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 phenomena, such as the dynamics of genes within populations. The questions I ask often involve how chance shapes the fate of genes.
Perhaps paradoxically, a biological population containing an infinite number of individuals is much easier to analyse than a finite population. In an infinite population there is no randomness in the frequency of a given type of gene (where frequency means the proportion of all genes that are the given type, which is a number between 0 and 1). In a finite population, there is randomness, because chance determines which offspring survive to the next generation. This leads to the frequency having random changes from one generation to the next — a process called random genetic drift. Much of my work involves mathematical treatments of this process. You can watch random genetic drift happen on the drift explorer page.
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
Effective decoupling of mutations and the resulting loss of biodiversity caused by environmental change
Journal of Theoretical Biology 616, 112277 (2026) with Ruixi Huang
An exact path-integral
representation of the Wright-Fisher model with mutation and selection
BMC Genomics 26, 1032 (2025)
A dynamical measure of algorithmically infused visibility
Royal Society Open Science 12, 242232 (2025) with Shaojing Sun and Zhiyuan Liu
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
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
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.