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Supporting Online Material
www.sciencemag.org/cgi/content/full/science.1177088/DC1
SOM Text
Figs. S1 to S8
Tables S1 and S2
References and Notes
1 June 2009; accepted 1 December 2009
Published online 10 December 2009;
10.1126/science.1177088
Include this information when citing this paper.
Rules for Biologically Inspired
Adaptive Network Design
Atsushi Tero,
1,2
Seiji Takagi,
1
Tetsu Saigusa,
3
Kentaro Ito,
1
Dan P. Bebber,
4
Mark D. Fricker,
4
Kenji Yumiki,
5
Ryo Kobayashi,
5,6
Toshiyuki Nakagaki
1,6
*
Transport networks are ubiquitous in both social and biological systems. Robust network performance
involves a complex trade-off involving cost, transport efficiency, and fault tolerance. Biological
networks have been honed by many cycles of evolutionary selection pressure and are likely to yield
reasonable solutions to such combinatorial optimization problems. Furthermore, they develop without
centralized control and may represent a readily scalable solution for growing networks in general. We
show that the slime mold Physarum polycephalum forms networks with comparable efficiency, fault
tolerance, and cost to those of real-world infrastructure networks—in this case, the Tokyo rail system.
The core mechanisms needed for adaptive network formation can be captured in a biologically
inspired mathematical model that may be useful to guide network construction in other domains.
T
ransport networks are a critical part of the
infrastruc ture needed to operate a modern
industrial society and facilitate efficient
movement of people, resources, energy, and
informati on. Despite their importance, most net-
works have emerged without clear global design
principles and are constrained by the priorities
impose d at thei r initia tion. Thus, the main motiva -
tion historical ly was to achieve high transpo rt
efficiency at reasonable cost, but with correspond-
ingly less emphasi s on makin g system s toler ant to
interruption or failure. Introducing robustness
inevit ably requ ires add ition al redund ant pathw ays
that are not cost-effective in the short term. In recent
years, the spectacular failure of key infrastructure
such as power grids (1, 2), financial systems (3, 4),
airline baggag e-handling systems (5), and railway
networks(6), as well asthepredictedvulnerabi lityof
systems such as information networks (7)orsupply
networks (8)toattack,havehighlightedtheneedto
develop networks with greater intrinsic resilience.
Some organisms grow in the form of an inter-
connected network as part of their normal forag-
ing strategy to discover and exploit new resources
(9–12 ). Such systems continuously adapt to their
environment and must balance the cost of produc-
ing an efficie nt network with the consequences of
even limited failure in a competitive world. Unlike
anthropogenic infrastructure systems, these biolog-
ical networks have been subjected to successive
rounds of evolutionary selection and are likely to
have reached a point at which cost, efficiency, and
resilience are appropriately balanced. Drawing in-
spiration from biology has led to useful approaches
to problem-solving such as neural networks, ge-
netic algorithms, and efficient search routines de-
veloped from ant colony optimization algorithms
(13). We exploited the slime mold Physarum
polycephalum to develop a biologically inspired
model for adaptive network development.
Physarum is a large, single-celled amoeboid
organism that forages for patchily distributed
food sources. The individual plasmodium ini-
tially explores with a relatively contiguous for-
aging margin to maximize the area searched.
However , behind the margin, this is resolved into
atubularnetworklinkingthediscoveredfood
sources through direct connections, additional in-
termediate junctions (Steiner points) that reduce
the overall length of the connecting network,
and the formation of occasional cross-links that
improve overall transport efficiency and resil-
ience (11, 12). The growth of the plasmodium is
influenced by the characteristics of the sub-
strate (14) and can be constrained by physical
barriers (15) or influenced by the light regime
(16), facilitating experiment al investigat ion of
the rules underlying network formation. Thu s,
for example, Physarum can find the shortest
path through a maz e (15–17) or connect dif-
ferent arrays of food sources in an efficient
manner with low total length (TL) yet short
average minimum distance (MD) between pairs
of food sources (FSs), with a high degree of
fault tolerance (FT) to accidental disconnection
(11, 18, 19). Capturing the essence of this sys-
tem in simple rules might be useful in guiding
the development of decentralized networks in
other domains.
We observed Physarum connecting a template
of 36 FSs that represented geographical locations
of cities in the T okyo area, and compared the result
with the actual rail network in Japan. The
Physarum plasmodium was allowed to grow from
Tokyo and initially filled much of the available
land space, but then concentrated on FSs by
thinning out the network to leave a subset of larger ,
interconnecting tubes (Fig. 1). An alternative
protocol, in which the plasmodium was allowed
to extend fully in the available space and the FSs
were then presented simultaneously, yielded sim-
ilar results. T o complete the network formation, we
allowed any excess volume of plasmodium to
1
Research Institute for Electronic Science, Hokkaido University,
Sapporo 060-0812, Japan.
2
PRESTO, JST, 4-1-8 Honcho,
Kawaguchi, Saitama, Japan.
3
Graduate School of Engineering,
Hokkai do Universi ty, Sapporo 060-8628, Japan.
4
Department of
Plant Sciences, University of Oxford, Oxford OX1 3RB, UK.
5
Department of Mathematical and Life Sciences, Hiroshima
University, Higashi-Hiroshima 739-8526, Japan.
6
JST, CREST, 5
Sanbancho, Chiyoda-ku, Tokyo, 102-0075, Japan.
*To whom correspondence should be addressed. E-mail:
nakagaki@es.hokudai.ac.jp
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