Neuronal coordination of cilia in marine zooplankton in moving and sensing

University of Regensburg
7 October 2026

Gáspár Jékely

Centre for Organismal Studies, Heidelberg University

@jekely@biologists.social

Ciliated zooplankton larvae

Larval behaviours



phototaxis

  • UV avoidance
  • thermosensing
  • settlement

startle response

  • ciliary coordination
  • chemosensing

crawling

  • flow sensing
  • pressure response

Coral larvae

Emelie Brodrick


Hannah Laeverenz



Rebecca Poon & Kirsty Wan

Platynereis dumerilii







  • breeding culture
  • genome sequence
  • microinjection, transgenesis
  • neuron-specific promoters
  • knock-out lines
  • neuronal connectome
  • neuronal activity imaging

Multiciliary bands form metachronal waves

with Kirsty Wan, LSI Exeter

  • basal-body spacing: 0.35-0.55 μm

Precise orientation of multiciliary arrays

expansion microscopy

Metachronal waves are robust

  • organised by short-range steric interactions

Modelling ciliated larvae

  • wave direction determined by azimuthal offset (azimuth - the horizontal angle from a cardinal direction)

Phototaxis

Helical swimming, sensing and turning are tightly linked

Global coordination of cilia

Closures = sinking

Calcium increase induces closures

Array tomography for vEM and connectomics



  • serial ultrathin sectioning (~5000 sections)
  • serial imaging by TEM or SEM
  • alignment of sections, tracing and annotation of neurons
  • whole-body reconstruction of all cells and the synaptic connectome

Whole-body vEM reconstruction

Whole-body annotation of all ciliary basal bodies

Nervous control of multiciliated cells

A pacemaker system for ciliary closures

Whole-body coordination of ciliary closures

Whole-body coordination of ciliary closures

Whole-body coordination of ciliary closures

  • wild type

  • mechanosensory mutant (pkd2)

Whole-body coordination during startle

Serotonergic ciliomotor neurons increase beat frequency

Ser-h1 neurons, EM reconstruction

Pressure response in Platynereis larvae

Precise control of pressure in the pressure chamber

Pressure response is graded

Ciliary beating increases under pressure

Pressure is sensed by photoreceptors with ramified cilia



Photoreceptor responses to pressure are graded

Defects of pressure response in c-opsin1 mutants

Reduced ciliary compartment in c-opsin1 mutants

Control of ciliary swimming in zooplankton

  • ongoing activity of cilia
  • ongoing activity of ciliomotor pacemaker system
  • modified by sensory cues (light, vibrations, pressure etc.)
  • motion as part of sensing (phototaxis, rheotaxis)
  • there are no ‘reflexes’
  • ongoing behaviour affects the senses – reafference

The non-bilaterian Metazoa

Ctenophora

Ernst Haeckel, Kunstformen der Natur (1904), plate 27: Ctenophorae

Ctenophora (Rippenquallen)


    1. Pleurobrachia bachei
    1. Mnemiopsis leidyi
    1. Beroe gracilis mit Pleurobrachia pileus im Magen
    1. Thalassocalyce inconstans
    1. Coeloplana astericola (yellow-brown, on a starfish)
  • Traditinally part of Coelenterata
  • Cteno-phora: from kteis = Comb [Genitiv ktenos] and pherein = carry

Bolinopsis

  • Iridescenc (from iris = rainbow)

Ctenophores

Bolinopsis mikado

Shimoda Marine Station, 2024

Ctenophora

Pleurobrachia Cydippea

  • marine hermaphrodites
  • develop through a freely-swimming cydippid larval stage
  • 8 comb rows
  • colloblasts (from collo = glue, blastos = bud), no stinging cells
  • subepithelial, syncytial nerve net

Ctenophora - Pleurobrachia

  • oral-aboral axis
  • two tentacles with side branches (Tentilla), with colloblasts
  • canal system for the spread of digested food
  • comb plates

Ctenophores - biradial symmetry

  • two orthogonal symmetry planes: sagittal and tentacular plane

Ctenophores - Comb plates

  • comb plates formed by hundreds of cilia glued together
  • “compartmenting lamellae” - protein complex to hold cilia together
  • individual cilia beat as a single unit - comb plate

  • efficient swimming
  • the largest organisms that can move with cilia (broke the low-Reynolds limit)
  • a ctneophore-specific protein complex glues the cilia together (e.g. CTENO64)

Ctenophores - Comb plates

Comb plates

Ctenophores - Nervous system

  • neurons in the subepithelial nerve net form a syncytium
  • also synapses between sensory cells and the nerve net and the nerve net and comb-row cells

Ctenophores – Statocyst (balancer organ)

  • Statocyst - balancer organ for graviorientation in the aboral organ
  • Statolith held by four bundles of balancer cilia

Balancer – four bundles of cilia supporting a statolyth

Balancer organ – reorientation after tilt

Kei Jokura

Sydney Tamm – balancer activity and graviorientation

  • differential load when body orientation changes
  • change in the beating of balancer cilia
  • pacemaker system for comb rows

The beating of balancer cilia depends on body orientation

Volume EM reconstruction of the aboral organ

  • 1011 cells
  • four quadrants
  • two symmetry planes

Kei Jokura

There syncytial nerve-net neurons

  • one large nerve net (all 4 quadrants)
  • two small nerve nets (two by two quadrants in sagittal plane)

Synaptic connectome of the aboral organ

  • synapses from nerve net to balancer cells
  • no synapses from balancer or other sensory cells to the nerve net
  • not a ‘reflex-like’ organisation
  • if not input-output, then what is the function of the nerve net?

High-speed imaging of balancer cilia

  • regular arrests of ciliary beating
  • syncronised in the sagittal plane

High-speed imaging of balancer cilia

  • regular arrests of ciliary beating
  • not syncronised in the tentacular plane

High-speed imaging of balancer cilia

  • re-beating is syncronised in both planes -> all balancers in sync

Synchronisation of balancer cilia

  • arrests only synchronised across 2 balancers in the sagittal plane
  • re-beats are synchronised across all 4 balancers

Synchronisation of balancer cilia

  • frequency of ciliary beating is synchronised across balancers

Connectome of the ctenophore balancer

  • ongoing ciliary activity
  • global neuronal coordination
  • synchronisation of arrests
  • synchronisation of re-beat
  • synchronisation of beat frequency
  • sensing and movement intimately linked

Acknowledgements

  • Alexandra Kerbl
  • Karel Mocaer
  • Sanja Jasek
  • David Hug
  • Jules Duruz
  • Mateusz Kostecki
  • Benedikt Dürr
  • Kata Szabó
  • Emily Savage
  • Simone Wolters
  • Anja Ciprianidis
  • Kevin Urbansky

Alumni

  • Kei Jokura (NIBB, Okazaki)
  • Luis Bezares (LBDV, Villefranche-sur-mer)
  • Luis Yanez-Guerra (Uni Sussex)
  • Emelie Brodrick (Sussex)
  • Csaba Verasztó (EPFL)

Facilities

  • Réza Shahidi
  • Charlotta Funaya
  • Ulrike Engel