Professor of Biology
Centre for Computational Systems Biology, ISTBI
Fudan University
220 Handan Road, Shanghai 200433
People's Republic of China
Room 2305 East Guanghua Tower
Handan Campus, Fudan University
Associate Editor, Theoretical Population Biology
Member, Board of Reviewing Editors, Science
I work in theoretical biology, combining mathematics and computer simulations to gain understanding of biological systems. Much of my work has been in evolution and population genetics, where I have focussed on random (or stochastic) phenomena, such as the dynamics of genes within populations. I also work in other fields where mathematical modelling is useful, such as pharmacokinetics and epidemiology.
Recent collaborators include:
Andy Overall (Brighton)
Greg Scutt (Brighton)
Hassan Shafiey (Halle)
Joel Peck (Cambridge)
Konstantinos Mavreas (Fudan)
Lei Zhao (ECNU)
Martin Lascoux (Uppsala)
Peng Ji (Fudan)
Pierre Nouvellet (Sussex)
Toni Gossmann (Dortmund)
Nikolas Vellnow (Dortmund)
Xinming Zhao (Fudan)
Ruixi Huang (Fudan)
Shaojing Sun (Fudan)
Changqian Rao (Fudan)
Wei Lin (Fudan)
Zhuoyi Song (Fudan)
For a complete list, click here.
Beyond two alleles: multiallelic genotypic selection and its estimation from time-series data
BMC Genomics 27, 837 (2026) with Nikolas Vellnow and Toni Gossmann
Inferring ancestral population type from limited samples
Frontiers in Ecology and Evolution 14, 1701295 (2026) with Andrew Overall
Effective decoupling of mutations and the resulting loss of biodiversity caused by environmental change
Journal of Theoretical Biology 616, 112277 (2026) with Ruixi Huang
A dynamical measure of algorithmically infused visibility
Royal Society Open Science 12, 242232 (2025) with Shaojing Sun and Zhiyuan Liu
Exact first-passage time distributions from time-dependent solutions of the chemical master equation,
Part I and
Part II
Journal of Chemical Physics 162, 224104 and 224105 (2025) with Changqian Rao, Wei Lin and Zhuoyi Song
Theoretically quantifying the direct and indirect benefits of vaccination against SARS-CoV-2 in terms of avoided deaths
Scientific Reports, Volume 12, Article number: 8833 (2022) with Greg Scutt and Mike Cross
Correcting bias in allele frequency estimates due to an observation threshold: A Markov chain analysis
Genome Biology and Evolution, Volume 14, Issue 4 (2022) with Toni Gossmann
Influence of dominance and drift on lethal mutations in human population
Frontiers in Genetics 11, 267 (2020) with Andrew Overall
What is adaptation and how should it be measured?
Journal of Theoretical Biology 447: 190-198 (2018) with Joel Peck
Evolutionary control: Targeted change of allele frequencies in natural populations using externally directed evolution
Journal of Theoretical Biology 419: 362-374 (2017) with Hassan Shafiey and Toni Gossmann
Complete numerical solution of the diffusion equation of random genetic drift
Genetics 194: 973-985 (2013) with Lei Zhao and Xingye Yue
Population growth enhances the mean fixation time of neutral mutations and the persistence of neutral variation
Genetics 191: 561-577 (2012)
A unified treatment of the probability of fixation when population size and the strength of selection change over time
Genetics 188: 907-913 (2011)
Is Life Impossible? Information, sex and the origin of complex organisms
Evolution 64: 3300-3309 (2010) with Joel Peck
Fundamental insights into the random movement of animals from a single distance-related statistic
American Naturalist 174: 506-514 (2009) with Pierre Nouvellet and Jonathan Bacon
Pleiotropic scaling of gene effects and the 'cost of complexity'
Nature 452: 470-472 (2008) with Günter Wagner, Jane Kenney-Hunt, Mihaela Pavlicev, Joel Peck and James Cheverud
Fisher's microscope and Haldane's ellipse
American Naturalist 166: 447-457 (2005) with John Welch
Sympatric speciation by sexual conflict
PNAS 99, 10533-10538 (2002) with Sergey Gavrilets
Mutation and sex in a competitive world
Nature 406: 399-403 (2000) with Joel Peck
Pleiotropy and the preservation of perfection
Science 279: 1210-1213 (1998) with Joel Peck
Why do asexual and self fertilizing populations tend to occur in marginal environments
Nature 391: 889-892 (1998) with Joel Peck and John Yearsley
For more details on my papers see publications.