Annals of Internal Medicine History of Medicine Whither ’ Liver? Greek Myth and the Science of Regeneration Carl Power, PhD, and John E.J. Rasko, MBBS, PhD

Stem-cell biologists and those involved in regenerative medicine are surgery, the beginnings of anatomical research, and the ancient art fascinated by the story of Prometheus, the Greek god whose im- of liver augury. In addition, the authors consider how the Greeks mortal liver was feasted on day after day by ’ eagle. This myth understood Prometheus’ immortal liver. Not only do the authors invariably provokes the question: Did the ancient Greeks know examine the general theme of regeneration in , about the liver’s amazing capacity for self-repair? The authors ad- they survey several scholarly interpretations of Prometheus’ torture. dress this question by exploring the origins of Greek myth and medicine, adopting a 2-fold strategy. First, the authors consider what opportunities the ancient Greeks had to learn about the liver’s Ann Intern Med. 2008;149:421-426. www.annals.org structure and function. This involves a discussion of early battlefield For author affiliations, see end of text.

holy grail of modern stem-cell research is the recre- liver regeneration. We then turn to the story of Aation of a functioning organ. The vital importance of Prometheus itself, placing it in the context of other Greek achieving this goal is all too clear. In the United States myths. After examining the general theme of regeneration alone, almost 9% of patients with liver failure die waiting in Greek mythology, we survey several plausible interpre- for a new organ. This is just 1 instance of a much broader tations of Prometheus’ unique torture. problem. With demand for transplantable organs far ex- ceeding supply, the need for regeneration therapies has FROM AUGURY TO ANATOMY never been greater. Like most Greek myths, that of Prometheus has ob- The story that has, above all others, captured the scure beginnings. The earliest literary account of imagination of those involved in regeneration research is Prometheus’ torture comes from Hesiod’s poem, The the Greek myth of Prometheus. Countless lectures and Theogony (eighth century BC), but the oral traditions on articles about regenerative medicine begin with a reference which Hesiod drew have uncertain origins (10). Because to this myth before turning to more scientific matters. Greek is an Indo-European language, one might expect Prometheus was the trickster god who tried to outwit Zeus that the Prometheus myth sprang from this linguistic fam- by stealing fire for mankind and, as punishment, was ily, but in fact its ancestors seem to have come from the chained to a crag in the Caucasus Mountains. Each day, an Near East. There are, for instance, close parallels between eagle would feast on his liver, and each night, his liver the myth of Prometheus and the Mesopotamian legend of would regrow in time for the eagle’s return (Figure 1). Enki (11, 12). The most plausible explanation is that the When hearing this tale, we cannot help but wonder if the mythical traditions of the Near East were transmitted to ancient Greeks had witnessed the amazing capacity of the Greece along trade routes either during the Late Bronze liver to restore itself. Although this possibility fascinates Age (1550 to 1200 BC) or during the period of renewed those engaged in regeneration research, they have not in- contact with the Near East that has become known as the vestigated the matter in any depth. Either they simply as- “orientalizing century” (750 to 650 BC)—the century in sume that the Greeks knew about the liver’s regenerative which Hesiod and lived (13). powers (1–4), or they adopt an agnostic attitude (5–8). The influence of the Near East can be seen in the Chen and Chen (9) are exceptions, arguing that the mythology of the Greeks as well as in their magical and very logic of the Prometheus myth presupposes knowledge medical practices. For instance, hepatoscopy was an East- of liver regeneration. In their view, the Greek mythmakers ern import. Archaeologic evidence suggests that the prac- needed a punishment for Prometheus that was brutal with- tice of liver augury spread west from Mesopotamia to out being fatal—one that the god could endure for a pro- Greece and Etruria (14). The famous bronze liver (circa longed period without sustaining permanent injury. Their solution: repeated attack on an organ known to be capable of self-repair. They offer 2 suggestions about how the Greeks could have acquired this knowledge. One concerns See also: hepatoscopy, the art of reading animal livers to divine the Print will of the gods. Another concerns the knowledge that Key Summary Points ...... 422 Greek healers might have gained by treating liver ailments. Although this case seems plausible prima facie, we think it Web-Only is necessary to take a much closer look at the evidence. We Conversion of graphics into slides examine how the Greeks may have glimpsed the truth of

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or color of the liver would be understood not as an effect of Key Summary Points past natural causes (for example, the sheep having previ- Many cite the myth of Prometheus as proof that the an- ously suffered and survived some injury to its liver), but as cient Greeks were familiar with liver regeneration, knowl- a sign of what the gods plan for the future. For this reason, edge that was lost for thousands of years. we very much doubt that a Greek seer would have been able to recognize the evidence of liver regeneration even if The ancient art of hepatoscopy may have taught the it were right before his eyes. Greeks much about the superficial structure of animal liv- If it is doubtful that hepatoscopy revealed the truth of ers, but it would not have revealed the livers’ function and liver regeneration, what about Greek medicine? The poets regenerative capacity. provide the earliest and best literary source of evidence on this topic. Particularly valuable is Homer’s Trojan War As Homer’s work attests, early Greek medicine yielded epic, the Iliad, in which he describes war wounds in strik- very limited knowledge of human anatomy. Healers were ing detail. Here we find a handful of liver injuries—all unable to undertake major surgery and were prohibited fatal. For instance, Tros is sliced open by Achilles and his from doing autopsies. liver tumbles out in a gush of black blood (16). Although An advanced knowledge of anatomy and physiology was some have imagined that Homer possessed the knowledge needed to reveal the fact of liver regeneration, which is of a skilled anatomist (17), a less extravagant view now why this discovery belongs to 19th-century scientists. prevails among medical historians (18–20). Homer and his contemporaries had a very limited knowledge of human Because the Greeks regarded their gods as immortal, their anatomy. They could identify and name the main bodily mythology abounds in tales of supernatural vitality. This organs, and they knew well enough how to injure them, fact makes it unlikely that the Prometheus myth expresses but little more. Their knowledge came from the battlefield, a special knowledge of liver regeneration. sacrificial altar, and kitchen, not the dissecting table. In fact, apart from those relatively rare occasions when the There are many explanations of why the Greek mythmak- Greeks broke their own code of conduct and desecrated the ers targeted Prometheus’ liver for terrible and repeated bodies of their enemies, they did not cut open cadavers. abuse. The best explanations do not depend on the idea Religious scruples, veneration for the dead, and an abiding that the Greeks actually knew about the liver’s regenera- horror of corpses formed a powerful taboo against human tive capacity. dissection (21). That is why, long after the Homeric era, one of the Greek pioneers of anatomy, Aristotle (384 to 322 BC), had to admit that “the inner parts...areforthe most part unknown—at least those of man are, and hence third century BC) discovered near Piacenza, Italy, bears a we have to...[examine] those of other animals, the nat- remarkable resemblance to more ancient clay livers (circa ural structure of whose parts those of man resemble” (22). 1900 BC) (Figure 2) of Mesopotamian origin. These mod- It was not until the third century BC that Greek physicians els of sheep livers were designed to help seers practice and living in the frontier colony of Alexandria dared to dissect teach their craft. Although similar liver models have not humans. Members of the Alexandrian school, such as actually been found in Greece, it is clear from their litera- Herophilus and Erasistratus, were less constrained by the ture and iconography that the early Greeks regarded hep- Greek taboos against interfering with the dead. Moreover, atoscopy as an important form of divination. their passion for anatomical research was encouraged by an It has been said that the study of anatomy began with Egyptian king who supplied them with bodies, including the careful and repeated inspection of livers for divinatory perhaps the living bodies of convicts. Not surprisingly, the purposes (15). Could hepatoscopy have taught the Greeks study of anatomy took a great leap forward, and we owe about the liver’s marvelous capacity for regeneration? In the first detailed description of the human liver to the absence of real evidence, we must rely on educated Herophilus. Its accuracy is in sharp contrast to those that guesses. Here, we should bear in mind the intrinsic limits came before, including those of the Hippocratics, who, like of the seer’s craft. Hepatoscopy focuses exclusively on the Aristotle, seem to have based their descriptions on animal, surface appearance of the liver. It does not delve into the rather than human, livers (23). This scientific advance oc- organ’s interior structure or cast light on the liver’s func- curred at least half a century after Aristotle pioneered com- tion, much less its regenerative capacity. This is not just parative anatomy, almost 2 centuries after Hippocrates es- because postmortem examinations are generally insufficient tablished rational medicine, 5 centuries after Hesiod for the study of physiology. The whole art of hepatoscopy composed his version of the Prometheus myth, and per- depends on not viewing the liver in naturalistic terms. haps a millennium or more after this myth was born. Divinatory expertise blocks scientific inquiry. In the eyes of One might argue that the discovery of liver regenera- a seer, the liver is a kind of divine text in which the will of tion did not require a systematic knowledge of anatomy the gods is written. An irregularity in the shape, structure, but merely surgical experience. We learn from Homer that

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the healing arts were well regarded by the ancient Greeks: Detail from Prometheus Bound, by Peter Paul “For a healer is worth many other men for the cutting out Figure 1. Rubens, 1612–8. of arrows and the sprinkling on of soothing herbs” (16). Yet Homer’s work suggests that the treatment of wounds was largely confined to the removal of foreign bodies, washing, and the application of drugs. The early Greek healers certainly knew how dangerous internal injuries could be. They knew, for instance, that wounds to the liver bled profusely and usually resulted in death, but they lacked the skills to undertake major surgery. In fact, they often refused to treat serious injuries, fearing that they would be blamed if the patient died while in their care (24). Nevertheless, some surgical procedures for curing liver ailments were developed. Hippocrates recommended the incision and drainage of liver abscesses (25), and Era- sistratus reportedly cut open patients in order to apply drugs directly onto the liver (26). Although these examples are firmly rooted in the era of rational medicine, it is at least conceivable that the Greeks did some minor liver sur- gery many centuries earlier, perhaps even when they were formulating their myths. Assuming this is true, what might such medical procedures have revealed about the liver? Witnessing cases in which patients survived liver disease and injury, early Greek healers would have learned that the liver can continue to function adequately when a portion is damaged. However, having a “functional reserve” is not unique to the liver. The lungs, stomach, and kidney—in Courtesy of the Philadelphia Museum of Art. fact, just about any internal organ—can tolerate some damage. The liver stands out from the rest of the splanchna we consider the painstaking, systematic, and collective la- (the Greek word for “innards”) because of its capacity to bor of knowledge required to reveal the liver’s secret, we regenerate. It is this special capacity, over and above the find it very difficult to believe that the ancient Greeks liver’s functional reserve, that the Prometheus myth is sup- could have stumbled on it through the serendipitous ob- posed to express. Surely, to discover this, one would need servation and treatment of injured and diseased livers. to do more than just treat nonfatal liver conditions. At the very least, one would also need to examine closely the livers of patients once they had recovered. As we have seen, how- THE LIVES AND LIVERS OF THE GODS ever, the early Greek healers had almost no opportunity to To properly understand the Prometheus myth, it must do this because they did not dare undertake major surgery be considered in its mythological and cultural context. Do- and postmortem examinations were forbidden. ing so provides further reasons to doubt that the Greeks Had the Greeks glimpsed the liver’s regenerative ca- knew about liver regeneration. pacity, they would have anticipated a discovery that, ac- In Hesiod’s Theogony, we learn that Zeus’ long-winged cording to medical historians, belongs to the 19th century eagle feasted daily on Prometheus’ “immortal liver,” but we (27). It is worth bearing in mind the great gulf of time— also learn that Mount Olympus trembled under Zeus’ “im- perhaps 3 millennia—separating the Greek mythmakers mortal feet,” and that it was from the “immortal flesh” of from the scientists who established the facts of liver regen- Uranus’ genitals—which Cronos had cut off and tossed eration. In the 1830s, Cruveilhier and Andral (28, 29) into the sea—that the goddess Aphrodite was born (10). both proposed the idea that the liver could restore dam- Greek gods enjoyed immortality, an attribute frequently aged tissue. These were tentative conjectures based on an extended to their body parts. They, and their divine kin- advanced knowledge of anatomy and physiology. Their folk, lived glorious, eternal lives (34). An immortal blood view spread, gaining further confirmation from micro- flowed in their veins; it might trickle from an open wound, scopic studies of the liver. Toward the end of the century, but it would not hemorrhage, coagulate, run dry, or pu- several scientists who were doing liver resections on rabbits trefy. The gods knew no hunger and had no need to eat; and other animals finally furnished positive proof of liver however, for the pleasure of it, they feasted on the food and regeneration. The first to publish significant results in this drink of immortality—“ambrosian” substances that did field was Tillmanns, in 1879, followed soon after by not decay. Although Prometheus may have had Near East- Gluck, Ponfick, von Meister, and others (30–33). When ern origins, his immortality was a Greek invention. Meso-

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emblematic images for the science of regeneration. Babylonian clay model of a sheep’s liver, Figure 2. Prometheus aside, there is the fearsome Hydra, a serpent- 1900–1600 BC. like creature with 9 heads. Every time 1 was cut off, 2 would grow back in its place. This mythical monster has a natural namesake, a freshwater coelenterate whose tentacles grow back when severed. There is also the Greek legend of the Graiae, 3 hags who shared a single eyeball that they passed between them so that, together, they remained ever- vigilant. This story reminded Goss (36) of experiments from the 1960s in which the eye of a newt was removed and later grafted back, resulting in the animal regaining its sight. Goss points out that although some myths seem to represent, in an exaggerated or distorted manner, things that their authors might have actually seen, other myths do not. We can imagine that the Greek mythmakers gained inspiration from such creatures as the freshwater hydra, but it is difficult to think of an observable phenomenon that could have inspired the legend of the Graiae. This very obvious point is worth stressing because if the legend of the Graiae sharing an eye does not suggest to us that the Greeks knew about organ transplantation, and if the story of being regrown from his own heart does not suggest some previous knowledge of cloning, why would Reproduced with permission from the British Museum. ௠The Trustees the myth of Prometheus express familiarity with liver re- of the British Museum. generation? No doubt the last hypothesis looks more plau- sible than the first 2, but surely this is not enough. There is potamian gods were not subject to natural death, but they nothing in a myth that tells us what its anonymous authors could be killed by violence (12). In contrast, Greek gods had actually seen or had merely imagined. In other words, could be wounded, weakened, tortured, imprisoned, and taken by itself, the myth of Prometheus provides no put to sleep, but they remained immortal. Zeus, for in- indication of whether the Greeks knew about the liver’s stance, was once overpowered by the monstrous Typhon regenerative powers. To make a sound judgment on this who, to render his opponent helpless, stripped the sinews issue, supporting evidence is required—evidence about from his legs and arms. Eventually, Zeus managed to re- the Greeks’ intellectual and technical accomplishments. trieve his sinews and, having slipped them back into place We presented such evidence in the first part of our ar- like suspenders, regained his strength. Another legend tells ticle, and none of it indicates that the Greeks knew of how Zeus drank down his wife, the goddess Metis, when about liver regeneration. she was in the form of a drop of water. He did this because An important question still needs consideration: Why he was afraid she would bear a child who might overthrow did the Greeks invent such an unusual punishment for him one day. So the story goes, Metis (whose name means Prometheus? Why did they single out his liver for abuse? “wisdom”) did not perish. Instead, her life force fused with Answers to this question are many and varied. that of Zeus so that he gained her wisdom (which, inci- Perhaps the best answer is also the most obvious. The dentally, enabled him to see through Prometheus’ trick- early Greeks regarded the liver as the seat of life. They ery). Dionysus also exemplifies the irrepressible vitality of granted the liver much the same privilege that later Greeks the Greek gods. According to the Orphic cult, the would grant the heart. Indeed, when early poets spoke of attacked Dionysus, tore him to pieces, and ate his flesh. someone being mortally wounded, they would often say Fortunately, the goddess Athena rescued his heart and that he or she was “struck in the liver” (15). This fact alone from it reformed a new Dionysus (35). Viewed in this may have made the liver a particularly attractive target for context, there seems nothing remarkable in the fact that revenge. Thus, when Zeus punishes Prometheus, he takes Prometheus’ liver could repair itself. On the contrary, it aim at the very core of his enemy’s being. Likewise, in the would have been astonishing if the Greeks decided that it Iliad, Hecabe declares that, if she could, she would avenge lacked this capacity. Whether or not the Greeks recognized her son’s death by devouring the liver of his killer, Achilles that the liver can regenerate itself in humans and other (16). She does not stop to consider whether Achilles, being animals, they were bound to grant this power to the liver of half-god, might have a liver capable of regeneration. She a god. simply wants to strike at his heart of hearts. Because the Greeks attributed marvelous vitality to This line of interpretation has the virtue of suggesting gods and monsters, their mythology provides a host of a very plausible explanation of liver augury. Like the myth

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of Prometheus, the art of hepatoscopy seems to depend on was sacrificed, the bones would go to the gods, and men the idea that the liver is the seat of life. Ancient seers would keep the edible share. This was a fortunate outcome probably believed that when they sacrificed an animal, its for humans, but it angered Zeus. So he deprived men of life force fused with that of the god to whom it was offered something previously available to them, something that the (15). At the moment of sacrifice, the 2 lives—one animal Greeks regarded as a hallmark of culture—fire. Men might and the other divine—were unified. It therefore made have meat, but they would have to eat it raw like wild sense to study the animal’s liver, the seat of its life, for signs animals. To avert this catastrophe, Prometheus committed of the god’s intentions. However, there is still a closer con- his second crime: He stole fire from the heavens to give to nection between hepatoscopy and the Prometheus myth. men, thereby incurring the wrath of Zeus. Because One of Prometheus’ characteristic powers was his ability to see Prometheus had acted as a mediator between the gods and into the future. For the Greeks, the name “Prometheus” men, Zeus had him bound to the summit of a mountain— literally meant “foresight.” By contrast, his dimwitted midway between heaven and earth—and because he had brother, Epimetheus, bore a name indicating that he would procured meat for men, he became meat for Zeus’ eagle. always “know too late.” Little wonder that some should But why did the eagle eat his liver? Vernant’s answer con- trace the art of fortunetelling back to Prometheus. In cerns the special role the liver played in Greek sacrificial Aeschylus’ play, Prometheus Bound, the captive god declares cuisine. Like Prometheus, the liver was a kind of mediator that, not only did he steal fire for humans, he taught them between man and god. It was that part of the sacrificed a host of useful skills, setting out for their benefit the animal in which the share of the gods tended to overlap “divers paths of prophecy,” including the art of liver au- with the share of humans. Having cut the animal open, the gury (37). Is it not fitting that the upstart god of foresight sacrificer would reach first for the liver. Its configuration, should suffer Zeus’ eagle to devour his liver, the mirror of sheen, grain, and color indicated whether the gods found the future? the sacrifice acceptable. Thus, before the liver could be Another explanation of Prometheus’ ordeal depends eaten, it had to be read: Before being a delicious morsel, it on the supposition that, for the Greeks, the liver was not so was a message from the gods. much the seat of life as the seat of the passions. On this It must be stressed that competing interpretations of a view, Prometheus’ theft of fire was a crime of passion and myth do not necessarily exclude one another. Just as a his punishment targeted the bodily source of his impulsive historical event may be the result of many causes, a myth behavior. Just which passion drove Prometheus to defy may be determined by many different factors. It is there- Zeus is a matter of dispute. Some scholars say it was spite, fore possible that several explanations of Prometheus’ tor- some say it was anger, some even say it was oedipal lust ture each contain a grain of truth. This does not mean, (38–40). Doubtless, this line of interpretation has its crit- however, that all such interpretations are equally credible, ics. West (10) argues that the Greeks only began to speak interesting, or insightful. In our opinion, the idea that the about the liver as the seat of the passions in the fifth cen- Greeks had a special knowledge of liver regeneration is less tury BC, long after the Prometheus myth was born (10). In plausible than any other alternative we surveyed here. response, Caldwell (40) insists that the passionate nature of When we began our research into the myth of the liver, if not explicitly stated, was at least suggested by Prometheus, we hoped to demonstrate that the Greeks had some Greek myths. For instance, the Tityos at- glimpsed the liver’s regenerative capacity. Unfortunately, tempted to rape the goddess . During his imprison- all the evidence points to the opposite conclusion—one we ment in , 2 vultures endlessly plucked at his liver—a find both disappointing and compelling. Although we have punishment similar to that of Prometheus’ (41). tried to use the best available evidence, we are mindful that Yet another reading of the Prometheus myth is offered by Vernant (42), who focuses on culinary matters. He our case is open to refutation. Far from fearing this possi- makes a very strong case that, as Hesiod tells it, the bility, we hope others explore this issue in greater historical Prometheus myth is centrally concerned with sacrificial depth and uncover new evidence about the myth of cuisine. Consider more closely the 2 crimes for which Prometheus and the origins of medicine. Prometheus was punished. Zeus gave Prometheus the task From the Centenary Institute, University of Sydney, and the Royal of slaughtering a great ox and dividing it into 2 portions: 1 Prince Alfred Hospital, Sydney, Australia. for the gods and the other for men (women did not yet exist). With characteristic cunning, Prometheus gathered all the bones together and hid them beneath a beautiful, Potential Financial Conflicts of Interest: None disclosed. shiny layer of fat. All the meat and tasty innards went into the other portion, which he wrapped in the ugly, slimy sac Requests for Single Reprints: John E.J. Rasko, MBBS, PhD, Centenary of the stomach. This was Prometheus’ first crime: He tried Institute, Locked Bag 6, Newtown, New South Wales 2042, Australia; to trick Zeus into choosing the bones, and Zeus (though e-mail, [email protected]. not really fooled) went along with the deceit, thereby es- tablishing a precedent. Henceforth, whenever an animal Current author addresses are available at www.annals.org.

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Current Author Addresses: Drs. Power and Rasko: Centenary Institute, Locked Bag 6, Newtown, New South Wales 2042, Australia.

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