Turing pattern - Wikipedia [Link]
org/wiki/Turing_pattern
Turing pattern
The Turing pattern is a concept introduced by
English mathematician Alan Turing in a 1952
paper titled "The Chemical Basis of
Morphogenesis", which describes how patterns
in nature, such as stripes and spots, can arise
naturally and autonomously from a
[1][2] Three examples of Turing patterns
homogeneous, uniform state. The pattern
arises due to Turing instability, which in turn
arises due to the interplay between differential
diffusion of chemical species and chemical
reaction.[3] The instability mechanism is
surprising because a pure diffusion, such as
molecular diffusion, would be expected to have a
stabilizing influence on the system (i.e., complete
mixing).
Six stable states from Turing equations, the last one
forms Turing patterns
Overview
In his paper,[1] Turing examined the behaviour of a system in which two diffusible substances
interact with each other, and found that such a system is able to generate a spatially periodic
pattern even from a random or almost uniform initial condition.[4] Prior to the discovery of this
instability mechanism arising due to unequal diffusion coefficients of the two substances,
diffusional effects were always presumed to have stabilizing influences on the system.
Turing hypothesized that the resulting wavelike patterns are the chemical basis of morphogenesis.
[4]Turing patterning is often found in combination with other patterns: vertebrate limb
development is one of the many phenotypes exhibiting Turing patterning overlapped with a
complementary pattern (in this case a French flag model).[5]
Before Turing, Yakov Zeldovich in 1944 discovered this instability mechanism in connection with
the cellular structures observed in lean hydrogen flames.[6] Zeldovich explained the cellular
structure as a consequence of hydrogen's diffusion coefficient being larger than the thermal
diffusion coefficient. In combustion literature, Turing instability is referred to as diffusive–thermal
instability.
Concept
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The original theory, a reaction–diffusion theory of
morphogenesis, has served as an important model in
theoretical biology.[7] Reaction–diffusion systems have
attracted much interest as a prototype model for pattern
formation. Patterns such as fronts, hexagons, spirals,
stripes and dissipative solitons are found as solutions of
Turing-like reaction–diffusion equations.[8]
Turing proposed a model wherein two homogeneously
distributed substances (P and S) interact to produce stable
patterns during morphogenesis. These patterns represent
regional differences in the concentrations of the two
A Turing bifurcation pattern
substances. Their interactions would produce an ordered
structure out of random chaos.[9]
In Turing's model, substance P promotes the production of
more substance P as well as substance S. However,
substance S inhibits the production of substance P; if S
diffuses more readily than P, sharp waves of concentration
differences will be generated for substance P. An important
feature of Turing's model is that particular wavelengths in
the substances' distribution will be amplified while other
wavelengths will be suppressed.[9]
The parameters depend on the physical system under
consideration. In the context of fish skin pigmentation, the
associated equation is a three field reaction–diffusion one
in which the linear parameters are associated with An example of a natural Turing pattern on
a giant pufferfish
pigmentation cell concentration and the diffusion
parameters are not the same for all fields.[10] In dye-doped
liquid crystals, a photoisomerization process in the liquid crystal matrix is described as a reaction–
diffusion equation of two fields (liquid crystal order parameter and concentration of cis-isomer of
the azo-dye).[11] The systems have very different physical mechanisms on the chemical reactions
and diffusive process, but on a phenomenological level, both have the same ingredients.
Turing-like patterns have also been demonstrated to arise in developing organisms without the
classical requirement of diffusible morphogens. Studies in chick and mouse embryonic
development suggest that the patterns of feather and hair-follicle precursors can be formed without
a morphogen pre-pattern, and instead are generated through self-aggregation of mesenchymal
cells underlying the skin.[12][13] In these cases, a uniform population of cells can form regularly
patterned aggregates that depend on the mechanical properties of the cells themselves and the
rigidity of the surrounding extra-cellular environment. Regular patterns of cell aggregates of this
sort were originally proposed in a theoretical model formulated by George Oster, which postulated
that alterations in cellular motility and stiffness could give rise to different self-emergent patterns
from a uniform field of cells.[14] This mode of pattern formation may act in tandem with classical
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reaction-diffusion systems, or independently to generate patterns in biological development.
Turing patterns may also be responsible for the formation of human fingerprints.[15]
As well as in biological organisms, Turing patterns occur in other natural systems – for example,
the wind patterns formed in sand, the atomic-scale repetitive ripples that can form during growth
of bismuth crystals, and the uneven distribution of matter in galactic disc.[16][17] Although Turing's
ideas on morphogenesis and Turing patterns remained dormant for many years, they are now
inspirational for much research in mathematical biology.[18] It is a major theory in developmental
biology; the importance of the Turing model is obvious, as it provides an answer to the
fundamental question of morphogenesis: "how is spatial information generated in organisms?".[4]
Turing patterns can also be created in nonlinear optics as demonstrated by the Lugiato–Lefever
equation. Reaction-diffusion models can be used to forecast the exact location of the tooth cusps in
mice and voles based on differences in gene expression patterns.[9] The model can be used to
explain the differences in gene expression between mice and vole teeth, the signaling center of the
tooth, enamel knot, secrets BMPs, FGFs and Shh. Shh and FGF inhibits BMP production, while
BMP stimulates both the production of more BMPs and the synthesis of their own inhibitors.
BMPs also induce epithelial differentiation, while FGFs induce epithelial growth.[19] The result is a
pattern of gene activity that changes as the shape of the tooth changes, and vice versa. Under this
model, the large differences between mouse and vole molars can be generated by small changes in
the binding constants and diffusion rates of the BMP and Shh proteins. A small increase in the
diffusion rate of BMP4 and a stronger binding constant of its inhibitor is sufficient to change the
vole pattern of tooth growth into that of the mouse.[19][20]
Experiments with the sprouting of chia seeds planted in trays have confirmed Turing's
mathematical model.[21]
Classic example: radiolarian shells
Turing wanted to advance the work D'Arcy Thompson published in 1917 called On Growth and
Form.[22] Bernard Richards, working under the supervision of Turing at Manchester as one of
Turing's last students, helped validate Turing's theory of morphogenesis as follows:[23][24][22][25]
So I set to work on seeking a solution to the Morphogenesis Equations on a
sphere. The theory was that a spherical organism was subject to diffusion across
its surface membrane by an alien substance, eg sea-water. The Equations were:
The function , taken to be the radius vector from the centre to any point on the
surface of the membrane, was argued to be representable as a series of
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normalised Legendre functions. The algebraic solution of the above equations ran
to some 30 pages in my Thesis and are therefore not reproduced here. They are
written in full in the book entitled "Morphogenesis" which is a tribute to Turing,
edited by P. T. Saunders, published by North Holland, 1992.[26]
The algebraic solution of the equations revealed a family of solutions,
corresponding to a parameter n, taking values 2, 4, 6. When I had solved the
algebraic equations, I then used the computer to plot the shape of the resulting
organisms. Turing told me that there were real organisms corresponding to what
I had produced. He said that they were described and depicted in the records of
the voyages of HMS Challenger in the 19th Century.
I solved the equations and produced a set of solutions which corresponded to the
actual species of Radiolaria discovered by HMS Challenger. That expedition to
the Pacific Ocean found eight variations in the growth patterns. These are shown
in the following figures (below). The essential feature of the growth is the
emergence of elongated "spines" protruding from the sphere at regular positions.
Thus the species comprised two, six, twelve, and twenty, spine variations.
— Bernard Richards, 2006 [27]
The images below show relevant spine variations of radiolarians as extracted from drawings made
by the German zoologist and polymath Ernst Haeckel in 1887.[27]
Cromyatractus Circopus sexfurcus with 6 Circopurus octahedrus
tetracelyphus with 2 spines with 6 spines and 8
spines faces
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Circogonia icosahedra Circorrhegma Cannocapsa
with 12 spines and 20 dodecahedra with 20 stethoscopium with 20
faces (incompletely drawn) spines
spines and 12 faces
Radiolarians are unicellular predatory protists encased in
elaborate globular shells (or "capsules"), usually made of
silica and pierced with holes. Their name comes from the
Latin for "radius". They catch prey by extending parts of
their body through the holes. As with the silica frustules of
diatoms, radiolarian shells can sink to the ocean floor
when radiolarians die and become preserved as part of the
ocean sediment. These remains, as microfossils, provide
valuable information about past oceanic conditions.[28]
Turing and radiolarian morphology
Some drawings by Ernst Haeckel of
radiolarian shells discovered by HMS
Challenger in the 19th Century
Shell of a spherical radiolarian Shell micrographs
Computer simulations of Turing patterns on a sphere
closely replicate some radiolarian shell patterns[29]
Biological application
A mechanism that has gained increasing attention as a generator of spot- and stripe-like patterns
in developmental systems is related to the chemical reaction-diffusion process described by Turing
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in 1952. This has been schematized in a biological "local
autoactivation-lateral inhibition" (LALI) framework by
Meinhardt and Gierer.[31] LALI systems, while formally
similar to reaction-diffusion systems, are more suitable to
biological applications, since they include cases where the
activator and inhibitor terms are mediated by cellular
"reactors" rather than simple chemical reactions,[32] and
spatial transport can be mediated by mechanisms in
addition to simple diffusion.[33] These models can be
applied to limb formation and teeth development among
other examples.
Simulations of effect of limb bud distal
expansion[30]
See also
▪ Evolutionary developmental biology
▪ Mathematical and theoretical biology
▪ Patterns in nature
▪ Reaction–diffusion system
▪ Spontaneous symmetry breaking
▪ Vermiculation
References
1. Turing, Alan (1952). "The Chemical Basis of Morphogenesis" ([Link]
ses/cs191/paperscs191/[Link]) (PDF). Philosophical Transactions of the Royal Society of
London B. 237 (641): 37–72. Bibcode:1952RSPTB.237...37T ([Link]
s/1952RSPTB.237...37T). doi:10.1098/rstb.1952.0012 ([Link]
12). JSTOR 92463 ([Link] S2CID 120437796 ([Link]
[Link]/CorpusID:120437796).
2. Stewart, Ian (1998). Life's Other Secret: The New Mathematics of the Living World. London:
Allen Lane. pp. 138–140, 142–146, 148, 149, 151, 152. ISBN 0-713-99161-5. OCLC 43126766
([Link]
3. "Turing patterns, 70 years later" ([Link] Nature
Computational Science. 2 (8): 463–464. 2022. doi:10.1038/s43588-022-00306-0 ([Link]
g/10.1038%2Fs43588-022-00306-0).
4. Kondo, Shigeru (7 February 2017). "An updated kernel-based Turing model for studying the
mechanisms of biological pattern formation" ([Link]
Journal of Theoretical Biology. 414: 120–127. Bibcode:2017JThBi.414..120K ([Link]
[Link]/abs/2017JThBi.414..120K). doi:10.1016/[Link].2016.11.003 ([Link]
6%[Link].2016.11.003). ISSN 0022-5193 ([Link]
PMID 27838459 ([Link]
6 of 10 2/6/2026, 11:46 PM
Turing pattern - Wikipedia [Link]
5. Sharpe, James; Green, Jeremy (2015). "Positional information and reaction-diffusion: two big
ideas in developmental biology combine" ([Link]
Development. 142 (7): 1203–1211. doi:10.1242/dev.114991 ([Link]
4991). hdl:10230/25028 ([Link] PMID 25804733 ([Link]
[Link]/25804733).
6. Zeldovich, Y. B. (1944). Theory of Combustion and Detonation of Gases. Selected Works of
Yakov Borisovich Zeldovich, Volume I: Chemical Physics and Hydrodynamics (pp. 162-232).
Princeton: Princeton University Press
7. Harrison, L. G. (1993). "Kinetic Theory of Living Pattern". Endeavour. 18 (4). Cambridge
University Press: 130–6. doi:10.1016/0160-9327(95)90520-5 ([Link]
0-9327%2895%2990520-5). PMID 7851310 ([Link]
8. Kondo, S.; Miura, T. (23 September 2010). "Reaction-Diffusion Model as a Framework for
Understanding Biological Pattern Formation". Science. 329 (5999): 1616–1620.
Bibcode:2010Sci...329.1616K ([Link]
doi:10.1126/science.1179047 ([Link] PMID 20929839
([Link] S2CID 10194433 ([Link]
CorpusID:10194433).
9. Gilbert, Scott F. (2014). Developmental Biology (Tenth ed.). Sunderland, Massachusetts:
Sinauer Associates. ISBN 978-0-87893-978-7. OCLC 837923468 ([Link]
clc/837923468).
10. Nakamasu, A.; Takahashi, G.; Kanbe, A.; Kondo, S. (11 May 2009). "Interactions between
zebrafish pigment cells responsible for the generation of Turing patterns" ([Link]
[Link]/pmc/articles/PMC2689028). Proceedings of the National Academy of Sciences. 106
(21): 8429–8434. Bibcode:2009PNAS..106.8429N ([Link]
S..106.8429N). doi:10.1073/pnas.0808622106 ([Link]
PMC 2689028 ([Link] PMID 19433782 (http
s://[Link]/19433782).
11. Andrade-Silva, Ignacio; Bortolozzo, Umberto; Clerc, Marcel G.; González-Cortés, Gregorio;
Residori, Stefania; Wilson, Mario (27 August 2018). "Spontaneous light-induced Turing
patterns in a dye-doped twisted nematic layer" ([Link]
6110868). Scientific Reports. 8 (1): 12867. Bibcode:2018NatSR...812867A ([Link]
[Link]/abs/2018NatSR...812867A). doi:10.1038/s41598-018-31206-x ([Link]
38%2Fs41598-018-31206-x). PMC 6110868 ([Link]
10868). PMID 30150701 ([Link]
12. Glover, James D.; Wells, Kirsty L.; Matthäus, Franziska; Painter, Kevin J.; Ho, William; Riddell,
Jon; Johansson, Jeanette A.; Ford, Matthew J.; Jahoda, Colin A. B.; Klika, Vaclav; Mort,
Richard L. (2017). "Hierarchical patterning modes orchestrate hair follicle morphogenesis" (http
s://[Link]/pmc/articles/PMC5507405). PLOS Biology. 15 (7) e2002117.
doi:10.1371/[Link].2002117 ([Link]
PMC 5507405 ([Link] PMID 28700594 (http
s://[Link]/28700594).
13. Shyer, Amy E.; Rodrigues, Alan R.; Schroeder, Grant G.; Kassianidou, Elena; Kumar, Sanjay;
Harland, Richard M. (2017). "Emergent cellular self-organization and mechanosensation
initiate follicle pattern in the avian skin" ([Link]
7). Science. 357 (6353): 811–815. doi:10.1126/science.aai7868 ([Link]
ence.aai7868). PMC 5605277 ([Link]
PMID 28705989 ([Link]
7 of 10 2/6/2026, 11:46 PM
Turing pattern - Wikipedia [Link]
14. Oster, G. F.; Murray, J. D.; Harris, A. K. (1983). "Mechanical aspects of mesenchymal
morphogenesis". Journal of Embryology and Experimental Morphology. 78: 83–125.
PMID 6663234 ([Link]
15. Gaines, James M. "Finally, Scientists Uncover the Genetic Basis of Fingerprints" ([Link]
[Link]/news-opinion/finally-scientists-uncover-the-genetic-basis-of-fingerprints-7098
3). TheScientist. Retrieved 27 February 2023.
16. Fuseya, Yuki; Katsuno, Hiroyasu; Behnia, Kamran; Kapitulnik, Aharon (8 July 2021).
"Nanoscale Turing patterns in a bismuth monolayer" ([Link]
21-01288-y). Nature Physics. 17 (9): 1031–1036. arXiv:2104.01058 ([Link]
4.01058). Bibcode:2021NatPh..17.1031F ([Link]
31F). doi:10.1038/s41567-021-01288-y ([Link]
ISSN 1745-2481 ([Link] S2CID 237767233 ([Link]
[Link]/CorpusID:237767233).
17. Smolin, Lee (3 December 1996). "Galactic disks as reaction-diffusion systems". arXiv:astro-
ph/9612033 ([Link]
18. Woolley, T. E., Baker, R. E., Maini, P. K., Chapter 34, Turing's theory of morphogenesis. In
Copeland, B. Jack; Bowen, Jonathan P.; Wilson, Robin; Sprevak, Mark (2017). The Turing
Guide. Oxford University Press. ISBN 978-0-19-874782-6.
19. Salazar-Ciudad, Isaac; Jernvall, Jukka (March 2010). "A computational model of teeth and the
developmental origins of morphological variation" ([Link]
8). Nature. 464 (7288): 583–586. Bibcode:2010Natur.464..583S ([Link]
abs/2010Natur.464..583S). doi:10.1038/nature08838 ([Link]
. ISSN 1476-4687 ([Link] PMID 20220757 ([Link]
[Link]/20220757). S2CID 323733 ([Link]
33).
20. Salazar-ciudad, Isaac; Jernvall, Jukka (January 2004). "How different types of pattern
formation mechanisms affect the evolution of form and development". Evolution and
Development. 6 (1): 6–16. doi:10.1111/j.1525-142x.2004.04002.x ([Link]
j.1525-142x.2004.04002.x). ISSN 1520-541X ([Link]
PMID 15108813 ([Link] S2CID 1783730 ([Link]
[Link]/CorpusID:1783730).
21. Riordon, James R. (26 March 2023). "Chia seedlings verify Alan Turing's ideas about patterns
in nature" ([Link]
ScienceNews.
22. Richards, Bernard (2017). "Chapter 35 – Radiolaria: Validating the Turing theory". In Copeland,
Jack; et al. (eds.). The Turing Guide. pp. 383–388.
23. Richards, Bernard (1954), "The Morphogenesis of Radiolaria", MSc thesis, Manchester, UK:
The University of Manchester
24. Richards, Bernard (2005). "Turing, Richards and morphogenesis" ([Link]
[Link]/[Link]). The Rutherford Journal. 1.
25. Copeland, Jack; Bowen, Jonathan; Sprevak, Mark; Wilson, Robin; et al. (2017). "Notes on
Contributors". The Turing Guide. Oxford University Press. p. 478. ISBN 978-0-19-874783-3.
26. Turing, Alan; Saunders, P. T. (1992). Morphogenesis (in Esperanto). Amsterdam: North-
Holland. ISBN 978-0-08-093405-1. OCLC 680063781 ([Link]
781).
27. Richards, Bernard (2006) "Turing, Richards and Morphogenesis" ([Link]
b/20060816084810/[Link] The Rutherford Journal,
Volume 1.
8 of 10 2/6/2026, 11:46 PM
Turing pattern - Wikipedia [Link]
28. Wassilieff, Maggy (2006) "Plankton – Animal plankton" ([Link]
h/5138/radiolarian-fossils), Te Ara – the Encyclopedia of New Zealand. Accessed: 2 November
2019.
29. Varea, C.; Aragon, J.L.; Barrio, R.A. (1999). "Turing patterns on a sphere". Physical Review E.
60 (4): 4588–92. Bibcode:1999PhRvE..60.4588V ([Link]
E..60.4588V). doi:10.1103/PhysRevE.60.4588 ([Link]
PMID 11970318 ([Link]
30. Zhu, Jianfeng; Zhang, Yong-Tao; Alber, Mark S.; Newman, Stuart A. (28 May 2010). Isalan,
Mark (ed.). "Bare Bones Pattern Formation: A Core Regulatory Network in Varying Geometries
Reproduces Major Features of Vertebrate Limb Development and Evolution" ([Link]
[Link]/pmc/articles/PMC2878345). PLOS ONE. 5 (5) e10892.
Bibcode:2010PLoSO...510892Z ([Link]
doi:10.1371/[Link].0010892 ([Link]
ISSN 1932-6203 ([Link] PMC 2878345 ([Link]
[Link]/pmc/articles/PMC2878345). PMID 20531940 ([Link]
531940).
31. Meinhardt, Hans (2008), "Models of Biological Pattern Formation: From Elementary Steps to
the Organization of Embryonic Axes", Multiscale Modeling of Developmental Systems, Current
Topics in Developmental Biology, vol. 81, Elsevier, pp. 1–63, doi:10.1016/
s0070-2153(07)81001-5 ([Link]
ISBN 978-0-12-374253-7, PMID 18023723 ([Link]
32. Hentschel, H. G. E.; Glimm, Tilmann; Glazier, James A.; Newman, Stuart A. (22 August 2004).
"Dynamical mechanisms for skeletal pattern formation in the vertebrate limb" ([Link]
[Link]/pmc/articles/PMC1691788). Proceedings of the Royal Society of London. Series
B: Biological Sciences. 271 (1549): 1713–1722. doi:10.1098/rspb.2004.2772 ([Link]
0.1098%2Frspb.2004.2772). ISSN 0962-8452 ([Link]
PMC 1691788 ([Link] PMID 15306292 (http
s://[Link]/15306292).
33. Lander, Arthur D. (January 2007). "Morpheus Unbound: Reimagining the Morphogen Gradient"
([Link] Cell. 128 (2): 245–256. doi:10.1016/
[Link].2007.01.004 ([Link] ISSN 0092-8674 ([Link]
[Link]/issn/0092-8674). PMID 17254964 ([Link]
S2CID 14173945 ([Link]
Further reading
▪ Ball, Philip (31 May 2012). "Turing Patterns" ([Link]
tterns/[Link]). Chemistry World. (See also extended version ([Link]
mages/stories/docs/pdf/Turing_long.pdf), June 2012.)
▪ Campagna, R.; Cuomo, S.; Giannino, F.; Severino, G.; Toraldo, G. (6 December 2017). "A
semi-automatic numerical algorithm for Turing patterns formation in a reaction-diffusion model"
([Link] IEEE Access. 6: 4720–4724.
doi:10.1109/ACCESS.2017.2780324 ([Link]
▪ Iber, Bagnar. "Turing Pattern" ([Link]
e/cobi-dam/documents/Spat-Temp_Modelling/Slides_Lecture3_SpatialModels.pdf) (PDF).
Computational Biology (CoBI). Switzerland: ETH Zurich. Retrieved 16 August 2018.
▪ Keim, Brandon (22 February 2011). "Alan Turing's Patterns in Nature and Beyond" ([Link]
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Turing pattern - Wikipedia [Link]
[Link]/2011/02/turing-patterns/). Wired.
▪ Ouellette, Jennifer (27 March 2013). "When Math Meets Nature: Turing Patterns and Form
Constants" ([Link]
e-turing-patterns-and-form-constants/). Scientific American.
▪ Thompson, D'Arcy Wentworth (1942) [1917]. On Growth and Form. Cambridge University
Press.
▪ Mirfendereski, Siamak; Gupta, Ankur (27 October 2025). "Imperfect Turing patterns:
Diffusiophoretic assembly of hard spheres via reaction-diffusion instabilities" ([Link]
0.1016%[Link].2025.102513). Matter (102513). doi:10.1016/[Link].2025.102513 ([Link]
rg/10.1016%[Link].2025.102513).
Retrieved from "[Link]
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