attitude towards cannibalism

ATTITUDE TOWARDS CANNIBALISM

Medical Research in Biblical Times from the Viewpoint of Contemporary Perspective

Liubov Ben-Nun

The present Book deals with cannibalism. Throughout history, humans have dined on human flesh. Whether it was part of war to gain the enemy's strength, or as a means to terrify opponents, cannibalism goes back a long way. Cannibalism has existed since the dawn of human history. Are there any description if cannibalism in the Bible? If so, what characters were involved? What were circumstances associated with this behavior? What is definition of cannibalism? What are types? What are its characteristics? History? Evolution? Is it present in modern times? What is non-human cannibalism? Are there any diseases than can be transmitted through cannibalistic behavior? What is the biblical attitude towards cannibalism? What can we learn from the biblical description of this behavior? All biblical texts were examined and verses dealing with cannibalism in the humans were studied closely. The approach to cannibalism was examined from the perspective of contemporary medicine. About the Author Dr. Liubov Ben-Nun ,the Author of dozens Books and Articles that have

been published in scientific journals worldwide. Professor Emeritus at Ben Gurion University of the Negev, Faculty of Health Sciences, Beer-Sheva, Israel. She has established the "LAHAV" International Forum for research into medicine in the Bible from the viewpoint of contemporary medicine.

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ATTITUDE TOWARDS CANNIBALISM Liubov Ben-Nun Professor Emeritus Ben-Gurion University of the Negev Faculty of Health Sciences, Dept. of Family Medicine Beer-Sheva, Israel th

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Scene of Cannibalism. From America Tertia Pars. 1562. Theodor Bry.

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CONTENTS MY VIEW PREFACE FOREWORD INTRODUCTION THE BIBLICAL DESCRIPTION OTHER VERSES DEFINITION OF CANNIBALISM MYTHS, LEGENDS, FOLKLORE, AND PAINTINGS HISTORICAL ASPECTS MODERN CANNIBALISM EVOLUTION OF CANNIBALISM CAUSES OF CANNIBALISM ADVANTAGES AND DISADVANTAGES OF CANNIBALISM NON-HUMAN CANNIBALISM CANNIBALISM IN DIFFERENT SPECIES CELL CANNIBALISM TRANSMISSION OF DISEASES PRION DISEASES KURU CREUTZFELD-JACOB DISEASE OTHER DISEASES SYPHILIS SCHIZOPHRENIA SADISTIC KILLER SELF-CANNIBALISM DOWRY CANNIBALISM TEARING CLOTHES THE BIBLICAL ATTITUDE TOWARDS CANNIBALISM SUMMARY

1 2 3 4 6 7 7 9 12 23 27 29 33 35 40 52 63 70 75 82 85 85 86 86 87 88 88 90 91

ABBREVIATIONS AMPK BSE CJD CNS CSF ER FFI G1UC G3UC IGP MRI PrP PrPres SSE TSE UC vCJD YOY WHO

AMP-activated protein kinase Bovine Spongiform Encephalopathy Creutzfeldt-Jakob disease Central nervous system Cerebrospinal fluid Endoplasmic reticulum Fatal familial insomnia Grade 1 urothelial carcinoma Grade 3 urothelial carcinoma Intraguild predation Magnetic resonance imaging Prion protein Protein-resistan prion protein Subacute spongiform encephalopathies Transmissible spongiform encephalopathy Urothelial carcinoma Variant Creutzfeldt-Jakob disease Young-of-the-year World Health Organization

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MY VIEW MEDICINE IN THE BIBLE AS A RESEARCH CHALLENGE This is a voyage along the well-trodden routes of contemporary medicine to the paths of the Bible, from the time of the first man to the period of the People of Israel. It covers the connection between body and soul, and the unbroken link between our earliest ancestors, accompanied by spiritual yearning and ourselves. Through the verses of the Bible flows a powerful stream of ideas for medical research combined with study of our roots and the Ancient texts. It would not be too adventurous to state that if there is one book in the world that all Jews are proud of, that is the Book of Books, the greatest classic among all literary works, whose original language is not Greek or Latin, but the Hebrew that I and other Israelis speak every day, our mother tongue, the language of Eliezer Ben Yehuda. The Bible exists as evidence in the Book of Books, open to all humankind. For thousands of years it has been placed before us, still as fresh as before, the history of peoples who have disappeared and of the Jewish people, which has survived with its Holy Text that has been translated into hundreds of languages and dialects, and remains our eternal taboo. Many people ask me about the connection between the Bible and medical science. My reply is simple: the roots of science are buried deep in the biblical period and I am just the archeologist and medical researcher. This scientific medical journey to the earliest roots of the nation in the Bible has been and remains moving, exciting and enjoyable. It has created a kind of meeting in my mind between the present and those Ancient times, through examining events frozen in time. Sometimes it is important to stop, to look back a little. In real time, it is hard to study every detail, because time is passing as they appear. However, when we look back we can freeze the picture and examine every detail, see many events that we missed during that fraction of a second when they occurred. The Book of Books, the Bible, is not just the identity card of the Jewish, but an essential source for the whole world.

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PREFACE The purpose of this research is to analyze the medical situations and conditions referred to in the Bible, as we are dealing with a contemporary medical record. These are scientific medical studies incorporating verses from the Bible, without no interpretation or historical descriptions of places. Fundamentally, this Research is constructed purely from an examination of passages from the Bible, exactly as written. The research is part of a long series of published studies on the subject of biblical medicine from a modern medical perspective. This is not a laboratory research. The Research is built entirely on a secular foundation. With due to respects to people faith, this Research takes a modern look at medical practices. Each to his own beliefs.

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FOREWORD Throughout history, humans have dined on human flesh. Whether it was an act of war to gain the enemy's strength, or as a means of terrifying opponents, cannibalism goes back a long way. There have been lone individuals who find eating people satisfying. They do it because they enjoy it, they have psychopathic personalities, and they are extremely lonely. Some cannibals are psychotic. Richard Trenton Chase was a serial killer who consumed his victims' blood because he believed space aliens were turning his blood to powder. After all, he had to replenish it. Obviously, Richard was a paranoid schizophrenic. However, most cannibals are not psychotic. They very well know what they are doing. The pattern is very clear. The killer lures an unsuspecting victim to his home and subdues him. Sometimes it is drugs. Other times, the offender simply shoots or stabs the victim. Then an elaborate fantasy plays out. Cutting up the meat is sexually exciting, according to cannibals. Feeling the flesh as it leaves the body brought many offenders to orgasm. They so enjoy the process of removing the meat from the skeleton. It makes them feel all-powerful and capable of something very few people have ever done. Even most serial killers do not eat their victims. Therefore, the cannibal is in the class by himself. Moreover, he knows it. This produces a euphoric state, which activates the pleasure center in the brain. Each cut brings more good feeling. Thus, it is common to find many smaller cuts on the body. The process is exciting. For someone who is isolated and resentful, it fills a void. Most cannibals are extreme loners. They do not have friends, and they are bitter about it. Killing and eating a victim ensures that the offender is never alone. He „has' the victims with him at all times. They can never leave. This helps the cannibal retain a sense of control over his life. To himself, he has demonstrated mastery over another human being. The victim is now part of him as a trophy. This is intoxicating and drives him to do it again. When looking at the victims, it was not anything the victims said or did wrong. It was just the predator's time to feed. That is how an offender sees it. It is simply a matter of being in the wrong place at the wrong time. There is no one type of cannibal victim. Unfortunately, it is very hard to predict who will become a cannibal. There may be signs in adolescence such as killing small animals and drinking their blood. However, offenders go to great lengths to hide this behavior. Someone with psychopathic tendencies who is drawn to blood and death is always of concern. The fascination with gore becomes all-consuming

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to the point where the individual pushes living people away. He would rather spend time focusing on blood and murder than anything else. Overall, cannibals are very proud of their actions. However, they are adamant about the fact that they are cannibals and nothing else. One offender was insulted that he had been accused of rape. He adamantly declared he had never raped anyone. He said, "I may kill them and eat them, but I never raped anyone! You make sure people know that!" The idea of rape was repugnant to him. Another offender asked his victim how he liked his meals prepared. The cannibal said, "I wouldn't want to insult him by cooking him the wrong way." Then, as if to justify what he had done, he said, "it didn't come easy you know. I had to work for it. It took a lot to kill him" (1). References 1. Schurman-Kauflin D. Why Cannibals Love Eating People. Available 12 December 2013 at www.psychologytoday.com/blog/.../why-cannibalslove-eating-people‎.

INTRODUCTION Very few topics in anthropology are as exotic as cannibalism. We might like to think of ourselves as open-minded and adventurous, but we still recoil at the thought of eating human flesh. Cannibalism, as much as any other practice, still represents a limit to cultural relativism. However, because of colonialism and evangelization, cannibalism is almost entirely outdated. Nowadays, the only way to study it is through either the testimony of witnesses who have eaten co-socials in funerals (endocannibalism), or victims of confrontations with neighboring groups (exocannibalism) (1). Cannibalism is a provocative interpretation put forth repeatedly for practices at various prehistoric sites, yet it has been so poorly supported by objective evidence that later, more critical reviews almost invariably reject the proposal. The basic data essential to a rigorous assessment of a cannibalism hypothesis include precise contextual information, analysis of postcranial and cranial remains of humans and animals, and detailed bone modification studies. Such data are available from the Neolithic levels of the Fontbrégoua Cave (southeastern France) where several clusters of human and animal bones have been excavated. The analysis of these bones strongly

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suggests that humans were butchered, processed, and probably eaten in a manner that closely parallels the treatment of wild and domestic animals at Fontbrbégoua (2). Drawing attention to the associations evoked in the process of food denomination, the traces of cannibalistic instinct in this realm are evaluated. By singling out some of the principal ways used in the semantic reticulum to name food, special attention is devoted to those names that allude to particular categories of enemies (devoured as wholes or as parts of the body). What is referred to as "anti-taboo" shows how the linguistic and more generally the anthropological 'substitution process' is a human reaction to the persistence of the cannibalistic taboo in the face of the belief that cannibalism is an impulse far removed from modern society. Finally, it is noted that cannibalistic metaphors have been analyzed in other fields, whereas in the area of food denomination they have been neglected. The suggestion is that the reason for such neglect may lie in an attempt to conceal our closeness to the cannibalistic instinct in the form of linguistic substitution (3). This essay discusses the image and practice of cannibalism in a wide range of studies. It also presents the anthropological research on kuru which led to the proposal that cannibalism had enabled transmission of the infectious agent, as well as doubts about the hypothesis, and the assertion by some that cannibalism as a socially approved custom did not exist. The figure of the cannibal as an icon of primitivism took form in the encounter between Europe and the Americas. Cannibalism was to become the prime signifier of "barbarism" for a language of essentialized difference that would harden into the negative racism of the nineteenth century. Anthropological and medical research now challenge the derogatory image of the cannibal as we learn more about the many past consumers of human flesh, including ourselves (4). Cannibalism has existed since the dawn of human history. Are there any descriptions of cannibalism in the Bible? If so, which characters were involved? What were the circumstances associated with this behavior? What is the definition of cannibalism? What are its types? What are its characteristics? History? Evolution? Is it present in the modern times? What is non-human cannibalism? Are there any diseases than can be transmitted through cannibalistic behavior? What is the biblical attitude towards cannibalism? What can we learn from the biblical description of this behavior? All biblical texts were examined and verses dealing with cannibalism

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in humans were studied closely. The approach to cannibalism was examined from the perspective of contemporary medicine. References 1. Aubry C. Consuming grief: compassionate cannibalism in an Amazonian society (review). Anthropological Quarterly. 2002;75(2):433-6. 2. Villa P, Bouville C, Courtin J, et al. Cannibalism in the neolithic. Science. 1986;233(4762):431-7. 3. Modena ML Traces of cannibalistic instinct in food denomination. Coll Antropol. 2004;28 Suppl 1:221-7. 4. Lindenbaum S. Cannibalism, kuru and anthropology. Folia Neuropathol. 2009;47(2):138-44.

THE BIBLICAL DESCRIPTION This is a biblical account of cannibalism: "And it came to pass after this, that Ben-Hadad king of Syria gathered all his host, and went up, and besieged Samaria. And there was a great famine in Samaria: and, behold, they besieged it, until an ass's head was sold for eighty pieces of silver, and the fourth part of a cab of dove's dung for five pieces of silver. And the king of Israel was passing by upon the wall, there cried a woman unto him, saying Help, my lord, O king. This [other] woman said to me. Give thy son, that we may eat him today, and we will eat my son tomorrow. So we boiled my son, and did eat him: and I said unto her [the other woman] on the next day, Give the son, that we eat him: and she hath hid her son. When the king heard the words of the woman, he rent his clothes; … he had sackcloth within upon his flesh" (II Kings 6:24-26,28-30). In this episode, Samaria was under siege by its enemies and an extreme famine afflicted the people and they paid very high prices for food such asses and birds in order to survive. The hunger was unbearable, so one miserable mother was ready to sacrifice and eat her own child. She made an agreement with another woman to eat their children and cooked her child. The second mother ate some of the child, but refused to reciprocate the next day by cooking her child. When the king heard this conversation, he was heartbroken, and he tore his clothes, and put on sackcloth as a sign of mourning.

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OTHER VERSES "Therefore fathers shall eat their sons in your midst, and sons shall eat their fathers. And I will execute judgments on you, and any of you who survive I will scatter to all the winds" (Ezekiel 5:10). "You shall eat the flesh of your sons, and you shall eat the flesh of your daughters" (Leviticus 26:29). "The hands of compassionate women have boiled their own children; they became their food during the destruction of the daughter of my people" (Lamentations 4:10). "And I will make them eat the flesh of their sons and their daughters, and everyone shall eat the flesh of his neighbor in the siege and in the distress, with which their enemies and those who seek their life afflict them" (Jeremiah 19:9). "Look, O Lord, and see! With whom have you dealt thus? Should women eat the fruit of their womb, the children of their tender care? Should priest and prophet be killed in the sanctuary of the Lord?" (Lamentations 2:20). These verses indicate that cannibalism was prevalent in biblical times.

DEFINITION OF CANNIBALISM Cannibal is defined as a human being who eats human flesh; an animal that eats its own kind. Cannibalistic is referred to Spanish Cannibals, the name of a West Indian tribe to be cannibals, from Arawakan Caniba, Carib, of Cariban origin (1). Consumption of a person from within the same community is called endocannibalism; ritual cannibalism of the recently deceased can be part of the grieving process (2), or a way of guiding the souls of the dead into the bodies of living descendants (3). Exocannibalism is the consumption of a person from outside the community, usually as a celebration of victory against a rival tribe (3). Group cannibalism refers to shared consumption of victims (4).

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Of the many known kinds of cannibalism, five are attested in archaeological remains. Three have independent and direct textual support - survival cannibalism (the Franklin material); ritualized, sacrifice-related cannibalism (the Aztec material); and psychopathic cannibalism (recognized in forensic archaeology and typically connected with serial killers). Reverential funerary cannibalism (Siberian Uyuk) has indirect textual support, and matches an anthropologically recognized category; while aggressive warfaredriven cannibalism (the Anasazi and Fremont Indians) is both known anthropologically and can be directly inferred from the bones and coprolite evidence (5). Filial cannibalism refers to the consumption of one's own offspring and co-occurs in many animals (6,7). Cellular cannibalism is defined when a large cell encloses a slightly smaller one within its cytoplasm (8). The term self-cannibalism or autophagy was coined to describe the ability of the cells to cannibalize their own damaged organelles or proteins (9). Autophagy is an evolutionarily preserved degradation process of cytoplasmic cellular constituents, which has been known for its role in protecting cells against stresses such as starvation and in eliminating defective subcellular structures. It is essentially a form of self-cannibalism, hence the name that means 'self-eating', in which the cell breaks down its own components (10). Xeno-cannibalism is a phagic process related to self-cannibalism. Xeno-cannibalism is described as the ability of certain cells, e.g. metastatic cells, to cannibalize their siblings as well as cells from the immune system (8). References 1. The Penguin English Dictionary. Robert Allen ed. Penguin Books. 2ND ed. London, England, 2003. 2. Andrew NW. Endocannibalism of the Yanomami. The Summit Times 1998;6(18-19). 3. James WD. Cannibalism. In Tenenbaum BA. Encyclopedia of Latin American History and Culture. Charles Scribner's Sons. New York. 1996, Vol. 1, pp. 535-7. 4. Volker HWR, Antonovics J. Disease transmission by cannibalism: rare event or common occurrence? Proc Biol Sci. 2007;274(1614):1205-10. 5. British Archaeology. The edible death. Available at 30 November 2013 at www.archaeologyuk.org/ ba/ba59/feat1.shtml‎. 6. Klug H, Bonsall MB. When to care for, abandon, or eat your offspring: the evolution of parental care and filial cannibalism. Am Nat. 2007;170(6): 886-901.

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7. Manica A. Filial cannibalism in teleost fish. Biol Rev Camb Philos Soc. 2002;77(2):261-77. 8. Sharma N, Dey P. Cell cannibalism and cancer. Diagn Cytopathol. 2011;39(3):229-33. 9. Matarrese P, Ciarlo L, Tinari A, et al. Xeno-cannibalism as an exacerbation of self-cannibalism: a possible fruitful survival strategy for cancer cells. Curr Pharm Des. 2008;14(3):245-52. 10. Vaccaro MI. Autophagy and pancreas disease. Pancreatology. 2008;8(4-5):425-9.

MYTHS, LEGENDS, FOLKLORE AND PAINTINGS In antiquity, Greek reports of cannibalism, (often called anthropophagy in this context) were related to distant non-Hellenic barbarians, or else relegated in Greek mythology to the 'primitive' chthonic world that preceded the coming of the Olympian gods: the explicit rejection of human sacrifice in the cannibal feast prepared for the Olympians by Tantalus of his son Pelops (1). The windigo is a creature appearing in the legends of the Algonquian people. It is thought variously as a malevolent cannibalistic spirit that could possess humans or a monster that humans could physically transform into. Those who indulged in cannibalism were at particular risk and the legend appears to have reinforced this practice as taboo (2). In mythology, religion, and literature there are many examples of cannibalism that have been passed down over the centuries and which do not strike us as shocking as long as they remain fixed in a symbolic context. Things only become problematic when cannibalistic impulses are taken literally and put into practice. Apart from situations of extreme emergency in which this rare phenomenon might enjoy certain sympathy, it also occurs within the context of serious sexual offences. In Germany, there was the case of a man who used the internet to find a person who wanted to have him eaten. The victim's consent unsettled not only the public at large, but also the judiciary, which at first did not know how the case was legally to be appropriately assessed. In a first trial in January 2004, the man was sentenced to a comparatively short prison term of only a few years, a sentence that was lifted by the Federal Supreme Court. In a fresh trial

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in May 2006, he was sentenced to life imprisonment for murder. In this essay, mythological, religious, and artistic models of cannibalism express something fundamentally anthropological and concrete examples should be assessed against this background (3). William Arens, author of The Man-Eating Myth: Anthropology and Anthropophagy (4), questions the credibility of reports of cannibalism and argues that the description by one group of people of another people as cannibals is a consistent and demonstrable ideological and rhetorical device to establish perceived cultural superiority. Arens bases his thesis on a detailed analysis of numerous "classic" cases of cultural cannibalism cited by explorers, missionaries, and anthropologists. He asserted that many were steeped in racism, unsubstantiated, or based on second-hand or hearsay evidence (5). Critognatus, the leader of the Celts, is mentioned only once in the extant ancient literature, namely in Caesar's description of the siege of Alesia in BG VII 77.2-78.2. Here he is portrayed as a determined patriot who wants to encounter the Roman invader bravely and at the risk of all available means. Nevertheless, crafty Caesar succeeds in stamping him by propagandistic pinches to an evil monster and cannibal. On the one hand, Caesar falls back on current Roman prejudices towards the Gauls. The endocannibalism practiced among Celts to a certain extent as a cult action has played a role. Caesar's propagandistic methods are transparent and at the same time so effective that the label of an ogre sticks to Critognatus until the present day. Caesar's portrayal aims at the justification of his Gallic War which he wages against uncivilized and inhuman opponents who are a menace to Rome and even to the culture itself (6). In 1819, Francisco Goya y Lucientes (1746-1828) bought a house west of Madrid called the “Quinta del Sordo” (“Villa of the Deaf Man”). A previous owner of the house was deaf, and the name remained apt as Goya himself had lost his hearing in his mid-forties. The artist painted directly on to the plaster walls of the Quinta the series of psychologically brooding images popularly known as the Black Paintings (1819-23). They were not intended to be shown to the public, and only later were the pictures lifted from the walls, transferred to canvas, and deposited in the Prado Museum. The haunting Saturn illustrates the myth of the Roman god Saturn who, fearing that his children would overthrow him ate them. Taking the myth as a starting point, the painting may be about God‟s wrath, the conflict between old age and youth, or Saturn as Time devouring all things. Goya, by then in his seventies and having survived two life-

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threatening illnesses, is likely to have been anxious about his own mortality. The artist may have been inspired by Peter Paul Rubens‟s Baroque portrayal of the myth, Saturn Devouring His Son (1636). Rubens' painting, also held at the Museum del Prado, is a brighter, more conventional treatment of the myth: his Saturn exhibits less of the cannibalistic ferocity portrayed in Goya's rendition. Goya‟s version, with its restricted palette and looser style, is much darker in all senses. The god‟s wide-eyed stare suggests madness and paranoia, and disturbingly he seems unselfconscious in carrying out his horrific act. However, some critics have suggested that Rubens' portrayal is the more horrific: the god is portrayed as a calculating remorseless killer, who fearing for his own position of power, murders his innocent child. Goya's vision, on the other hand, shows a man driven mad by the act of killing his own son. In addition, the body of the son in Goya's picture is that of an adult, not the helpless baby depicted by Rubens. There is also evidence to show that in the original image Saturn had a partially erect phallus. In 1823, Goya moved to Bordeaux. After a brief return to Spain, he went back to France, where he died in 1828 (7). Ugolino and his sons in their cell were painted by William Blake circa 1826. Ugolino della Gherardesca was an Italian nobleman who, together with his sons Gaddo and Uguccione and his grandsons Nino and Anselmuccio were detained in the Muda, in March 1289. The keys were thrown into the Arno River and the prisoners left to starve. According to Dante, the prisoners were slowly starved to death and before dying Ugolino's children begged him to eat their bodies (8). References 1. Cannibals and Kings. Available 15 December 2013 at www.geni.com/projects/Cannibals-and-Kings/1667‎. 2. Brightman, Robert A. The Windigo in the material world. Ethnohistory. 1998;35(4):337-79. 3. Pfäfflin F. Good enough to eat. Arch Sex Behav. 2008;37(2):286-93. Erratum in Arch Sex Behav. 2008;37(2):360. 4. New York: Oxford University Press, 1979. ISBN 0-19-502793-0. 5. William A. The Man-Eating Myth: Anthropology and Anthropophagy. Oxford University Press US. 1981, p. 165. ISBN 978-0-19-502793-8. 6. Moog FP. Was cannibalism practiced in ancient Gaul? Critognatus, the leader of the Celts, in the focus of Caesar's propaganda. Wurzbg Medizinhist Mitt. 2011;30:204-27. 7. Paintings: Francisco Goya y Lucientes – Saturn. Available 30 November 2013 at favourite-paintings.blogspot.com/.../francisco-goya-ylucientes-saturn.ht.‎

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8. Amin M. The Crusades Through Arab Eyes. New York: Schocken Books. 1985, pp. 37–40. ISBN 0-8052-0898-4.

HISTORICAL ASPECTS Archeological evidence indicates that cannibalism was present in ancient times (1). In Britain, the only other prehistoric (as opposed to modern forensic) material comes from Gough's (New) Cave in the Cheddar Gorge, dating to between 14,000 and 12,500 years ago. In continental Europe, examples include the extensively butchered remains of Homo antecessor from the Gran Dolina at Atapuerca in Spain, dated to sometime before 780,000 years ago. Claims for cannibalism among Neanderthals (notably at Krapina in Croatia) and modern humans have been advanced. Perhaps the most compelling evidence comes from the transitional Mesolithic-Neolithic site at Fontbr"goua Cave in south-eastern France, excavated by Paola Villa, where cut-marked human and deer bones occur in the same assemblages, apparently butchered in closely analogous ways. In the Americas, work on Anasazi and Fremont Indian sites dating to just 1,000 years ago are considered by scholars such as Tim White, Christy and Jacqueline Turner and Shannon Novak to display a pattern of systematic butchery and cooking of human bodies. A distinctive protein, human myoglobin, has also been identified in a coprolite (or fossilised turd) from one of these sites. Archaeological evidence for documented cannibalism associated with the mass ritual killings of the Aztecs is coming to light. Accounts of the Aztec ceremonies describe how choice elements, such as the heart and good leg meat, were eaten ritually at the élite level, with the remainder being cycled down through descending social levels. Numerous examples of small, spilled and weathered fragments of human bone consistent with these historical accounts have been found widely scattered outside the swept-clean temple precincts. One of those who objected to the presentation of the evidence in Cannibal was the prehistorian and writer Paul Bahn. Although Bahn accepts some of the hominid evidence as 'survival cannibalism' or nutritional cannibalism of a sort, he strongly disputes the arguments for the modern-human material, in both Europe and the Americas. Along with him, the Australian archaeologist Michael Pickering has argued that the Fontbrégoua evidence could as well be the product

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of ritual treatment of the dead, while Bahn argues that animals can be treated ritually no less than humans. The claim of Anasazi cannibalism, and even the coprolite, have been similarly attacked. The alternative explanations are ritual activity on the one hand and, on the other, that the turd was either not of human origin or was excreted by someone suffering intestinal bleeding (i.e., that it was the person's own myoglobin that had been preserved). There have, indeed, been anthropological observations of defleshing as part of non-cannibalistic funerary rites. However, this behavior makes sense, only against knowledge of human edibility put bluntly, to assuage the fear that if you do not deflesh your relatives, then other humans or animals will scavenge them. To backproject, this sophisticated routine of ritual funerary defleshing onto Neanderthal communities is perverse. Anthropologists have long recognized cases where the label 'cannibal' was denigratory and untrue. However, alongside them they accepted that there were classic cultural forms of cannibalism reverential funerary endo-cannibalism (eating one's own); and aggressive, gustatory exo-cannibalism (eating one's enemies, having first branded them 'animals'). Ultimately, human flesh was cooked on board the Resolution and eaten as part of a demonstration on the quarterdeck by a group of Maoris in the shocked presence of the crew. Cook wrote an apologia for the Maoris, stressing their honesty, the difficulty that any nation has in breaking from tradition, and the fact that it was enemies that were eaten: 'I firmly believe they eat the flesh of no others,' he wrote. Even Herodotus's man-eaters find archaeological support, as Jim Mallory and EM Murphy of the University of Belfast have shown. Herodotus's account of funerary cannibalism among the Massagetae of the Iron Age steppes in the 5th century BC is supported by cut-marks in a 3rd century burial of the Uyuk culture in southern Siberia. By a curious chain of association, not previously noted, Herodotus seems to have been ultimately responsible for the name 'cannibal' too. When Columbus was told of the existence of people eaters in the Caribbean with an ethnic name variously given as Caribes and Canibales, he seems to have taken it as confirmation that he had indeed reached the Indies, understanding Khan-ibales as people of the Great Khan, the Central Asiatic descendants of the Androphagi. Therefore, the name stuck. Returning to the late prehistoric practices at Eton and Alveston, Margaret Cox of Bournemouth University examined both bone

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assemblages and there were deliberate cut-marks and some splitting of long bones. Sadly, it is far from clear what the standard funerary rite was for Britain at this time. The 1st century BC/AD geographer Strabo mentions reverential funerary cannibalism among the Irish, but abnormal behavior cannot be rejected out of hand either. If, for example, a band of local brigands controlled the Alveston bone cave, they could have split up their victims' bones to reward their dogs. The evidence is open to interpretation, but such a behavior is perhaps less likely than cannibalism in some form. At Gough's Cave, the human bones interpreted as cannibalized appear among an assemblage of hunted wild fauna, and display a pattern consistent with butchery for meat, including the removal of tongues. The evidence can be interpreted as warfare cannibalism, but survival cannibalism should not be ruled out as a response to hardship years. Unfortunately, as with the Iron Age, the standard funerary pattern for Britain in the Late Paleolithic is unknown, so a comparison between potentially cannibalized and non-cannibalized human remains is not possible. Individual instances of cut-marked human bones will always remain open to case-by-case analysis. They will not all be examples of cannibalism, but we should now move beyond the unconstructive forand-against debate. It is clear that the edibility of human beings has led in a systematic way to their being eaten on every continent in nearly every period of human existence (1). Human remains belonging to at least six individuals were found in an exploratory excavation made at the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These remains were recovered from the Aurora Stratum of Unit TD6. This stratum has a thickness of approximately 30 cm. The area of the exploratory excavation is about 7 m(2). According to palaeomagnetic analyses, Unit TD6 shows reversed polarity, which is considered to belong to the Matuyama chron. This unit is immediately below TD7, where the MatuyamaBrunhes boundary has been detected, indicating an age of around 780,000 years BP. There is no specific distribution, treatment, or arrangement of the human remains, which were found randomly mixed with abundant faunal remains and stone tools. Most of the faunal and human fossil bones from the Aurora Stratum have human induced damage. Stone tool cut marks are frequent, and peeling (a type of fracture similar to bending a fresh twig between the hands) provides a specific breakage pattern together with percussion and chop marks. Both nonhuman and human remains show similar intensive exploitation. Slight differences, however, have been

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observed between fauna and humans (e.g., peeling frequent in humans, rare in fauna) that appear related to different musculature, weight, and bone structure. The characteristics of this fossil assemblage suggest that it is solely the result of consumptive activities as there is no evidence of ritual or other intention. In addition, there is the possibility of distinguishing between dietary vs. survival cannibalism (2).

Cannibalism among the Tupi-Inbas Indias of Brazil. Theodore de Bry copper engravings from J. de Léry, Le Voyage au Brézil de Jean de Léry 1556-1558 (La Rochelle, 1578).

Results of a somewhat preliminary bioanthropological study on the skeletal material from the Pitted ware site of Jettböle, excavated in the parish of Jomala, Aland (Ahvenanmaa) Islands, some 85-90 years ago indicate that there is the possibility of cannibalism (3). The Iberomaurusian necropolis of Taforalt (Morocco, 11-12,000 BP), excavated by Roche in the 1950s, contains 28 multiple graves. In the absence of the excavation records of the necropolis, these funerary practices were investigated through the analysis of the contents of each grave and the distribution of intentionally modified specimens (ochre-dyeing, cut marks). Previous research has drawn particular attention to Grave XII (containing three male adults and two juveniles), where many intentionally modified specimens were identified. The present study focused specifically on the human remains recovered from Grave XII. Analysis of these remains has provided evidence of interventions, such as dismemberment and defleshing of the cadaver, and the use of ochre to color the bones. The presence of lesions on two skulls suggests the possibility of intentional killing and cannibalism among the Taforalt population. This study

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further supports the previous impression of the complex and diversified funerary practices, characterizing the social life of the Iberomaurusian population of Taforalt (4). A charnel pit that contained the disarticulated and intentionally damaged remains of eight incomplete adult and subadult Anasazi skeletons was found and excavated in 1926 by F.H.H. Roberts, Jr., at an AD 900 ruin he named Small House, located in Chaco Canyon, northwestern New Mexico. Damage includes extensive perimortem cranial and postcranial bone breakage, cut marks, anvil-hammerstone abrasions, burning, many missing vertebrae, and fragment end polishing. Together, these six types of perimortem damage are believed to be the taphonomic signature of prehistoric Anasazi cannibalism (5). The existence of cannibalism is one of the most controversial issues in the archaeology of the American Southwest. Disarticulated, cut-marked and heat-altered human remains from non-burial contexts at prehistoric Puebloan (Anasazi) archaeological sites in the Four Corners region of the American Southwest have been interpreted by some scholars as evidence of cannibalism. Osteological studies indicate that many of the disarticulated bodies found at these sites were processed in a manner consistent with food preparation. Opponents of this interpretation point out that non-cannibalistic practices such as secondary interment, corpse mutilation and ritualized witch executions might account for the assemblages. Osteological evidence alone does not document the actual ingestion of human flesh. Consumption of human flesh did occur as demonstrated in preserved human waste containing identifiable human tissue remains from a site with osteological evidence of cannibalism (6). St. Jerome, in his letter against Jovinianus, discusses how people come to their present condition because of their heritage, and then lists several examples of peoples and their customs. In the list, he mentions that Atticoti eat human flesh and that Massagetae and Derbices (a people on the borders of India) kill and eat old people (7). In 2001, archaeologists at the University of Bristol found evidence of Iron Age cannibalism in Gloucestershire (8). Cannibalism was practiced as recently as 2000 years ago in Great Britain (9). In Germany, Emil Carthaus and Dr. Bruno Bernhard have observed 1,891 signs of cannibalism in the caves at the Hönne (1,000-700 BC) (10). During excavations of the Bronze Age levels at El Mirador Cave, a hole containing human remains was found. Taphonomic analysis revealed the existence of cut marks, human tooth marks, cooking

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damage, and deliberate breakage in most of the remains recovered, suggesting a clear case of gastronomic cannibalism. The piled distribution of the remains, the uneven skeletal representation, and the chronological difference between the pit and the remains suggest that these bones were subsequently buried by a human group that inhabited the cave later. Evidence of gastronomic cannibalism has already been documented in Gran Dolina, another site in the Sierra de Atapuerca, on remains of Homo antecessor (2,11,12). Human remains belonging to at least 6 individuals were found in an exploratory excavation at the site of Gran Dolina (Sierra de Atapuerca, Burgos, Spain). These remains were recovered from the Aurora Stratum of Unit TD6. This stratum has a thickness of approximately 30 cm. The area of the exploratory excavation is about 7 m(2). According to palaeomagnetic analyses, Unit TD6 shows reversed polarity, which is considered to belong to the Matuyama chron. This unit is immediately below TD7, where the MatuyamaBrunhes boundary has been detected, indicating an age of around 780,000 years BP. There is no specific distribution, treatment, or arrangement of the human remains, which were found randomly mixed with abundant faunal remains and stone tools. Most of the faunal and human fossil bones from the Aurora Stratum have human induced damage. Stone tool cutmarks are frequent, and peeling (a type of fracture similar to bending a fresh twig between the hands) provides a specific breakage pattern together with percussion marks and chopmarks. Both nonhuman and human remains show similar intensive exploitation. Slight differences, however, have been observed between fauna and humans (e.g., peeling frequent in humans, rare in fauna) that appear related to different musculature, weight, and bone structure. The characteristics of this fossil assemblage suggest that it is solely the result of consumptive activities as there is no evidence of ritual or other intention. There is the possibility of dietary vs. survival cannibalism (2). Eshkaft-e Gavi is a cave located in the southern Zagros Mountains of Iran and is one of the few archaeological sites in the region to preserve both Middle Paleolithic and Upper Paleolithic occupations. Excavation of the site in the 1970s yielded an assemblage of lithic and faunal remains, including 10 hominin specimens: a mandibular molar, 4 cranial fragments, a clavicular diaphysis, the proximal half of a metacarpal, a fragment of os coxa, the proximal diaphysis of a juvenile femur, and a patella. The bones were derived from a minimum of 4 individuals, including 2 juveniles. Although many of these remains could be Epi-Paleolithic in age, in 1 of the juvenile

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specimens the mandibular molar occurred at the base of the cave's Upper Paleolithic sequence. The remains are very fragmentary, but those that preserve diagnostic morphology indicate that they represent modern humans. The molar is taxonomically diagnostic, thus confirming the association of the Aurignacian-like Baradostian Industry with modern humans. Four of the specimens-a piece of frontal bone, the clavicle, the juvenile femur, and the patella-display evidence for intentional butchery in the form of stone-tool cut marks. These cut marked specimens, along with a fragment of parietal bone, were burned. Although this evidence is consistent with cannibalism, the small sample makes it difficult to say whether the individuals represented by the hominin remains were butchered and cooked for consumption. Nevertheless, the cutmarked Eshkaft-e Gavi specimens add to a growing sample of hominin remains extending back into the Plio-Pleistocene that display evidence of intentional defleshing (13). Reconstruction of early Pleistocene hominin carcass acquisition and processing behaviors are necessarily based at least in part on butchered fossil bones. This paper provides zooarchaeological and taphonomic analyses and behavioral interpretations of 3 approximately 1.5 million-year-old archaeofaunas from areas 1A and 103 in the Okote Member of the Koobi Fora Formation, northern Kenya: FwJj14A, FwJj14B, and GaJi14. These sites are all located in similar paleoenvironmental contexts, near shallow water with swampy, seasonally flooded areas, and some evidence for more wooded or gallery forest settings. Both individual specimen and assemblage-level analyses of butchery-marked bones indicate that the hominins appear to have practiced similar butchery strategies at all of these sites, with butchery (defleshing, disarticulation, and marrow extraction) of both high- and low-ranked skeletal elements with no apparent preference for prey size, skeletal region, limb class, or limb portion. Only 4 tooth-marked specimens, including 1 likely crocodiletooth-marked bone, are preserved in all 3 archaeofaunas. A paucity of limb epiphyses suggests that bone-crunching hyenids may have deleted these portions subsequent to hominin butchery. Strangely, there were no stone tools preserved with the 292 cut-marked and 27 percussion-marked faunal specimens (out of 6,039 specimens), suggesting that raw material availability may have conditioned hominin lithic discard patterns at these locales. These assemblages increase our knowledge of the dietary behavior and ecology of Homo erectus, and provide support for variability in early Pleistocene hominin carcass foraging patterns (14).

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Cannibalism in chimpanzees, modern humans, and in archaeological cases was compared with cannibalism inferred from evidence from the Early Pleistocene assemblage of level TD6 of Gran Dolina (Sierra de Atapuerca, Spain). The cannibalism documented in level TD6 mainly involves the consumption of infants and other immature individuals. The human induced modifications on Homo antecessor and deer remains suggest that butchering processes were similar for both taxa, and the remains were discarded on the living floor in the same way. This finding implies that a group of hominins that used the Gran Dolina cave periodically hunted and consumed individuals from another group. However, the age distribution of the cannibalized hominins in the TD6 assemblage is not consistent with that from other cases of exo-cannibalism by human/hominin groups. Instead, it is similar to the age profiles seen in cannibalism associated with intergroup aggression in chimpanzees. For this reason, an analogy with chimpanzees was used to propose that the TD6 hominins mounted low-risk attacks on members of other groups to defend access to resources within their own territories and to try to expand their territories at the expense of neighboring groups (15). The hypothesis that the human remains from the Navatu midden (50 BC to AD 1900) in Fiji represent cannibalized individuals was tested by an analysis of the skeletal remains. The site includes formal human burials and a separate, contemporaneous midden containing commingled fragmentary human and nonhuman bones. All remains were examined for a variety of modifications. The medium mammal and human remains in the midden have similar modifications: ancient breaks (92% of midden human specimens and 88% of medium mammal specimens), burning (29% and 11%), crushing (1% and 1%), cut marks (9% and 8%), peeling (4% and 1%), and percussion pits (1% and 1%). The human burials (for which cannibalism had not been hypothesized) are essentially unmodified except for some breakage. The pattern of element representation and low incidence of animal bite marks rules out carnivores and rodents as major modifiers of the assemblage. The breakage patterns, element representation, light weathering, and rarity of random striae indicate that sediment pressure, wave action, weathering, and trampling did insignificantly alter the remains. The modifications of the midden human and nonhuman remains generally correspond in type and frequency. The evaluation of the assemblage's taphonomic history suggests that most of the modifications were caused by humans. The Navatu midden human sample does not resemble assemblages interpreted as secondary burials with defleshing, nor does it resemble violence-

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derived assemblages. The burials at Navatu and other Fijian sites indicate that the various non-cannibalistic Fijian mortuary rituals do not mimic butchery and consumption. Therefore, the hypothesis of cannibalism at Navatu is supported (16). The Carib tribe in the Lesser Antilles, from whom the word cannibalism derives, acquired a long-standing reputation as cannibals following the recording of their legends in the 17th century (17). Some controversy exists over the accuracy of these legends and the prevalence of actual cannibalism in the culture. Cannibalism was widespread in the past among humans in many parts of the world, continuing into the 19th century in some isolated South Pacific cultures, and to the present day in parts of tropical Africa. In a few cases in insular Melanesia, indigenous flesh-markets existed (18). Cannibalism has been documented around the world, from Fiji to the Amazon Basin to the Congo to Māori in New Zealand (19). Neanderthals are believed to have practiced cannibalism (20,21) and may have been eaten by anatomically modern humans (22). The cave site of Moula-Guercy, 80 meters above the modern Rhone River, was occupied by Neanderthals approximately 100,000 years ago. Excavations since 1991 have yielded rich paleontological, paleobotanical, and archaeological assemblages, including parts of 6 Neanderthals. These parts are contemporary sites with stone tools and faunal remains in the same tightly controlled stratigraphic and spatial contexts. The inference of Neanderthal cannibalism at Moula-Guercy is based on comparative analysis of hominid and ungulate bone spatial distributions, modifications by stone tools, and skeletal part representations (23). Portuguese accounts of cannibalism in sixteenth-century southeast Africa reflect important but mostly unrecognized elements of the region's political and cultural history. Descriptions of the Zimba cannibals in Ethiopia Oriental, written by the Portuguese priest Joo dos Santos, were analyzed. Dos Santos's evidence figures significantly in scholarship for this period, and while many historians include his colorful descriptions of cannibalism, none has taken them seriously, largely dismissing them as a product of European myth making. In focusing on the question of cannibalism, the article asks not whether the Zimba ate human flesh, nor why they might have, but how dos Santos came to believe that they did. Early modern European cultural outlooks had a role in producing such accounts, but the argument here focuses on how claims of cannibalism reflected African, rather than European, perspectives. Such claims were a vernacular expression of beliefs about, and critiques of political power in the threatening and

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unsettled political environment of the time. In transmitting descriptions of cannibalism from African informants, dos Santos acted as an unwitting vehicle for this vernacular critique, conveying its meaning quite imperfectly to his readers (24). It was considered a great triumph among the Marquesans to eat the body of a dead man. They treated their captives with great cruelty. They broke their legs to prevent them from attempting to escape before being eaten, but kept them alive so that they could brood over their impending fate. With this tribe, as with many others, the bodies of women were in great demand (25). In China during the Tang Dynasty, cannibalism was supposedly resorted to by rebel forces early in the period (who were said to raid neighboring areas for victims to eat), as well as both soldiers and civilians besieged during the rebellion of An Lushan. Eating an enemy's heart and liver was claimed to be a feature of both official punishments and private vengeance. References to cannibalizing the enemy has also been seen in poetry written in the Song Dynasty, (for example, in Man Jiang Hong) although the cannibalizing is perhaps poetic symbolism, expressing hatred towards the enemy (26). Assessment: since the dawn of human history, in different cultures cannibalism has been a prevalent human behavior. References 1. British Archaelogy. The edible death. Available at 30 November 2013 at www.archaeologyuk.org/ ba/ba59/feat1.shtml‎. 2. Fernández-Jalvo Y, Carlos Díez J, Cáceres I, Rosell J. Human cannibalism in the Early Pleistocene of Europe (Gran Dolina, Sierra de Atapuerca, Burgos, Spain). J Hum Evol. 1999;37(3-4):591-622. 3. Nunez M, Lidén K. Taking the 5000 year old "Jettböle skeletons" out of the closet: a palaeo-medical examination of human remains from the Aland (Ahvenanmaa) Islands. Int J Circumpolar Health. 1997;56(1-2):30-9. 4. Belcastro MG, Condemi S, Mariotti V. Funerary practices of the Iberomaurusian population of Taforalt (Tafoughalt, Morocco, 11-12,000 BP): the case of Grave XII. J Hum Evol. 2010;58(6):522-32. 5. Turner CG 2nd. Cannibalism in Chaco Canyon: the charnel pit excavated in 1926 at Small House ruin by Frank HH Roberts. Jr. Am J Phys Anthropol. 1993;91(4):421-39. 6. Marlar RA, Leonard BL, Billman BR, et al. Biochemical evidence of cannibalism at a prehistoric Puebloan site in southwestern Colorado. Nature. 2000;407(6800):74-8.

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7. Against Jovinianus-Book II. A Select Library of Nicene and PostNicene Fathers of the Christian Church. 2nd 6. New York: The Christian Literature Company. 1893, c.393. p. 394. 8. Cannibalism - The Way The Truth And The Life. Available 23 December 2013 at thewaythetruthandthelife.net/index/7.../7.../7.../6-2-326_cannibalism. 9. Druids Committed Human Sacrifice, Cannibalism?. National Geographic. Available 14 December 2013 at news.nationalgeographic. com/.../090320-druids-sacrifice-cannibalism.‎ 10. Steiger B. Real Zombies, the Living Dead, and Creatures of the Apocalypse. Visible Ink Press. 2010, pp. 201. ISBN 978-1-57859-296-8. 11. Fernández-Jalvo Y, Diez CJ, de Castro BJM, et al. Evidence of early cannibalism. Science. 1996;271:277-8. 12. Cáceres I, Lozano M, Saladié P. Evidence for bronze age cannibalism in El Mirador Cave (Sierra de Atapuerca, Burgos, Spain). Am J Phys Anthropol. 2007;133(3):899-917. 13. Scott JE, Marean CW. Paleolithic hominin remains from Eshkaft-e Gavi (southern Zagros Mountains, Iran): description, affinities, and evidence for butchery. J Hum Evol. 2009;57(3):248-59. 14. Pobiner BL, Rogers MJ, Monahan CM, Harris JW. New evidence for hominin carcass processing strategies at 1.5 Ma, Koobi Fora, Kenya. J Hum Evol. 2008;55(1):103-30. 15. Saladié P, Huguet R, Rodríguez-Hidalgo A, et al. Intergroup cannibalism in the European Early Pleistocene: the range expansion and imbalance of power hypotheses. J Hum Evol. 2012;63(5):682-95. 16. Degusta D. Fijian cannibalism: osteological evidence from Navatu. Am J Phys Anthropol. 1999;110(2):215-41. 17. Salisbury DF. Brief history of cannibal controversies. Exploration, Vanderbuilt University. 2001. 18. Knauft BM. From primitive to post-colonial in Melanesia and anthropology. University of Michigan Press. 1999, p. 104. ISBN 0-47206687-0. 19. Rubinstein WD. Genocide: a history. Pearson Education. 2004, pp. 17-8. ISBN 0-582-50601-8. 20. Culotta, E. Neanderthals Were Cannibals, Bones Show. Science. 1999;286(5437):18b. 21. Gibbons A. Archaeologists Rediscover Cannibals. Science. 1997; 277(5326):635-7. 22. McKie R. How Neanderthals met a grisly fate: devoured by humans. The Observer (London). 2009. 23. Defleur A, White T, Valensi P, et al. Neanderthal cannibalism at Moula-Guercy, Ardèche, France. Science. 1999;286(5437):128-31. 24. Allina E. The Zimba, the Portuguese, and other cannibals in late sixteenth-century Southeast Africa. J South Afr Stud. 2011;37(2):211-27. 25. Rubinstein WD. Genocide: a history. Pearson Education. 2004, pp. 17-8. ISBN 0-582-50601-8.

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26. Charles B. China's Golden Age: Everyday Life in the Tang Dynasty. Oxford University Press. 2002, pp. 123-4. ISBN 0-19-517665-0.

MODERN CANNIBALISM Cannibalism is still practiced as a ritual and in war in various Melanesian tribes. Historically, allegations of cannibalism were used by the colonial powers as a tool of empire to justify the subjugation of what were seen as primitive peoples (1). Cannibalism tests the bounds of cultural relativism as it challenges anthropologists "to define what is or is not beyond the pale of acceptable human behavior" (2). Cannibalism is rare and is legal in most countries. People who eat human flesh are usually charged with crimes other than cannibalism, such as murder or desecration of a body (3). During Europe's Great Famine of 1315-1317, there were many reports of cannibalism among the starving populations. In North Africa, as in Europe, there are references to cannibalism as a last resort in times of famine (4). The Korowai is one of the few cannibal tribes that are still around today. As of 2012, the tribe lives in southeastern Papua New Guinea. There are also reports of cannibals in the Congo and Liberia (5). Reports of cannibalism were recorded during the First Crusade, as Crusaders were alleged to have fed on the bodies of their dead opponents following the Siege of Ma'arrat al-Numan. Amin Maalouf also alleges further cannibalism incidents on the march to Jerusalem, and to the efforts made to delete mention of these from the western history (6). Today's practice of ritual killings in Liberia still exists mainly because of a combination of traditional beliefs which inspire meneating and modern-day criminal behavior of unscrupulous politicians who consider their ambitions worth more than the life of their victims. During the 14-year civil war (1989-2003), there were many cases of gunmen, some of them child soldiers, eating their victims' hearts and other body parts that the Catholic Church issued a formal denunciation of these practices (7). In the 1980s, Médecins Sans Frontières, the international medical charity, supplied photographic and other documentary evidence of ritualized cannibal feasts among the participants in Liberia's internecine strife to representatives of Amnesty International who

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were on a fact-finding mission to the neighboring state of Guinea. However, Amnesty International declined to publicize this material; the Secretary-General of the organization, Pierre Sane, said at the time in an internal communication that "what they do with the bodies after human rights violations are committed is not part of our mandate or concern". The existence of cannibalism on a wide scale in Liberia was subsequently verified (8). Francis, Baron Dhanis (1861–1909), a Belgian civil servant was given command of an expedition to the Upper Nile. His troops, largely composed of the Batetela tribes who had only been recently enlisted, and who had been irritated by the execution of some of their chiefs for indulging their cannibal proclivities, mutinied and murdered many of their white officers in what has become known as the Batetela Rebellion (9). Nearly all the tribes in the Congo Basin either are or have been cannibals; and among some of them, the practice is on the increase. Races who until lately do not seem to have been cannibals, though situated in a country surrounded by cannibal races, have from increased intercourse with their neighbors learned to eat human flesh. There is not the doubt that they prefer human flesh to any other. Cannibalism has existed in the Democratic Republic of the Congo for generations and still takes place in the bush. The latest practice of cannibalism morphed into a demoralization technique employed by rebel armies during the first and second Congo Wars in order to keep the villagers subservient and was directly related to the number of skirmishes between rebel and governmental forces in the Eastern part of the country. No one who has spent time in East Africa would deny the existence of cannibalism in the Congo but most individuals consider it a delicate topic, like incest and politely refrain from discussing it unless specifically asked to do so. Yet the latest concern with the practice of cannibalism has extended beyond the question of morality and now centers around its connection to a fatal disease called Kuru; the implications of which have just begun to be understood by the WHO. It seems that this disease “bites back” by killing the living who prey upon the dead (10). Numerous reports from North Korea indicate that starving residents of the isolated regime, where famine is an all-too-regular occurrence, now are resorting to cannibalism (11). India is a land of various sect and religions. Out of its vastness belief, the sect known as Aghori or Aghori sadhus (sadhu means sage) is unique. Aghori Sadhu is associated with cannibalism and rituals using human skulls and animal sacrifices. Aghoriss are distinguished

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from other Hindu sects by their alcoholic and cannibalistic rituals. Aghoris in India are beyond human imagination with their rituals and mysterious practices. Aghoris eat human dead bodies on the banks of river Ganges. They drink in a bowl made of human skull. They apply ash from the human pyre. They are the followers of Hindu God – Lord Shiva, they are the “Aghoris”. A place considered dreadful by others is home for Aghoris, the Hindu cremation ground. Out of the various sect of Hindu priest, the most extreme and the most feared of all are the Aghoris. The term “Aghori” is derived from the Sanskrit word “Aghor” which has various meanings. Aghor means not to forget or non-terrible in one perspectives. On the other hand, it means absence of darkness. Agor implies a simple and a natural state of consciousness in which there is no fear or disgust. On the contrary, the Aghoris have rituals that have seen as being disgustful and feared by common people. The practice of cannibalism, animal and human sacrifices are mostly related to tantric rites of the worshipers of the Mother goddess in any of the worship form – Kali, Dhurga, Kamakhya or Chammunda. The life of an Aghori Sadhu is not easy, to become like them one has to meditate about twelve years and complete certain rituals under the guidance of an Aghori guru in order to enhance one spiritual strength. Aghoris use woods from pyre, clothes from dead bodies, and ash of burnt bodies for the various rituals. There are certain rules one must follow to become an aghori. Firstly, an Aghori must find a teacher and do what the teacher tells him to do. While Aghori must find a human skull known as “kappala” using only this as a ritual tool before initiation. An Aghori must apply pyre on his body to symbolically show the nature of the Lord Shiva. The final part of ritual requires eating of rotten human flesh and meditating sitting on a dead corpse. It symbolizes the rise from Savak (human nature) to Siva (12). In Ukraine, survival was a moral as well as a physical struggle. A woman doctor wrote to a friend in June 1933 that she had not yet become a cannibal, but was "not sure that I shall not be one by the time my letter reaches you." The good people died first. Those who refused to steal or to prostitute themselves died. Those who gave food to others died. Those who refused to eat corpses died. Those who refused to kill their fellow man died. At least 2,505 people were sentenced for cannibalism in the years 1932 and 1933 in Ukraine, though the actual number of cases was certainly much greater (13). Human cannibal cases are relatively rare in modern civilized societies. A single case of familial cannibalism occurred in Oklahoma

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in 1977. The investigation posed unique problems in pathology, psychiatry, criminology, and serology (14). Assessment: in contemporary days, there are places in the world where cannibalism is practiced. References 1. Francis B, Peter H, Margaret I (ed.) Cannibalism and the Colonial World. Cambridge University Press. 1998. ISBN 9780521629089. 2. Salisbury DF. Brief history of cannibal controversies. Exploration. Vanderbuilt University. 2001. 3. Cusack C. Placentaphagy and Embryophagy. J Law Social Deviance. 2011, Vol. 1. 4. Cannibalism in Early Modern North Africa. British J Middle Eastern Studies. Available 30 November 2013 at lib.convdocs.org/docs/index119348.html. 5. Where Do Cannibals Live Today? Available 10 December 2013 at www.ask.com › Q&A › Society › Other‎. 6. Amin M. The Crusades Through Arab Eyes. New York: Schocken Books. 1984, pp. 37–40. ISBN 0-8052-0898-4. 7. Francis Dhanis. Available 15 December 2013 at en.wikipedia.org/wiki/ Francis_Dhanis‎. 8. Cannibalism during the civil war – Liberia. Available 20 December 2013 at www.liberiapastandpresent.org/RitualKillingsCivilWar.htm‎. 9. Gillian G. From cannibalism to genocide: the work of denial. J Interdiscip Hist. 2006;(3):395-414. 10. Cannibalism in the Congo: A New Take on an Ancient Practice . Available 20 December 2013 at katsafrica.wordpress.com/.../cannibalism-inthe-congo- a-new-take-on-an.‎ 11. Cannibalism Reported in Famine-Stricken North Korea. Available 5 December 2013 at www.livescience.com/26677-north-korea-cannibalism. html‎. 12. The Dark Side of Hinduism: Aghori: Human Flesh Eaters ~ An Indian. Available 13 December 2013 at anindianchristian.blogspot.com/ .../dark-side-of-hinduism-aghori-human. 13. Snyder T. Bloodlands: Europe Between Hitler and Stalin. Basic Books. 2010, pp.50–51. ISBN 0-465-00239-0. 14. McClain JL, Jordan FB, Blakeney R. Human cannibalism: a case report. Am J Forensic Med Pathol. 1986;7(2):172-3.

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EVOLUTION OF CANNIBALISM The propensity for cannibalism varies considerably both within and between species. Currently we have little understanding of both the causes of this variation and its evolutionary consequences for other life-history traits. How different levels of spatial structure affect the evolution of cannibalism and how cannibalism in turn drives the evolution of dispersal were examined. Cannibalism can easily evolve in spatially structured populations as long as some dispersal exists. For a wide range of intermediate levels of spatial structure, the possibility of evolutionary branching leading to polymorphism in cannibalism was found. Cannibalism itself can have important evolutionary consequences and select for increased dispersal rates, thus helping to determine the spatial structure of populations. The coevolution of cannibalism and dispersal results in the evolution of various alternative life-history strategies with different dispersal and cannibalism regimes. Which strategy evolves depends on the environmental conditions that determine initial cannibalism rates. These results therefore suggest that differences in spatial structure could explain variation in the propensity for cannibalism and cannibalistic polyphenism. Cannibalism can drive the evolution of other life-history traits and determine the spatial structure of natural populations (1). The evolution of cannibalistic traits in consumer populations is studied with the approach of adaptive dynamics theory. The model is kept at its minimum complexity by eliminating some environmental characteristics, like heterogeneity and seasonality, and by hiding the size-structure of the population. Evolutionary dynamics are identified through numerical bifurcation analysis, applied both to the ecological (resident-mutant) model and to the canonical equation of adaptive dynamics. The result is a rich catalog of evolutionary scenarios involving evolutionary stable strategies and branching points both in the monomorphic and dimorphic dynamics. The possibility of evolutionary extinction of highly cannibalistic populations is also ascertained. This allows one to explain why cannibalism can be a transient stage of evolution (2). A rational explanation for cannibalism is that it would be favored under conditions of crowding of conspecific individuals and/or low availability of alternative prey with the fear of starvation, to maximize individual fitness. Cannibalism has, however, not evolved and is not maintained by a simple individual optimization, while it has evolved

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and is maintained as a game among population members. The attainable state of an evolutionary cannibalism game within a framework reflects the minimum essence of cause-effect in the cannibalism phenomenon. Cannibalism is predator-prey interaction among conspecifics. Immediate direct payoffs (survival in the interaction among conspecifics) and indirect payoffs (growth results in potential productivity and survival against the threat of starvation) would be included. No morphological specialization and no size priority of cannibalism individuals are assumed as conservative situations in which the possibility of cannibalism were analyzed. Cannibalism would be possible under the conservative condition, if initially the wild population's cannibalism rate were insufficiently lower than a threshold value. Crowding and/or low availability of alternative prey with the fear of starvation facilitates cannibalism evolution. Energy gain from nonspecific prey would be attenuated by costs of counterattacks by nonspecific victims and by challenge cost of its own. Discounting net intake energy required in the arms race for cannibalism challenge results in a relative disadvantage of having a high cannibalism rate and makes an evolutionary equilibrium of low cannibalism rate, even when potential profitability of nonspecific prey is high (3). A new study of genetic variation in the human prion protein gene suggests that balancing selection has operated on an amino acid sequence polymorphism in the gene during the last five hundred thousand years. Is this a legacy of widespread cannibalism by our ancestors? (4). References 1. Rudolf VH, Kamo M, Boots M. Cannibals in space: the coevolution of cannibalism and dispersal in spatially structured populations. Am Nat. 2010; 175(5):513-24. 2. Dercole F, Rinaldi S. Evolution of cannibalistic traits: scenarios derived from adaptive dynamics. Theor Popul Biol. 2002;62(4):365-74. 3. Nishimura K, Isoda Y. Evolution of cannibalism: referring to costs of cannibalism. J Theor Biol. 2004 Feb 7;226(3):293-302. Erratum in J Theor Biol. 2004;228(2):291. 4. Brookfield JF. Human evolution: a legacy of cannibalism in our genes? Curr Biol. 2003;13(15):R592-3.

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CAUSES OF CANNIBALISM The human eating of human beings is not confined to exotic peoples. It is part of the human condition. It occurs as a preferred form of protein consumption or as the result of extreme necessity, to absorb the virtues of others, to facilitate conception, for magicoreligious reasons, in warfare, famine, revenge, filial piety, and justice. Unique is the case of cannibalism for cargo reported here in which the eater suffered an overt psychosis before, during, and after the consumption, and whose goal was the acquisition of real and symbolic cargo as exemplified in the Melanesian cargo cult (1). Parental care is expected to increase the likelihood of offspring survival at the cost of investment in future reproductive success. However, alternative parental behaviors, such as filial cannibalism, can decrease current reproductive success and consequently individual fitness. The role of among-offspring relatedness on the evolution of parental care and filial cannibalism was evaluated. The evolution of care is influenced by the effect of among-offspring relatedness on both the strength of competition and filial cannibalism. When there is a positive relationship between among-offspring competition and relatedness, parental care will be favored when among-offspring relatedness is relatively low, and the maintenance of both care and nocare strategies is expected. If the relationship between amongoffspring competition and relatedness is negative, parental care is most strongly favored when broods contain highly related offspring. Further, the range of conditions over which the level of this amongoffspring relatedness can affect the co-occurrence of different care/no care and cannibalism/no cannibalism strategies. Coexistence of multiple strategies is independent of the effects of among-offspring relatedness on cannibalism but more likely when among-offspring relatedness and competition are positively associated (2). A cannibal feast on Tanna, Vanuatu, c. 1885-9 indicates that in some societies, especially tribal societies, cannibalism is a cultural norm (3). Endocannibalism and exocannibalis can also be fueled by the belief that eating a person's flesh or internal organs will endow the cannibal with some of the characteristics of the deceased (4). As in some other Papuan societies, the Urapmin people are engaged in cannibalism in war. Notably, the Urapmin had a system of food taboo wherein dogs could not be eaten and had to be kept from breathing on food, unlike humans who could be eaten and with whom food could be shared (5).

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Cannibalism was practice in some Pacific Islands in the Second World War by many Japanese soldiers, and was often a systematic activity conducted by whole squads under the command of officers. Written reports and testimonials collected by the Australian War Crimes Section of the Tokyo tribunal and investigated by prosecutor William Webb (the future Judge-in-Chief) indicate that Japanese personnel in many parts of Asia and the Pacific committed acts of cannibalism against Allied prisoners of war (6). Japanese troops practiced cannibalism on enemy soldiers and civilians in the last war, sometimes cutting flesh from living captives, according to documents discovered by a Japanese academic in Australia. In most cases, the motive was apparently not shortage of food, but 'to consolidate the group feeling of the troops'. The revelation adds more evidence to the toll of atrocities carried out by Japanese soldiers during the Second World War, only weeks before Japanese troops were due to be posted overseas for the first time in five decades as part of the UN peacekeeping operation in Cambodia. Japan's Asian neighbors have expressed strong reservations about the use of the troops. In some cases, the (Japanese) soldiers were suffering from starvation, but in many other cases they were not starving at all. The Japanese soldiers were fit and strong, and had potatoes, rice and dried fish. It was not a result of a breakdown in morale: however, morale was good. Often it was in a group under instruction of a commander. It was to get a feeling for victory, and to give the soldiers nerves of steel. It helped the soldiers to bond because the whole troop broke the taboo of cannibalism together. A Pakistani, who was captured when Japan overran Singapore and taken to New Guinea, testified that in his area Japanese soldiers killed and ate one prisoner a day for 'about 100' days. The corporal said he saw flesh being cut from prisoners who were still alive (7). Another well-documented case occurred in Chichijima in February 1945, when Japanese soldiers killed and consumed five American airmen. This case was investigated in 1947 in a war crimes trial, and of 30 Japanese soldiers prosecuted, five (Maj. Matoba, Gen. Tachibana, Adm. Mori, Capt. Yoshii, and Dr. Teraki) were found guilty and hanged (8). In his book Flyboys: A True Story of Courage, James Bradley details several instances of cannibalism of World War II Allied prisoners by their Japanese captors (9). The author claims that this included not only ritual cannibalization of the livers of freshly killed prisoners, but also the cannibalization-for-sustenance of living prisoners over the course of several days, amputating limbs only as needed to keep the meat fresh (10).

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Following the German surrender in January 1943, roughly 100,000 German soldiers were taken prisoners of war. Almost all of them were sent to prisoner of war camps in Siberia or Central Asia where, due to being chronically underfed by their Soviet captors, many resorted to cannibalism. Fewer than 5,000 of the prisoners taken at Stalingrad survived captivity. The majority, however, died early in their imprisonment due to exposure or sickness brought on by conditions in the surrounded army before the surrender (11). During the 872-day Siege of Leningrad, reports of cannibalism began to appear in the winter of 1941–1942, after all birds, rats and pets were eaten by survivors. Leningrad police even formed a special division to combat cannibalism. During January 1944, 800,000 people, nearly a third of the population at the siege‟s beginning, starved to death. Roughly one in three. Many of them in the streets. People were concealing deaths in the family, hiding the bodies so that the deceased‟s ration card could be used until it expired. Husbands and fathers helped to feed their families posthumously. It was a very severe winter - temperatures of minus 35 degrees. Trams froze in their tracks. Buildings burned for days - fire services ceased to function. Factories closed, hospitals were overwhelmed, and cemeteries could not keep pace. Bodies, shrouded but uncoffined, were dragged through the streets on sleds. At one cemetery gate, a corpse propped upright with a cigarette in its mouth extended a frozen arm and finger as a sign post to the newest mass graves. There was a crime wave, mainly of adolescent muggers thieving food and ration cards. One 18-year-old killed his two younger brothers for their cards. Another murdered his granny with an axe and boiled her liver. A 17-year-old stole a corpse from a cemetery and put it through a mincer. Rumors of cannibalism abounded. Amputated limbs disappeared from hospital theatres. Police records released years later showed that 2,000 people were arrested for cannibalism; 586 of them were executed for murdering their victims. Most people arrested however were women. Mothers smothered very young children to feed their older ones (12). In his book, The Gulag Archipelago, Soviet writer Alexander Solzhenitsyn described cases of cannibalism in 20th-century in USSR. Of the famine in Povolzhie (1921–1922) he wrote: "That horrible famine was up to cannibalism, up to consuming children by their own parents - the famine, which Russia had never known even in Time of Troubles [in 1601–1603] ..." (13,14).

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Assessment: cannibalism occurred as a preferred form of protein consumption or as the result of extreme necessity, to absorb the virtues of others, to facilitate conception, for magicoreligious reasons, in warfare, famine, revenge, filial piety, and justice. References 1. Burton-Bradley BG. Cannibalism for cargo. J Nerv Ment Dis. 1976; l63(6):428-31. 2. Bonsall MB, Klug H. Effects of among-offspring relatedness on the origins and evolution of parental care and filial cannibalism. J Evol Biol. 2011;24(6):1335-50. 3. Woznicki, Andrew N. Endocannibalism of the Yanomami. The Summit Times. 1998;6:18-19. 4. Goldman, Laurence, ed. The Anthropology of Cannibalism. Greenwood Publishing Group. 1999, p. 16. ISBN 0-89789-596-7. 5. Joel R. Properties of Nature, Properties of Culture: Ownership, Recognition, and the Politics of Nature in a Papua New Guinea Society. In Aletta B, James G. Reimagining Political Ecology. Duke University Press. 2006, pp. 176–7. ISBN 0-8223-3672-3. 6. Japanese Cannibalism Against Allied & Other POW's! Available 20 December 2013 at www.ww2f.com › ... › War in the Pacific › Land Warfare in the Pacific‎. 7. Japanese troops 'ate flesh of enemies and civilians'. Available 14 December 2013 at www.independent.co.uk › News › World‎. 8. Welch JM. Cannibalism without a hangman, without a rope: Navy war crimes trials after World War II. Intern J Naval Hist. 2002;1(1). 9. Flyboys JB. A True Story of Courage (1st ed.). Little, Brown and Company (Time Warner Book Group). 2003. ISBN 0-316-10584-8. 10. Flyboys JB: A True Story of Courage, first ed. Boston, Massachusetts: Back Bay Books. 2003,2004. pp. 229–230, 311, 404. ISBN 0-316-15943-3. 11. Antony B. Stalingrad: The Fateful Siege. Penguin Books, 1999. 12. Reid A. Leningrad: tragedy of a city under siege. 1941-44. Lewis P. Beyond horror: They ate cats, sawdust, wallpaper paste...even their own babies. Leningrad's agony as the Nazis tried to starve it into submission. Available 20 December 2013 at www.theguardian.com › Culture › Books › History‎. 13. Solzhenitsyn A. The Gulag Archipelago, Part I, comments to Chapter 5. 14. Solzhenitsyn A. The Gulag Archipelago, Part III, Chapter 15.

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ADVANTAGES AND DISADVANTAGES OF CANNIBALISM Cannibalism is an interaction between individuals that can produce counter-intuitive effects at the population level. A striking effect is that a population may persist under food conditions such that the noncannibalistic variant is doomed to go extinct. This so-called lifeboat mechanism has received considerable attention. Implicitly, such studies sometimes suggest, that the lifeboat mechanism procures an evolutionary advantage to the cannibalistic trait. In the context of a size structured population model, the conditions under which the lifeboat mechanism works were compared with those that guaranteed that a cannibalistic mutant can invade successfully under the steady environmental conditions as set by a non-cannibalistic resident. Qualitative agreement and quantitative difference were found. In particular, a prerequisite for the lifeboat mechanism is that cannibalistic mutants are successful invaders. Roughly speaking, these results show that cannibalism brings advantages to both the individuals and the population when adult food is limited (1).

Cannibalism is characterized by four aspects: killing victims, gaining energy from victims, size-dependent interactions and intraspecific competition. In the mathematical models of cannibalistic populations, the predicted population dynamic consequences of cannibalism to its four defining aspects were related. Five classes of effects of cannibalism were distinguished: (i) regulation of population size; (ii) destabilization resulting in population cycles or chaos; (iii) stabilization by damping population cycles caused by other interactions; (iv) bistability such that, depending on the initial

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conditions, the population converges to one of two possible stable states; and (v) modification of the population size structure. Different combinations of cannibalism aspects may cause the same effects. By contrast, the same combination of aspects may lead to different effects. For particular cannibalistic species, the consequences of cannibalism will depend on the presence and details of the four defining aspects (2). Drawing attention to the associations evoked in the process of food denomination, attempts were made to reveal the traces of cannibalistic instinct in this realm. By singling out some of the principal ways used in the semantic reticulum to name food, special attention is devoted to those names that allude to particular categories of enemies (devoured as wholes or as parts of the body). What is referred to as "anti-taboo" shows how the linguistic and more generally the anthropological 'substitution process' is a human reaction to the persistence of the cannibalistic taboo in the face of the belief that cannibalism is an impulse far removed from modern society. Cannibalistic metaphors have been analyzed in other fields, whereas in the area of food denomination they have been neglected. The suggestion is that the reason for such neglect may lie in an attempt to conceal our closeness to the cannibalistic instinct in the form of linguistic substitution (3). Assessment: cannibalism is an interaction between individuals that can produce counter-intuitive effects at the population level. A prerequisite for the lifeboat mechanism is that cannibalistic mutants are successful invaders. Cannibalism brings advantages to both the individuals and the population when adult food is limited. Four aspects characterize cannibalism: killing victims, gaining energy from victims, size-dependent interactions and intraspecific competition. For particular cannibalistic species, the consequences of cannibalism will depend on the presence and details of these four defining aspects. References 1. Getto P, Diekmann O, de Roos AM. On the (dis) advantages of cannibalism. J Math Biol. 2005;51(6):695-712. 2. Claessen D, de Roos AM, Persson L. Population dynamic theory of size-dependent cannibalism. Proc Biol Sci. 2004;271(1537):333-40. 3. Modena ML. Traces of cannibalistic instinct in food denomination. Coll Antropol. 2004;28 Suppl 1:221-7.

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NON-HUMAN CANNIBALISM Ecological interactions determine the structure and dynamics of communities and their responses to the environment. Understanding the community-level effects of ecological interactions, such as intraand interspecifc competition, predation, and cannibalism, is therefore central to ecological theory and ecosystem management. Here, the community-level consequences of cannibalism in populations with density-dependent maturation and reproduction were investigated. A stage-structured consumer population with an ontogenetic diet shift was modeled to analyze how cannibalism alters the conditions for the invasion and persistence of stage-specific predators and competitors. Thus, cannibalistic interactions can facilitate coexistence with other species at both trophic levels. This effect of cannibalism critically depends on the food dependence of the demographic processes. The underlying mechanism is a cannibalism-induced shift in the biomass distribution between the consumer life stages. These findings suggest that cannibalism may alter the structure of ecological communities through its effects on species coexistence (1). Cannibalism can play a prominent role in the structuring and dynamics of ecological communities. Previous studies have emphasized the importance of size structure and density of cannibalistic species in shaping short- and long-term cannibalism dynamics, but our understanding of how predators influence cannibalism dynamics is limited. This is despite widespread evidence that many prey species exhibit behavioral and morphological adaptations in response to predation risk. This study examined how the presence and absence of predation risk from larval dragonflies Aeshna nigroflava affected cannibalism dynamics in its prey larval salamanders Hynobius retardatus. Feedback dynamics between size structure and cannibalism depended on whether dragonfly predation risk was present. In the absence of dragonfly risk cues, a positive feedback between salamander size structure and cannibalism through time occurred because most of the replicates in this treatment contained at least one salamander larvae having an enlarged gape (i.e. cannibal). By contrast, this feedback and the emergence of cannibalism were rarely observed in the presence of the dragonfly risk cues. Once salamander size divergence occurred, experimental reversals of the presence or absence of dragonfly risk cues did not alter existing cannibalism dynamics as the experiment progressed. Thus, the effects of risk on the mechanisms driving cannibalism

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dynamics likely operated during the early developmental period of the salamander larvae. The effects of dragonfly predation risk on behavioral aspects of cannibalistic interactions among hatchlings may prohibit the initiation of dynamics between size structure and cannibalism. Encounter rates among hatchlings and biting and ingestion rates of prospective prey by prospective cannibals were significantly lower in the presence vs. absence of dragonfly predation risk even though the size asymmetry between cannibals and victims was similar in both risk treatments. These results suggest that dragonfly risk cues first suppress cannibalism among hatchlings and then prevent size variation from increasing through time. The positive feedback dynamics between size structure and cannibalism and their modification by predation risk may also operate in other systems to shape the population dynamics of cannibalistic prey species as well as overall community dynamics (2). Cannibalistic and asymmetrical behavioral interactions between stages are common within stage-structured predator populations. Such direct interactions between predator stages can result in density- and trait-mediated indirect interactions between a predator and its prey. A set of structured predator-prey models is used to explore how such indirect interactions affect the dynamics and structure of communities. Analyses of the separate and combined effects of stage-structured cannibalism and behavior-mediated avoidance of cannibals under different ecological scenarios show that both cannibalism and behavioral avoidance of cannibalism can result in short- and long-term positive indirect connections between predator stages and the prey, including "apparent mutualism." These positive interactions alter the strength of trophic cascades such that the system's dynamics are determined by the interaction between bottom-up and top-down effects. Contrary to the expectation of simpler models, enrichment increases both predator and prey abundance in systems with cannibalism or behavioral avoidance of cannibalism. The effect of behavioral avoidance of cannibalism, however, depends on how strongly it affects the maturation rate of the predator. Behavioral interactions between predator stages reduce the short-term positive effect of cannibalism on the prey density, but can enhance its positive long-term effects. Both interaction types reduce the destabilizing effect of enrichment. These results suggest that inconsistencies between data and simple models can be resolved by accounting for stage-structured interactions within and among species (3). Theory and empirical studies suggest that cannibalism in agestructured populations can regulate recruitment depending on the

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intensity of intraspecific competition between cannibals and victims and the nature of the cannibalism window, i.e., which size classes interact as cannibals and victims. A series of experiments quantify that window for age-structured populations of salamander larvae and paedomorphic adults. Body size limits on cannibalism in microcosms and then the consumptive and nonconsumptive (injuries, foraging and activity, diet, and growth) effects on victims in mesocosms with seminatural levels of habitat complexity and alternative prey were determined. Cannibalism by the largest size classes (paedomorphs and ≥ age 3+ years, larvae) occurs mainly on YOY victims. Surviving YOY and other small larvae had increased injuries, reduced activity levels, and reduced growth rates in the presence of cannibals. Data on YOY survival in an experiment in which the density of paedomorphs combined with historical data on the number of cannibals in natural populations were manipulated indicate that dominant cohorts of paedomorphs can cause observed recruitment failures. Dietary data indicate that ontogenetic shifts in diet should preclude strong intraspecific competition between YOY and cannibals in this species. These results are consistent with previous empirical and theoretical work that suggests that recruitment regulation by cannibalism is most likely when YOY are vulnerable to cannibalism but have low dietary overlap with cannibals. Understanding the role of cannibalism in regulating recruitment in salamander populations is timely, given the widespread occurrences of amphibian decline. Previous studies have focused on extrinsic (including anthropogenic) factors that affect amphibian population dynamics, whereas the data presented here combined with long-term field observations suggest the potential for intrinsically driven population cycles (4). Direct and indirect interactions between two prey species can strongly alter the dynamics of predator-prey systems. Most predators are cannibalistic, and as a consequence, even systems with only one predator and one prey include two prey types: conspecifics and heterospecifics. The effects of the complex direct and indirect interactions that emerge in such cannibalistic systems are still poorly understood. This study examined how the indirect interaction between conspecific and heterospecific prey affects cannibalism and predation rates and how the direct interactions between both species indirectly alter the effect of the cannibalistic predator. These effects using larvae of the stream salamanders Eurycea cirrigera (prey) and Pseudotriton ruber (cannibalistic predator) by manipulating the relative densities of the conspecific and heterospecific prey in the presence and absence of the predator in experimental streams were tested. The rates of

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cannibalism and heterospecific predation were proportional to the respective densities and negatively correlated, indicating a positive indirect interaction between conspecific and heterospecific prey, similar to "apparent mutualism." Direct interactions between prey species did not alter the effect of the predator. Although both types of prey showed a similar 30% reduction in night activity and switch in microhabitat use in response to the presence of the predator, cannibalism rates were three times higher than heterospecific predation rates irrespective of the relative densities of the two types of prey. Cumulative predation risks differed even more due to the 48% lower growth rate of conspecific prey. Detailed laboratory experiments suggest that the 3:1 difference in cannibalism and predation rate was due to the higher efficiency of heterospecific prey in escaping immediate attacks. However, no difference was observed when the predator was a closely related salamander species, Gyrinophilus porphyriticus, indicating that this difference is species specific. This demonstrates that cannibalism can result in the coupling of predator and prey mortality rates that strongly determines the dynamics of predator-prey systems (5). Occurrence of cannibalism and inferior competitive ability of predators compared to their prey have been suggested to promote coexistence in size-structured IGP systems. The intrinsic size-structure of fish provides the necessary prerequisites to test whether the above mechanisms are general features of species interactions in fish communities where IGP is common. Whether Arctic char (Salvelinus alpinus) was more efficient as a cannibal than as an interspecific predator on the prey fish ninespine stickleback (Pungitius pungitius) and whether ninespine stickleback were a more efficient competitor on the shared zooplankton prey than its predator, Arctic char were experimentally tested. Secondly, a literature survey was evaluated if piscivores in general are more efficient as cannibals than as interspecific predators and whether piscivores are inferior competitors on shared resources compared to their prey fish species. Both controlled pool experiments and outdoor pond experiments showed that char imposed a higher mortality on YOY char than on ninespine sticklebacks, suggesting that piscivorous char is a more efficient cannibal than interspecific predator. Estimates of size dependent attack rates on zooplankton further showed a consistently higher attack rate of ninespine sticklebacks compared to similar sized char on zooplankton, suggesting that ninespine stickleback is a more efficient competitor than char on zooplankton resources. The literature survey showed that piscivorous top consumers generally selected

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conspecifics over interspecific prey, and prey species are competitively superior compared to juvenile piscivorous species in the zooplankton niche. In conclusion, the observed selectivity for cannibal prey over interspecific prey and the competitive advantage of prey species over juvenile piscivores are common features in fish communities and the observed selectivity for cannibalism over interspecific prey has the potential to mediate coexistence in size structured intraguild predation systems (6). Cases when several adults kill and consume an infant together have been recorded in social mammals. This occurs in lions, for example, when a group of new males acquires another male's harem (7). It also occurs in chimpanzees where male groups commonly attack conspecifics (8-10). References 1. Ohlberger J, Langangen O, Stenseth NC, Vøllestad LA. Communitylevel consequences of cannibalism. Am Nat. 2012;180(6):791-801. 2. Kishida O, Trussell GC, Ohno A, et al. Predation risk suppresses the positive feedback between size structure and cannibalism. J Anim Ecol. 2011;80(6):1278-87. 3. Rudolf VH. Consequences of stage-structured predators: cannibalism, behavioral effects, and trophic cascades. Ecology. 2007; 88(12):2991-3003. 4. Rudolf VH. Impact of cannibalism on predator-prey dynamics: sizestructured interactions and apparent mutualism. Ecology. 2008;89(6):165060. 5. Wissinger SA, Whiteman HH, Denoël M, et al. Consumptive and nonconsumptive effects of cannibalism in fluctuating age-structured populations. Ecology. 2010;91(2):549-59. 6. Byström P, Ask P, Andersson J, Persson L. Preference for cannibalism and ontogenetic constraints in competitive ability of piscivorous top predators. PLoS One. 2013;8(7):e70404. 7. Bertram B.C. Social factors influencing reproduction in wild lions. J. Zool. 1975;177:463-82. 8. Arcadi A.C, Wrangham R.W. Infanticide in chimpanzees: review of cases and a new within-group observation from the Kanyawara study group in Kibale National Park. Primates. 1999;40:337-51. 9. Mitani J.C, Watts D.P, Muller M.N. Recent developments in the study of wild chimpanzee behavior. Evol Anthropol Issues, News, Rev. 2002;11:925. 10. Wilson M.L, Wallauer W.R, Pusey A.E. New cases of intergroup violence among chimpanzees in Gombe National Park, Tanzania. Int J Primatol. 2004;25:523-49.

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CANNIBALISM IN DIFFERENT SPECIES Cannibalism is widespread in the animal kingdom and is a major mortality factor in the biology of numerous species (1-4). Cannibalism differs from other predator–prey interactions, in that both the prey and the predator belong to the same species, and this intraspecific predation can result in population dynamics that are very different from that found in systems without cannibalism (5).

Cannibalistic lions

Cannibalism is quite common among animals. Lions eat other lions, bears eat other bears, and chimpanzees eat other chimpanzees. This is not necessarily the result of a food shortage. Such incidents occur even when there is a plentiful supply of food from other sources. In the case of lions, for example, a male lion sometimes kills and eats the cubs of a female lion without a partner to protect her. The male lion does this because the female lion will not mate as long as she has cubs, but she will mate after the cubs are dead. For some species, cannibalism is extremely common. Approximately one fifth of the diet of a tiger salamander, for example, consists of other tiger salamanders (6). Male chimpanzees at the Gombe National Park were twice seen to attack 'stranger' females and seize their infants. One infant was then killed and partially eaten: the other was 'rescued' and carried by 3 different males. Once several males were found eating a freshly killed 'stranger' infant. A similar event was observed in Mahali Mountains, Tanzania. A different kind of killing occurred at Gombe when a female and her daughter killed and ate 3 infants of other females of the same community during a 2-year period. There is evidence

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suggesting that other infants may have died in this way. These are puzzling aspects of infant killing and cannibalism in chimpanzees (7). Observations of ape cannibalism have been limited to chimpanzees (Pan troglodytes) where it is associated with infanticide and consumption by unrelated individuals (Watts and Mitani, Primates 41(4):357-365, 2000). Two unrelated female Sumatran orangutans (Pongo abelii) cannibalized the remains of their infants on different occasion, a behavior never before reported in any ape species. The two orangutans were wild-born rehabilitated individuals, and had been reintroduced to an area hosting a largely unregulated primate tourism industry and experienced restricted ranging conditions. Though it is possible that this is a strategy to regain energy and nutrients or a result of individual history, comparative data suggest that this is an aberrant behavior which may be linked to environmental stressors within the area (8). The cannibalization of an infant bonobo (circa 2.5 years old) at Lui Kotale, in the Democratic Republic of Congo is described. The infant died of unknown causes and was consumed by several community members including its mother and an older sibling one day after death. Certain features concerning the pattern of consumption fit in with previously observed episodes of cannibalism in Pan, whereas others, such as the mother's participation in consuming the body, are notable. This incident suggests that filial cannibalism among apes need not be the result of nutritional or social stress and does not support the idea that filial cannibalism is a behavioral aberration (9). Why should animals knowingly consume their own young? It is difficult to imagine many circumstances in which eating one's own young (i.e., filial cannibalism) actually increases an individual's fitness; however, filial cannibalism commonly co-occurs with parental care in fishes. The evolutionary significance of filial cannibalism remains unclear. The most commonly accepted explanation is that filial cannibalism is a mechanism by which caring males gain energy or nutrients that they reinvest into future reproduction, thereby increasing net reproductive success. There is mixed support for this hypothesis and, at best, it can only explain filial cannibalism in some species. A recent alternative hypothesis suggests that filial cannibalism improves the survivorship of remaining eggs by increasing oxygen availability, and thus increases current reproductive success. This theory has received little attention as of yet. The hypothesis of oxygen-mediated filial cannibalism in the sand goby examined the effect of oxygen and egg density on the occurrence of filial cannibalism, and evaluated the effects of partial clutch

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cannibalism on the survivorship of remaining eggs, comparing potential costs and benefits of filial cannibalism related to the net number of eggs surviving. Oxygen level and egg density affected the occurrence of cannibalism and simulated partial clutch cannibalism improved survivorship of the remaining eggs. Additionally, because increased egg survivorship, stemming from partial egg removal, compensated for the cost of cannibalism (i.e., number of eggs removed) at a range of cannibalism levels, filial cannibalism potentially results in no net losses in reproductive success. However, oxygen did not affect egg survivorship. Thus, it has been suggested in a more general hypothesis that filial cannibalism is mediated by density-dependent egg survivorship (10). References 1. Fox LR. Cannibalism in natural populations. Annu Rev Ecol Syst. 1975;6:87-106. 2. Polis GA. The evolution and dynamics of intraspecific predation. Annu Rev Ecol Syst. 1981;12:225-51. 3. Elgar MA, Crespi BJ, editors. Cannibalism: Ecology and Evolution among Diverse Taxa. Oxford University Press; New York, NY. 1992. 4. Van Schaik CP, Janson CH. Infacticide by males and its implications. Cambridge University Press; Cambridge, UK. 2000. 5. Claessen D, De Roos AM, Persson L. Population dynamic theory of size-dependent cannibalism. Proc R Soc B. 2004;271:333-40. 6. Cannibalism in the animal kingdom. Available 29 November 2013 at ar.vegnews.org/cannibalism.html‎. 7. Goodall J. Infant killing and cannibalism in free-living chimpanzees. Folia Primatol (Basel). 1977;28(4):259-89. 8. Dellatore DF, Waitt CD, Foitova I. Two cases of mother-infant cannibalism in orangutans. Primates. 2009;50(3):277-81. 9. Fowler A, Hohmann G. Cannibalism in wild bonobos (Pan paniscus) at Lui Kotale. Am J Primatol. 2010;72(6):509-14. 10. Klug H, Lindström K, St Mary CM. Parents benefit from eating offspring: density-dependent egg survivorship compensates for filial cannibalism. Evolution. 2006;60(10):2087-95.

INSECTS Cannibalism is important in the collective motion of mass migratory bands of insects, such as locusts and Mormon crickets. These mobile groups consist of millions of individuals and are highly destructive to vegetation. Individuals move in response to attacks from

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approaching conspecifics and bite those ahead, resulting in further movement and encounters with others. Despite the importance of cannibalism, the way in which individuals make attack decisions and how the social context affects these cannibalistic interactions is unknown. This can be understood by examining the decisions made by individuals in response to others. A field investigation shows that adult Mormon crickets were more likely to approach and attack a stationary cricket that was side-on to the flow than either head- or abdomen-on, suggesting that individuals could reduce their risk of an attack by aligning with neighbors. Strong social effects on cannibalistic behavior encounters lasted longer that were more likely to result in an attack, and attacks were more likely to be successful if other individuals were present around a stationary individual. This local aggregation appears to be driven by positive feedback whereby the presence of individuals attracts others, which can lead to further crowding. This work improves our understanding of the local social dynamics driving migratory band formation, maintenance and movement at the population level (1). Generation cycles, population cycles with a period of approximately one generation, have been observed in a variety of field and laboratory studies. Such dynamics are predicted to arise through the effects of resource competition and cannibalism or involve consumer-natural enemy interactions. In a new highly simplified model, generation cycles are a very common outcome of strongly agestructured intraspecific interactions involving cannibalism. Unique long-term time-series of ladybeetle (Coccinellidae) abundances from tropical Indonesia were analyzed. Some of the time-series display clear generation cycles, and these are caused by intraspecific cannibalism (2). References 1. Bazazi S, Ioannou CC, Simpson SJ, et al. The social context of cannibalism in migratory bands of the Mormon cricket. PLoS One. 2010; 5(12):e15118. 2. Nakamura K, Hasant N, Abbas I, et al. Generation cycles in Indonesian lady beetle populations may occur as a result of cannibalism. Proc Biol Sci. 2004;271 Suppl 6:S501-4.

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SPIDER CANNIBALISM Cannibalism among generalist predators has implications for the dynamics of terrestrial food webs. Spiders are common, ubiquitous arthropod generalist predators in most natural and managed terrestrial ecosystems. Thus, the relationship of spider cannibalism to food limitation, competition, and population regulation has direct bearing on basic ecological theory and applications such as biological control. This review first briefly treats the different types of spider cannibalism and then focuses in more depth on evidence relating cannibalism to population dynamics and food web interactions to address the following questions: Is cannibalism in spiders a foraging strategy that helps to overcome the effects of a limited supply of calories and/or nutrients? Does cannibalism in spiders reduce competition for prey? Is cannibalism a significant density-dependent factor in spider population dynamics? Does cannibalism dampen spider-initiated trophic cascades? (1). Cannibalism is considered an adaptive foraging strategy for animals of various trophic positions, including carnivores. However, previous studies on wolf spiders have questioned the high nutritional value of cannibalism. Two different aspects of nutritional quality of conspecifics in the wolf spider Pardosaprativaga were analyzed: their value for survival, growth and development; and the growth efficiency of feeding on conspecifics. The propensity for cannibalistic attacks and the consumption rate of conspecifics were measured in an experiment where hunger level and nutrient balance were manipulated. In all experiments, cannibalism was compared with predation on fruit flies as control prey. The growth experiment gave ambiguous results regarding the nutritional quality of conspecifics. Spiders on pure cannibalistic diets split into two distinct groups, one performing much better and the other much worse than spiders on fruit fly diets. There is the possibility that the population is dimorphic in its cannibalistic propensity, with the latter group of individuals showing a high level of inhibition against cannibalistic attacks in spite of a high nutritional value of cannibalism. The food utilization experiment confirmed the high nutritional quality of conspecifics, as cannibalistic spiders had the same growth rate as spiders fed insect prey in spite of a much lower consumption rate. Inhibition against cannibalistic attacks was demonstrated in medium-sized juveniles: only half of the spiders attacked a prescribed victim of 50% the size of their opponents, and the latency for those that did attack was more

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than half an hour, compared with a few minutes for spiders fed fruit flies. Nutrient-imbalanced spiders utilized an alternative insect diet less efficiently than balanced spiders, whereas no difference was present in efficiency of utilizing conspecifics. This result indicates that spiders can remedy at least part of a nutrient imbalance through cannibalism. As spiders can escape nutritional imbalance as well as restore energy reserves through cannibalism, both nutrient imbalance and hunger to stimulate cannibalism can be predicted. This prediction was confirmed only with respect to hunger. Nutrient-imbalanced spiders had reduced cannibalistic consumption, perhaps due to lowered predatory aggressiveness because of bad condition (2).

Spider versus spider

Sexual cannibalism may be a form of extreme sexual conflict in which females benefit more from feeding on males than mating with them, and males avoid aggressive, cannibalistic females in order to increase net fitness. A thorough understanding of the adaptive significance of sexual cannibalism is hindered by our ignorance of its prevalence in nature. Furthermore, there are serious doubts about the food value of males, probably because most studies that attempt to document benefits of sexual cannibalism in the female have been conducted in the laboratory with non-natural alternative prey. Thus, to understand more fully the ecology and evolution of sexual cannibalism, field experiments are needed to document the prevalence of sexual cannibalism and its benefits to females. Field experiments with the Mediterranean tarantula (Lycosa tarantula), a burrowing wolf spider, and to address these issues were conducted. At natural rates of encounter with males, approximately a third of L. tarantula females cannibalized the male. The rate of sexual cannibalism increased with male availability, and females were more likely to kill and consume an approaching male if they had previously mated with another male. Females benefit from feeding on a male by breeding earlier, producing

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30% more offspring per egg sac, and producing progeny of higher body condition. Offspring of sexually cannibalistic females dispersed earlier and were larger later in the season than spiderlings of noncannibalistic females. In conclusion, in nature a substantial fraction of female L. tarantula kill and consume approaching males instead of mating with them. This behavior is more likely to occur if the female has mated previously. Cannibalistic females have higher rates of reproduction, and produce higher-quality offspring than noncannibalistic females. These findings further suggest that female L. tarantula is nutrient-limited in nature and that males are high-quality prey. The results of these field experiments support the hypothesis that sexual cannibalism is adaptive to females (3). Cannibalism is hypothesized to have evolved as a way to obtain a high-quality meal. The extraction of lipid and protein by female wolf spiders, Hogna helluo, during sexual cannibalism of males and predation of crickets was examined. Most food-limited females did not cannibalize males but immediately consumed a size-matched cricket. When consuming male H. helluo and crickets, female H. helluo only consumed 51% of the male body while they consumed 72% of the cricket body. While males had higher protein content in their bodies than crickets and other insects, female H. helluo ingested similar amounts of protein from male H. helluo and crickets. Female H. helluo extracted 47% of the protein present in male H. helluo and 67% of the protein present in crickets. Females were able to extract nearly the entire lipid present in male H. helluo and crickets. However, crickets and other insects had almost 4 times higher lipid content than male H. helluo. The ratio of lipid to protein consumed from crickets appeared more similar to the nutritional requirements of egg production than that of males. Taken together, female hesitancy to engage in cannibalism, low extraction of nutrients from males and a low ratio of lipid to protein in the food extracted from males suggest that males may be poor-quality prey items compared to common insects such as crickets (4). References 1. Wise DH. Cannibalism, food limitation, intraspecific competition, and the regulation of spider populations. Annu Rev Entomol. 2006;51:441-65. 2. Mayntz D, Toft S. Nutritional value of cannibalism and the role of starvation and nutrient imbalance for cannibalistic tendencies in a generalist predator. J Anim Ecol. 2006;75(1):288-97.

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3. Rabaneda-Bueno R, Rodríguez-Gironés MA, Aguado-de-la-Paz S, et al. Sexual cannibalism: high incidence in a natural population with benefits to females. PLoS One. 2008;3(10):e3484. 4. Wilder SM, Rypstra AL. Males make poor meals: a comparison of nutrient extraction during sexual cannibalism and predation. Oecologia. 2010;162(3):617-25.

LARVAE COMPLEX

OF

ANOPHELES

GAMBIA

Among the aquatic developmental stages of the Anopheles gambiae complex (Diptera: Culicidae), both inter- and intra-specific interactions influence the resulting densities of adult mosquito populations. For three members of the complex, An. arabiensis Patton, An. quadriannulatus (Theobald) and An. gambiae Giles sensu stricto, some aspects of this competition under laboratory conditions were investigated. First-instar larvae were consumed by fourth-instar larvae of the same species (cannibalism) and by fourth-instar larvae of other sibling species (predation). Even when larvae were not consumed, the presence of one fourth-instar larva caused a significant reduction in development rate of first-instar larvae (1). Reference 1. Koenraadt CJ, Takken W. Cannibalism and predation among larvae of the Anopheles gambiae complex. Med Vet Entomol. 2003;17(1):61-6.

TUCURANES CANNIBALISM Individuals of its own genus were the main food item of two species of tucunares (Cichla cf. ocellaris and Cichla monoculus) introduced into the Volta Grande Reservoir. The abundance of adult tucunares may cause intra-specific competition, possibly leading to the high cannibalism rates (1). Reference 1. Gomiero LM, Braga FM. Cannibalism as the main feeding behaviour of tucunares introduced in southeast Brazil. Braz J Biol. 2004;64(3B):625-

32.

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BACTERIAL POPULATION Bacterial populations can evolve and adapt to become diverse niche specialists, even in seemingly homogeneous environments. One source of this diversity arises from newly 'constructed' niches that result from the activities of the bacteria themselves. Ecotypes specialized to exploit these distinct niches can subsequently coexist via frequency-dependent interactions. Here, a novel form of niche construction that is based upon differential death and cannibalism, and which evolved during 20,000 generations of experimental evolution in Escherichia coli in a seasonal environment with alternating growth and starvation is described. In one of 12 populations, two monophyletic ecotypes, S and L, evolved that stably coexist with one another. When grown and then starved in monoculture, the death rate of S exceeds that of L, whereas the reverse is observed in mixed cultures. As shown by experiments and numerical simulations, the competitive advantage of S cells is increased by extending the period of starvation, and this advantage results from their cannibalization of the debris of lysed L cells, which allows the S cells to increase both their growth rate and total cell density. At the molecular level, the polymorphism is associated with divergence in the activity of the alternative sigma factor RpoS, with S cells displaying no detectable activity, while L cells show increased activity relative to the ancestral genotype. These results extend the repertoire of known cross-feeding mechanisms in microbes to include cannibalism during starvation, and confirm the central roles for niche construction and seasonality in the maintenance of microbial polymorphisms (1). Cannibalism and fratricide refer to the killing of genetically identical cells (siblings) that was recently documented in two Grampositive species, Bacillus subtilis and Streptococcus pneumoniae, respectively. Cannibalism occurs during the early stages of sporulation in Bacillus subtilis, whereas fratricide occurs in Streptococcus pneumoniae during natural genetic transformation. Are these processes fundamentally different from the more traditional 'chemical warfare' among bacteria? (2). References 1. Rozen DE, Philippe N, Arjan de Visser J, et al. Death and cannibalism in a seasonal environment facilitate bacterial coexistence. Ecol Lett. 2009; 12(1):34-44. 2. Claverys JP, Håvarstein LS. Cannibalism and fratricide: mechanisms and raisons d'être. Nat Rev Microbiol. 2007;5(3):219-29.

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EGG CANNIBALISM In laboratory experiments, third and early fourth instar larvae of Toxorhynchites amboinensis, Tx. splendens and Tx. brevipalpis, previously starved 24 hours, rapidly cannibalized eggs of their own species, or ate the eggs of other species present on the water surface in small containers. Toxorhynchites amboinensis and Tx. splendens larvae of either instar cannibalized eggs somewhat more rapidly than Tx. brevipalpis, probably because brevipalpis eggs distributed themselves around the edge of the container and were less accessible. When offered heterospecific eggs, fourth instar larvae of all three species ate them as efficiently as they cannibalized their own except that eggs of Tx. brevipalpis were eaten very slowly. Toxorhynchites amboinensis larvae were offered nonspecific hatched eggs, and these also were consumed (1). Parental care and filial cannibalism (the consumption of one's own offspring) co-occur in many animals. While parental care typically increases offspring survival, filial cannibalism involves the killing of one's young. Using an evolutionary ecology approach, the importance of a range of factors on the evolution of parental care and filial cannibalism were evaluated. Parental care, no care/total abandonment, and filial cannibalism evolved often coexisted over a range of parameter space. While no single benefit was essential for the evolution of filial cannibalism, benefits associated with adult or offspring survival and/or reproduction facilitated the evolution of cannibalism. This model highlights the plausibility of a range of alternative hypotheses. Specifically, the evolution of filial cannibalism was enhanced if [1] parents could selectively cannibalize lowerquality offspring, [2] filial cannibalism increased egg maturation rate, [3] energetic benefits of eggs existed, or [4] cannibalism increased a parent's reproductive rate (e.g., through mate attractiveness). Densitydependent egg survivorship alone did not favor the evolution of cannibalism. However, when egg survival was density dependent, filial cannibalism invaded more often when the density dependence was relatively more intense. These results suggest that populationlevel resource competition potentially plays an important role in the evolution of both parental care and filial cannibalism (2).

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References 1. Linley JR, Duzak D. Egg cannibalism and carnivory among three species of Toxorhynchites. J Am Mosq Control Assoc. 1989;5(3):359-62. 2. Klug H, Bonsall MB. When to care for, abandon, or eat your offspring: the evolution of parental care and filial cannibalism. Am Nat. 2007;170(6): 886-901.

CANNIBALISM IN FISHES Filial cannibalism (the eating of one's own offspring) occurs in a variety of taxa, but is especially prevalent in fishes with parental care. Recent research supports a central tenet of parental-investment theory; that is, parents consume their offspring when it maximizes their lifetime reproductive success (1).

Shark eats shark

This review summarizes information on filial cannibalism in teleost fish. Cannibalistic parents can either consume their whole brood (total filial cannibalism), or eat only some of the eggs in the nest (partial filial cannibalism). Offspring consumption has been argued to be adaptive under the assumption that offspring survival is traded against feeding, and that offspring can act as an alternative food source for the parents. The evidence supporting the basic predictions formulated under these assumptions is summarized for both total and partial filial cannibalism. These two forms of cannibalism differ significantly since the former represents an investment only in future reproductive success, whereas the latter can affect both present and future reproductive success. Despite a few inconsistencies in the data

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from laboratory and field studies, the energy-based explanation appears valid for both forms of cannibalism. Alternative non-energybased explanations are considered, but they are unable to account for the wide distribution of this behavior in teleosts. The intersexual conflict arise from attempts of the non-cannibal sex to minimize the cost of filial cannibalism together with the potential effect of this behavior on the operational sex ratio at a population level (2). References 1. Fitzgerald GJ. Filial cannibalism in fishes: Why do parents eat their offspring? Trends Ecol Evol. 1992;7(1):7-10. 2. Manica A. Filial cannibalism in teleost fish. Biol Rev Camb Philos Soc. 2002;77(2):261-77.

CANNIBALISTIC GIANTS IN ARCTIC CHAR Recent theoretical studies on the population dynamic consequences of cannibalism have focused on mechanisms behind the emergence of large cannibals (giants) in size-structured populations. Theoretically, giants emerge when a strong recruiting cohort imposes competition induced mortality on stunted adults, but also provides a profitable resource for a few adults that accelerate in growth and reach giant sizes. The effects of a recruitment pulse on the individual and population level in an allopatric Arctic char population have been studied over a 5-year period and these results were contrasted with theoretical model predictions for the conditions necessary for the emergence of cannibalistic giants. The recruitment pulse had negative effects on invertebrate resource abundance, and the decrease in body condition and increase in mortality of adult char suggested that strong intercohort competition took place. The frequency of cannibalism increased and a few char accelerated in growth and reached 'giant' sizes. The main discrepancy between model predictions and field data was the apparently small effect the recruited cohort had on resources and adult char performance during their first summer. Instead, the effects became pronounced when the cohort was 1 year old. This mismatch between model predictions and field observations was suggested to be due to the low per capita fecundity in char and the restricted nearshore habitat use in YOY char. This study provides empirical evidence that the emergence of giants is associated with the

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breakthrough of a strong recruiting cohort and the claimed stable char populations with large cannibals may instead be populations with dynamic size structure that results in intermittent breakthroughs of recruitment pulses, providing the conditions necessary for char to enter the cannibalistic niche. The data suggest that increased recruit survival through restricted habitat use may destabilize dynamics and cause the emergence of giants. However, they also suggest that this does not necessarily develop into populations with bi-modal size structure in populations with low per capita fecundity and size- and density-dependent habitat use of recruiting cohorts (1). Assessment: non-human cannibalism is a widely prevalent behavior in different species. Reference 1. Byström P. Recruitment pulses induce cannibalistic giants in Arctic char. J Anim Ecol. 2006;75(2):434-44.

CELL CANNIBALISM The term self-cannibalism, or autophagy, was coined to describe the ability of the cells to cannibalize their own damaged organelles or proteins. It was morphologically described as the presence of doublemembraned autophagic vesicles filled with diverse cellular materials or debris inside the cells. The presence of autophagic vacuoles has been associated with cell survival, including cell senescence and cancer and appears to be activated by nutrient deprivation. The occurrence of autophagic processes can also lead, as final event, to the death of the cell. The xeno-cannibalism was described as the ability of certain cells, e.g. metastatic cells, to cannibalize their siblings as well as cells from the immune system. Interestingly, metastatic tumor cells are also able to engulf and digest living cells, including autologous lymphocytes that should kill them, i.e. CD8(+) cytotoxic lymphocytes. This can represent a formidable opportunity for metastatic cells to survive in adverse conditions such as those they encounter in their "journey" towards the target organ to establish a colony. Altogether, these findings seem to suggest a pathogenetic role for cannibalic behavior in human pathology and point at this surprising cellular aggressiveness as an innovative pharmacological target in the clinical management of metastatic disease (1).

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Autophagy works as a cellular housekeeper by eliminating defective proteins

Cellular cannibalism, defined as a large cell enclosing a slightly smaller one within its cytoplasm. The morphology, mechanism, and the cytology of cannibalism in relation to malignancy are significant. Cannibalism is a completely different entity than phagocytosis, entosis, and emeriopoliosis. It is an important morphologic feature to distinguish benign from malignant lesions. Cannibalism has been described in various cancers such as, bladder cancer, breast cancer, lung cancer, etc, and this is related with the aggressiveness of the malignancy (2). Autophagy plays an important role in cellular survival by resupplying cells with nutrients during starvation or by clearing misfolded proteins and damaged organelles and thereby preventing degenerative diseases. Conversely, the autophagic process is also recognized as a cellular death mechanism. The circumstances that determine whether autophagy has a beneficial or a detrimental role in cellular survival are currently unclear. Autophagy induction is detrimental in neurons that lack a functional AMPK enzyme and suffer from severe metabolic stress. Autophagy and AMPK are interconnected in a negative feedback loop that prevents excessive and destructive stimulation of the autophagic process. A new survival mechanism in AMPK-deficient neurons - cell cannibalism is uncovered (3). Autophagy is a process in which subcellular membranes undergo dynamic morphological changes that lead to the degradation of cellular proteins and cytoplasmic organelles. This process is an important cellular response to stress or starvation. Many studies have shed light on the importance of autophagy in cancer, but it is still unclear whether autophagy suppresses tumorigenesis or provides

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cancer cells with a rescue mechanism under unfavorable conditions. What is the present state of our knowledge about the role of autophagy in cancer development, and in response to therapy? How can the autophagic process be manipulated to improve anticancer therapeutics? (4).

Neutrophil-tumor cell phagocytosis (cannibalism) in human tumors.

Macroautophagy, often refers to as autophagy, and designates the process by which portions of the cytoplasm, intracellular organelles and long-lived proteins are engulfed in double-membraned vacuoles (autophagosomes) and sent for lysosomal degradation. Basal levels of autophagy contribute to the maintenance of intracellular homoeostasis by ensuring the turnover of supernumerary, aged and/or damaged components. Under conditions of starvation, the autophagic pathway operates to supply cells with metabolic substrates, and hence represents an important pro-survival mechanism. Moreover, autophagy is required for normal development and for the protective response to intracellular pathogens. Conversely, uncontrolled autophagy is associated with a particular type of cell death (termed autophagic, or type II) that is characterized by the massive accumulation of autophagosomes. Regulators of apoptosis (e.g. Bcl-2 family members) also modulate autophagy, suggesting an intimate cross talk between these two degradative pathways. It is still unclear whether autophagic vacuolization has a causative role in cell death or whether it represents the ultimate attempt of cells to cope with lethal stress. For a multicellular organism, autophagic cell death might well represent a pro-survival mechanism, by providing metabolic supplies during whole-body nutrient deprivation. Alternatively, type II cell death might contribute to the disposal of cell corpses when heterophagy is deficient (5).

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A breast cancer cell engages in a major bout of autophagy - "self eating". The bright spots, fluorescent labels, show the cell's garbage collection and recycling plants that are called autophagosomes.

Autophagy or self-eating by mostly morphological studies has been linked to a variety of pathological processes such as neurodegenerative diseases and tumorigenesis, which highlights its biological and medical importance. However, whether autophagy protects from or causes disease is unclear. Actually, acute pancreatitis is one of the earlier pathological processes where autophagy has been described in a human tissue. Autophagy, autodigestion and cell death are early cellular events in acute pancreatitis (6,7). Autodigestion of the pancreas by its own prematurely activated digestive proteases is thought to be an important event in the onset of acute pancreatitis. Although lysosomal hydrolases, such as cathepsin B, play a key role in intrapancreatic trypsinogen activation, it remains unclear where and how trypsinogen meets these lysosomal enzymes. Autophagy is an intracellular bulk degradation system in which cytoplasmic components are directed to the lysosome/vacuole by a membrane-mediated process. To analyze the role of autophagy in acute pancreatitis, a conditional knockout mouse that lacks the autophagy-related (Atg) gene Atg5 in the pancreatic acinar cells was produced. The severity of acute pancreatitis induced by cerulein is greatly reduced in these mice. In addition, Atg5-deficient acinar cells show a significantly decreased level of trypsinogen activation. These data suggest that autophagy exerts a detrimental effect in pancreatic acinar cells by activation of trypsinogen to trypsin. Autophagy accelerates trypsinogen activation by lysosomal hydrolases under acidic conditions, triggering acute pancreatitis in its early stage (8). The autophagy pathway is the major degradation pathway of the cell for long-lived proteins and organelles. Dysfunction of autophagy has been linked to several neurodegenerative disorders that are associated with an accumulation of misfolded protein aggregates.

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Alzheimer's disease, the most common neurodegenerative disorder, is characterized by 2 aggregate forms, tau tangles and amyloid-beta plaques. Autophagy has been linked to Alzheimer's disease pathogenesis through its merger with the endosomal-lysosomal system, which has been shown to play a role in the formation of the latter amyloid-beta plaques. However, the precise role of autophagy in Alzheimer's disease pathogenesis is still under contention. One hypothesis is that aberrant autophagy induction results in an accumulation of autophagic vacuoles containing amyloid-beta and the components necessary for its generation, whereas other evidence points to impaired autophagic clearance or even an overall reduction in autophagic activity playing a role in Alzheimer's disease pathogenesis. Autophagy is linked with Alzheimer's disease. However, there is the uncertainty over the exact role and level of autophagic regulation in the pathogenic mechanism of Alzheimer's disease (9). Autophagy is a major intracellular degeneration pathway involved in the elimination and recycling of damaged organelles and long-lived proteins by lysosomes. Many of the pathological factors, which trigger neurodegenerative diseases, can perturb the autophagy activity, which is associated with misfolded protein aggregates accumulation in these disorders. Alzheimer's disease, the first neurodegenerative disorder between dementias, is characterized by two aggregating proteins, βamyloid peptide (plaques) and τ-protein (tangles). In Alzheimer's disease autophagosomes dynamically form along neurites within neuronal cells and in synapses but effective clearance of these structures needs retrograde transportation towards the neuronal soma where there is a major concentration of lysosomes. Maturation of autophago-lysosomes and their retrograde trafficking are perturbed in Alzheimer's disease, which causes a massive concentration of autophagy elements along degenerating neurites. Transportation system is disturbed along defected microtubules in Alzheimer's disease brains. τ-protein controls the stability of microtubules, however, phosphorylation of τ-protein or an increase in the total level of τ-protein can cause dysfunction of neuronal cells microtubules. Autophagy is developing in Alzheimer's disease brains because of ineffective degradation of autophagosomes, which hold amyloid precursor protein-rich organelles and secretases important for βamyloid peptides generation from amyloid precursor. The combination of raised autophagy induction and abnormal clearance of β-amyloid peptide-generating autophagic vacuoles creates circumstances helpful for β-amyloid peptide aggregation and

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accumulation in Alzheimer's disease. However, the key role of autophagy in Alzheimer's disease development is still under consideration today. One point of view suggests that abnormal autophagy induction causes a concentration of autophagic vacuoles rich in amyloid precursor protein, β-amyloid peptide and the elements crucial for its formation, whereas other hypothesis points to marred autophagic clearance or even decrease in autophagic effectiveness playing a role in maturation of Alzheimer's disease. In conclusion, the recent evidence links autophagy to Alzheimer's disease and autophagic regulation has the role in the development of full-blown Alzheimer's disease (10). Cannibalism of tumors is an old story for pathologists, but it remained a mystery for at least one century. Recent data highlighted tumor cannibalism as a key advantage in tumor malignancy, possibly involved in resistance of tumors to the specific immune reaction. However, new data suggests also that metastatic tumor cells may use this peculiar function to feed in conditions of low nutrient supply. This makes malignant cancer cells more similar to microorganisms, rather than to normal cells undergoing malignant transformation. In cytological or histological samples of human tumors it is common to detect cells with one or many vacuoles, possibly containing cells under degradation, that push the nucleus to the periphery giving it the shape of a crescent moon. The cannibal cells may feed on sibling tumor cells, but also of the lymphocytes that should kill them. Cannibal cells eat everything without distinguishing between the feeding materials, with a mechanism that mostly differ from typical phagocytosis. Despite such phenomenon is considered mainly nonselective, a molecular framework of factors that contribute to cannibalism has been described. This machinery includes the presence of an acidic environment that allows a continuous activation of specific lytic enzymes, such as cathepsin B. Cannibalism occurs in apparently well defined structures whose main actors are big caveolarlike vacuoles and a connection between caveolin-1 and the actin cytoskeleton through the actin-linker molecule ezrin. Each of the components of the cannibal framework may represent specific tumor targets for future new strategies against cancer (11). Aneuploidy is a hallmark of human cancers originating from abnormal mitoses. Many aneuploid cancer cells also have greaterthan-diploid DNA content, suggesting that polyploidy is a common precursor to aneuploidy during tumor progression. Polyploid cells can originate from cell fusion, endoreplication, and cytokinesis failure. Cell cannibalism by entosis, a form of cell engulfment involving live

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cells, also leads to polyploidy, as internalized cells disrupt cytokinesis of their engulfing cell hosts. By this mechanism, cannibalistic cell could promote tumor progression by leading to aneuploidy (12). The phenomenon of cell cannibalism, which generally refers to the engulfment of cells within other cells, was described in malignant tumors, but its biological significance is still largely unknown. In the present study, the occurrence, the in vivo relevance, and the underlying mechanisms of cannibalism in human melanoma were investigated. As first evidence, tumor cannibalism was clearly detectable in vivo in metastatic lesions of melanoma and often involved T cells, which could be found in a degraded state within tumor cells. Then, in vitro experiments confirmed that cannibalism of T cells was a property of metastatic melanoma cells but not of primary melanoma cells. In particular, morphologic analyses, including timelapse cinematography and electron microscopy, revealed a sequence of events, in which metastatic melanoma cells were able to engulf and digest live autologous melanoma-specific CD8(+) T cells. Importantly, this cannibalistic activity significantly increases metastatic melanoma cell survival, particularly under starvation condition, supporting the evidence that tumor cells may use the eating of live lymphocytes as a way to "feed" in condition of low nutrient supply. The mechanism underlying cannibalism involved a complex framework, including lysosomal protease cathepsin B activity, caveolae formation, and ezrin cytoskeleton integrity and function. In conclusion, this study shows that human metastatic melanoma cells may eat live T cells, which are instead programmed to kill them, suggesting a novel mechanism of tumor immune escape. Cannibalism may represent a sort of "feeding" activity aimed at sustaining survival and progression of malignant tumor cells in an unfavorable microenvironment (13). In this study, the cytological relevance of cannibalism as a dependable feature of malignancy in effusion and urine cytology was investigated. A total of 40 cases consisting of 10 each of malignant effusion, benign effusion, malignant urine samples, and benign urine samples were selected. These smears were assessed for the presence of cell cannibalism. The number of cannibalistic cells/100 tumor cells was counted. The cannibalistic cells were seen more commonly in malignant effusion cases (3.4/100 cells) compared with malignant urine cases (2/100 cells). There was not a single cannibalistic cell in benign conditions. The finding of an increased number of cannibalistic cell was highly significant in malignant versus benign samples. The present study highlights the significance of cannibalism in malignant

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urine and effusion cytology. In conclusion, cell cannibalism is a dependable cytological feature of malignancy (14). Cytological examination of effusion fluid is a relatively easy and quick method for the diagnosis of primary or secondary malignancy. The main aim of this study was to analyze the cytological significance of cell cannibalism in malignant effusion samples. A retrospective review of 100 cases of malignant effusion was conducted. These 100 cases included 50 cases of contiguous, local spread to pleural/ascitic fluid. The remaining 50 cases were of disseminated malignancy. Effusions due to hematolymphoid malignancies were excluded. Smears from these cases were assessed for the presence of cell cannibalism, tumor cell within a tumor cell. The cannibalistic cells were more common in effusions with disseminated malignancy (9 of 50 cases, 18%) compared with cases of contiguous, local spread (2 of 50 cases, 4%). This difference was statistically significant (p=0.025). The majority of the cases were of carcinoma lung (6/11). Cytomorphologically, histiocytes displaying phagocytosis can simulate tumor cells and need to be distinguished. In conclusion, presence of cell cannibalism in malignant effusions is more often an indicator of disseminated malignancy with secondary and higher tumor stage. Cannibalism may provide a reliable predictor of progression of tumor from primary to the metastatic site (15). The main objective of this study was to clarify whether the 3 parameters of cell clusters, cell cannibalism and nucleus-fragmented cells could improve diagnostic accuracy for G1UC. A total of 52 voided urine samples from 31 patients histologically diagnosed as having G1UC were reviewed. In addition, 10 voided urine samples from cases with G3UC and 30 voided urine samples from 25 patients with a histological diagnosis of chronic inflammation of the bladder were evaluated for comparison. Areas of tumor cells with cannibalism were measured. Cell cannibalism was evident in 12 of 31 G1UC cases (38.7%), significantly less often than with G3UC, but never identified in the control group. Mean areas of tumor cells featuring cannibalism were significantly smaller in G1 UC than in G3UC cases. Nucleusfragmented cells were also less frequent in G1UC than in G3UC, but more common than in the control group. In conclusion, cell cannibalism and nucleus-fragmented cells in voided urine with special attention to areas of tumor cell with cannibalism could be applied as a parameter to improve diagnostic accuracy for G1UC (16). Non-apoptotic forms of programmed cell death are targets for novel approaches in anticancer therapy. Indeed, cancer cells often present with mutations in the apoptotic machinery that result in

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resistance to most anticancer therapies and contribute to a relatively low response rate to therapies based on the use of pro-apoptotic strategies. (Macro-)autophagy can be a highly efficient mode of cell death induction by excessive self-digestion as demonstrated by experiments that studied the effect of radiation to induce autophagy cell death in apoptosis-deficient cells. Despite current controversies on the possible role of autophagy in the process of carcinogenesis and cancer progression by promoting cell survival, autophagy is a backup cell death mechanism, when other cell death mechanisms fail (17). Macroautophagy (autophagy) is a cellular catabolic process which can be described as a self-cannibalism. It serves as an essential protective response during conditions of ER stress through the bulk removal and degradation of unfolded proteins and damaged organelles; in particular, mitochondria (mitophagy) and ER (reticulophagy). Autophagy is genetically regulated and the autophagic machinery facilitates removal of damaged cell components and proteins; however, if the cell stress is acute or irreversible, cell death ensues. Despite these advances in the field, very little is known about how autophagy is initiated and how the autophagy machinery is transcriptionally regulated in response to ER stress. Some three dozen autophagy genes have been shown to be required for the correct assembly and function of the autophagic machinery; however; little is known about how these genes are regulated by cellular stress (18). Breast cancer cells often respond to an endocrine therapy by altering expression of specific estrogen-responsive genes and inducing autophagy, a cannibalistic lysosomal pathway. Autophagy eliminates damaged or other organelles, allowing the recovery of the energy stored in their macromolecules to attempt restoration of metabolic homeostasis. Induction of autophagy can result from activation of the unfolded protein response following metabolic stress, the final cell fate often being determined by the extent and duration of autophagy. One study builds upon this extensive knowledge, adding HSPB8 to the list of altered genes associated with endocrine resistance in breast cancer and describing the ability of HSPB8 to regulate autophagy and confer tamoxifen resistance (19,20). Assessment: the term self-cannibalism, or autophagy, was coined to describe the ability of the cells to cannibalize their own damaged organelles or proteins. The occurrence of autophagic processes can lead to the death of the cell. The xeno-cannibalism refers to the ability of certain cells, e.g. metastatic cells, to cannibalize their siblings as well as cells from the immune system. Metastatic tumor cells are able

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to engulf and digest living cells, including autologous lymphocytes that should kill them. Cannibalism has been described in various cancers such as, bladder cancer, breast cancer, lung cancer, etc, and is related to the aggressiveness of the malignancy. Autophagy plays an important role in cellular survival by resupplying cells with nutrients during starvation or by clearing misfolded proteins and damaged organelles and thereby preventing degenerative diseases. The autophagic process is recognized as a cellular death mechanism. Autophagy is a process in which subcellular membranes undergo dynamic morphological changes that lead to the degradation of cellular proteins and cytoplasmic organelles. It is a major intracellular degeneration pathway involved in the elimination and recycling of damaged organelles and long-lived proteins by lysosomes. Many of the pathological factors, which trigger neurodegenerative diseases, can perturb the autophagy activity, which is associated with misfolded protein aggregates accumulation in these disorders. The recent evidence links autophagy to Alzheimer's disease and autophagic regulation has the role in the development of full-blown Alzheimer's disease. The phenomenon of cell cannibalism, which generally refers to the engulfment of cells within other cells, was described in malignant tumors, but its biological significance is still largely unknown. Tumor cannibalism was clearly detectable in vivo in metastatic lesions of melanoma and often involved T cells, which could be found in a degraded state within tumor cells. Then, vitro experiments confirmed that cannibalism of T cells was a property of metastatic melanoma cells but not of primary melanoma cells. Cell cannibalism is a dependable cytological feature of malignant urine and effusion cytology. The cannibal cells may feed on sibling tumor cells, but also on the lymphocytes that should kill them. Cannibal cells eat everything without distinguishing between the feeding materials, with a mechanism that mostly differ from typical phagocytosis. References 1. Matarrese P, Ciarlo L, Tinari A, et al. Xeno-cannibalism as an exacerbation of self-cannibalism: a possible fruitful survival strategy for cancer cells. Curr Pharm Des. 2008;14(3):245-52. 2. Sharma N, Dey P. Cell cannibalism and cancer. Diagn Cytopathol. 2011;39(3):229-33.

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3. Poels J, Spasić MR, Callaerts P, Norga KK. An appetite for destruction: from self-eating to cell cannibalism as a neuronal survival strategy. Autophagy. 2012;8(9):1401-3. 4. Kondo Y, Kanzawa T, Sawaya R, Kondo S. The role of autophagy in cancer development and response to therapy. Nat Rev Cancer. 2005;5(9): 726-34. 5. Galluzzi L, Morselli E, Vicencio JM, et al. Life, death and burial: multifaceted impact of autophagy. Biochem Soc Trans. 2008;36(Pt 5):78690. 6. Vaccaro MI. Autophagy and pancreas disease. Pancreatology. 2008; 8(4-5):425-9. 7. Helin H, Mero M, Markkula H, Helin M. Pancreatic acinar ultrastructure in human acute pancreatitis. Virchows Arch A Pathol Anat Histol. 1980;387(3):259-70. 8. Ohmuraya M, Yamamura K. Autophagy and acute pancreatitis: a novel autophagy theory for trypsinogen activation. Autophagy. 2008; 4(8):1060-2. 9. Funderburk SF, Marcellino BK, Yue Z. Cell "self-eating" (autophagy) mechanism in Alzheimer's disease. Mt Sinai J Med. 2010;77(1):59-68. 10. Ułamek-Kozioł M, Furmaga-Jabłońska W, Januszewski S, et al. Neuronal autophagy: self-eating or self-cannibalism in Alzheimer's disease. Neurochem Res. 2013;38(9):1769-73. 11. Fais S. Cannibalism: a way to feed on metastatic tumors. Cancer Lett. 2007;258(2):155-64. 12. Krajcovic M, Overholtzer M. Mechanisms of ploidy increase in human cancers: a new role for cell cannibalism. Cancer Res. 2012; 72(7):1596-60. 13. Lugini L, Matarrese P, Tinari A, et al. Cannibalism of live lymphocytes by human metastatic but not primary melanoma cells. Cancer Res. 2006;66(7):3629-38. 14. Gupta K, Dey P. Cell cannibalism: diagnostic marker of malignancy. Diagn Cytopathol. 2003;28(2):86-7. 15. Bansal C, Tiwari V, Singh U, et al. Cell Cannibalism: a cytological study in effusion samples. J Cytol. 2011;28(2):57-60. 16. Hattori M, Nishino Y, Kakinuma H, et al. Cell cannibalism and nucleus-fragmented cells in voided urine: useful parameters for cytologic diagnosis of low-grade urothelial carcinoma. Acta Cytol. 2007;51(4):547-51. 17. Moretti L, Yang ES, Kim KW, Lu B. Autophagy signaling in cancer and its potential as novel target to improve anticancer therapy. Drug Resist Updat. 2007;10(4-5):135-43. 18. Deegan S, Saveljeva S, Gorman AM, Samali A. Stress-induced selfcannibalism: on the regulation of autophagy by endoplasmic reticulum stress. Cell Mol Life Sci. 2013;70(14):2425-41. 19. Gonzalez-Malerva L, Park J, Zou L, Hu Y, et al. High-throughput ectopic expression screen for tamoxifen resistance identifies an atypical

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kinase that blocks autophagy. Proc Natl Acad Sci U S A. 2011;108(5):205863. 20. Clarke R. Cannibalism, cell survival, and endocrine resistance in breast cancer. Breast Cancer Res. 2011;13(4):311.

TRANSMISSION OF DISEASES Although many pathogenic agents have the potential for transmission from prey to predator, diseases transmitted predominantly by cannibalism are rare because the epidemiological conditions necessary for its spread, especially group cannibalism, are rarely met in natural populations. However, such transmission occurs in animals such as chimpanzees where cannibalism occurs by socially organized groups of males (1-3). Cannibalistic transmission may therefore play an important role in the maintenance of blood-borne infections like simian immunodeficiency virus, or different types of hepatitis in these populations. Given the importance of social primates as disease reservoirs for humans (7), this possibility deserves further investigation (7). Cannibalism is uncommon in most species despite being taxonomically widespread. This rarity is surprising, because cannibalism can confer important nutritional and competitive advantages to the cannibal. A general, but untested, explanation for why cannibalism is rare is that cannibals may be especially likely to acquire pathogens from conspecifics, owing to greater genetic similarity among conspecifics and selection for host specificity and resistance to host immune defenses among pathogens. This hypothesis was tested by contrasting the fitness consequences of intra versus interspecific predation of diseased and non-diseased prey. Cannibalistic tiger salamander (Ambystoma tigrinum) larvae diseased conspecifics, healthy conspecifics, diseased heterospecifics (a sympatric congener, small-mouthed salamanders, A. texanum), or healthy heterospecifics were fed. Cannibals that ate diseased conspecifics were significantly less likely to survive to metamorphosis and grew significantly less than those that ate diseased heterospecifics, but none of the other groups differed. Tiger salamander larvae also preferentially preyed on heterospecifics when given a choice between healthy conspecifics and heterospecifics. These results suggest that pathogen transmission is an important cost of cannibalism and provide

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a general explanation for why cannibalism is infrequent in most species (8). Transovarial transmission was not detectable among Blastocrithidia triatomae-infected Triatoma infestans. Rather, B. triatomae was transmitted directly between triatomines by cannibalism and coprophagy. Cannibalism conditions that excluded coprophagy always resulted in an infection of Dipetalogaster maxima. The efficiency of transmission was not influenced by the blood source mice or chickens - fed to the infected donor bugs although chicken blood lyses the epimastigotes of the stomach population. Triatoma infestans was infected by coprophagy only if fed, not if unfed. Blastocrithidia triatomae in dry feces was taken up only if the feces were redissolved in fresh feces. Infections also appeared in groups of bugs fed on chickens previously used for feeding infected bugs (9). In a herd of approximately 1,000 hogs, evaluation of muscle specimens collected at various intervals during a 12-year period (1973 to 1985) indicated continuous transmission of Trichinella spiralis. The farm's rat population and the incidence of trichinosis in the rats was high during 1974, but diminished markedly by 1978. In January 1984, a longitudinal investigation, using tracer pigs, was performed to determine whether rodents and/or other wild animals were involved in transmission of Trichinella spiralis on this farm. Tracer pigs exposed to rodents and wild animals did not become infected with Trichinella spiralis. The rodent population on the farm was small and none of the rodents trapped and examined were infected. Hog cannibalism also was evaluated as a mode of Trichinella spiralis transmission. Results of the investigation indicate that hog cannibalism was the mode of transmission for trichinosis in the herd (10). Hamsters developed scrapie 100-160 days after eating either scrapie-infected hamsters or infected brain. The clinical signs and neuropathology of scrapie transmitted by cannibalism were identical to those observed after intracerebral or intraperitoneal inoculation of the agent. Oral transmission of scrapie appears to be extremely inefficient. Cannibalism requires a dose of the scrapie agent of approximately 10(9) times greater than that needed to produce the disease by intracerebral injection for comparable periods of incubation. These results provide compelling evidence for oral transmission of scrapie and may offer new insights into the spread of kuru by cannibalism among the Fore people and their neighbors. The extreme inefficiency of oral infection with scrapie might also have implications for understanding the sporadic occurrence and worldwide distribution of CJD (11).

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Cannibalism in the past has been a common practice in many human societies worldwide (12-16), in spite of some of the cases being exaggerated (17). The connection between cannibalism and disease has received considerable attention in the case of Kuru, a degenerative prion disease transmitted through cannibalism and necrophagy among the Fore people in Papua New Guinea (18,19). Cannibalism has been documented as a possible disease transmission route in several species, including humans. However, the dynamics resulting from this type of disease transmission are not well understood. Using a theoretical model, how cannibalism (i.e. killing and consumption of dead conspecifics) and intraspecific necrophagy (i.e. consumption of dead conspecifics) affect host-pathogen dynamics were explored. Group cannibalism, i.e. shared consumption of victims, is a necessary condition for disease spread by cannibalism in the absence of alternative transmission modes. Thus, endemic diseases transmitted predominantly by cannibalism are likely to be rare, except in social organisms that share conspecific prey. These results are consistent with a review of the literature showing that diseases transmitted by cannibalism are infrequent in animals, even though both cannibalism and trophic transmission are common (20). A literature review was carried out to determine instances in which cannibalism has been documented as a mechanism of disease transmission. Using Web of Science and Biological Abstracts, a search based on various combinations of the roots of the keywords „disease‟, „cannibalism‟, „intraspecific predation‟ or „necrophagy‟ was carried out. Then it has been noted whether a disease was recorded as being transmitted through cannibalism, if there were alternative transmission modes and, if so, whether cannibalism was the major mode of disease transmission in the system. This review revealed that cannibalism was documented as the predominant transmission mode in very few species, even though specific instances of cannibalistic transmission had been noted in many groups of organisms (mammals, reptiles, amphibians, fishes, insects and crustaceans), as well as in many pathogen types (including prions, viruses, bacteria and microsporidia). Indeed, cannibalism was implicated as the major transmission mode for only two cases: prion transmission in humans (19) and transmission of the protozoan Sarcocystis in lizards (21). Group cannibalism by multiple individuals on one victim is a necessary (albeit not always sufficient) precondition for disease spread through cannibalism. However, in the animal kingdom, cannibalism is generally a one-on-one interaction in which a larger and stronger individual kills and consumes a smaller and weaker conspecific (22).

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Under these conditions, cannibalism is likely to be an ineffective mode of disease transmission. The model is clearly simplified in that it assumes there is no cannibalism-independent population regulation, that infection itself does not influence the probability of being cannibalistic or being cannibalized and that there is no age structure. However, intuitively, it is clear that in one-on-one cannibalism, the R0 of a cannibalistically transmitted disease has to be less than unity unless the infected individuals have some compensatory advantage: an infected individual can only transmit the disease to one other individual and this victim is killed in the transmission process. This is somewhat analogous to the case of vertical (maternal) transmission, in which one female parent leaves on average only one offspring, such that the spread of vertically transmitted parasites only occurs under special circumstances (e.g. where the infected individual has a higher fitness than the uninfected). This is consistent with the finding that diseases spread through cannibalism are rare in natural populations, and cannibalistically transmitted parasites or pathogens usually have one or more alternative transmission modes. This rarity of cannibalistic transmission is even more notable because trophic transmission itself (disease transmission from prey to predator) is a commonplace in parasites with life cycles involving alternate hosts, especially in the digenean trematodes and cestodes (23), as well as in many nematodes. It would seem that within-species trophic transmission via cannibalism would evolve more easily than crossspecies trophic transmission because in the former case the host species and hence the internal environment for the pathogen would remain unchanged (24). Humans have also had the required social structure and social practices that promote disease spread by cannibalism. Historically, members of families or a village often shared captured individuals in ritualized meals (17,25,26), and the group size of individuals sharing one victim was often very large. Additionally, some human societies practiced cannibalism across groups and necrophagy within the group (25,27). This seems to have been the case for the Fore people of Papua New Guinea, in which both intraspecific necrophagy and cross-group cannibalism were common (28). Thus, in the case of Kuru, necrophagy could maintain and spread the disease within a village, while cross-group cannibalism could be the mechanism that promoted the disease spread on a larger meta-population scale. Cannibalism in humans has been a common and widespread practice that dates back at least to the Neanderthals (14,15). Several authors have argued that

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cannibalism has been a part of the natural ecology of human societies owing to the substantial nutritional gain (29). The Neanderthals were a Eurasian human species of the genus Homo that disappeared approximately 30,000 years ago. The cause or causes of their extinction continue to intrigue specialists and nonspecialists alike. Here a contributory role for TSEs is suggested. TSEs could have infected Neanderthal groups because of general cannibalistic activity and brain tissue consumption in particular. Further infection could then have taken place through continued cannibalistic activity or via shared used of infected stone tools. A modern human hunter-gatherer proxy has been developed and applied as a hypothetical model to the Neanderthals. This hypothesis suggests that the impact of TSEs on the Neanderthals could have been dramatic and have played a large part in contributing to the processes of Neanderthal extinction (30). Public health and agricultural policy attempts to keep BSE out of North America using infectious disease containment policies. Inconsistencies of the infectious disease model as it applies to the spongiform encephalopathies may result in failure of these policies. Review of historical, political and scientific literature determines the appropriate disease model of spongiform encephalopathy. Spongiform encephalopathy has always occurred sporadically in man and other animals. Hippocrates may have described it in goats and cattle. Transmission of spongiform encephalopathy between individuals is too uncommon for it to be usefully considered an infection. Spongiform encephalopathy is a somatic disorder whose dissemination within a host or transmission between individuals is more like cancer than infectious disease. Spongiform encephalopathy transmission within a species is facilitated in comparison to transmission between species so that cannibalism may amplify the prevalence of the disease. In conclusion, agricultural policy should be directed toward an absolute prohibition on occult cannibalism and away from surveillance, quarantine and slaughter, the principal measures of infectious disease containment used to control BSE (31). Assessment: pathogen transmission is an important cost of cannibalism. Diseases transmitted predominantly by cannibalism are rare because the epidemiological conditions necessary for its spread, especially group cannibalism, and are rarely met in natural populations. Such transmission occurs in animals such as chimpanzees where cannibalism occurs in socially organized groups of males. Cannibalistic transmission may play an important role in the

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maintenance of blood-borne infections like simian immunodeficiency virus or different types of hepatitis. The connection between cannibalism and disease has received considerable attention in the case of Kuru, a degenerative prion disease transmitted through cannibalism and necrophagy among the Fore people in Papua New Guinea. Cannibalism was documented as the predominant transmission mode in few species, even though specific instances of cannibalistic transmission had been noted in many groups of organisms (mammals, reptiles, amphibians, fishes, insects and crustaceans), as well as in many pathogen types (including prions, viruses, bacteria and microsporidia). Indeed, cannibalism was implicated as the major transmission mode for only two cases: prion transmission in humans and transmission of the protozoan Sarcocystis in lizards. References 1. Arcadi AC, Wrangham RW. Infanticide in chimpanzees: review of cases and a new within-group observation from the Kanyawara study group in Kibale National Park. Primates. 1999;40:337-51. 2. Mitani JC, Watts DP, Muller MN. Recent developments in the study of wild chimpanzee behavior. Evol. Anthropol. Issues, News, and Rev. 2002; 11:9-25. 3. Wilson ML, Wallauer WR, Pusey AE. New cases of intergroup violence among chimpanzees in Gombe National Park, Tanzania. Int. J. Primatol. 2004;25:523-49. 4. Santiago ML, Rodenburg CM, Kamenya S, et al. SIVcpz in wild chimpanzees. Science. 2002;295(5554):465. 5. Weiner A, Erickson AL, Kansopon J, et al. Persistent hepatitis-C virusinfection in a chimpanzee is associated with emergence of a cytotoxic Tlymphocyte escape variant. Proc. Natl Acad. Sci. USA. 1995;92:2755-9. 6. Birkenmeyer LG, Desai SM, Muerhoff AS, et al. Isolation of a GB virus-related genome from a chimpanzee. J. Med. Virol. 1998;56:44-51. 7. Wolfe ND, Escalante AA, Karesh WB, et al. Wild primate populations in emerging infectious disease research: the missing link? Emerg. Infect. Dis. 1998;4:149-58. 8. Pfennig DW, Ho SG, Hoffman EA. Pathogen transmission as a selective force against cannibalism. Anim Behav. 1998;55(5):1255-61. 9. Schaub GA, Böker CA, Jensen C, Reduth D. Cannibalism and coprophagy are modes of transmission of Blastocrithidia triatomae (Trypanosomatidae) between triatomines. J Protozool. 1989; 36(2):171-5. 10. Hanbury RD, Doby PB, Miller HO, Murrell KD. Trichinosis in a herd of swine: cannibalism as a major mode of transmission. J Am Vet Med Assoc. 1986;188(10):1155-9.

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11. Prusiner SB, Cochran SP, Alpers MP. Transmission of scrapie in hamsters. J Infect Dis. 1985;152(5):971-8. 12. Volhard E. Strecker und Schroeder Verlag; Stuttgart, Germany: Kannibalismus. Studien zur Kulturkunde. 1968. 13. White TD. Princeton NJ. Prehistoric cannibalism in Mancos. Princeton University Press. 1992, 5MTUMR-2346. 14. Defleur A, White T, Valensi P, et al. Neanderthal cannibalism at Moula-Guercy, Ardeche, France. Science. 1999;286:128-31. 15. Marlar RA, Leonard BL, Billman BR, et al. Biochemical evidence of cannibalism at a prehistoric Puebloan site in southwestern Colorado. Nature. 2000;407:74-8. 16. Lindenbaum S. Thinking about cannibalism. Annu Rev Anthropol. 2004;33:475-98. 17. Wendt A. Kannibalismus in Brasilien: Eine Analyse europaeischer Reiseberichte in Amerika-Darstellungen fuer die Zeit zwischen. Eurpaeische Hochschulschriften: Reihe 19, Volkskunde, Ethnologie. Verlag Peter Lang GmbH; Frankfurt, Germany. 1989, 1500 und 1654. 18. Collinge J, Whitfield J, McKintosh E, et al. Kuru in the 21st century an acquired human prion disease with very long incubation periods. Lancet. 2006;2006:2068-74. 19. Lindenbaum S. Kuru sorcery: disease and dangers in the New Guinean highlands. Mayfield Publishing Company; Mountain Few, CA. 1979. 20. Volker HWR, Antonovics J. Disease transmission by cannibalism: rare event or common occurrence? Proc Biol Sci. 2007;274(1614):1205-10. 21. Matuschka FR, Bannert B. Recognition of cyclic transmission of Sarcocystis stehlinii n-sp in the Gran Canarian giant lizard. J. Parasitol. 1989; 75:383-7. 22. Polis GA. The evolution and dynamics of intraspecific predation. Annu. Rev. Ecol. Syst. 1981;12:225-51. 23. Odening K. Conception and terminology of hosts and parasitology. Adv. Parasitol. 1976;14:1-93. 24. Anderson R.C. Nematode parasites of vertebrates. CABI publishing; Wallington, UK. 2000. 25. Volhard E. Kannibalismus. Studien zur Kulturkunde. Strecker und Schroeder Verlag. Stuttgart, Germany. 1968. 26. Sahlins M. Raw women, cooked men, and other “great things” of the Fiji Islands. In: Brown P, Tuzin D, editors. The ethnography of cannibalism. Society for Psychological Anthropology; Washington, DC. 1983, pp. 72-93. 27. Conklin B.A. Consuming grief. Compassionate cannibalism in an Amazonian society. University of Texas Press; Austin, TX. 2001. 28. Rumsey A. The white man as cannibal in the New Guinean highlands. In: Goldman LR, editor. The Anthropology of Cannibalism. Bergin and Garvey; Westport, CT. 1999, p. 168. 29. Darnstreich M, Moren G. Does New Guinean cannibalism have a nutritional value. Human Ecol. 1974;2:1-12.

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30. Underdown S. A potential role for transmissible spongiform encephalopathies in Neanderthal extinction. Med Hypotheses. 2008;71(1):47. 31. McAlister V. Sacred disease of our times: failure of the infectious disease model of spongiform encephalopathy. Clin Invest Med. 2005;28(3): 101-4.

PRION DISEASES Prions (proteinaceous infectious particles) are responsible for SSE in man and animals. Recent outbreak of bovine SSE (BSE), or mad cow disease in UK provoked concerns on its possible human hazards. A statement of the British Government in March 1996 upset the world, which was based on 10 cases of "new variant" form of CJD. Prion diseases in animals are often epizootic and may spread to different species through various routes including ingestion of contaminated meats. It is possible that mad cow disease causes a CJDlike malady in man through a dietary route. Human SSE include; 1] Classical CJD, which represents over 95% of the cases with SSE, dispersedly sporadic in occurrence in 1 per million a year, with monotonously rising incidence rates with age, of which the origin of the prion is totally unknown. 2] Gerstmann-Sträussler-Scheinker's disease which is inherited dominantly due to various genetic variants of the prion. 3] Transmitted cases of CJD occurring due to the prion introduced through contaminated objects either directly to the brain or its coverings, or indirectly to the brain through peripheral organs. 4] Kuru occurring among the Fore people in Papua New Guinea, probably due to the prion spread by a dietary route in cannibalism. New vCJD now totals 14 cases all observed in UK and is different from all of four, is encountered in a dispersedly sporadic manner, is far younger at onset than classical cases, showing kuru-plaques. Prions from vCJD have physical-chemical properties similar to those from BSE. Countermeasures in Japan against human hazards of BSE include, 1] agent controls by means of quarantine, and 2] host controls. So far, no vCJD-like case is observed in Japan. An emergent surveillance for CJD was introduced by the Ministry of Health and Welfare in July 1996. It obtained 2,637 answers from 4,027 departments of neurology, psychiatry, etc throughout the country and

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identified 766 cases including 51 familial cases, but no case with vCJD (1).

Prion diseases Prions represent a group of proteins with a unique capacity to fold into different conformations. One isoform is rich in beta-pleated sheets and can aggregate into amyloid that may be pathogenic. This abnormal form propagates itself by imposing its confirmation on the homologous normal host cell protein. Pathogenic prions cause lethal neurodegenerative diseases in humans and animals. These diseases are sometimes infectious and hence referred to as TSEs. The origin of this phenomenon is based on information transfer between homologous proteins, without the involvement of nucleic acid-encoded mechanisms. Historically, kuru and CJD were the first infectious prion diseases to be identified in man. It was their relationship to scrapie in sheep and experimental rodents that allowed an unraveling of the particular molecular mechanism underlying the disease process. Transmission between humans has been documented to have occurred in particular contexts, including ritual cannibalism, iatrogenic transmission because of pituitary gland-derived growth hormone or the use in neurosurgical procedures of dura mater from cadavers, and the temporary use of a prion-contaminated protein-rich feed for cows. The latter caused a major outbreak of BSE, which spread to man by human consumption of contaminated meat, causing approximately 200 cases of vCJD. All these epidemics now appear to be over because of measures taken to curtail further spread of prions. The mechanism of protein aggregation may apply to a wider range of diseases in and possibly outside the brain, some of which are relatively common such as Alzheimer's and Parkinson's diseases. Furthermore, it has become apparent that the phenomenon of prion

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aggregation may have a wider physiological importance, but a full understanding remains to be defined. It may involve maintaining neuronal functions and possibly contributing to the establishment of long-term memory (2). The transmissible spongiform encephalopathies or prion diseases represent a new group of diseases with unique clinical and neuropathological features, the transmission of which is both genetic and infectious. The responsible agent is unconventional and appears to be largely composed of a glycoprotein, the PrP. This is normally present on different cells. In prion diseases, it becomes converted to the pathogenic form PrPres which is resistant to proteinase and accumulates within the brain and this process is accompanied by the development of spongiform change, gliosis and neuronal loss. The human prion diseases include Kuru a progressive cerebellar degeneration with late dementia affecting Fore tribes in New-Guinea, now almost extinct, regarded as being related to cannibalism. CJD is the more frequent human prion disease. Its incidence is approximately one case per million per year. Four variants are recognized: sporadic, familial, iatrogenic, and the new variant. The latter represents a distinct clinico-pathological entity. It is now widely accepted that it is due to the same agent responsible for BSE in cattle. GerstmannSträussler-Scheinker disease is a very rare inherited disorder due to a number of different mutations in the PRP gene, characterized by abundant deposits of plaque PrPres in the cerebral grey matter. FFI is another inherited disorder due to a mutation at codon 178 of the PRP gene associated with methionine on codon 129 of the mutant allele. The main neuropathological change is neuronal loss in the thalamus with little or no spongiosis and usually no PrPres deposition. Following the emergence of new variant CJD in 1996, surveillance of all forms of prion diseases has now been actively introduced in many European nations in order to determine the true incidence and geographic distribution of these rare disorders in humans (3). Although prion diseases, such as CJD in humans and scrapie in sheep, have long been recognized, our understanding of their epidemiology and pathogenesis is still in its early stages. Progress is hampered by the lengthy incubation periods and the lack of effective ways of monitoring and characterizing these agents. Protease-resistant conformers of the PrP, known as the "scrapie form", are used as disease markers, and for taxonomic purposes, in correlation with clinical, pathological, and genetic data. In humans, prion diseases can arise sporadically or genetically, caused by mutations in the PrP gene, or as a foodborne infection, with the agent of BSE causing vCJD.

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Person-to-person spread of human prion disease has only been known to occur following cannibalism (kuru disease in Papua New Guinea) or through medical or surgical treatment (iatrogenic CJD). By contrast, scrapie in small ruminants and chronic wasting disease in cervids behave as infectious diseases within these species. Recently, however, so-called atypical forms of prion diseases have been discovered in sheep (atypical/Nor98 scrapie) and in cattle, BSE-H and BSE-L. These maladies resemble sporadic or genetic human prion diseases and might be their animal equivalents. This hypothesis also raises the significant public health question of possible epidemiological links between these diseases and their counterparts in humans (4).

Neurodegenerative diseases

Prion diseases are fatal neurodegenerative disorders in which an abnormal isoform of prion protein (PrPSc) accumulates in brain. Prion disease is either inherited as an autosomal dominant disorder with very high penetrance, or sporadic, where no epidemiological association with other human or animal prion diseases can be demonstrated or acquired, where disease results from contamination with PrPSc from another case of prion disease. No human case of prion disease is known to have been acquired from an animal but about 50 cases have been acquired from contaminated growth hormone or gonadotrophin prepared from human pituitaries, from human meningeal transplants or other neuro-surgical procedures. Several thousand cases of the prion disease, kuru, occurred in Papua New Guinea, mainly during the first half of this century. This epidemic probably started with a sporadic case of CJD but was transmitted through funerary practices which are thought to have involved endo-cannibalism. Inherited cases of prion disease are associated with mutations in the PrP gene. There are at least five point

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mutations and a further five mutations involving 5 to 9 extra repeats of the five octapeptide repeat sequence in the PrP gene, all of which may lead to one of the forms of human prion disease i.e. CJD, GerstmannSträussler syndrome and atypical prion dementia. In addition, there is a common polymorphism at codon 129 of the PrP gene. This may affect age at onset and duration of illness in inherited and sporadic cases (5). Human prion diseases may be sporadic, genetic, or acquired. The human Prion protein gene is located to chromosome 20 (20p12-ter). Mutations and polymorphisms in the Prion protein gene are associated with prion disease. Genetic prion diseases are inherited in an autosomal dominant trait, and examination of the penetrance is restricted to mutation E200K (59-89%). Mutations can be substitutions or insertions. Genetic prion diseases are classified according to the clinicopathological phenotype and comprise genetic CJD, Gerstmann-Sträussler-Scheinker disease and FFI. Base pair insertions may resemble CJD or Gerstmann-Sträussler-Scheinker disease phenotypes, however, their unique clinicopathological presentations are also emphasized. Among the polymorphisms of the Prion protein gene, the one at codon 129 is the most important, where methionine or valine may be encoded. This polymorphism is known to influence the phenotype of disease forms. Molecular classification of sporadic CJD depends on the codon 129 polymorphisms in addition to the Western blot pattern of the protease resistant prion protein. According to this, at least 6 well-characterized forms of sporadic CJD are known. Influence of other genes were also investigated. Contrasting results are reported regarding the role of apolipoprotein E allele epsilon4, but presence of allele epsilon2 seems to influence the prognosis. Polymorphisms in the doppel gene or ADAM10 could not be clearly associated with CJD. Polymorphisms in the upstream and intronic regulatory region of the Prion protein gene may be a risk factor for CJD. The Prion protein gene codon 129 polymorphism was examined in non-prion diseases. Some studies suggest that this polymorphism may have influence on the cognitive decline and early onset Alzheimer's disease (6). References 1. Kondo K. The prion diseases. Hokkaido Igaku Zasshi. 1997;72(1):2736. 2. Norrby E. Prions and protein-folding diseases. J Intern Med. 2011;270 (1):1-14.

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3. Keohane C. The human prion diseases. A review with special emphasis on new variant CJD and comments on surveillance. Clin Exp Pathol. 1999; 47(3-4):125-32. 4. Tranulis MA, Benestad SL, Baron T, Kretzschmar H. Atypical prion diseases in humans and animals. Top Curr Chem. 2011;305:23-50. 5. Ridley RM, Baker HF. Genetics of human prion disease. Dev Biol Stand. 1993;80:15-23. 6. Kovács GG. Genetic background of human prion diseases. Ideggyogy Sz. 2007;60(11-12):438-46.

KURU Kuru disease is linked with the name of Carleton Gajdusek who was the first to show that this human neurodegenerative disease can be transmitted to chimpanzees and subsequently classified as a TSE, or slow unconventional virus disease, and was reported to Western world in 1957 (1-2), and in 1975 (3). "Kuru" in the Fore language in Papua New Guinea means to shiver from fever and cold. The disease spreads through ritualistic cannibalism and is an invariably fatal cerebellar ataxia accompanied by tremor, choreiform and athetoid movements. Neuropathologically, kuru is characterized by the presence of amyloid "kuru" plaques (4). A well-known case of mortuary cannibalism is that of the Fore tribe in New Guinea which resulted in the spread of the prion disease kuru. Although the Fore's mortuary cannibalism was well documented, the practice had ceased before the cause of the disease was recognized. However, some scholars argue that although postmortem dismemberment was the practice during funeral rites, cannibalism was not. Marvin Harris theorizes that it happened during a famine period coincident with the arrival of Europeans and was rationalized as a religious rite (5). Kuru is a subacute neurodegenerative disease presenting with limb ataxia, dysarthria, and a shivering tremor. The disease progress to complete motor and mental incapacity and death within 6 to 24 months. Neuropathologically, a typical pattern of neuronal loss, astrocytic and microglial proliferation, characteristic "kuru-type" amyloid plaques, and PrP deposits in the cerebral cortex and cerebellum are observed. Kuru is the prototype of a group of human TSEs, or "prion" diseases, that include hereditary, sporadic and infectious forms. The latest member of this group, the vCJD, linked to

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transmission of BSE to humans, shows features similar to kuru. Kuru has emerged at the beginning of the 1900s in a small indigenous population of New-Guinean Eastern Highlands, reached epidemic proportions in the mid-1950s and disappeared progressively in the latter half of the century to complete absence at the end of the 1990s. Early studies made infection, the first etiologic assumption, seem unlikely and led to a hypothesis that kuru might be a genetically determined or genetically mediated illness. After transmissibility of kuru had been discovered and all major epidemiologic phenomena adequately explained by the spread of an infectious agent with long incubation period through the practice of cannibalism, the pattern of occurrence still continued to suggest a role for genetic predisposition. Recent studies indicate that individuals homozygous for methionine at a polymorphic position 129 of the PrP were preferentially affected during the kuru epidemic. The carriers of the alternative 129Met/Val and 129Val/Val genotypes had a longer incubation period and thus developed disease at a later age and at a later stage of the epidemic. Observations made during the kuru epidemic are helpful in the understanding of the current vCJD outbreak, and vice versa clinical and experimental data accumulated in studies of other TSE disorders contributed to better understanding of the documented kuru phenomena (6). Kuru is a fatal TSE restricted to the Fore people and their neighbors in a remote region of the Eastern Highlands of Papua New Guinea. When first investigated in 1957 it was present in epidemic proportions, with approximately 1,000 deaths in the first 5 years, 1957-1961. The changing epidemiological patterns and other significant findings such as the transmissibility of kuru are described in their historical progression. Monitoring the progress of the epidemic has been carried out by epidemiological surveillance in the field for 50 years. From its peak, the number of deaths from kuru declined to 2 in the last 5 years, indicating that the epidemic is approaching its end. The mode of transmission of the prion agent of kuru was the local mortuary practice of transumption. The prohibition of this practice in the 1950s led to the decline in the epidemic, which prolonged into the present century by incubation periods that may exceed 50 years. Currently, the epidemiological surveillance is maintained and studies on human genetics and the past mortuary practices are conducted in the kuru-affected region and in communities beyond it (7).

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The occurrence of a disease such as Kuru in humans is most probably the result of the frequent occurrence of group cannibalism and/or group necrophagy. Additionally, the persistence of Kuru into modern times in part is explained by the pathogenic agent's resilience to cooking, although, in many of the cannibalistic rituals, raw human flesh was often consumed (8). Because cannibalism is no longer a regular feature of human populations, it is not possible to assess the degree to which other diseases may have had this transmission mode. However, the transmission and establishment of tapeworms in humans may have been aided by cannibalism, and it is quite conceivable that a number of blood-borne infections may have been regularly transmitted in the same way (9). The epidemics of kuru have been transmitted among the Fore by ritual consumption of infected organs from deceased relatives. As government patrol officers suppressed cannibalism during the 1950s, most transmission had ceased by 1957, when the kuru research program first commenced. As predicted in the 1960s, the epidemic has waned, with progressive ageing of kuru-affected cohorts over the years to 2007. The few cases seen in the twenty-first century, with the longest incubation periods, were almost certainly exposed as children prior to 1960. Although the research program had almost no role in ending the kuru epidemic, it did provide important knowledge that was to help the wider world in controlling the later epidemics of iatrogenic and vCJD and BSE (10). Kuru was the first human TSE or prion disease identified, occurring in the Fore linguistic group of Papua New Guinea. Kuru was a uniformly fatal cerebellar ataxic syndrome, usually followed by choreiform and athetoid movements. Kuru imposed a strong balancing selection on the Fore population, with individuals homozygous for the 129 Met allele of the gene (Prion protein gene) encoding for PrP being

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the most susceptible. The decline in the incidence of kuru in the Fore has been attributed to the exhaustion of the susceptible genotype and ultimately by discontinuation of exposure via cannibalism. Neuropathologically, kuru-affected brains were characterized by widespread degeneration of neurons, astroglial and microglial proliferation, and the presence of amyloid plaques. These early findings have been confirmed and extended by recent immunohistochemical studies for the detection of the TSE-specific PrP. Confocal laser microscopy showed the concentration of glial fibrillary acidic protein-positive astrocytic processes at the plaque periphery. The fine structure of plaques corresponds to that described earlier by light microscopy. The successful experimental transmission of kuru led to the awareness of its similarity to CJD and GerstmannSträussler-Scheinker disease forming a background against which the recent epidemics of iatrogenic and variant CJD could be studied (11). Kuru provides the principal experience of epidemic human prion disease. Its incidence has steadily fallen after the abrupt cessation of its route of transmission (endocannibalism) in Papua New Guinea in the 1950s. The onset of vCJD, and the unknown prevalence of infection after the extensive dietary exposure to BSE prions in the UK, has led to renewed interest in kuru. Possible incubation periods, pathogenesis, and genetic susceptibility factors in kuru patients in Papua New Guinea were investigated. Active kuru surveillance in 1996 with an expanded field team was strengthened to investigate all suspected patients. Detailed histories of residence and exposure to mortuary feasts were obtained together with serial neurological examination, if possible. Eleven patients with kuru from July 1996, to June 2004, all living in the South Fore were identified. All patients were born before the cessation of cannibalism in the late 1950s. The minimum estimated incubation periods ranged from 34 to 41 years. However, likely incubation periods in men ranged from 39 to 56 years and could have been up to 7 years longer. Prion protein gene analysis showed that most patients with kuru were heterozygous at polymorphic codon 129, a genotype associated with extended incubation periods and resistance to prion disease. Incubation periods of infection with human prions can exceed 50 years. In human infection with BSE prions, species-barrier effects, which are characteristic of cross-species transmission, would be expected to further increase the mean and range of incubation periods, compared with recycling of prions within species. These data should inform attempts to model variant CJD epidemiology (12).

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Kuru is so far the principal human epidemic prion disease. While its incidence has steadily declined since the cessation of its route of transmission, endocannibalism, in Papua New Guinea in the 1950s, the arrival of vCJD, also thought to be transmitted by dietary prion exposure, has given kuru a new global relevance. Suspected cases of kuru from July 1996 to June 2004 were investigated and 11 kuru patients were identified. There were four females and seven males, with an age range of 46-63 years at the onset of disease, in marked contrast to the age and sex distribution when kuru was first investigated 50 years ago. Detailed histories of residence and exposure to mortuary feasts were obtained and serial neurological examination and genetic studies were performed where possible. All patients were born a significant period before the mortuary practice of transumption ceased and their estimated incubation periods in some cases exceeded 50 years. The principal clinical features of kuru in the studied patients showed the same progressive cerebellar syndrome. Two patients showed marked cognitive impairment well before preterminal stages, in contrast to earlier clinical descriptions. In these patients, the mean clinical duration of 17 months was longer than the overall average in kuru but similar to that previously reported for the same age group, and this may relate to the effects of both patient age and Prion protein gene codon 129 genotype. Importantly, no evidence for lymphoreticular colonization with prions, seen uniformly in vCJD, was observed in a patient with kuru at tonsil biopsy (13). Kuru is a devastating epidemic prion disease that affected a highly restricted geographic area of the Papua New Guinea highlands; at its peak, it predominantly affected adult women and children of both sexes. Its incidence has steadily declined since the cessation of its route of transmission, endocannibalism. Genetic and selected clinical and genealogic assessments of more than 3,000 persons from Eastern Highland populations, including 709 who participated in cannibalistic mortuary feasts, 152 of whom subsequently died of kuru, were performed. Persons who were exposed to kuru and survived the epidemic in Papua New Guinea were predominantly heterozygotes at the known resistance factor at codon 129 of the Prion protein gene. A novel Prion protein gene variant- G127V was found exclusively in people who lived in the region in which kuru was prevalent and was present in half of the otherwise susceptible women from the region of highest exposure who were homozygous for methionine at Prion protein gene codon 129. Although this allele is common in the area with the highest incidence of kuru, it is not found in patients with kuru and in unexposed population groups worldwide. Genealogic analysis

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reveals a significantly lower incidence of kuru in pedigrees that harbor the protective allele than in geographically matched control families. In conclusion, the 127V polymorphism is an acquired prion disease resistance factor selected during the kuru epidemic, rather than a pathogenic mutation that could have triggered the kuru epidemic. Variants at codons 127 and 129 of Prion protein gene demonstrate the population genetic response to an epidemic of prion disease and represent a powerful episode of recent selection in humans (14). The acquired prion disease kuru was restricted to the Fore and neighboring linguistic groups of the Papua New Guinea highlands and largely affected children and adult women. Oral history documents the onset of the epidemic in the early twentieth century, followed by a peak in the mid-twentieth century and subsequently a welldocumented decline in frequency. In the context of these strong associations (gender, region and time), the genetic factors associated with susceptibility and resistance to kuru were considered. Heterozygosity at codon 129 of the human prion protein gene (PRNP) is known to confer relative resistance to both sporadic and acquired prion diseases. In kuru, heterozygosity is associated with older patients and longer incubation times. Elderly survivors of the kuru epidemic, who had multiple exposures at mortuary feasts, are predominantly Prion protein gene (PRNP) codon 129 heterozygotes and this group show marked Hardy-Weinberg disequilibrium. The deviation from Hardy-Weinberg equilibrium is most marked in elderly women, but is also significant in a slightly younger cohort of men, consistent with their exposure to kuru as boys. Young Fore and the elderly from populations with no history of kuru show HardyWeinberg equilibrium. An increasing cline in 129V allele frequency centers on the kuru region, consistent with the effect of selection in elevating the frequency of resistant genotypes in the exposed population. The genetic data are thus strikingly correlated with exposure. Considering the strong coding sequence conservation of primate prion protein genes, the number of global coding polymorphisms in man is surprising. By intronic resequencing in a European population, haplotype diversity at PRNP comprises two major and divergent clades associated with 129M and 129V. Kuru may have imposed the strongest episode of recent human balancing selection, which may not have been an isolated episode in human history (15).

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References 1. Gajdusek DC, Zigas V. Degenerative disease of the central nervous system in New Guinea; the endemic occurrence of kuru in the native population. N Engl J Med. 1957;257(20):974-8. 2. Zigas V, Gajdusek DC. Kuru: clinical study of a new syndrome resembling paralysis agitans in natives of the Eastern Highlands of Australian New Guinea. Med J Aust. 1957;44(21):745-54. 3. Gajdusek DC, Alpers MP. Recent data on the properties of the viruses of kuru and transmissible virus dementias. P N G Med J. 1975;18(4):207-13. 4. Liberski PP, Sikorska B, Brown P. Kuru: the first prion disease. Adv Exp Med Biol. 2012;724:143-53. 5. Lindenbaum S. Understanding kuru: the contribution of anthropology and medicine. Philos. Trans R Soc. Lond B Biol Sci. 2008;363(1510): 371520. 6. Goldfarb LG, Cervenakova L, Gajdusek DC. Genetic studies in relation to kuru: an overview. Curr Mol Med. 2004;4(4):375-84. 7. Alpers MP. Review. The epidemiology of kuru: monitoring the epidemic from its peak to its end. Philos Trans R Soc Lond B Biol Sci. 2008; 363(1510):3707-13. 8. Lindenbaum S. Kuru sorcery: disease and dangers in the New Guinean highlands. Mayfield Publishing Company; Mountain Few, CA. 1979. 9. Hoberg EP, Alkire NL, De Queiroz A, Jones A. Out of Africa: origins of the taenia tapeworms in humans. Proc. R. Soc. B. 2001;268:781-7. 10. Mathews JD. Review. The changing face of kuru: a personal perspective. Philos Trans R Soc Lond B Biol Sci. 2008;363(1510):3679-84. 11. Liberski PP, Sikorska B, Lindenbaum S, et al. Kuru: genes, cannibals and neuropathology. J Neuropathol Exp Neurol. 2012;71(2):92-103. 12. Collinge J, Whitfield J, McKintosh E, et al. Kuru in the 21st century an acquired human prion disease with very long incubation periods. Lancet. 2006;367(9528):2068-74. 13. Collinge J, Whitfield J, McKintosh E, et al. A clinical study of kuru patients with long incubation periods at the end of the epidemic in Papua New Guinea. Philos Trans R Soc Lond B Biol Sci. 2008;363(1510):3725-39. 14. Mead S, Whitfield J, Poulter M, et al. A novel protective prion protein variant that colocalizes with kuru exposure. N Engl J Med. 2009;361(21): 2056-65. 15. Mead S, Whitfield J, Poulter M, et al. Genetic susceptibility, evolution and the kuru epidemic. Philos Trans R Soc Lond B Biol Sci. 2008; 363(1510):3741-6.

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CREUTZFELD-JACOB DISEASE CJD is one of several related disorders collectively called prion diseases. These disorders affect man and animals and caused by the abnormal configuration of a naturally occurring protein, PrP(c). By mechanisms still not well understood, this natural protein is converted into a pathologic variant, PrP(sc). The disease is 'acquired' spontaneously perhaps by posttranslational conversion of a PrP(c) into a PrP(sc) population. This sporadic form of CJD has been reported worldwide with a frequency of 1/million. Other modes of acquisition include the following: ingestion of brain tissue from deceased victims through ritual cannibalism at burial ceremonies formerly (and no longer) practiced by New Guinea Highlanders; iatrogenically, through corneal transplants from infected donors, inoculation of human growth hormone and gonadotropin prepared from infected human pituitary glands; from inadequately sterilized depth electrodes introduced neurosurgically into the brain during workups of patients with epilepsy, and applications of infected dura mater in neurosurgical procedures. Most recently, an infected bovine source (BSE) has been implicated and produces a new vCJD. Clusters of CJD in families in some populations have been recognized which are inherited in Mendelian fashion. These clusters are related to mutations of the PRNP gene in specific codons (e.g. codon 200). Homozygosity for these mutations increases the chances of manifesting the disease. Other potential methods of acquisition, such as by blood transfusion, surgical sutures, tonometers, consumption of hog brain or other organs and tissue, remain unproven (1).

Creutzfeldt-Jakob disease

CJD is a rapidly progressive dementia syndrome which is probably caused by prions. The annual incidence of this disease is 1/1,000,000.

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Most cases are sporadic in type, although 10-15% are familial. The total incidence of CJD has not changed following the epidemic due to a new variant (nv-CJD); however, this has led to greater awareness of the subject. The main objective of this study was to review current scientific knowledge of CJD. A search was made for relevant literature using MEDLINE. The criteria proposed for diagnosis of CJD include the presence of progressive dementia and at least two of the following characteristics: 1. Myoclonias, 2. Cortical blindness, 3. Pyramidal, extrapyramidal or cerebellar signs, 4. Akinetic mutism, or 5 Abnormal EEG. Laboratory and neuroimaging investigations may also help in diagnosis of CJD, although neuropathological confirmation is necessary for definite diagnosis. Two promising methods of pre mortem diagnosis of CJD are determination of 14-3-3 protein in the CSF, in the case of sporadic CJD, and biopsy of the palatine tonsil in the case of nv-CJD. The physiopathology of CJD seems to be centered on the proteins forming prions, which are glycoproteins found in the plasmatic membrane and are very often expressed in the neurons, particularly at neuromuscular junctions and synapses. The pathological form resists proteolytic degradation, so that they accumulate in the CNS. The precise neurotoxic mechanism of these proteins is still not clear. In conclusion, there is still no treatment for CJD. Further studies of the physiopathological mechanisms of prion diseases may help in the development of treatment to delay the progress of this disease (2). vCJD is an acquired prion disease causally related to BSE that has occurred predominantly in young adults. All clinical cases studied have been methionine homozygotes at codon 129 of the prion protein gene (PRNP) with distinctive neuropathological findings and molecular strain type (PrP(Sc) type 4). Modeling studies in transgenic mice suggest that other prion protein gene (PRNP) genotypes will also be susceptible to infection with BSE prions but may develop distinctive phenotypes. The main objective of this study was to describe the histopathological and molecular investigation in a young British woman with atypical sporadic CJD and valine homozygosity at prion protein gene (PRNP) codon 129. Case report, autopsy, and molecular analysis were conducted at specialist neurology referral center, together with the laboratory services of the Medical Research Council Prion Unit. Autopsy findings were atypical of sporadic CJD, with marked gray and white matter degeneration and widespread PrP deposition. Lymphoreticular tissue was not available for analysis. Molecular analysis of PrP(Sc) (the scrapie isoform of PrP) from cerebellar tissue demonstrated a novel PrP(Sc) type similar to that

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seen in vCJD (PrP(Sc) type 4). However, this could be distinguished from the typical vCJD pattern by an altered protease cleavage site in the presence of the metal ion chelator EDTA. In conclusion, further studies will be required to characterize the prion strain seen in this patient and to investigate its etiologic relationship with BSE. This case illustrates the importance of molecular analysis of prion disease, including the use of EDTA to investigate the metal dependence of protease cleavage patterns of PrP(Sc) (3).

Variant Creutzfeldt-Jakob disease

The MRI scans of 103 patients with CJD referred to the National Prion Clinic since 2007 and the presence of CJD-associated changes were reviewed, comparing these findings with the formal report from the referring centre and the types of sequence performed were reviewed. In sCJD, CJD-associated MRI changes in 83 of 91 cases (91% sensitivity) were found. However, the referring centers documented CJD-associated MRI changes in 43 of the sCJD cases (47% sensitivity). The most common region not documented by referring centers was the cortex (23 of 68 sCJD cases), but there was a statistically significant discrepancy in all regions (p