The yeast Saccharomyces cerevisiae is one of the most studied model systems for molecular and cellular biology. In 1996, it became the first eukaryotic organism to have a completely sequenced genome (Dujon, 1996; Goffeau et al., 1996), which led to a number of valuable and widely accessed databases. Among its features is the short generation time (usually 90–120 minutes) and the ability to grow at various temperatures in relatively inexpensive media. Moreover, many of its genes are well characterized, thanks in part to its amenability to modifications such as gene disruption, gene marking, mutations, or gene-dosage modifications. Because of these advantageous features, it has become the model organism of choice for many investigators in fields ranging from basic biology to biomedical research.

THE NURSING SHE-WOLF AND THE GODDESS OF SUCKLING

In the infancy narrative of romulus and remus, the wolf does not prey on any of the protagonists. The true predator, in this legend, is the war god Mars, who raped a virgin in her sleep, impregnated her, and then took off. To be fair to this otherwise deadbeat father, he sends a couple of animal associates – the she-wolf and the woodpecker – to watch over his two sons. It is bloody business to rape a virgin, and bloodier still must have been Rhea Silvia's delivery of the twins. Blood flows at the twins' conception: A virgin, whether raped or willing, usually bleeds at the moment of deflowering. Blood flows at the twins' birth: Human birth is always bloody. Blood flows again, more unpredictably this time, when the city is founded: Romulus, it is known, slayed his own brother Remus and remained Rome's sole founder. Blood signifies violence (sometimes) and pain (usually) but also the historical necessity of Rome's birth through the hurt of the woman who facilitated the event and the death of our hero's antagonist – who also happens to be, alas, his only brother. Blood, however, is not the only bodily fluid that visibly flows in this story of early Rome: Without the milk of the she-wolf – milk meant for wolf cubs but eagerly sucked down by two ravenous human babies – Rome's founder surely would have succumbed.

RECYCLING THE SHE-WOLF AND THE DIPTYCH OF RAMBONA

Rome is a palimpsest, it is often said: as the shapes of old writing remain faintly visible under more recent layers of text on a manuscript repeatedly inscribed, traces of the city's past also may be read underneath its current appearance. Elizabeth Barrett Browning said it well in 1854: “It's a palimpsest Rome – a watering place written over the antique” (2: 165). The Eternal City destroys nothing, preferring to transform objects instead to suit current beliefs and customs: Temples become churches when a cross is placed at their heart, pagan monuments are turned Christian by the addition of holy people's statues, images of gods are allegorized into the likeness of saints, and so on. Neither the words telling of the she-wolf nor the images that picture the Roman beast have escaped this process of reconstruction and reinterpretation through verbal and visual representation. Rome and the she-wolf play at writing and rewriting their own past; Rome and the she-wolf constantly recycle themselves – like sheets of parchment too precious to discard after just one use. The city's layers, like the she-wolf's, are physical as well as literary, artistic, and – more generally – cultural; they embrace objects and texts as well as beliefs and practices.

“It is curious to note in Rome how many a modern superstition has its root in an ancient one,” noticed William Wetmore Story in 1863, “and how tenaciously customs still cling to their old localities.”

THE NEOCLASSICAL SHE-WOLVES OF VALADIER AND PINELLI

The roman she-wolf is a cultural monument. whether made of stone or words, metal or pigment, the picture of a wild beast nursing two human babies is an enduring one and continues to speak of the past – especially of a great Roman past – like few other objects. In the course of the nineteenth century, the she-wolf became an icon of the desire for Italian national unity: Her indigenous form united nature and culture, present and past, and two different babies under one protector. Naturally, then, images of the she-wolf appear in architectural structures of modern Rome. There are she-wolves, for example, on that most visible of its buildings, the Vittoriano of Piazza Venezia, the controversial monument to the first king of united Italy, Victor Emmanuel II, situated at the foot of the Capitoline Hill. An allegorical she-wolf accompanies the Tyrrhenian Sea in one of the Fountains of the Two Seas and, at the top of the monument, the bronze allegories of Liberty and Unity ride triumphal chariots on the hubs of which wheels are the heads of she-wolves. In a monument such as the Vittoriano, which repeatedly summons images of past incarnations of Rome – a monument that has been described as “a rhetorical device for manipulating public memory” (Atkinson and Cosgrove 31) – the she-wolf carries out her multiple political tasks.

THE SHE-WOLF AND THE MEANING OF HISTORY IN WORDSWORTH AND MACAULAY

Literally speaking, there is no history of the she-wolf. Ancient writers, historians included, did not usually pretend to believe in a flesh-and-blood beast stumbling across a set of whimpering baby boys and offering them suckle. Almost from the beginning, figurative interpretations were devised to account for the story of Romulus's serendipitous rescue from certain death by the intervention of a wild animal. “I have no intention either to affirm or refute,” wrote Livy; “Some say,” echoes Plutarch. Although the she-wolf has no history that today we would promptly recognize as such, this same animal has repeatedly compelled writers to examine the meanings of the discourse we call history. Livy, Dionysius, and Plutarch prefaced the she-wolf's tale with reminders of the unreliability of their sources. Propertius hoped to tell of Rome's origins by imitating the she-wolf's task. Through the memory of the she-wolf, Petrarch could nostalgically relate to the grandeur of Roman antiquity. For du Bellay, the she-wolf's death allegorized the historic fall of Rome itself, the fate of an entire nation epitomized by that of a single beast. The rhetorical figure of allegory, as well as the language of misogyny, entertains a close relationship with the discourse of history. This relationship may be adversarial, with history proving allegory obsolete, or it may be complementary: Allegorical figures, in this sense, continue to help us interpret the meaning of history.

THE LUPA AS SYMBOL OF POWER AT THE LATERAN

Maxime Formont's 1912 historical novel, The She-Wolf: A Romance of the Borgias, provides imaginative insight to the impression that the bronze beast must have made during her Lateran residence – the sense of her might heightened by physical and historical distance (she was positioned atop the column and reminiscent of an ancient founder), as well as by the belief in the Lupa's origin on the Capitoline Hill, her endurance across the ages, and the sharp fierceness of her eyes, ears, teeth, and jaws. Although Formont's novel is set during the high Renaissance, the Lupa stayed before the Lateran Palace only until 1471.

MISOGYNY AND THE SHE-WOLF FROM TERTULLIAN TO MASUCCIO

Unlike their pagan predecessors discussed inchapter 4, early Christian writers were not as tolerant of sexual promiscuity, no matter how generous – with milk or money – were the women who practiced it. In his tract titled To the Nations, Tertullian (ca. 155–230), the first great writer of Latin Christianity, rather predictably thundered against the “abominable cases” of shameless veneration of unworthy divinities – Acca Larentia, first and foremost. She is honored, Tertullian inveighed, even though “she was a hired prostitute, whether as the nurse of Romulus, and therefore called Lupa, because she was a prostitute, or as the mistress of Hercules, now deceased, that is to say, now deified” (486). Tertullian described this woman's two major identities: He saw Larentina (i.e., Acca Larentia, her name having several variations) in either the lupa who rescued and nursed Romulus (no great honor; Tertullian had little respect for Rome's founder, in his view a trickster, fratricide, and rapist) or in the wealthy prostitute who had received great riches through Hercules's help and at her death bequeathed her fortune to the Roman people. In gratitude, the Romans deified her. However, what could be the distinction of accessing heaven for heathens and of becoming divine if prostitutes “mount it in all directions” too? Another early Christian writer, Lactantius (third–fourth century ce) – probably from North Africa like Tertullian – likewise ironizes, in his discussion of the Romans' divinization of the she-wolf: “Now how great must that immortality be thought which is attained even by harlots!” (54).

MAMMARY MEANINGS IN THE SHE-WOLF'S VISUAL REPRESENTATION

With a spare compositional structure of three naked figures – one adult animal and two juvenile humans – the statue of the Lupa Capitolina recounts a foundation legend far more intricate than her own shape. Hers is the story that Romans have been telling for centuries about their own past, so that it is enough to see a wolf give suck to a set of twins to know exactly who she is, what she has done, and what will happen a few years hence. In a compact frame, the bronze image at the Capitoline Museum expresses ancient legends of divine intervention in human history. This is the story that the Lupa has long been believed to be telling; the statue, after all, may have had nothing to do with the twins when the bronze was first cast. Then there are the stories of the bronze She-Wolf herself, tales of varying reception and changing usage, including accounts of both worldly and supernatural power struggles – ancient as well as medieval, early modern, and contemporary. Of course, the complexity of the bronze Lupa can best be understood in the context of other visual as well as verbal she-wolves and by being mindful of the interpretations attached to all these beasts.

The wolf's much-sung teats and tongue dramatize attitudes of and toward mothers, of and toward poets.

The word troia in italian may describe a female animal – a sow – or a human sex worker: troia is a derogatory term for a female prostitute. Capitalize the word, however, and it becomes, instead, the illustrious ancient city of Troy, Rome's urban ancestor. Likewise, the Latin word lupa has two meanings: It indicates, metaphorically, a prostitute; yet lupa is also the she-wolf who suckled Romulus and Remus. It is from the fusion of two separate legends embedded in each of these two ambiguous, grammatically feminine words – troia and lupa – that the story of Rome's birth derives: the fifth-century bce Greek legend, which attributed Rome's founding to the Trojan hero Aeneas around the twelfth century bce, and the later indigenous legend of the she-wolf, dating from 300 bce and according to which a native founder and his twin brother were saved by a local beast four and a half centuries earlier. By the first century bce, the two legends had blended harmoniously, making Romulus a descendant of Aeneas as a compromise and with a few hundred years elapsing between the births of the two alleged (and, until then, competing) founders. However crucial Aeneas's arrival on Italian soil, though, and no matter how necessary Romulus's building of Rome's first wall and his murder of his brother, it is the scene of the she-wolf nursing two baby boys – and the narrative that this image tells – that remains the most visible and most frequently represented moment in the story of Rome's foundation.

A SHE-WOLF AMONG ROME'S ANIMALS

When considering the presence of animals in the history and the geography of Rome, there is little doubt that the she-wolf comes first: There is no Rome before her and there would be no Rome without her. Rome is a she-wolf and the she-wolf is – in so many ways – Rome. As Francesco Domenico Guerrazzi, writer and politician, wrote in 1863, “Rome has the nature of the she-wolf” (572). The beast is maternal: She rescues and feeds twins of a different species the way Rome, since its beginnings, has hosted visitors from other lands and settlers of diverse backgrounds; the wolf god was the protector of fugitives and exiles, and these outcasts were indeed the earliest inhabitants of Romulus's city. (Of course, one never knows whether Rome and the she-wolf will protect their vulnerable charges or gobble them up.) The she-wolf is a wild predator that attacks those weaker than herself; she prefers lambs above all else. Rome, too, has done its share of preying on the weakness of others; any history book easily confirms the city's proverbial greed: “The she-wolf of Rome devoured many sheep. Carthage, Corinth, Jericho, Jerusalem, Londinum (London), Piraeus, Seleucia, Troy – these are but a few of the cities that the Romans destroyed” (Schneider 133). The she-wolf, perhaps, was not a wild animal at all but rather a very human prostitute, her profession indicated by the Latin word for she-wolf, lupa.

She-wolves, in figurative language, are usually human females. Whether fictional or historical, powerful women have long been described with this animal name to suggest their political ambition, sexual insatiability, maternal obsessions, or illicit desires: Marie Antoinette, Hillary Rodham Clinton, Reynard's Hersent, Masuccio's unnamed lady, De Sade's Madame Saint Ange, and Verga's Pina (see Chapters 5 and 6) are some examples of this metaphorical practice. The she-wolf's voracious hunger, in misogynous discourse, can have a variety of objects – primarily politics, sex, and motherhood. Revenge is one of its motivating forces. One such human she-wolf, maternal and vengeful, appears in Victor Hugo's acclaimed 1831 novel, Notre-Dame de Paris. This wolfish woman is a walled-up recluse who rejoices at the news that a gypsy woman has been executed because gypsies had long ago abducted her only daughter. Her fierce motherhood mixes with a blind hunger for revenge, turning the recluse into the caged beast that best embodies such characteristics: “And she began pacing up and down before the bars of her window, disheveled, eyes blazing, banging her shoulder against the wall, with the wild look of a caged she-wolf which has long been hungry and feels feeding time draw near” (360). I describe Victor Hugo's recluse as a way to introduce Rome's own caged she-wolves: Unlike their Parisian counterparts, Rome's captives were not only allegorical animals but also flesh and blood.