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To travel forth so far

Shakespeare, exported to new stages

Rachel Rodman 19 April 2015

Wild type: pushing the boundaries of the Bard

For Shakespeare, there would be no biological immortality. His genetic line ended in 1670, with the death of his granddaughter. In the centuries after his death, only his work persisted

Shakespeare wrote in a rather different sort of universe. His solar system contained no Uranus or Neptune. His natural world – meadows, forests, human bodies – consisted of units no smaller than could be perceived by eye.

For Shakespeare, there would be no biological immortality. His genetic line ended in 1670, with the death of his granddaughter. In the centuries after his death, only his work persisted.

Shakespeare's characters – his heirs – would come to inhabit many new environments. From their birthplace, in England, they would rapidly disperse. From their mouths, new languages would emerge. On their bodies, they would wear new costumes, adapted to new climates.

These unique bodies – defined by Shakespeare's pen, and not by his loins – influenced the mode of this dispersal. In some ways, they were considerably less flexible than genetic descendants would have been. Their central nervous systems, composed of ink, were best suited to repetition...not to learning. Over and over again, they would confront the same obstacles, with precisely the same flawed response. Night after night (performance after performance), Othello would be guided by Iago's lies. King Lear would reward the wrong daughters. Romeo would kill himself, while embracing a living Juliet. (Upon them, also, natural selection would operate in a rather different way. Variants of Othello and Lear and Romeo who made more sensible choices would not displace the self-destructive parental types.)

In other ways, of course, they were much more adaptable than any biological heir could have been. Fewer physical rules constrained them. With a few small strokes, their existing bodies could be entirely overhauled. Human engineers could remove (previously essential) structures, or implant new ones. They could also dramatically re-imagine what a “body” might consist of.

To the Solar System:

In our solar system, Uranus is the seventh planet from the sun. It is named for the Greek god of the sky. For many years, Uranus the god was the lover of Gaia, the earth goddess. Eventually, of course, this relationship ended, when Uranus' penis was cut off by their son, Cronus (Saturn).

As a planet, Uranus shares several features with the god. It is very large (four times the width of earth). It also lacks a penis. Its position, in addition, retells the god's genealogy. Uranus (planet 7) is the father of Saturn (planet 6), who is the father of Jupiter (planet 5), who is the father of Mars (planet 4).

About Uranus orbit twenty-seven moons. Twenty-four are named after characters from Shakespeare's plays. (The remaining three come from Alexander Pope). None bears any particular relationship to the Greek god. They do not appear in any common stories. They do not have shared friends. As neighbors, their interactions may be marked by a certain cultural awkwardness: Who are you, anyway...?


In the late 16th century, Shakespeare wrote A Midsummer Night's Dream. It featured a forest kingdom, ruled by two fairies, Titania and Oberon. They were decadent rulers. Assisted by their servants, they pursued a range of bizarre whims. In Act 3, for instance, Titania directed a servant “to fan the moonbeams” from the eyes of a donkey-headed visitor. In the same act, Oberon directed a servant to alter the weather (“overcast the night...with drooping fog”) as part of a plot to re-engineer the relationships of four strangers.

In 1787, Titania and Oberon adopted very different roles. At the behest of the astronomer William Herschel, they undertook a harrowing journey – out of their forest, off of earth, and to the very edges of the solar system. At the same time, they abandoned their fairy bodies – arms, legs, heads. In place, each took on a spherical costume, consisting of ice and rock. This new identity also required a dramatic expansion. From a likely starting mass of less than 100 kg, each ballooned more than 1019-fold, taking on a final mass of over 3 x 1021 kg.

This physical transformation may perhaps be summarized by modifying one of Titania's own speeches (Act 3):

I will purge thy...[humanoid anatomy] thou shalt like an...[Uranian satellite]...go.

In later centuries, other Shakespearean characters would make the same transition. Upon arrival, each was presented with a new set of instructions. These scripts, though thick, were repetitive and easily summarized. They stipulated: revolve around Uranus again and again and again and again.

To the Genome:

Drosophila melanogaster is a model insect. It is small – just a few millimeters. It feeds on rotting plants. Its juveniles are limbless and earthbound; its adults are winged.

For many scientists, Drosophila is an attractive tool. It is easy to grow and easy to breed. It has a small genome, which is easy to manipulate.

Within Drosophila – encoding it, defining it – there are many genes. Some act to built the body – the stomach, the brain. Others support the action of existing body parts. They help to digest food, or to relay nervous impulses.

In describing these genes, scientists have given each a unique name. Some are quite colorful. The tinman gene (which is essential for heart formation) is named for a character in The Wizard of Oz, who lacks a physical heart. The methuselah gene (which influences longevity) is named for a character in the Bible, reported to have died at age 969.

With these names, Drosophila has become very human. Cultural references pattern its interior. Each cell contains a copy of Groucho and Smaug, present on chromosome 3. Thor, with his thunderbolts, resides on chromosome 2. Lava Lamp and Ariadne add their own color, stationed on the X chromosome.


In the early 17th century, Shakespeare wrote The Tempest. It featured a strange island, inhabited by a magician, Prospero, and his daughter, Miranda. The humanoid Caliban (“half a fish and half a monster”) served as their slave.

With magic spells (“his art is of such power”), Prospero shaped the play's major events. Re-directing the course of a ship, he staged a showdown with his old enemies. With additional magic, he regained the dukedom of Milan, and engineered an engagement between his daughter and the Prince of Naples.

In 1991, Prospero adopted a new role. Biologists, studying the development of the nervous system, reassigned him as a Drosophila gene. With this new identity, he exercised his power over a very different kind of story. Using new tools – but no magic – he helped to coordinate the transition from an immature cell, a neuroblast, to a mature cell, a neuron.

When a character becomes a gene, part of the challenge is physical. Familiar structures, consisting of flesh, are replaced with a single (or double) string of molecules. This body is, in addition, measured in new units, consisting of the billionth part of a meter.

This type of transition also requires existential adjustments. A dramatic character is already fairly schizoid in his understanding of himself. (He is present simultaneously on many different stages; he is embodied simultaneously by many different actors.) In becoming a gene, however, his identity is fragmented manyfold further. He exists at the center of every cell, within every member of the species. During each cell division, he gives birth, again, precisely to himself.

Prospero's own transition, from character to gene, may perhaps be summarized by modifying some of his own lines, from Act 5:

But this...[macroscopic existence]
I here abjure...I'll break my staff, Bury it certain fathoms in the...[ Drosophila genome]. And deeper than did ever plummet sound; I'll...[incorporate myself into this genetic] book.

In the following decades, additional Shakespearean characters joined Prospero. Miranda and Caliban arrived in 1997 and 2005, each assigned to embody a protein with which Prospero (or pieces of Prospero) had been shown to interact directly.

In 1995, Malvolio, the cranky butler from Twelfth Night, also entered Drosophila. Once described to “taste with a distempered appetite,” he would re-direct his talents as one of the genes that underpinned the fly's preference for sugary foods.

In 2002, finally, Hamlet underwent a similar miniaturization surgery, cast to oversee a developmental decision in the IIB cell lineage. His assignment referenced his famous “to be or not to be...” soliloquy (IIB = two B = to be). As in the soliloquy, Hamlet would govern a fundamental choice: whether to pursue a future as a nerve cell with a single dendrite (an extension, for receiving sensory input), or as one possessing multiple dendrites.

As in the play, Hamlet the molecule would also reflect upon questions of life and death. When Hamlet function was disrupted, the balance of these reflections would tip prematurely death-wards. hamlet mutants would die as larvae. Moulting “off this mortal coil,” they would enter “the undiscover'd country.”

“If This Were Played Upon a Stage”:

After Shakespeare's death, there were many scientific discoveries. Each opened a new potential habitat. Units of culture – including Shakespeare's characters – were rapidly modified, in order to populate them.

In 1939, for instance, Othello's Iago would acquire fins, and swap a lung-based gas exchange system for a gill-based one. With these accoutrements, he would descend into the ocean, assigned to embody a genus of shark. Here, he would participate in new – perhaps more violent – dramas, whose victims included bony fishes and cephalopods.

In the orbit of Uranus, and in the genome of the fly, additional stories were initiated. These were chimeric dramas. They consisted, in part, of the characters' own backstories, as Shakespeare first imagined them. They consisted, also, of events in the physical world – stories, of another sort, which had been told and retold for many millions of years.

In 1986, heroines from eight plays were exported into space, following discoveries by the NASA probe Voyager 2. Startled by the cold and dark, Rosalind (later, moon 8) may have spoken for the group (As You Like It, 1.3):

Alas, what danger will it be to us,
Maids as we are, to travel forth so far!

As Uranus' eight innermost moons, they framed a new drama. At speeds of more than 20,000 mph, they formed brief alignments, one with the other, until the inner moons again surged ahead. Tragic characters networked cyclically with comedic characters. Females forged new friendships, in the absence of the males that had previously defined them. Hamlet's Ophelia (moon 2) responded to slight gravitational influences from King Lear's Cordelia (moon 1) and The Taming of the Shrew's Bianca (moon 3) Prince Hamlet, however, who was not a moon, she could no longer be physically perturbed.

On earth, on a much smaller set of stages, Hamlet also adapted to a very new script. In the cell of a fly, he underwent a series of profound identity shifts – from DNA, to RNA, and then to Protein. In this final avatar, he served as a transcription factor – himself assisting in the identity shifts (DNA to RNA), as required by a series of non-Shakespearean characters. In this complex theatrical space, events on both sides of the curtain (backstage and forestage; nucleus and cytoplasm) served, equally, as performance.

In this same space, a heavily adapted version of The Tempest was also staged. Gentle Miranda guided her father's protein-avatar, ensuring that he would be inherited only by the cells that required him. To escort him, she grasped him by a specialized lapel – the asymmetric localization domain (ALD). Sinister Caliban, meanwhile, also lay in wait, eager for an opportunity to grasp a specialized hem of Prospero's garment (the nuclear export signal, or NES).

On another section of this stage stood Malvolio. In Twelfth Night, he had been the victim of a cruel prank. Duped into believing that the Countess loved him, he altered his dress (“in yellow stockings, and cross-gartered”) and behavior. One of Malvolio's tormentors (while howling with laughter) described this transformation as “an improbable fiction.”

In Drosophila – encoding an ion transporter – Malvolio may have experienced mixed feelings. Stung by his earlier humiliations, he may have wished to understand the degree to which this new assignment was a joke...and, if it were a joke, upon which side of it he fell. Did his new role – in the service of a fly – simply expose him to a new kind of laughter? (Had his tormentors, in fact, engineered it?) Or was it, instead, a great honor? As a gene, he was the subject of educated (and increasingly wide-ranging) scrutiny...compared, for instance, with immune cell proteins (Nramps) in other organisms, and invoked during discussions of labor division in honey bees. In this new light, Malvolio may have again reflected upon a mocking letter he had once received, and wondered whether, contrary to the wishes of those who had penned it, it might have constituted a real prediction:

..but be not afraid of greatness: some
are born great, some achieve greatness, and some
have greatness thrust upon 'em.

“An Improbable Fiction”:

Certain ambiguities, familiar to Malvolio (Absurdity or Greatness? Joke or Honor?), attend many efforts to export Shakespeare's characters. The relationship between Shakespeare and the scientific communities that have organized these exportations can be difficult to dissect. Within these transactions, upon whom, predominantly, is the honor/joke placed? Upon Shakespeare? Upon the team of scientists? Upon the non-human phenomena that receive the Shakespeare-inspired names?

Temporal complexities also attend these projects. Each straddles Shakespeare's lifetime...somewhat excluding the man himself. Based upon knowledge that was acquired only after Shakespeare's death, they connect Shakespeare to narratives that began long before his birth (initiated, respectively, by events in the early solar system, and in the evolution of life). They are not concerned with biography. They serve, instead, as speculative exercises – opportunities to experiment, as Shakespeare might have done, had he been forced (privileged?) to employ a non-human cast, and an alternatively-sized set of stages.

What the real Shakespeare might have thought – an English playwright, at the turn of the 16th/17th century – is, of course, impossible to determine. We can hope that he would have felt the honor, or been amused by the joke, or, at least, been not too profoundly repelled, by the somewhat awkward stories that resulted from his characters' reassignments. In articulating a response to these performances, he might perhaps have availed himself of some of the words of Twelfth Night's Clown. To these weird new scripts – which he had, to a certain degree, incited, but whose existence, afterwards, he could not entirely credit – he may simply have wished to emphasize his surprise:

If this were played upon a stage now, I could
 condemn it as an improbable fiction.

Additional Reading:

Doe et al. The prospero gene specifies cell fates in the Drosophila central nervous system. Cell, 1991.

Shen et al. Miranda is required for the asymmetric localization of Prospero during mitosis in Drosophila. Cell, 1997.

Bi et al. Drosophila caliban, a nuclear export mediator, can function as a tumor suppressor in human lung cancer cells. Oncogene, 2005.

Moore et al. hamlet, a binary genetic switch between single- and multiple- dendrite neuron morphology. Science, 2002.

Rodrigues et al. malvolio, the Drosophila homologue of mouse NRAMP-1 (Bcg), is expressed in macrophages and in the nervous system and is required for normal taste behaviour. EMBO J, 1995.

Ben-Shahar et al. Phenotypic deconstruction reveals involvement of manganese transporter malvolio in honey bee division of labor. J Exp Biol, 2004.