Research Activities

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Joe Yuichiro Wakano

School of Interdisciplinary Mathematical Sciences (IMS)
Graduate School of Advanced Mathematical Sciences (AMS)
Meiji Institute for Advanced Study of Mathematical Sciences (MIMS)

Meiji University
4-21-1 Nakano, Nakano-ku, Tokyo, Japan 164-8525

How to visit Nakano campus

E-mail: joe [at]
E-mail: joewakano [at]

TEL/PH +81-3-5343-8351



Two email addresses (gmail and go to the same mailbox. I usually send from gmail just because of convenience.

My old email address ( joe [at] ) is obsolete.

Never forget the 0311 disaster (the 2011 Tohoku Earthquake):
We, Japanese people, are very grateful for many aids (physical, financial and most importantly spiritual) from all over the world.

Research Interests

Mathematical Biology
(evolutionary game theory, behavioral ecology, evolution of social/individual learning, evolution of cooperation, pattern formation, reaction diffusion equations, etc.)
See my publication list

Unified understanding of two major theories of biological evolution

Inclusive Fitness Theory (IFT) and Adaptive Dynamics Theory (ADT) are two major theories of evolution. IFT has contributed toward understanding the evolution of cooperation since 1960s, while ADT has contributed toward understanding the mechanism of speciation since 1990s. In the 21st century, IFT has developed very rapidly and now IFT is applicable to various phenomena that have been studied by ADT and other specific theories. This research project aims at the unified understanding of IFT and ADT, with the ultimate goal of constructing the unified theory of evolution [28, 31, 33, 34, 37].

The project is supported by JST PRESTO from 2009 to 2012.

Evolution of Cooperation

Although Darwinian evolution is the maximization of individual fitness, cooperative behavior is widely observed from bacteria to human society. Thus, evolution of cooperation has been a major topic in evolutionary biology. I mainly target at public goods games where multiple individuals share the common resource and large-scale cooperation is required to maximize the society's benefit [2, 7, 14, 17, 19, 21, 23, 27, 31, 33].

Evolution of Learning Abilities in Human

What characterizes human being? Among several candidates such as bipedalism or naked skin, our outstanding learning ability supported by increased brain size is the most impressive. Social learning (imitation of behaviors from parents/social members) is the fundamental ability to form human culture. Successive accumulation of individual inventions enables us to establish our culture, including arts, science and technology. Individual learning ability is also quite important, when we observe archeaological records of Neanderthals who have faithfully inherited Mousterian culture for long years and of modern humans who show an amazing explosion of cultural evolution. Evolutionary origin of such learning ability is of great interest [10, 11, 15, 18, 26, 29, 30, 32, 35, 36].

This study is a part of the JSPS Scientific Research on Innovative Areas "Replacement of Neanderthals by Modern Humans".

Pattern Formation

Complex and beautiful patterns are often observed in nature. Especially, living creatures show various patterns, which is one of the signs of life. Understaning such pattern formation is an important study in both biology and physics. I have studied the pattern formation in bacterial colony and in spatial games. Recent studies include more mathematical topics including Turing-Hopf bifurcation of a subcritial focus in a reaction-diffusion equation and non-continuous trait distribution in a framework of Adaptive Dynamics [4, 5, 6, 9, 14, 17, 21, 24, 25, 28, 37].

Behavioral Ecology

The detailed study on a particular behavior of a model species is very important since hypothesss in the theory of evolution must be tested by experiments. Closely collaborating with experimental and field researchers, I have studied the division of labor in ant society [1] and cannibalism in salamanders [2, 8, 20] so far. Another interesting topic is the evolution of mating system and sex ratio [12, 13, 16].

Curriculum Vitae

PDF version of CV and Research, updated on 22nd May 2013
Date of Birth: 06 May 1973
Nationality: Japanese
Language: Japanese (mother tongue), English (written and spoken)

Meiji University, Professor, Apr 2018 -
Meiji University, Associate Professor, Apr 2013 - Mar 2018
Meiji University, Specially Appointed Associate Professor(*), Sep 2007 - Mar 2013
JSPS (Japan Society for the Promotion of Science), Reseach fellow, Apr 2006 - Sep 2007
University of Tokyo, COE Reseach fellow, Jun 2003 - Mar 2006
University of Tokyo, Technical officer, Apr 2001 - May 2003

(*)Specially Appointed (Associate) Professor is a non-tenured professor. Recently, Japanese Universities tend to hire more professors by limited term contract because the research fund of limited term is increasing while the number of tenure positions is decreased by the policy of the Japanese government.

Doctor of Science, Kyoto University, supervised by Prof. Norio Yamamura, 23 Mar 2001
(Thesis: Adaptation and evolutionary dynamics of social traits of creatures in game-theoretical situations)
Master of Science, Kyoto University, 31 Mar 1998
Bachelor of Science, Kyoto University, 31 Mar 1996

Japanese Society for Mathematical Biology (JSMB)
Japan Physical Society (JPS)
Japan Society for Industrial and Applied Mathematics (JSIAM)
Ecological Society of Japan (ESJ)
The Anthropological Society of Nippon (ASN)
Affiliated Scientist of Center for Ecological Research (Kyoto University)

The 1st JSMB Young Scholar Award (Japanese-Korean joint meeting for mathematical biology, Sep 16-18, 2006, Kyushu Univ)
Marcus W. Feldman Prize 2020, Theoretical Population Biology

Research Grants:

Administrative positions:
Chair of Mathematical Sciences Program, Graduate School of Advanced Mathematical Sciences, Meiji Univ (2014 Apr - 2017 Mar)
Chief secretariat of Japanese Society for Mathematical Biology (2013 Jan - 2014 Dec)

Reseach experiences and skills

Academic background:
Physics (mechanics, electromagnetics, statistical mechanics, field theory, elementary quantum mechanics, elementary general theory of relativity)
Biology (ecology, ethology, evolutionary game theory, population genetics)
Mathematics (linear algebra, complex analysis, dynamical system, bifurcation analysis, center manifold theory, reaction diffusion systems, Markov chain, stochastic differential equation)

Computer skills:
Numerical integration and visualization of reaction diffusion systems (mainly by the original C code, somtimes by FreeFEM++)
Individual- or agent-based simulations
Programming language (mainly by C, sometimes by JAVA, C++, VisualBasic)
Parallel computing by MPI/Open-MP
UNIX based network administration (SMTP, POP, IMAP, WWW, DNS and Samba servers)
LaTeX and MS-Word Equation editor

Long-term (2+ weeks) study experiences in foreign countries:
University of British Columbia, Visiting Associate Professor, Apr 2018 - Mar 2019
University of British Columbia, Mar 2014
Stanford University, Aug 2013
University of British Columbia, Jul-Aug 2012
Universite de Lausanne, Jul 2011
University of British Columbia, Mar 2011
Universite Pierre et Marie CURIE, Universite de Neuchatel, Oct-Nov 2010
University of British Columbia, Apr-May 2009
Program for Evolutionary Dynamics, Harvard University, Jun-Jul 2007
Program for Evolutionary Dynamics, Harvard University, Jun-Jul 2006
Murray State University, Mar 2006
Stanford University, Jan-Feb 2004

Contribution as a reviewer:
Physical Review Letters, The American Naturalist, Journal of Theoretical Biology, Theoretical Population Biology, Evolutionary Ecology, Mathematical Biosciences, Journal of Ethology, Population Ecology, BMC Evolutionary Biology, Current Anthropology, Physica A, International Journal of Dynamical Systems and Differential Equations, Evolution and Human Behavior, Behavioral Ecology, Proceedings B, PNAS, Journal of Royal Society Interface, Journal of Physical Society of Japan, Journal of Mathematical Biology, Chaos Solitons & Fractals, Discrete & Continuous Dynamical System A, Theoretical Ecology, PLOS ONE, Royal Society Oepn Science, Nature, Journal of Archaological Research

Published Papers (peer-reviewed journal)

See Japanese page for publications written in Japanese.
  1. Wakano JY, Nakata K & Yamamura N (1998)
    Dynamic model of optimal age polyethism in social insects under stable and fluctuating environments.
    Journal of Theoretical Biology 193:153-165
    Abstract | Full Text PDF

  2. Wakano JY & Yamamura N (2001)
    A simple learning strategy which realizes robust cooperation better than Pavlov.
    Journal of Ethology 19:1-8
    Abstract | Full Text PDF | Presentation PDF

  3. Wakano JY, Kohmatsu Y & Yamamura N (2002)
    Evolutionary dynamics of frequency-dependent growth strategy in cannibalistic amphibians.
    Evolutionary Ecology Research 4:719-736
    Abstract | Full Text PDF | Presentation PDF

  4. Eiha N, Komoto A, Maenosono S, Wakano JY, Yamamoto K & Yamaguchi Y (2002)
    The mode transition of the bacterial colony.
    Physica A 313:609-624
    Abstract | Full Text PDF

  5. Komoto A, Hanaki K, Maenosono S, Wakano JY, Yamaguchi Y & Yamamoto K (2003)
    Growth dynamics of Bacillus circulans colony.
    Journal of Theoretical Biology 225:91-97.... DOI 10.1016/S0022-5193(03)00224-8
    Abstract | Full Text PDF

  6. Wakano JY, Maenosono S, Komoto A, Eiha N & Yamaguchi Y (2003)
    Self-organized pattern formation of bacteria colony modelled by reaction diffusion system and nucleation theory
    Physical Review Letters 90:258102.... DOI 10.1103/PhysRevLett.90.258102
    Abstract | Full Text PDF

  7. Yamamura N, Higashi M, Behera N & Wakano JY (2004)
    Evolution of mutualism through spatial effects
    Journal of Theoretical Biology 226:421-428.... DOI 10.1016/j.jtbi.2003.9.016
    Abstract | Full Text PDF

  8. Wakano JY (2004)
    Drastic growth effect may explain sympatric cannibalistic polymorphism
    Journal of Theoretical Biology 226:69-77.... DOI 10.1016/j.jtbi.2003.8.005
    Abstract | Full Text PDF | Presentation PDF

  9. Wakano JY, Komoto A & Yamaguchi Y (2004)
    Phase transition of traveling waves in bacterial colony pattern
    Physical Review E 69:051904 .... DOI 10.1103/PhysRevE.69.051904
    Abstract | Full Text PDF

  10. Wakano JY, Aoki K & Feldman MW (2004)
    Evolution of social learning: a mathematical analysis
    Theoretical Population Biology 66:249-258 .... DOI 10.1016/j.tpb.2004.06.005
    Abstract | Full Text PDF

  11. Aoki K, Wakano JY & Feldman MW (2005)
    The emergence of social learning in a temporally changing environment: A theoretical model
    Current Anthropology 46:334-340.... DOI 10.1086/428791
    (Abstract and Full Text not available)

  12. Wakano JY & Ihara Y (2005)
    Evolution of male parental care and female multiple mating: game-theoretical and two-locus diploid models
    The American Naturalist 166:E32-44
    Abstract | Full Text

  13. Wakano JY (2005)
    Evolution of extraordinary female-biased sex ratios: The optimal schedule of sex ratio in local mate competition
    Journal of Theoretical Biology 237:193-202.... DOI 10.1016/j.jtbi.2005.04.006
    Abstract | Full Text PDF

  14. Wakano JY (2006)
    A mathematical analysis on public goods games in the continuous space
    Mathematical Biosciences 201:72-89.... DOI 10.1016/j.mbs.2005.12.015
    Abstract | Full Text PDF

  15. Wakano JY & Aoki K (2006)
    A mixed strategy model for the emergence and intensification of social learning in a periodically changing natural environment
    Theoretical Population Biology 70:486-497.... DOI 10.1016/j.tpb.2006.04.003
    Abstract | Full Text PDF

  16. Seki M, Wakano JY & Ihara Y (2007)
    A theoretical study on the evolution of male parental care and female multiple mating: Effects of female mate choice and male care bias
    Journal of Theoretical Biology 247:281-296.... DOI 10.1016/j.jtbi.2007.03.010

  17. Wakano JY (2007)
    Evolution of cooperation in spatial public goods games with common resource dynamics
    Journal of Theoretical Biology 247:616-622.... DOI 10.1016/j.jtbi.2007.04.008

  18. Wakano JY & Aoki K (2007)
    Do Social Learning and Conformist Bias Coevolve? Henrich and Boyd Revisited
    Theoretical Population Biology 72:504-512.... DOI 10.1016/j.tpb.2007.04.003

  19. Hauert C, Wakano JY & Doebeli M (2008)
    Ecological Public Goods Games: cooperation and bifurcation
    Theoretical Population Biology 73:257-263.... DOI 10.1016/j.tpb.2007.11.007

  20. Wakano JY & Whiteman HH (2008)
    Evolution of polyphenism: the role of density and relative body size on morph determination
    Evolutionary Ecology Research 10:1157-1172....

  21. Wakano JY, Nowak MA & Hauert C (2009)
    Spatial dynamics of ecological public goods
    Proceedings of the National Academy of Sciences of the United States of America 106:7910-7914.... DOI 10.1073/pnas.0812644106
    Full Text PDF | Supplement PDF

    Visit VirtualLabs and let your PC run 2D simulations!

  22. Zu J, Mimura M & Wakano JY (2010)
    The evolution of phenotypic traits in a predator-preysystem subject to Allee effect
    Journal of Theoretical Biology 262:528-543.... DOI 10.1016/j.jtbi.2009.10.022

  23. Kurokawa S, Wakano JY & Ihara Y (2010)
    Generous cooperators can outperform non-generous cooperators when replacing a population of defectors
    Theoretical Population Biology 77:257-262.... DOI 10.1016/j.tpb.2010.03.002

  24. Wakano JY & Hauert C (2011)
    Pattern formation and chaos in spatial ecological public goods games
    Journal of Theoretical Biology 268:30-38.... DOI 10.1016/j.jtbi.2010.09.036

  25. Wakano JY, Ikeda K, Miki T & Mimura M (2011)
    Effective dispersal rate is a function of habitat size and corridor shape: mechanistic formulation of a two-patch compartment model for spatially continuous systems
    Oikos 120: 1712-1720.... DOI 10.1111/j.1600-0706.2011.19074.x

  26. Wakano JY, Kawasaki K, Shigesada N & Aoki K (2011)
    Coexistence of individual and social learners during range-expansion
    Theoretical Population Biology 80:132-140.... DOI 10.1016/j.tpb.2011.06.001

  27. Wakano JY (2012)
    Spatiotemporal dynamics of cooperation and spite behavior by conformist transmission
    Communications on Pure and Applied Analysis 11:375-386.... DOI 10.3934/cpaa.2012.11.375

  28. Mirrahimi S, Perthame B & Wakano JY (2012)
    Evolution of species trait through resource competition
    Journal of Mathematical Biology 64:1189-1223.... DOI 10.1007/s00285-011-0447-z

  29. Kobayashi Y & Wakano JY (2012)
    Evolution of social versus individual learning in an infinite island model
    Evolution 66:1624-1635.... DOI 10.1111/j.1558-5646.2011.01541.x

  30. Aoki K, Wakano JY & Lehmann L (2012)
    Evolutionarily stable learning schedules and cumulative culture in discrete generation models
    Theoretical Population Biology 81:300-309.... DOI 10.1016/j.tpb.2012.01.006

  31. Wakano JY & Lehmann L (2012)
    Evolutionary and convergence stability for continuous phenotypes in finite populations derived from two-allele models
    Journal of Theoretical Biology 310:206-215.... DOI 10.1016/j.jtbi.2012.06.036

  32. Nakahashi W, Wakano JY & Henrich J (2012)
    Adaptive social learning strategies in temporally and spatially varying environments
    Human Nature 23:386-418.... DOI 10.1007/s12110-012-9151-y

  33. Wakano JY & Iwasa Y (2013)
    Evolutionary branching in a finite population: Deterministic branching versus stochastic branching
    Genetics 193:229-241.... DOI 10.1534/genetics.112.144980
    Full Text PDF
    OPEN ACCESS, full text available through the DOI link

  34. Wakano JY, Ohstuki H & Kobayasi Y (2013)
    A mathematical description of the inclusive fitness theory
    Theoretical Population Biology 84:46-55.... DOI 10.1016/j.tpb.2012.11.007

  35. Lehmann L, Wakano JY & Aoki K (2013)
    On optimal learning schedules and the marginal value of cumulative cultural evolution
    Evolution 67:1435-1445.... DOI 10.1111/evo.12040

  36. Lehmann L, & Wakano JY (2013)
    The handaxe and the microscope: individual and social learning in a multidimensional model of adaptation
    Evolution and Human Behavior 34:109-117.... DOI 10.1016/j.evolhumbehav.2012.11.001

  37. Mirrahimi S, Perthame B, & Wakano JY (2014)
    Direct competition results from strong competiton for limited resource
    Journal of Mathematical Biology 68:931-949.... DOI 10.1007/s00285-013-0659-5

  38. Wakano JY & Miura C (2014)
    Trade-off between learning and exploitation: the Pareto-optimal versus evolutionarily stable learning schedule in cumulative cultural evolution
    Theoretical Population Biology 91:37-43.... DOI 10.1016/j.tpb.2013.09.004

  39. Wakano JY & Lehmann L (2014)
    Evolutionary branching in deme-structured populations
    Journal of Theoretical Biology 351:83-95.... DOI 10.1016/j.jtbi.2014.02.036

  40. Scotti T, Mimura M & Wakano JY (2015)
    Avoiding Toxic Prey May Promote Harmful Algal Blooms
    Ecological Complexity 21:157-165.... DOI 10.1016/j.ecocom.2014.07.004

  41. Kobayashi Y, Wakano JY, Ohtsuki H (2015)
    A paradox of cumulative culture
    Journal of Theoretical Biology 379:79-88.... DOI 10.1016/j.jtbi.2015.05.002

  42. Kobayashi Y, Ohtsuki H, & Wakano JY (2016)
    Population size vs. social connectedness - A gene-culture coevolutionary approach to cumulative cultural evolution
    Theoretical Population Biology 111:87-95.... DOI 10.1016/j.tpb.2016.07.001

  43. Fogarty L, Wakano JY, Feldman MW & Aoki K (2017)
    The driving forces of cultural complexity: Neanderthals, modern Humans, and the question of population size
    Human Nature 28:39-52.... DOI 10.1007/s12110-016-9275-6

  44. Parvinen K, Ohtsuki H, & Wakano JY (2017)
    The effect of fecundity derivatives on the condition of evolutionary branching in spatial models
    Journal of Theoretical Biology 416:129-143.... DOI 10.1016/j.jtbi.2016.12.019

  45. Ohtsuki H, Wakano JY, Kobayashi Y (2017)
    Inclusive fitness analysis of cumulative cultural evolution in an island-structured population
    Theoretical Population Biology 115:13-23.... DOI 10.1016/j.tpb.2017.03.001

  46. Wakano JY, Funaki T, Yokoyama S (2017)
    Derivation of replicator-mutator equations from a model in population genetics
    Japan Journal of Industrial and Applied Mathematics 34:473-488... DOI 10.1007/s13160-017-0249-9
    OPEN ACCESS, full text available through the DOI link

  47. Wakano JY, Gilpin W, Kadowaki S, Feldman MW, Aoki K (2018)
    Ecocultural range-expansion scenarios for the replacement or assimilation of Neanderthals by modern humans
    Theoretical Population Biology 119:3-14... DOI 10.1016/j.tpb.2017.09.004

  48. Kobayashi Y, Wakano JY, Ohtsuki H (2018)
    Genealogies and ages of cultural traits: An application of the theory of duality to the research on cultural evolution
    Theoretical Population Biology 123:18-27... DOI 10.1016/j.tpb.2018.04.007

  49. Parvinen K, Ohtsuki H, Wakano JY (2018)
    Spatial heterogeneity and evolution of fecundity-affecting traits
    Journal of Theoretical Biology 454:190-204... DOI 10.1016/j.jtbi.2018.06.005

  50. Kurokawa S, Wakano JY, Ihara Y (2018)
    Evolution of groupwise cooperation: generosity, paradoxical behavior, and non-linear payoff functions
    Games 9:100... DOI 10.3390/g9040100
    OPEN ACCESS, full text available through the DOI link

  51. Kobayashi Y, Wakano JY, Ohtsuki H (2019)
    Evolution of cumulative culture for niche construction
    Journal of Theoretical Biology 472:67-76... DOI 10.1016/j.jtbi.2019.04.13

  52. Kadowaki S, Tamura T, Sano K, Kurozumi T, Maher LA, Wakano JY, Omori T, Kida R, Hirose M, Massadeh S, Henry DO (2019)
    Lithic technology, chronology, and marine shells from Wadi Aghar, southern Jordan, and Initial Upper Paleolithic behaviors in the southern inland Levant
    Journal of Human Evolution 135:102646... DOI 10.1016/j.jhevol.2019.102646

  53. Nakamura M, Wakano JY, Aoki K, Kobayashi Y (2020)
    The popularity spectrum applied to a cross-cultural question
    Theoretical Population Biology 133:104-116... DOI 10.1016/j.tpb.2019.10.003
    OPEN ACCESS, full text available through the DOI link

  54. Parvinen K, Ohtsuki H, Wakano JY (2020)
    Evolution of dispersal in a spatially heterogeneous population with finite patch sizes
    Proceedings of the National Academy of Sciences of the United States of America 117:7290-7295 ... DOI 10.1073/pnas.1915881117
    OPEN ACCESS, full text available through the DOI link

  55. Ohtsuki H, Rueffler C, Wakano JY, Parvinen K, Lehmann L (2020)
    The components of directional and disruptive selection in heterogeneous group-structured populations
    Journal of Theoretical Biology 507:110449... DOI 10.1016/j.jtbi.2020.110449
    OPEN ACCESS, full text available through the DOI link

  56. Wakano JY, Kadowaki S (2021)
    Application of the ecocultural range expansion model to modern human dispersals in Asia
    Quaternary International 596:171-184... DOI 10.1016/j.quaint.2020.12.019
    OPEN ACCESS, full text available through the DOI link

  57. Kobayashi Y, Kurokawa S, Ishii T, Wakano JY (2021)
    Time to extinction of a cultural trait in an overlapping generation model
    Theoretical Population Biology 137:32-45... DOI 10.1016/j.tpb.2021.01.002
    OPEN ACCESS, full text available through the DOI link

  58. Mullon C, Wakano JY, Ohtsuki H (2021)
    Coevolutionary dynamics of genetic traits and their long-term extended effects under non-random interactions
    Journal of Theoretical Biology 525:110750... DOI 10.1016/j.jtbi.2021.110750
    OPEN ACCESS, full text available through the DOI link

  59. Aoki K, Wakano JY (2022)
    Hominin forager technology, food sharing, and diet breath
    Theoretical Population Biology 144:37-48... DOI
    OPEN ACCESS, full text available through the DOI link

  60. Aoki K, Takahata N, Oota H, Wakano JY, Feldman MW (2023)
    Infectious diseases may have arrested the southward advance of microblades in Upper Palaeolithic East Asia
    Prceedings B 290:20231262... DOI 10.1098/rspb.2023.1262

  61. Parvinen K, Ohtsuki H, Wakano JY (2023)
    Evolution of dispersal under spatio-temporal heterogeneity
    Journal of Theoretical Biology 547:111612... DOI 10.1016/j.jtbi.2023.111612

Book chapter (peer-reviewed)

Reports, Reviews, etc. (not peer-reviewed)

Secondary appearance

Submitted Papers

For Fun

My Erdos number = 4
  1. Davies, R. O.; Erdo"s, P.: Splitting almost-disjoint collections of sets into subcollections admitting almost-transversals. Infinite and finite sets (Colloq., Keszthely, 1973; dedicated to P. Erdo"s on his 60th birthday), Vol. I, pp. 307--322. Colloq. Math. Soc. Janos Bolyai, Vol. 10, North-Holland, Amsterdam, 1975.
  2. Davies, R. O.; Sessa, S.: A common fixed point theorem of Gregu\v s type for compatible mappings. Facta Univ. Ser. Math. Inform. No. 7 (1992), 99--106.
  3. Higashi, Masahiko; Di Nola, Antonio; Sessa, Salvatore; Pedrycz, Witold: Ordering fuzzy sets by consensus concept and fuzzy relation equations. Internat. J. Gen. Systems 10 (1984), no. 1, 47--56.
  4. Yamamura, Norio; Higashi, Masahiko; Behera, Narayan; Wakano, Joe Yuichiro: Evolution of mutualism through spatial effects. J. Theoret. Biol. 226 (2004), no. 4, 421--428.

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