effect of perceived threat [Bode et al.,
Another often observed phenomena regarding col- 2010]. This latter circumstance was taken into ac-
64 5 MODELING ACTUAL SYSTEMS
count by relating it to higher updating frequency, and lective motions as the function of the system pa-
it resulted a more synchronized group regarding its rameters. In this framework, the individuals obey
speed and nearest neighbor distribution. The book to the following basic rules: (i) they continually at-
of Parrish and Hamner [1997] provides an excellent tempt to maintain a certain distance among them-
review on this topic. selves and their mates, (ii) if they are not performing
Models proposed by biologists tend to take into ac- an avoidance manoeuvre (described by rule i), then
count many of the biological details of the modeled they are attracted towards their mates, and (iii) they
animals. A good example for this kind of approach is align their direction to their neighbors. Their percep-
the very interesting work of Heppner and Grenander tion zone (in which they interact with the others) is
[1990], who proposed a system of stochastic differen- divided into three non-overlapping regions, as illus-
tial equations with 15 parameters. Huth and Wissel trated in Fig. 60. The radius of these spheres (zone of
[1992] used a similar approach for schools of fish. repulsion, zone of orientation and zone of attraction)
Some other models included a more realistic represen- are Rr , Ro and Ra , respectively. Thus, the width
tation of body size and shape [Kunz and Hemelrijk, of the two outer annulus are Ro = Ro Rr and
2003]. Ra = Ra Ro . denotes the field of perception,
Regarding the methodology of most of the simula- thus, the blind volume is behind the individual,
tions, the agent-based (or individual-based ) approach with interior angle (360 ).
proved to be very popular (although there are al- In order to explore the global system behavior, the
ternative approaches as well, e.g., Hayakawa [2010]). authors analyzed the consequences of varying certain
The reason behind this is that this approach pro- system parameters, like the number of individuals,
vides a link between the behavior of the individuals preferred speed, turning rate, width of the zones, etc.
and the emergent properties of the swarm as a whole, For every case, the following two global properties
thus appropriate to investigate how the properties of were calculated:
the system depend on the actual behavior of the in-
dividuals. The most common rules applied in these
N
1 X
~u
models are: (i) short-range repulsive force aiming to (t) = vi (t) , (54)
N
avoid collision with mates and with the borders, (ii)
i=1
adjusting the velocity vector according to the direc-
where N is the number of individuals within the
tion of the neighboring units, (iii) a force avoiding
group, (i = 1, 2, . . . , N ), and v~iu (t) is the unit direc-
being alone, e.g., moving towards the center of the
tion vector of the ith animal at time t. (Since v~iu (t)
swarms mass, (iv) noise, (v) some kind of drag force
is a unit vector, the expression defined by Eq. (54) is
if the medium is considered in which the individuals
equivalent with the order parameter defined by Eq.
move (which is usually not taken into account, in case
(1).)
of birds and fish). Then, the concrete models differ in
the rules they apply (usually most of the above ones), The other measure, group angular momentum,
in their concrete form and in the system parameters. is the sum of the angular momenta of the group-
Some biologically more realistic, yet still sim- members about the center of the group, ~rGr . This ex-
ple individual-based models were also suggested pression measures the degree of rotation of the group
[Couzin et al., 2002, Hemelrijk and Hildenbrandt, about its center
2008]. In a computer model called StarDisplay,
N
Hildenbrandt et al. [2010] combined the above men- 1 X
mGr (t) = ~riGr (t) v~i (t) ,
u (55)
tioned usual rules with system specific ones in or- N
i=1
der to generate patterns that resemble to the aerial
displays of starlings, recorded by the Starflag project where ~riGr = ~ri ~rGr , is the vectorial difference of
(see Sec. 3.7). the position of individual i, r~i , and the position of
Couzin et al. [2002] categorized the emergent col- the group-center, ~rGr .
5.3 Models inspired by animal behavioral patterns 65
and demonstrate that they reproduce the observed within the interaction range . If so, the desired di-
behavior. (On the other hand, Couzin et al. [2002] rection is defined as
demonstrated that vice versa the same rule and
parameter set may result in different collective behav-
X ~rj (t) ~ri (t) X ~vj (t)
ior in the very same system, depending on its recent d~i (t + t) = + . (58)
past, history.) Yates et al. [2009] note that many |~rj (t) ~ri (t)| |~vj (t)|
j6=i j6=i
models have weak predictive power concerning the
various relevant aspects of collective motion, includ- We will use the corresponding unit vector, di (t) =
ing, for example, the rate at which groups suddenly ~
di (t)/|d~i (t)|.
change their direction.
Until this point the algorithm is very similar to
the one described in Sec. 5.3.2. In order to study
5.4 The role of leadership in consen- the influence of informed individuals, a portion of
the group, p, is given information about a preferred
sus finding
direction, described by the unit vector ~g . The rest of
Animals traveling together have to develop a method the group is naive, without any preferred direction.
to make collective decisions regarding the places of Informed individuals balance their social alignment
foraging, resting and nesting sites, route of migration, and their preferred direction with the weighting fac-
etc. By slightly modifying (typically extending) mod- tor
els (such as the one described in Sec. 5.3.2), a group
of individuals can get hold of such abilities. Accord- di (t + t) + ~gi
d~i (t + t) = . (59)
ingly, Couzin et al. [2005] suggested a simple model di (t + t) + ~gi
in which individuals were not required to know how
many and which individuals had information, they ( can exceed 1; in this case the individual is influ-
did not need to have a signaling mechanism and no enced more heavily by its own preferences than by
individual recognition was required from the group its mates.) The accuracy of the group (describing
members. Informed individuals were not necessitated the quality of information transfer) is characterized
to know anything about the information-level of their by the normalized angular deviation of the group di-
mates either and that how the quality of their infor- rection around the preferred direction ~g, similarly to
mation was compared to that of others. The model the term given in Eq. 55.
looks as follows: The authors found that for fixed group size, the
N individuals compose the group. The position accuracy increases asymptotically as the portion p of
of the ith particle is described by the vector ~ri , and the informed members increased, see Fig. 62. This
it is moving in the direction ~vi . The group mem- means, that the larger the group, the smaller the por-
bers endeavor to continually maintain a minimum tion of informed members is needed, in order to guide
distance, , among themselves, by turning away from the group towards a preferred direction.
the neighbors j which are within this range towards However, informed individuals might also differ in
the opposite direction, described by the desired di- their preferred direction. If the number of individuals
rection d~i preferring one or another direction is equal, than the
group direction will depend on the degree to which
X ~rj (t) ~ri (t)
~
di (t + t) = (57) these preferred directions differ from each other: if
|~rj (t) ~ri (t)| these preferences are similar, then the group will go
j6=i
in the average preferred direction of all informed in-
This rule has the highest priority. If there are dividuals. As the differences among the preferred di-
no mates within this range, than the individual will rections increase, individuals start to select randomly
attempt to align with those neighbors j, which are one or another preferred direction. If the number of
5.5 Relationship between observations and models 67
teristics of flocking. In fact, the aim of these models theoretical/modeling interpretations can be estab-
is to give a minimal model, i.e., a description, includ- lished.
ing those and only those rules which are inevitable
for the emergence of collective motion. All other as-
pects are neglected, by design. The question these 6 Summary and conclusions
studies aim to clarify is the following: What are the
necessary conditions for collective motion to appear From the continuously growing number of exciting
and how the emergent behavior is affected by these new publications on flocking we are tempted to con-
terms? clude that collective motion can be regarded as an
On the other hand, Sec. 5 comprises models which emerging field on the borderline of several scientific
aim to apprehend a well-described observation as disciplines. Thus, it is a multidisciplinary area with
adequately as possible. In fact, a model is con- many applications, involving statistical physics, tech-
sidered to describe a phenomenon properly if it is nology and branches of life sciences. Because of the
capable of giving predictions, that is, the behav- nature of the problem (treating many similar enti-
ior of the observed system can be foretold by ap- ties) studies in this field make quantitative compari-
plying the model. Since models ensure a deep in- son with observations possible even for living systems
sight into the observed phenomenon, many authors and there is a considerable potential for constructing
prefer to propose straight away a model accompa- theoretical approaches.
nying their experimental findings ([Keller and Segel, The results we have presented support a deep anal-
1971, Czirok et al., 1996, Wu et al., 2009, 2007, ogy with equilibrium statistical physics. The essen-
Czirok et al., 2001, Rappel et al., 1999, Szabo et al., tial deviation from equilibrium is manifested in the
2006, Couzin and Franks, 2003, Buhl et al., 2006, collision rule: since the absolute velocity of the par-
Rose et al., 2009]) while other models give account ticles is preserved and in most cases an alignment of
for previously reported observations. As such, the direction of motion after interaction is preferred,
Wolgemuth [2008] developed a two-phase model in the total momentum increases both during individ-
order to describe the jet-like patterns and vor- ual collisions and, as a result, gradually in the whole
tices appearing in dense colonies of Bacillus subtilis system of particles as well.
([Mendelson et al., 1999, Dombrowski et al., 2004]), The observations we have discussed can be suc-
Belmonte et al. [2008] proposed a model which gives cessfully interpreted in terms of simple simulational
account for the cell-sorting phenomenon observed in models. Using models based on simplified units (also
Hydra cells by Rieu et al. [2000], or we can men- called particles) to simulate the collective behavior
tion Vab and Skaret [2008] who modeled the unique of large ensembles has a history in science, especially
forms of herring schools observed by Axelsen et al. in statistical physics, where originally particles rep-
in 2000. resented atoms or molecules. With the rapid increase
Sometimes the theoretical calculations come be- of computing power and a growing appreciation for
fore the observations, i.e., the given model predicts understanding through simulations, models based
a previously unreported phenomenon. As an exam- on a plethora of complex interacting units, nowadays
ple, Toner and Tu predicted the phenomenon of giant widely called agents, have started to emerge. Agents,
number fluctuation in their 1995, 98 paper, which was even those that follow simple rules, are more complex
observed more than a decade later by Narayan et al. entities than particles because they have a goal they
[2007]. These cases lend strong support for the mod- intend to achieve in an optimal way (for example,
els giving the predictions. using as little amount of resources as possible).
Overall, we conclude that a considerable number As a rule, models of increasing complexity are
of evidences have accumulated each demonstrating bound to be born in order to account for the inter-
that both qualitative and quantitative agreements esting variants of a fundamental process. However,
between observations (of collective motion) and their there is a catch. A really good model must both re-
69
produce truly life-like behavior and be as simple as Critical (flocks of all sizes moving coherently in
possible. If the model has dozens of equations and different directions. The whole system is very
rules, and correspondingly large numbers of param- sensitive to perturbations)
eters, it is bound to be too specific and rather like
an imitation than a model that captures the few Quasi-long range velocity correlations and rip-
essential features of the process under study. ples
Thus, if a model is very simple, it is likely to be ap- Jamming
plicable to other phenomena for which the outcome is
dominated by the same few rules. On the other hand, A few further, less widely-occurring patterns are also
simplification comes with a price, and some of the possible, for example in systems made of two or more
exciting details that distinguish different phenomena distinctively different types of units.
may be lost. But, as any fan of natural life movies The following types of transitions between the
knows, collective behavior in nature typically involves above collective motion classes are possible:
sophisticated, occasionally amazing techniques aimed
Continuous (second order, accompanied with
at coordinating the actions of the organisms to max-
large fluctuations and algebraic scaling)
imize success. The art of designing models of reality
is rooted in the best compromise between oversimpli- Discontinuous (first order)
fication and including too many details (eventually
preventing the location of the essential features). No singularity in the level of directedness
On the basis of the numerous observations and Jamming (transition to a state in which mobility
models/simulations we have discussed above, the fol- is highly restricted)
lowing conclusions can be made concerning the gen-
eral features of systems exhibiting collective motion: The above transitions usually take place as i) a
function of density, or, ii) the changing magnitude
Most patterns of collective motion are universal of perturbations the units are subject to. The role of
(the same patterns occur in very different sys- noise is essential; all systems are prone to be strongly
tems) influenced by perturbations. In some cases noise can
have a paradoxical effect and, e.g., facilitate ordering.
Simple models can reproduce this behavior
This could be understood, for example, as a result of
A simple noise term can account for numerous perturbations driving the system out from an inef-
complex deterministic factors ficient deterministic regime (particles moving along
trajectories systematically (deterministically) avoid-
Global ordering is due to non-conservation ing each other) into a more efficient one, character-
of the momentum during individual colli- ized by an increased number of interactions.
sions/interactions of pairs of units After reviewing the state of the art regarding col-
lective motion, we think that some of the most ex-
The universally occurring patterns can be divided citing challenges in this still emerging field can be
into a few classes of motion patterns: summarized as: i) Additional, even more precise data
about the positions and velocities of the collectively
Disordered (particles moving in random direc-
moving units should be obtained for establishing a
tions)
well defined, quantitative set of interaction rules typ-
Fully-ordered (particles moving in the same di- ical for most of the flocks. ii) the role of leadership
rection) in collective decision making should be further ex-
plored. Is it hierarchical? Can a leadership-driven
Rotational (within a rectangular or circular decision-making mechanism be scalabe up to huge
area) group sizes? iii) The problem of a coherently-moving,
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