„Discovery”
[IMAGE: Title photo — New York, USA – November 25, 2021: 95th Macy’s Annual Thanksgiving Day Parade celebration near Radio City Music Hall. Ronald McDonald clown addresses the crowd from big red shoe car in Manhattan. Viktor_IS – Shutterstock.]
I have published many times, claiming and showing that the ancients encoded biology in everything. Literally everything.
In the stone of temples and the walls of fortresses. In the plans of cities. In coats of arms and orders. In paintings and sculptures. In gowns, corsets and hairstyles.
This code did not vanish with those who wrote it down. It survived because it was passed on — from generation to generation, from hand to hand. Often by people who no longer knew what they were passing on. They copied the shape because “that’s how it has always been done”.
It survived best where nobody looks for it. In what we consider entertainment.
In this post I will show it once again — through the example of Mr. Clown.
New York. Winter coats, crowds behind the barriers, police officers at the intersection.
And suddenly — a shoe. Red, gleaming, as long as a car. Because it is a car. On the licence plate, a single word: REDSHOE. Red shoe.
A clown stands inside. White face, red wig, a smile from ear to ear. He waves to the children. The children wave back.
Nobody on that street asks why a clown rides in a shoe. It is obvious, after all. Clowns and giant shoes have been inseparable for generations.
But it is precisely the obvious that deserves a second look.
This time, the Wardrobe of the Genome will not feature a gown. No corset. No hairstyle. This time it will be one more complete human figure — from the top of the head to the tips of the shoes. Mr. Clown. With balloons in his hand, pompoms on his belly and a ruff beneath his chin.
And with shoes that are far too big.
Everyone who has ever seen a clown thinks the giant shoes are a joke. That they are there to make us laugh.
But what if they are no joke at all… but the most precise technical drawing ever to walk into a circus arena?
1. Mr. Clown with Balloons
He stands in the middle of the circus arena. Upright, calm, smiling. He does nothing funny. He simply stands there — and waits for us to look at him.
[IMAGE: A clown with balloons in a circus ring — stock illustration. Shutterstock AI.]
Let us start at the top.
Balloons. Above his head floats a bunch of colourful balloons: blue, red, yellow, green. Together they form one large, round cloud.
Strings. Thin white strings run down from the balloons. The clown gathers them in his right hand. Their ends hang loosely below his fist.
Head. A bald, white skull. On either side, two thick tufts of orange hair sticking out sideways like two flames. A face covered in white make-up, black arched eyebrows, a round red nose, a wide red smile.
Ruff. Beneath the chin, a wide, white, frilled collar with a red trim. It encircles the neck and falls over the shoulders.
Bow tie. In the centre of the ruff — a large blue bow tie.
Jumpsuit. A single grey, baggy garment from neck to ankles. Wide sleeves, wide trouser legs, the fabric falling loosely in folds.
Pompoms. On the front of the jumpsuit, one below the other, three orange pompoms.
Gloves and cuffs. White gloves. At the wrists, white frills with a red edge. The left arm hangs loosely by the hip.
Ankle frills. The trouser legs are gathered above the feet, and from beneath them emerge white frills — like little skirts above the shoes.
Shoes. And finally, them. Two enormous, red, gleaming shoes. Each many times larger than an ordinary foot.Round, bulging shoe tips and a black sole. The shoe tips point outwards — the left shoe tip to the left, the right shoe tip to the right.
Ring floor. Arena floor. The clown stands in a round circus arena whose floor has a radial red-and-white pattern.
A Short History of the Clown
The figure of the jester has accompanied humanity forever:fools at royal courts, comic performers at fairs, and the characters Harlequin and Pierrot in the Italian commedia dell’arte.
The modern circus was born in London in 1768. Philip Astley marked out a round circus arena — just like the one our clown stands in.
At the beginning of the 19th century, Joseph Grimaldi (1778–1837) created the white-faced clown. After his name, clowns in England were long called “Joey”.
In the second half of the 19th century, the Auguste appeared — a clumsy clown with a red nose, in oversized clothes and oversized shoes. Legend links his birth to a Berlin circus.
Orange wig, ruff, pompoms and giant shoes — this costume is stitched together from all these traditions. Each element has its own lineage. None is accidental.
Balloons are the youngest addition. The rubber balloon was invented in 1824 by Michael Faraday — for his experiments with hydrogen. A year later, Thomas Hancock began making toy balloons from rubber.
2. The Protein — Kinesin-1 (PDB: 3KIN)
[IMAGE: 3D illustration “3d rendered illustration of a motor protein” — 3dMediSphere / Shutterstock, no. 1215278068. Kinesin carrying cargo. A pink molecule stands on two lobes on a green tube built of beads. A long, thin stalk rises upwards. At its top, two lobes press against a large blue sphere covered with fine bristles and red dots. Thin red fibres in the background.]
What Is a Motor Protein?
Most proteins in the cell work in one place. A channel stands in the membrane like a door in a wall. An enzyme sits and cuts or joins.
Motor proteins are different. They move. They turn chemical energy into motion — just as a car engine turns petrol into the turning of wheels.
Their fuel is ATP (adenosine triphosphate) — a small molecule that acts like a charged battery. When the protein removes one phosphate group from it, the battery discharges and releases a portion of energy. That portion is enough for one movement.
Our bodies contain three great families of motor proteins: myosins (muscle work), dyneins and kinesins. Dyneins and kinesins walk along the same roads, but in opposite directions.
What Is Kinesin?
[VIDEO: YouTube video embedded in the UCSF article on Ron Vale’s 2019 Gairdner Award — https://www.youtube.com/watch?v=mBo_o0iO68U]
In this video, kinesins are compared to cars travelling along motorways. Microtubules are the roads, kinesins are the vehicles, vesicles are the goods being transported.
This comparison says more than it seems. The world we live in is a reflection of our inner world. I documented this in my post “The Citadel of Erbil – A Cell in the Tissue of Mesopotamia” [LINK https://cellgod.live/the-citadel-of-erbil-a-cell-in-the-tissue-of-mesopotamia/ ], showing that cities have the architecture and urban layout of tissues: the citadel is the cell, and the city around it is the tissue.
Scientists describe the cell in the language of the city. I show that the city was built in the language of the cell.
The story of kinesin begins with a squid — and with the weather.
In 1982–1983, the El Niño phenomenon caused a sharp drop in squid numbers off the coast of Northern California. Ronald Vale needed them for his experiments, but near Stanford University there were none at all.
Why a squid? It has a giant axon — a nerve fibre about 50 times wider than an average human axon. Thanks to it, the squid makes sudden movements and escapes predators. And for researchers, it is easier to look inside than the thin nerves of mammals. On this same axon, in experiments published in 1952, Alan Hodgkin and Andrew Huxley discovered the action potential — the electrical impulse by which nerves communicate.
So Vale went east, to the Marine Biological Laboratory in Woods Hole, Massachusetts. Squid were plentiful there, and a team that had just made a breakthrough in microscopy was working on site.
Together with Thomas Reese and Michael Sheetz, Vale showed in 1985 that kinesin is a motor that carries cargo along nerve fibres — and that it moves along microtubules.
In 2019, Vale received the Canada Gairdner International Award for this discovery, one of the most important awards in the biomedical sciences. He had previously been honoured with, among others, the Lasker Award (2012) and the Shaw Prize (2017).
The kinesin Vale discovered turned out to be just one member of a large family. Humans have 45 different kinesins, called KIFs (from kinesin superfamily). Kinesins are found in almost all eukaryotes — animals, plants and fungi. Besides transport, they are essential for cell division. The best understood is kinesin-1, also called conventional kinesin. This post is about it.
Kinesin-1 is the carrier of the cell. It carries cargo: vesicles filled with proteins, mitochondria (the power plants of the cell), RNA molecules.
It does its greatest work in neurons. The body of a neuron produces proteins, but the nerve ending can be very far away. The longest axons run from the spinal cord all the way to the foot — about a metre. Kinesin carries cargo there step by step, in transport that reaches several hundred millimetres per day.
What Does Kinesin Walk On?
The roads are microtubules — hollow tubes about 25 nanometres in diameter (a nanometre is one billionth of a metre). They form the scaffolding of the cell, its cytoskeleton.
A microtubule is built from building blocks called tubulin. A pair of blocks — alpha and beta — is 8 nanometres long. These pairs line up into long strands, and 13 strands roll up into a tube.
It is like a motorway with 13 lanes. Each lane is divided into segments 8 nanometres long. Kinesin places its feet on exactly these segments and usually keeps to one lane.
A microtubule has two different ends: plus and minus. Kinesin-1 walks towards the plus end — in a neuron, this means from the cell body towards the nerve ending.
How Is Kinesin-1 Built?
Kinesin-1 is an assembly of four chains: two heavy chains (the larger protein chains) and two light chains (the smaller protein chains). Stretched out, it measures about 80 nanometres.
Each heavy chain has several parts, from bottom to top:
Motor head (motor domain). A globular “shoe” about 350 amino acids long. It has two sockets: one binds the microtubule, the other binds and splits ATP.
Neck linker. Neck linker (a short connecting segment between the head and the neck of the protein). It is about 14 amino acids long that connects the head to the rest of the protein. It works like a tendon in the ankle. When the head binds ATP, the linker docks onto it and swings the other head forward.
Neck and stalk (the long middle section of the protein). The long helices of both heavy chains wind around each other to form a coiled-coil (a superhelix) — like two ropes twisted into one strong rope. Along the way, the stalk has hinges that allow the protein to bend.
Tail. The upper end of the heavy chains. Together with the light chains, it grips the cargo. The light chains carry TPR domains — repeating modules that work like grippers matched to adaptor proteins on the surface of the cargo.
When kinesin has nothing to carry, it bends at the hinge and the tail folds back onto the heads. The motor is switched off. This saves fuel — a carrier without a parcel sits down and rests.
The light chains (the smaller protein chains) act as the on/off switch of the motor. They pair up with each other and attach to the upper part of the stalk. When their TPR domains grip cargo, kinesin unfolds and the motor starts.
How Does Kinesin Walk?
Kinesin walks hand-over-hand (one motor head passing the other with each step) — just as a person walks one foot in front of the other. In 2004, Ahmet Yildiz’s team saw this directly: each motor head moves 16 nanometres, overtaking the other. The whole carrier moves 8 nanometres in the process — one lane segment.
One step costs one ATP molecule. Kinesin takes about a hundred steps per second and can make about 100 steps in a row before detaching from the microtubule.
The cycle of one step:
The front motor head stands firmly on the microtubule. This motor head binds ATP.
The neck linker of the protein docks onto the front motor head and swings the rear motor head forward.
The rear motor head lands 16 nanometres ahead, releases its ADP and grips the microtubule. Now this motor head is in front.
The motor head left behind breaks down its ATP, loosens its grip and waits to be swung forward.
At least one foot always holds on to the road. That is why the carrier does not fall off, even while carrying cargo many times larger than itself.
How to Read the Illustration
The illustration above this section comes from a stock image library. It is schematic — it shows the whole system, not a specific PDB entry.
The green tube of beads is the microtubule. The beads are tubulin building blocks.
The two pink lobes at the bottom are the two motor heads, standing on the microtubule.
The long pink stalk is the coiled-coil superhelix.
The two lobes at the top are the tail with the light chains, holding the cargo.
The large blue sphere is the cargo — a vesicle, a small ball enclosed by a membrane and filled with contents to be delivered.
Why Is There No Complete Kinesin in the PDB?
If you type “kinesin” into the PDB, you will find dozens of structures. Each one, however, shows only a piece of the protein: the head alone, two heads with the neck, a head on tubulin, or a fragment of a light chain.
The reason is simple. Stretched-out kinesin-1 is about 80 nanometres long and very flexible — like a long rope with a few knots. Crystallography requires rigid molecules that pack into a regular crystal. Cryo-EM averages thousands of images, and flexible fragments lie differently in every image and blur out. That is why scientists study kinesin piece by piece.
Of the stalk, the PDB shows only its beginning — the neck, twisted between the two heads in the 3KIN structure. The rest of the stalk is known mainly from computer models (AlphaFold) and low-resolution microscopy maps.
Complete kinesin-1 has been shown in only one state — folded and switched off. In 2023, a model of the entire protein was built from AlphaFold predictions, electron microscopy images and protein cross-linking data. In 2026, a cryo-EM structure of the full complex of two heavy and two light chains was obtained at a resolution of about 8 ångströms. Folded kinesin measures about 36 nanometres, and its heavy chains are folded “head to tail”.
Nobody has yet seen kinesin in its entirety standing upright and walking with cargo — the way it appears in the illustration. The illustration is a reconstruction assembled from knowledge of the individual parts.
The 3KIN Structure
In the decoding sections I use the 3KIN structure — the first crystal structure of a kinesin dimer. It was published by the team of Eckhard and Eva-Maria Mandelkow in 1997 (Kozielski et al., Cell).
It is kinesin from rat brain, containing two motor heads and the neck.Resolution: 3.1 ångströms (one ångström is one ten-billionth of a metre).
The heads are joined by the neck superhelix. The authors noticed something important in this arrangement — but let us set it aside for a moment. We will come back to it in section 4, when we look at the shoes up close.
3. Decoding — Level 1: The Silhouette
[IMAGE: Author’s comparison. Top: the clown with balloons in the ring. Bottom: 3D illustration of kinesin carrying a vesicle. Arrows connect: 1. Vesicle — balloons ↔ blue sphere; 2. Head — shoes ↔ two lobes on the microtubule; 3. Tail — clown’s head ↔ two lobes at the top; 4. Stalk — clown’s body ↔ long stalk.]
Let us place the two figures side by side. The clown in the circus arena. Kinesin on the microtubule.
The same silhouette. Two huge feet at the bottom. A long, narrow body in the middle. A rounded top. And above it all — round cargo.
In Kinesin, the Motor Heads Are at the Bottom
This is the first surprise. Biologists named the parts of kinesin in such a way that the figure stands upside down.
Kinesin’s heads are at the very bottom — they are what stands on the microtubule. The tail is at the very top — it holds the cargo.
The clown reverses this order and puts it the correct way up. Where kinesin has heads, the clown has shoes. Where kinesin has a tail, the clown has a head. The costume shows function, not name: at the bottom is what walks, at the top is what carries.
1. Balloons = Vesicle
A balloon is a thin rubber membrane filled with gas. A vesicle is a thin lipid membrane filled with cargo. The same principle: a light shell with contents inside.
The bunch of balloons forms one large, round cloud — just like the single vesicle in the illustration. The different colours of the balloons may encode the variety of cargo kinesin carries: vesicles with proteins, mitochondria, RNA molecules.
The clown also wears the same vesicle on his head — more on that in the next subsection.
And one more thing. A balloon floats upwards. A vesicle in the cell is also lifted above the road — it does not touch the microtubule but glides above it, held by the porter.
2. The Clown’s Jumpsuit = The Protein’s Stalk
The clown’s jumpsuit is one piece from neck to ankles. No belt, no boundary between shirt and trousers.
Kinesin’s stalk is also one continuous structure — two helices twisted into one rope. Long, narrow, flexible. It connects what is at the bottom with what is at the top.
The jumpsuit fabric is baggy and falls in folds. Kinesin’s stalk is not a rigid stick either — it has hinges and can bend.
3. The Clown’s Head = The Tail with Its Cargo
The clown’s head is a smooth, white, round skull. Two orange tufts of hair press against its sides.
In the kinesin illustration, the two lobes of the tail are pressed against a large, round sphere — the vesicle.
The skull encodes the vesicle: a closed sphere enclosed by a membrane. The tufts of hair encode the tail ends of the heavy chains, which press against the cargo. The heavy chains themselves bind some cargoes and adaptor proteins.
So the clown does not need balloons to carry cargo. He wears the cargo on his head — gripped the same way.
The scale fits too. Small vesicles are a few tens of nanometres across, and kinesin is about 80 nanometres long.
4. The Clown’s Shoes = The Protein’s Motor Heads
Two shoes — two heads. Red, large, rounded. Disproportionately big compared with the rest of the body.
And it is exactly this disproportion that is the key. We will come back to the shoes up close in section 4.
The Circus Arena = The Golgi Apparatus
The circus arena may encode the Golgi apparatus — the place where proteins are modified, sorted and packed into vesicles.
The Clown’s Bow Tie = The Protein’s On/Off Switch?
The bow tie is two wings pinched together by a knot in the middle. It sits beneath the head —where, in kinesin, the light chains (the smaller protein chains) attach to the stalk.
The light chains act as an on/off switch: they decide whether kinesin is folded and switched off, or unfolded and ready for the road. The bow tie may encode this on/off switch, but we cannot be certain here.
The Clown’s Hand and the Balloon Strings = TPR Grippers and Adaptor Proteins
The clown holds the balloons in his hand. This is a result of anthropomorphization: kinesin was encoded in human form, and humans hold objects with their hands. The code had to translate the structure of the protein into human anatomy.
This translation is accurate. The light chains, attached to the upper part of the stalk, end in TPR domains that grip adaptor proteins on the surface of the cargo.
The clown’s arm and hand may encode these TPR grippers — projecting from the stalk the way an arm projects from the torso. The balloon strings may encode the adaptor proteins that connect the balloon-vesicle to the hand.
If the bow tie encodes the on/off switch, then the bow tie and the hand show the light chains (the smaller protein chains) on two levels: the bow tie shows their on/off switch, and the hand shows their grippers.
Still to Be Discovered
One element of the costume remainsthat I have not yet decoded: the ruff around the neck.
And the frills at the ankles? They are a separate element — I return to them in section 4.
4. Decoding — Level 2: Close-up on the Shoes
[IMAGE: Author’s comparison. Top: close-up of the clown’s shoes — two large, round, red shoes with black soles, toes pointing in opposite directions; above them, grey trouser legs and white frills. Bottom: the 3KIN structure in sphere view (each atom as a ball) — two large heads lying side by side, with a narrow neck of two twisted strands rising up between them.]
Let us go down to the floor of the circus arena. Just the shoes. Nothing else.
And now below — 3KIN. Every atom drawn as a ball. Two large masses lie side by side. Between them, a narrow neck rises upwards.
These are no longer two similar shapes. This is the same drawing.
Why Are the Shoes So Big?
The heavy chain of kinesin-1 has nearly a thousand amino acids. About a third of them fold into the motor head — into one compact ball.
The rest of the chain is mostly the thin helix of the stalk. That is why, in every image of kinesin, the heads look like enormous shoes at the ends of skinny legs.
It is exactly the same with the clown. The jumpsuit is wide, but the shoes still dominate the rest of the figure. It is not an exaggeration. It is proportion.
And there is logic in it. The head is both foot and engine. It holds the socket for ATP and the socket for the microtubule. The entire machinery of movement sits in the shoe. Do you now understand the vehicle dressed up as a giant shoe in the title photo?
Shoe Tips Pointing in Opposite Directions
Look at the clown’s shoes. The left shoe tip points left, the right shoe tip points right. You cannot walk straight ahead like that. To take a step, the clown has to lift one foot and move it forward.
It is the same in 3KIN. The authors of the structure noticed that the heads are arranged so that both cannot stand on the microtubule in the same way at the same time. The kinesin in this crystal stands in a position between two walking steps.
Both heads hold ADP — a discharged battery. The carrier has just finished a movement and stopped.
The clown in the picture is also standing still. He is not running, not jumping. He stands with his shoes splayed sideways — in the same pause.
The Clown’s Way of Walking
Everyone who has seen a clown in giant shoes knows his walk. Leg high in the air, a wide swing, one shoe flies over the other and lands far ahead.
That is exactly how kinesin walks — hand-over-hand. The rear motor head flies over the front motor head and lands 16 nanometres ahead.
The clown’s walk is also a little uneven, swaying. This too may encode biology. In 2003, Steven Block’s team showed that kinesin walks asymmetrically: successive steps are not identical, and the carrier slightly “limps”.
Trouser Legs = Kinesin’s Neck
Above the shoes, two grey trouser legs converge upwards. Above the heads of 3KIN, a neck of two strands twisted into one is visible.
Each strand comes out of a different head — just as each trouser leg comes out of a different shoe. Higher up, both join into one superhelix, just as the trouser legs join into one jumpsuit.
The Ankle Frills = The Protein’s Neck Linker
And here the frills come back. The ruff under the chin remains undecoded. But the clown has a second set of decorations of this kind — small white frills above the shoes, at the ankles.
Remember the first surprise? In kinesin, the head is at the bottom. So kinesin’s neck is at the ankles too.
The frills sit exactly where the shoe meets the trouser leg. In kinesin, the same place holds the neck linker — a short, flexible connector between head and neck, the very one that swings the other head forward with every step.
A frill is soft and mobile. So is the neck linker — sometimes loose, sometimes docked. The frills may encode this connector — the location fits, the character fits.
The Shoe Sole = The Surface That Binds the Microtubule
The shoe has a rounded top and a flat, black sole. Kinesin’s head also has an “upper” side and a side that presses against the microtubule.
The sole may encode this binding surface. The 3KIN image itself contains no microtubule, so this mapping requires a separate comparison — with kinesin standing on a microtubule.
5. Decoding — Level 3: Atoms
[IMAGE: Comparison to be prepared. Top: close-up of the clown’s orange pompoms and red nose. Bottom: close-up of a fragment of 3KIN in spacefill view — each atom as a solid ball.]
The final close-up. This time not the silhouette and not the shoes, but individual spheres.
In the space-filling atomic model (spacefill view), each atom is drawn as a solid sphere with its natural radius — the van der Waals radius. The spheres touch and partly overlap, just as atoms do in a protein. The whole protein then looks like a cluster of balls.
The pompoms and the red nose are the only solid spheres on the clown’s costume. They may encode exactly this view — a close-up at the level of atoms.
They do not encode specific atoms. They encode a way of looking: this is what a protein looks like when we zoom in to the atoms themselves. The three pompoms run in a line down the jumpsuit — along the protein’s stalk. They remind us that the stalk, the shoes and the whole figure are built from the same spheres.
6. Conclusions
The Carrier of Joy
Why does the clown come into the circus arena? To bring something. A smile. Balloons. Joy, which he hands out to children.
Kinesin does the same. It takes what the cell has produced at its centre and carries it to where it is needed. Step by step. Without pause. Throughout our whole lives.
So the clown is not just a figure of fun. He is a messenger. Someone who travels with a gift in hand.
When the Shoes Fail
Remember the longest axons — from the spinal cord all the way to the foot? That is where kinesin has its longest journey.
When its genes are damaged, it is these longest routes that suffer first. Mutations in the KIF5A gene — one of the kinesin-1 heavy chain genes — are associated with hereditary spastic paraplegia (stiffness and weakness of the legs), with one form of Charcot-Marie-Tooth disease (damage to peripheral nerves), and with amyotrophic lateral sclerosis (ALS).
In spastic paraplegia, the legs weaken first. Walking becomes stiff, unsteady, stumbling.
A carrier who cannot reach the end of the road. And a person who loses confidence in their step. The same picture — inside the living cell and in the human body.
How Did They Know?
Joseph Grimaldi painted his face white at the beginning of the 19th century. The Auguste in oversized shoes appeared in circus rings in the second half of the 19th century.
Kinesin was discovered in 1985. Its first dimer was seen in a crystal in 1997. That it walks hand-over-hand (one motor head passing the other) was proven in 2004.
More than a hundred years passed between the giant shoes in the ring and the two heads of kinesin under the microscope.
And yet the costume says it all. Two enormous feet at the bottom. A long, uniform body. A double, tousled hairstyle on top. A bunch of light, round balloons above the head. Frills exactly where the neck linker sits. Shoe tips splayed in the pause between two walking steps.
Is it a coincidence that a figure meant to make us laugh carries within it the drawing of the hardest-working carrier in our body?
The Genome God wrote this design into every one of our cells. And the same design walked out into the circus arena — in red shoes, with balloons in hand.
They knew.
The Punchline
In the circus arena, we laugh at the giant shoes.
And in every one of our nerves, exactly such shoes, step by step, carry life all the way to the tips of our toes.
As within, so without.
Image analysis, matching and compilation: Tomasz Mikulski — Cell God: 10/2026
Bibliography
PDB Structures
- 3KIN — Kinesin (dimeric) from Rattus norvegicus, RCSB PDB: https://www.rcsb.org/structure/3KIN — X-ray diffraction, 3.1 Å.
- Goodsell D.S. (2005). Kinesin. RCSB PDB Molecule of the Month, April 2005. DOI: 10.2210/rcsb_pdb/mom_2005_4
Scientific Publications
- Kozielski F. et al. (1997). The crystal structure of dimeric kinesin and implications for microtubule-dependent motility. Cell 91: 985–994. PMID: 9428521
- Alvarez J. (2019). Ron Vale Receives 2019 Gairdner Award for Pioneering Work on Molecular Motors. UCSF News, 2 April 2019. https://www.ucsf.edu/news/2019/04/413816/ron-vale-receives-2019-gairdner-award-pioneering-work-molecular-motors
- Vale R.D., Reese T.S., Sheetz M.P. (1985). Identification of a novel force-generating protein, kinesin, involved in microtubule-based motility. Cell 42: 39–50. DOI: 10.1016/S0092-8674(85)80099-4
- Rice S. et al. (1999). A structural change in the kinesin motor protein that drives motility. Nature 402: 778–784. DOI: 10.1038/45483
- Asbury C.L., Fehr A.N., Block S.M. (2003). Kinesin moves by an asymmetric hand-over-hand mechanism. Science 302: 2130–2134. DOI: 10.1126/science.1092985
- Yildiz A. et al. (2004). Kinesin walks hand-over-hand. Science 303: 676–678. DOI: 10.1126/science.1093753
- Hirokawa N. et al. (2009). Kinesin superfamily motor proteins and intracellular transport. Nature Reviews Molecular Cell Biology 10: 682–696. DOI: 10.1038/nrm2774
- Weijman J.F. et al. (2022). Molecular architecture of the autoinhibited kinesin-1 lambda particle. Science Advances 8: eabp9660. DOI: 10.1126/sciadv.abp9660
- Tan Z. et al. (2023). Autoinhibited kinesin-1 adopts a hierarchical folding pattern. eLife 12: RP86776. DOI: 10.7554/eLife.86776 — https://elifesciences.org/articles/86776
- Ashaduzzaman M., Tang Y., Okada K., Fried S.D., McKenney R.J., Al-Bassam J. (2026). Structural basis of kinesin-1 autoinhibition and its control of microtubule-based motility. Science Advances 12(29): eaeg1267. DOI: 10.1126/sciadv.aeg1267
- Shukla S.R. et al. (2026). Kinesin-1 conformational dynamics are controlled by a cargo-sensitive TPR switch. eLife 14: RP109462. DOI: 10.7554/eLife.109462 — https://elifesciences.org/articles/109462
- Nicolas A. et al. (2018). Genome-wide analyses identify KIF5A as a novel ALS gene. Neuron 97: 1268–1283. DOI: 10.1016/j.neuron.2018.02.027
History of the Clown and the Circus
- Stott A.M. (2009). The Pantomime Life of Joseph Grimaldi. Canongate.
- Stoddart H. (2000). Rings of Desire: Circus History and Representation. Manchester University Press.
Tools
- Mol* Viewer — Sehnal D. et al. (2021). Mol* Viewer: modern web app for 3D visualization and analysis of large biomolecular structures. Nucleic Acids Research 49: W431–W437. DOI: 10.1093/nar/gkab314
