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<!DOCTYPE html>
<html lang="en">
<head>
<meta charset="UTF-8">
<meta name="viewport" content="width=device-width, initial-scale=1.0">
<meta name="description" content="An interactive timeline of sixty years of clathrin and endocytosis research, leading up to a new kinetic Monte Carlo study of curvature generation.">
<meta name="author" content="Johannes Dreckhoff">
<title>How Cells Pull Things In — A Clathrin Timeline | Johannes Dreckhoff</title>
<link rel="stylesheet" href="css/main.css?v=20260714">
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</style>
</head>
<body>
<!-- Header & Navigation -->
<header>
<div class="container">
<a href="index.html" class="logo">Johannes Dreckhoff</a>
<nav>
<a href="index.html#about">About</a>
<a href="professional.html">Professional</a>
<a href="projects.html">Projects</a>
<a href="index.html#gallery">Gallery</a>
<a href="places.html">Places</a>
<a href="blog/index.html">Blog</a>
<a href="index.html#contact">Contact</a>
</nav>
</div>
</header>
<!-- Intro -->
<section class="timeline-hero">
<div class="container">
<div class="section-header">
<h2>How Cells Pull Things In</h2>
<p>A sixty-year story of clathrin, curvature, and a question only just being answered</p>
</div>
<div class="timeline-intro">
<p>
Every one of your cells constantly pulls material in from the outside world. To do it, the cell
wraps tiny patches of its own surface into bubbles and swallows them. The machine that builds
these bubbles is a protein called <strong>clathrin</strong>, which assembles into a soccer-ball-like
cage that bends a flat membrane into a sphere.
</p>
<p>
How a flat sheet decides to curve into a ball — and why it sometimes fails — is a puzzle
scientists have chased for sixty years. Here's the story.
</p>
</div>
<p class="timeline-hint">Click any milestone to expand it. Read along the snake: left to right, then back again.</p>
</div>
</section>
<!-- Timeline -->
<!--
NOTE TO OWNER: the references at the bottom of each card are field-standard
citations (author, journal, year). Please verify and refine the exact
journals/volumes you prefer to credit. The Naming-clathrin card kept the
original wording you provided.
-->
<section class="timeline-section">
<div class="container">
<ol class="timeline" aria-label="Milestones in clathrin and endocytosis research">
<!-- 1964 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-1">
<span class="timeline-number" aria-hidden="true">1</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">1964</div>
<h3 class="timeline-title">The first glimpse</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-1">
<p>
Roth & Porter, peering into mosquito cells with an electron microscope, spot strange
"bristle-coated" pits forming at the cell surface and pinching off into the cell.
Nobody yet knows what the bristly coat is made of — but they correctly guess that
these pits are how the cell takes things in.
</p>
<span class="timeline-attribution">
Roth, T.F. & Porter, K.R., <em>J. Cell Biol.</em>, 1964.
</span>
</div>
</article>
</li>
<!-- 1969 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-2">
<span class="timeline-number" aria-hidden="true">2</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">1969</div>
<h3 class="timeline-title">"A vesicle in a basket"</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-2">
<p>
Kanaseki & Kadota describe the coat's striking geometry: a lattice of hexagons and
pentagons, like a tiny geodesic basket cradling each vesicle. The shape itself hints that
the coat is what curves the membrane.
</p>
<span class="timeline-attribution">
Kanaseki, T. & Kadota, K., <em>J. Cell Biol.</em>, 1969.
</span>
</div>
</article>
</li>
<!-- 1973 — Helfrich -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-3">
<span class="timeline-number" aria-hidden="true">3</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">1973</div>
<h3 class="timeline-title">The physics of bending membranes</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-3">
<p>
In parallel, physicist Wolfgang Helfrich writes down the mathematics of how thin
membranes resist bending — treating the cell membrane like an elastic sheet with a
stiffness and a preferred shape. This framework becomes the language for everything that
follows.
</p>
<span class="timeline-attribution">
Helfrich, W., <em>Z. Naturforsch. C</em>, 1973.
</span>
</div>
</article>
</li>
<!-- 1975–76 — Pearse -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-4">
<span class="timeline-number" aria-hidden="true">4</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">1975–76</div>
<h3 class="timeline-title">Naming clathrin</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-4">
<p>
Barbara Pearse purifies these coated vesicles and discovers that a single protein
builds the entire cage. She names it <strong>clathrin</strong>, from the Latin for
"lattice." The central player finally has a name.
</p>
<span class="timeline-attribution">
Purification: Pearse, B.M.F., <em>J. Mol. Biol.</em>, 1975.
Naming & hexagon/pentagon network description: Pearse, B.M.F., <em>PNAS</em>, 1976.
</span>
</div>
</article>
</li>
<!-- 1980 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-5">
<span class="timeline-number" aria-hidden="true">5</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">1980</div>
<h3 class="timeline-title">Catching the cage in the act</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-5">
<p>
John Heuser uses rapid-freezing electron microscopy to capture clathrin lattices frozen
mid-assembly on the membrane, seeing flat sheets and curved baskets side by side. This
raises the question at the heart of the field: how does a flat lattice become a curved
one?
</p>
<span class="timeline-attribution">
Heuser, J., <em>J. Cell Biol.</em>, 1980.
</span>
</div>
</article>
</li>
<!-- ~1990s -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-6">
<span class="timeline-number" aria-hidden="true">6</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">~1990s</div>
<h3 class="timeline-title">Mapping the machine</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-6">
<p>
Through the work of Tom Kirchhausen and many others, the molecular cast assembles:
the <em>triskelion</em> (clathrin's three-legged building block), the adaptor proteins
that recruit it, and the picture of clathrin as a self-assembling, programmable cage.
The parts list is now known — but the mechanism of bending is still debated.
</p>
<span class="timeline-attribution">
e.g. Ungewickell & Branton, <em>Nature</em>, 1981;
Kirchhausen, T., <em>Annu. Rev. Cell Dev. Biol.</em>, 1999.
</span>
</div>
</article>
</li>
<!-- 2018 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-7">
<span class="timeline-number" aria-hidden="true">7</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">2018</div>
<h3 class="timeline-title">Flat or curved from the start?</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-7">
<p>
A long-running controversy: does clathrin first lay down a flat sheet and then bend
it, or does it curve as it grows? Bucher et al. and others show that the balance of
clathrin and adaptors tips the outcome, reframing the flat-to-curved transition as
something tunable rather than fixed.
</p>
<span class="timeline-attribution">
Bucher, D. et al., <em>Nat. Commun.</em>, 2018.
</span>
</div>
</article>
</li>
<!-- 2021 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-8">
<span class="timeline-number" aria-hidden="true">8</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">2021</div>
<h3 class="timeline-title">The coat is under tension</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-8">
<p>
Two key experiments. Sochacki et al. map the spontaneous curvature built into flat
lattices, and Tagiltsev et al. perform a remarkable trick: they cut a clathrin lattice
and watch it spring to a tighter curl — proving the coat is an elastically loaded
spring, storing mechanical energy. The coat doesn't just sit there; it pushes.
</p>
<span class="timeline-attribution">
Sochacki, K.A. et al., <em>Dev. Cell</em>, 2021;
Tagiltsev, G. et al., <em>eLife</em>, 2021.
</span>
</div>
</article>
</li>
<!-- 2023 -->
<li class="timeline-item">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="false"
aria-controls="milestone-body-9">
<span class="timeline-number" aria-hidden="true">9</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">2023</div>
<h3 class="timeline-title">Watching curvature happen, live</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-9">
<p>
Mund et al. use super-resolution microscopy to follow individual coats bending in
real cells, supporting a "cooperative" picture where assembly and curvature progress
together. The flat-to-curved question is now answerable with data — but a unifying
physical explanation is still missing.
</p>
<span class="timeline-attribution">
Mund, M. et al., <em>Nat. Cell Biol.</em>, 2023.
</span>
</div>
</article>
</li>
<!-- Final: This work -->
<li class="timeline-item timeline-final">
<article class="timeline-card" role="button" tabindex="0" aria-expanded="true"
aria-controls="milestone-body-10">
<span class="timeline-number" aria-hidden="true">10</span>
<div class="timeline-card-header">
<div class="timeline-card-heading">
<div class="timeline-year">This work</div>
<h3 class="timeline-title">One model, three fates</h3>
</div>
<span class="timeline-chevron" aria-hidden="true">+</span>
</div>
<div class="timeline-body" id="milestone-body-10">
<p>
This paper builds a computer simulation of a growing clathrin coat from the bottom up
and finds something surprising: <strong>curvature doesn't need a special trigger.</strong>
As the cage grows, its own geometry makes it progressively stiffer, and this stiffening
alone can drive a flat sheet to curl into a sphere.
</p>
<p>
The very same rules spontaneously produce all three outcomes seen in cells — flat
plaques, stalled "aborted" pits, and fully sealed vesicles — depending only on how
the coat grows. It even reproduces the spring-back seen in the 2021 cutting experiments.
A sixty-year-old question, answered by geometry.
</p>
<a href="research/cme/index.html" class="btn btn-primary timeline-cta">Read the paper →</a>
<span class="timeline-attribution">
Dreckhoff, J.H.H., Schwarz, U.S. & Lettermann, L., arXiv:2606.30267, 2026.
</span>
</div>
</article>
</li>
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