In a multitude of life’s processes, cilia and eukaryotic flagella are ubiquitous and indispensable. In the human body, lung and ependymal cilia coordinate the pumping of fluid, but the very same cellular appendages are used by diverse species of protists and algae as microscale analogues of limbs to swim through aqueous environments. In our group, we use a combination of theory, experiment, and computation to investigate the physics of motile cilia and the mechanistic origins of motility and basal cognition.

K.Y. Wan, Essays in Biochemistry 62(6), 829-838 (2018) [pdf]

1. Self-propulsion of microorganisms

How do cells control their direction of movement?

Locomotion achieved through the actuation of multiple appendages, is inherently combinatorial. Even simple microorganisms display a fascinating behavioural heterogeneity, transitioning between different swimming modes, or fast and slow dynamics. We use high-resolution spatiotemporal information obtained from quantitative live-cell imaging to characterise and model stereotyped states or gaits, revealing an unprecedented complexity in the non-nervous control of locomotor behaviour in different species of single-celled eukaryotes.

Distinct gaits of self-propelled quadriflagellate microswimmers
D. Cortese & K. Y. Wan, biorxiv (2022) [online]

Control of helical navigation by three-dimensional flagellar beating
D. Cortese & K. Y. Wan, Physical Review Letters 126, 088003 (2021) [online] [Exeter News]
* Selected as Editors’ Suggestion
* See also Physics synopsis

Intracellular coupling mediates flagellar synchrony
H. Guo, Y. Man, K.Y. Wan & E. Kanso, Journal of the Royal Society Interface 18, 20200660 (2020) [online] [arxiv]

Synchrony and symmetry-breaking in active flagellar coordination
K. Y. Wan, Philosophical Transactions of the Royal Society B 375, 20190393 (2019) [online] [ScienceNews]

Antiphase Synchronization in a Flagellar-Dominance Mutant of Chlamydomonas
K.C. Leptos*, K.Y. Wan*, M. Polin, I. Tuval, A.I. Pesci  & R.E. Goldstein, Physical Review Letters 111, 158101 (2013) [online]

2. Genesis and control of motile cilia

Functional exploration of heterotrimeric kinesin-II in IFT and ciliary length control in Chlamydomonas
S. Li, K. Y. Wan, W. Chen, H. Tao, X. Liang, J. Pan, eLife, 9, e58868 (2020) [online]

Reorganisation of complex ciliary flows around regenerating Stentor coeruleus
K. Y. Wan, S. K. Hürlimann, A. M. Fenix, R. M. McGillivary, T. Makushok, E. Burns, J. Y. Sheung, W. F. Marshall, Philosophical Transactions of the Royal Society B 375, 20190167 (2019) [online]

Coordination of eukaryotic cilia and flagella
K.Y. Wan, Essays in Biochemistry 62(6), 829-838 (2018) [online]

Coordinated Beating of Algal Flagella is Mediated by Basal Coupling
K.Y. Wan & R.E. Goldstein, Proceedings of the National Academy of Sciences USA 113, E2784-93 (2016) [online] [Cambridge News] [Fierceroller Blog] [galloping algae]

Flagellar Synchronization Through Direct Hydrodynamic Interactions
D.R. Brumley*, K.Y. Wan*, M. Polin & R.E. Goldstein, eLife 3, e02750 (2014) [online] [eLife Insight] [Cambridge News

Cilia often exhibit synchronization phenomena, including phase-locking and metachronal waves. It has long been hypothesised that synchrony arises from fluid dynamical coupling. By controlling the distance of separation between pairs of pipette-held somatic cells of Volvox, we proved that hydrodynamic interactions were sufficient to produce synchrony in ciliary arrays. In contrast, many unicellular organisms rely on a different mechanism – which we have termed intracellular or basal coupling, to orchestrate ciliary coordination. The correct length and placement of cilia is critical to achieve proper function.

3. Stochasticity and behavioural stereotypy 

Like the human heartbeat, the Chlamydomonas flagellum is a complex biological oscillator, exhibiting strong rhythmicity, yet also responsiveness to certain environmental or physiological cues. Extracting and digitising flagellar beat patterns and frequencies from long-time, high-speed recordings, we devise novel measures of shape and waveform stochasticity. When complex spatiotemporal dynamics are projected onto a low-dimensional state-space, macroscopic breaking of detailed balance is revealed.

Phenotyping single-cell motility in microfluidic confinement
S.A. Bentley, H. Laeverenz-Schlogelhofer, V. Anagnostidis, J. Camman, M.G. Mazza, F. Gielen, K.Y. Wan, in press, elife (2022) [biorxiv]

Time-irreversibility and criticality in the motility of a flagellate microorganism
K.Y. Wan & R.E. Goldstein, Physical Review Letters 121, 058103 (2018) [online]

Lag, Lock, Sync, Slip: The Many ‘Phases’ of Coupled Flagella
K.Y. Wan, K.C. Leptos & R.E. Goldstein, Journal of the Royal Society Interface 11, 20131160 (2014) [online]

Rhythmicity, Recurrence, and Recovery of Flagellar Beating
K.Y. Wan & R.E. Goldstein, Physical Review Letters 113, 238103 (2014) [online]

4. Ciliated larvae of marine invertebrates

Ciliary propulsion and metachronal coordination in reef coral larvae
R.N. Poon, T.A. Westwood, H. Laeverenz-Schlogelhofer, E. Broderick, J. Craggs, E.E. Keaveny, G. Jékely, K.Y. Wan, bioRxiv (2022) [online] [tweetorial]

5. Protists and the origin of eukaryotes

Origins of eukaryotic excitability
K.Y. Wan & G. Jékely, in press, Philosophical Transactions of the Royal Society B 376, 20190758 (2021) [online] [tweetorial]

Single-celled eukaryotes are capable of surprisingly complex behaviour.

We propose a new scenario for eukaryogenesis in which eukaryotes harnessed their larger size, improved complement of ion channels, and novel locomotor organelles to access new biophysical and sensory regimes.

6. Bio-inspired robotics

A minimal robophysical model of quadriflagellate self-propulsion
K. Diaz, T.L. Robinson, Y. Ozkan-Aydin, E. Aydin, D.I. Goldman & K.Y. Wan, Bioinspiration and Biomimetics 16, 066001 (2021) [online]

We developed a novel autonomous (internally powered) robot capable of swimming at low Reynolds numbers, modelled on quadriflagellate algae.

7. Book chapter

Physics and mechanics of ciliary beating
L. Woodhams, D. Cortese, P. Bayly, K.Y. Wan, The Chlamydomonas Sourcebook (Chapter 12, 3rd Edition, 2022), to appear

8. Editorials and Commentaries

Isolating the beat of cilia
K.Y. Wan, Nature Physics (2022) [online]

Flagella: a new kind of beat
K.Y. Wan, elife 10, e67701 (2021) [online]

On the unity and diversity of cilia (Foreword to Special Issue)
K.Y. Wan & Gáspár Jékely, Philosophical Transactions of the Royal Society B 375, 20190148 (2019) [online]

Ciliate Biology: the Graceful Hunt of a Shape-Shifting Predator
K.Y. Wan, Current Biology 29, R1174-R1176 (2019) [online] [pdf]

Link to my google scholar profile.