Kommentiertes Referat
Controlling defecation: to be (predator) or not to be (prey), that is the question… Räuber) oder Nichtsein (Beute), das ist die Frage … Z Gastroenterol 2015; 53: 460–462 G. Bassotti, S. Müller-Lissner He who can no longer pause and stand rapt in awe is as good as dead; his eyes are closed. To raise new questions, new possibilities, to regard old problems from a new angle, requires creative imagination and marks real advance in science. Logic will get you from A to B. Imagination will take you everywhere. (Albert Einstein)
Introduction The process of defecation in human beings involves a series of physiological events that, after the digestive and absorptive processes occurring in the upper part of the digestive tract have been completed, cause the residual more or less compacted contents (fecal matter) to be transported through the colon and eventually expelled by relaxation of the anal sphincter [1]. Although defecation occurs almost daily in normal subjects, the very act is under strict cultural taboos, at least in most societies, and it is rarely if ever thought or spoken of, apart from practical jokes and medical evaluation. However, it is worth noting that defecation is subject to a strict and voluntary central control [2, 3] that, depending on the cultural environment, allows the individual to decide if, when and where it is socially convenient to evacuate [4]. Moreover, the control of defecation is not an innate process, since it is acquired in infancy by toilet training [5], usually in the family environment; recognition of the need to defecate is usually well established by age four, and the ability to hold back defecation improves with age [6]. Thus, controlling defecation is an important aspect of the human being’s life and, provided all the appropriate reflexes are intact, bowel motions may be (within reasonable limits) kept under control until the subject decides it is appropriate to evacuate his/her gut contents.
Since it is likely that our ancestors (hominins) were subject to the same ecological pressures as other primates, particularly predation risk and competition for resources [7], in this article it will be hypothesized why men control defecation, and the possible evolutionary role of controlling defecation as a method for hominins to increase foraging strategies, but also as a tool to avoid predation.
To be a predator… Predation is an important, sometimes essential method to assure survival. However, predation implies that the predator’s abilities overwhelm that of prey, and this may be difficult when the prey is highly mobile. One of the main points, besides agility and force, is therefore that the predator must be able to reach the prey or to approach it in order to use tools. For this purpose, camouflage and speed are of paramount importance, even though other factors are equally important. In fact, we could assume that predation for hominins would hardly have involved camouflage (scarcely effective on moving targets [8]) and speed, whereas stealth probably played a major role for hunting. Thus, predators have to be silent and, more importantly, to offer little or no clues (in particular, olfactory traces) of their approach. This is particularly important for mammals, whose nose is a powerful chemosensor, capable of detecting and distinguishing a myriad of chemicals [9]. Therefore, it is not surprising that olfactory hunting is very common in predators [10] since many of them must locate resources in environments in which sensory signals are noisy and without directional information [11]. The scents of mammals are complex blends of natural products that reveal a wealth of individual information [12]; under this light, fecal scents are particularly effective, as revealed by the analysis of fecal odorgrams [13, 14]. Indeed, selective delimitation of territorial areas with fecal (and/or urinary) amounts is a method employed by several mammal species, primates and non-primates, to mark their territory and suggests that control of defecation and targeted deployment of excrements is widely present in animals [15 – 17]. In addition, there is evidence that several species (including some primates) use latrines, i. e., repeated specific defecation/urination sites for territory demarcation, resource defense, signaling of repro-
ductive state, social bonding, and mate defense [18]. However, concerning predator-prey relationships, the importance of fecal odors may have double-faced implications. On the one hand, in fact, the predator follows the scents of the prey, and the fecal ones are outstanding under this light; on the other hand, the reverse is equally important, in that preys are usually able to detect the scents of predators, and those related to feces are extremely useful to avoid predation (see below).
…or not to be prey There is presently evidence that hominins may have been the prey of several species of large carnivores [19 – 22], as also shown by the recent demonstration of transmission of Helicobacter spp from early humans to preying felids [23]. Thus, as it happens in other animal species [24], our ancestors had to develop a series of adaptive evolutionary strategies to avoid predation, strategies that included morphological defenses as well as behavioral traits. Indeed, experimental animal studies suggest that large-brained prey are likely to be more effective at evading predators because they can effectively alter their behavioral responses to specific predator encounters [25]. Predator recognition and avoidance are important behaviors in allowing prey to mitigate the impacts of their predators; for instance, some authors hypothesized that the development of erect stature in hominins, regardless of locomotion proficiency, actually enhanced both active and passive defense against predators [26]. However, several other and effective mechanisms may play a role in this context. For instance, many prey species try to avoid predation also by learning the predators’ odors [27, 28]. Data derived from experience in animal studies show that several prey species avoid the odors of their predators (including that of feces) [29, 30], and that compounds isolated from the feces elicit a series of fear-related responses, including avoidance and risk assessment behavior [31, 32].
…the control of defecation is the question On the basis of the above considerations, it might be hypothesized that learning to control defecation, before becoming a so-
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
Downloaded by: Collections and Technical Services Department. Copyrighted material.
460
cial necessity, may have had some evolutionary significance for hominins as a means both not to advertise their presence to a possible prey and to mask an odoriferous trail to a possible predator [33]. This may also be inferred by the fact that other primate species are able to control defecation and deposit their feces in specific sites such as latrines [18]. Of interest, the development of erect stature might have helped the control upon defecation by our ancestors by means of specific attachments and functions of the structure of the pelvic floor [34, 35]. In fact, humans are thought to be the first purely bipedal mammals [36], and the erect position contributes significantly to the maintenance of fecal continence by sharpening the anorectal angle [37]; species that live at least part of their life in trees have fewer demands on their pelvic structures for support than bipedal terrestrial animals that maintain a vertical posture for longer periods of time [38]. For instance the gorilla, that only occasionally is bipedal, is known to have poor control of defecation, due to a different conformation of the pelvic floor [39]. There is no doubt that among all the living creatures humans have achieved a lot in evolutionary terms. The main propulsive force leading to the differentiation toward the other species was the emergence of human cognition, and the researchers are still actively investigating this phenomenon [40]. The process of creative thinking, the ability to develop new tools, the never-ending struggle toward knowledge allowed our ancestors to graduate from an almost animal state to the present one (although someone would argue that this had not necessarily been for the better…), allowing human hunter-gatherer bands to function as unique and highly competitive predatory organisms [41]. In this respect, among the various strategies to improve both foraging and survival, it might be hypothesized that the control of defecation (even though diversified in the various species) may have played a role, allowing a minor detection by the possible preys and a greater likelihood of escaping predation by not leaving around easily detectable spoors.
Kommentierte Referate können von jedem interessierten Kollegen verfasst werden. Sie sollten zuvor beim Koordinator angemeldet werden. Die formale Gestaltung orientiert sich an den publizierten Referaten (Kurzzitate im Text!). Einreichung nicht über die Online-Manuskripteinreichung des Thieme Verlags, sondern via E-mail an den Koordinator der Kommentierten Referate. Prof. Dr. Stefan Müller-Lissner Park-Klinik Weißensee Schönstraße 80, 13086 Berlin Tel. + 49 30 96 28 36 00, Fax 36 05, [email protected]
References 01 Bassotti G, de Roberto G, Castellani D et al. Normal aspects of colorectal motility and abnormalities in slow transit constipation. World J Gastroenterol 2005; 11: 2691 – 2696 02 Brading AF, Ramalingam T. Mechanisms controlling normal defecation and the potential effects of spinal cord injury. Prog Brain Res 2006; 152: 345 – 358 03 Bittorf B, Ringler R, Forster C et al. Cerebral representation of the anorectum using functional magnetic resonance imaging. Br J Surg 2006; 93: 1251 – 1257 04 Bajwa A, Emmanuel A. The physiology of continence and evacuation. Best Pract Res Clin Gastroenterol 2009; 23: 477 – 485 05 Kiddoo DA. Toilet training children: when to start and how to train. CMAJ 2012; 184: 511 – 512 06 Kyrklund K, Koivusalo A, Rintala RJ et al. Evaluation of bowel function and fecal continence in 594 Finnish individuals aged 4 to 26 years. Dis Colon Rectum 2012; 55: 671 – 676 07 Rose L, Marshall F. Meat eating, hominid sociality, and home bases revisited. Curr Anthropol 1996; 37: 307 – 338 08 Hall JR, Cuthill IC, Baddeley R et al. Camouflage, detection and identification of moving targets. Proc Biol Sci 2013; 280 20130064 09 Liberles SD. Trace amine-associated receptors are olfactory receptors in vertebrates. Ann N Y Acad Sci 2009; 1170: 168 – 172 10 Hughes NK, Price CJ, Banks PB. Predators are attracted to the olfactory signals of prey. PLoS One 2010; 5: e13114 11 Hein AM, McKinley SA. Sensing and decisionmaking in random search. Proc Natl Acad Sci USA 2012; 109: 12070 – 12074 12 Ferrero DM, Liberles SD. The secret codes of mammalian scents. Wiley Interdiscip Rev Syst Biol Med 2010; 2: 23 – 33 13 Moore JG, Krotoszynski BK, O'Neill HJ. Fecal odorgrams. A method for partial reconstruction of ancient and modern diets. Dig Dis Sci 1984; 29: 907 – 911 14 Moore JG, Jessop LD, Osborne DN. Gas-chromatographic and mass-spectrometric analysis of the odor of human feces. Gastroenterology 1987; 93: 1321 – 1329
15 Charles-Dominique P. Urine marking and territoriality in Galago alleni (Waterhouse, 1837 – Lorisoidea, Primates) – a field study by radio-telemetry. Z Tierpsychol 1977; 43: 113 – 138 16 Brashares JS, Arcese P. Scent marking in a territorial African antelope: II. The economics of marking with faeces. Anim Behav 1999; 57: 11 – 17 17 Bekoff M. Observations of scent-marking and discriminating self from others by a domestic dog (Canis familiaris): tales of displaced yellow snow. Behav Processes 2001; 55: 75 – 79 18 Dröscher I, Kappeler PM. Maintenance of familiarity and social bonding via communal latrine use in a solitary primate (Lepilemur leucopus). Behav Ecol Sociobiol 2014; 68: 2043 – 2058 19 Brain CK. New finds at the Swartkrans australopithecine site. Nature 1970; 225: 1112 – 1119 20 Lee-Thorp J, Thackeray JF, van der Merwe N. The hunters and the hunted revisited. J Hum Evol 2000; 39: 565 – 576 21 Pickering TR. Taphonomy of the Swartkrans hominid postcrania and its bearing on issues of meat-eating and fire management. In Stanford CB, Bunn HT eds. Meat-eating and human evolution. Oxford: Oxford University; 2001: 33 – 51 22 Blumenschine RJ, Stanistreet IG, Njau JK et al. Environments and hominin activities across the FLK Peninsula during Zinjanthropus times (1.84 Ma), Olduvai Gorge, Tanzania. J Hum Evol 2012; 63: 364 – 383 23 Eppinger M, Baar C, Linz B et al. Who ate whom? Adaptive Helicobacter genomic changes that accompanied a host jump from early humans to large felines. PLoS Genet 2006; 2: e120 24 Zu J, Takeuchi Y. Adaptive evolution of antipredator ability promotes the diversity of prey species: critical function analysis. Biosystems 2012; 109: 192 – 202 25 Shultz S, Dunbar RI. Chimpanzee and felid diet composition is influenced by prey brain size. Biol Lett 2006; 2: 505 – 508 26 Walter M. Defence of bipedalism. Human Evol 2004; 19: 19 – 44 27 Ferrero DM, Lemon JK, Fluegge D et al. Detection and avoidance of a carnivore odor by prey. Proc Natl Acad Sci USA 2011; 108: 11235 – 11240 28 Anson JR, Dickman CR. Behavioral responses of native prey to disparate predators: naiveté and predator recognition. Oecologia 2013; 171: 367 – 377 29 Apfelbach R, Blanchard CD, Blanchard RJ et al. The effects of predator odors in mammalian prey species: a review of field and laboratory studies. Neurosci Biobehav Rev 2005; 29: 1123 – 1144 30 Parsons MH, Blumstein DT. Familiarity breeds contempt: kangaroos persistently avoid areas with experimentally deployed dingo scents. PLoS One 2010; 5: e10403 31 Takahashi LK, Nakashima BR, Hong H et al. The smell of danger: a behavioral and neural analysis of predator odor-induced fear. Neurosci Biobehav Rev 2005; 29: 1157 – 1167 32 Takahashi LK, Chan MM, Pilar ML. Predator odor fear conditioning: current perspectives and new directions. Neurosci Biobehav Rev 2008; 32: 1218 – 1227
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
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Downloaded by: Collections and Technical Services Department. Copyrighted material.
Kommentiertes Referat
Kommentiertes Referat 33 Bassotti G, Villanacci V. The control of defecation in humans: an evolutionary advantage? Tech Coloproctol 2013; 17: 623 – 624 34 Porges RF, Porges JC, Blinick G. Mechanisms of uterine support and the pathogenesis of uterine prolapse. Obstet Gynecol 1960; 15: 711 – 726 35 Dubrovsky B, Filipini D. Neurobiological aspects of the pelvic floor muscles involved in defecation. Neurosci Biobehav Rev 1990; 14: 157 – 168 36 Rosenberg K, Trevathan W. Birth, obstetrics and human evolution. Br J Obstet Gynaecol 2002; 109: 1199 – 1206 37 Altomare DF, Rinaldi M, Veglia A et al. Contribution of posture to the maintenance of anal
38
39
40
41
continence. Int J Colorectal Dis 2001; 16: 51 – 54 Schimpf M, Tulikangas P. Evolution of the female pelvis and relationships to pelvic organ prolapse. Int Urogynecol J 2005; 16: 315 – 320 Abitbol MM. Evolution of the ischial spine and of the pelvic floor in the Hominoidea. Am J Phys Anthropol 1988; 75: 53 – 67 Somel M, Liu X, Khaitovich P. Human brain evolution: transcripts, metabolites and their regulators. Nat Rev Neurosci 2013; 14: 112 – 127 Whiten A, Erdal D. The human socio-cognitive niche and its evolutionary origins. Phi-
los Trans R Soc Lond B Biol Sci 2012; 367: 2119 – 2129
Prof. Dr. Gabrio Bassotti Department of Clinical & Experimental Medicine, Gastroenterology & Hepatology Section Via Enrico Dal Pozzo 06100 Perugia Italy Tel.: ++ 39/075/5784423 Fax.: ++ 39/0 75/5 84/75 70 [email protected]
Downloaded by: Collections and Technical Services Department. Copyrighted material.
462
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
Controlling defecation: to be (predator) or not to be (prey), that is the question… Räuber) oder Nichtsein (Beute), das ist die Frage … Z Gastroenterol 2015; 53: 460–462 G. Bassotti, S. Müller-Lissner He who can no longer pause and stand rapt in awe is as good as dead; his eyes are closed. To raise new questions, new possibilities, to regard old problems from a new angle, requires creative imagination and marks real advance in science. Logic will get you from A to B. Imagination will take you everywhere. (Albert Einstein)
Introduction The process of defecation in human beings involves a series of physiological events that, after the digestive and absorptive processes occurring in the upper part of the digestive tract have been completed, cause the residual more or less compacted contents (fecal matter) to be transported through the colon and eventually expelled by relaxation of the anal sphincter [1]. Although defecation occurs almost daily in normal subjects, the very act is under strict cultural taboos, at least in most societies, and it is rarely if ever thought or spoken of, apart from practical jokes and medical evaluation. However, it is worth noting that defecation is subject to a strict and voluntary central control [2, 3] that, depending on the cultural environment, allows the individual to decide if, when and where it is socially convenient to evacuate [4]. Moreover, the control of defecation is not an innate process, since it is acquired in infancy by toilet training [5], usually in the family environment; recognition of the need to defecate is usually well established by age four, and the ability to hold back defecation improves with age [6]. Thus, controlling defecation is an important aspect of the human being’s life and, provided all the appropriate reflexes are intact, bowel motions may be (within reasonable limits) kept under control until the subject decides it is appropriate to evacuate his/her gut contents.
Since it is likely that our ancestors (hominins) were subject to the same ecological pressures as other primates, particularly predation risk and competition for resources [7], in this article it will be hypothesized why men control defecation, and the possible evolutionary role of controlling defecation as a method for hominins to increase foraging strategies, but also as a tool to avoid predation.
To be a predator… Predation is an important, sometimes essential method to assure survival. However, predation implies that the predator’s abilities overwhelm that of prey, and this may be difficult when the prey is highly mobile. One of the main points, besides agility and force, is therefore that the predator must be able to reach the prey or to approach it in order to use tools. For this purpose, camouflage and speed are of paramount importance, even though other factors are equally important. In fact, we could assume that predation for hominins would hardly have involved camouflage (scarcely effective on moving targets [8]) and speed, whereas stealth probably played a major role for hunting. Thus, predators have to be silent and, more importantly, to offer little or no clues (in particular, olfactory traces) of their approach. This is particularly important for mammals, whose nose is a powerful chemosensor, capable of detecting and distinguishing a myriad of chemicals [9]. Therefore, it is not surprising that olfactory hunting is very common in predators [10] since many of them must locate resources in environments in which sensory signals are noisy and without directional information [11]. The scents of mammals are complex blends of natural products that reveal a wealth of individual information [12]; under this light, fecal scents are particularly effective, as revealed by the analysis of fecal odorgrams [13, 14]. Indeed, selective delimitation of territorial areas with fecal (and/or urinary) amounts is a method employed by several mammal species, primates and non-primates, to mark their territory and suggests that control of defecation and targeted deployment of excrements is widely present in animals [15 – 17]. In addition, there is evidence that several species (including some primates) use latrines, i. e., repeated specific defecation/urination sites for territory demarcation, resource defense, signaling of repro-
ductive state, social bonding, and mate defense [18]. However, concerning predator-prey relationships, the importance of fecal odors may have double-faced implications. On the one hand, in fact, the predator follows the scents of the prey, and the fecal ones are outstanding under this light; on the other hand, the reverse is equally important, in that preys are usually able to detect the scents of predators, and those related to feces are extremely useful to avoid predation (see below).
…or not to be prey There is presently evidence that hominins may have been the prey of several species of large carnivores [19 – 22], as also shown by the recent demonstration of transmission of Helicobacter spp from early humans to preying felids [23]. Thus, as it happens in other animal species [24], our ancestors had to develop a series of adaptive evolutionary strategies to avoid predation, strategies that included morphological defenses as well as behavioral traits. Indeed, experimental animal studies suggest that large-brained prey are likely to be more effective at evading predators because they can effectively alter their behavioral responses to specific predator encounters [25]. Predator recognition and avoidance are important behaviors in allowing prey to mitigate the impacts of their predators; for instance, some authors hypothesized that the development of erect stature in hominins, regardless of locomotion proficiency, actually enhanced both active and passive defense against predators [26]. However, several other and effective mechanisms may play a role in this context. For instance, many prey species try to avoid predation also by learning the predators’ odors [27, 28]. Data derived from experience in animal studies show that several prey species avoid the odors of their predators (including that of feces) [29, 30], and that compounds isolated from the feces elicit a series of fear-related responses, including avoidance and risk assessment behavior [31, 32].
…the control of defecation is the question On the basis of the above considerations, it might be hypothesized that learning to control defecation, before becoming a so-
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
Downloaded by: Collections and Technical Services Department. Copyrighted material.
460
cial necessity, may have had some evolutionary significance for hominins as a means both not to advertise their presence to a possible prey and to mask an odoriferous trail to a possible predator [33]. This may also be inferred by the fact that other primate species are able to control defecation and deposit their feces in specific sites such as latrines [18]. Of interest, the development of erect stature might have helped the control upon defecation by our ancestors by means of specific attachments and functions of the structure of the pelvic floor [34, 35]. In fact, humans are thought to be the first purely bipedal mammals [36], and the erect position contributes significantly to the maintenance of fecal continence by sharpening the anorectal angle [37]; species that live at least part of their life in trees have fewer demands on their pelvic structures for support than bipedal terrestrial animals that maintain a vertical posture for longer periods of time [38]. For instance the gorilla, that only occasionally is bipedal, is known to have poor control of defecation, due to a different conformation of the pelvic floor [39]. There is no doubt that among all the living creatures humans have achieved a lot in evolutionary terms. The main propulsive force leading to the differentiation toward the other species was the emergence of human cognition, and the researchers are still actively investigating this phenomenon [40]. The process of creative thinking, the ability to develop new tools, the never-ending struggle toward knowledge allowed our ancestors to graduate from an almost animal state to the present one (although someone would argue that this had not necessarily been for the better…), allowing human hunter-gatherer bands to function as unique and highly competitive predatory organisms [41]. In this respect, among the various strategies to improve both foraging and survival, it might be hypothesized that the control of defecation (even though diversified in the various species) may have played a role, allowing a minor detection by the possible preys and a greater likelihood of escaping predation by not leaving around easily detectable spoors.
Kommentierte Referate können von jedem interessierten Kollegen verfasst werden. Sie sollten zuvor beim Koordinator angemeldet werden. Die formale Gestaltung orientiert sich an den publizierten Referaten (Kurzzitate im Text!). Einreichung nicht über die Online-Manuskripteinreichung des Thieme Verlags, sondern via E-mail an den Koordinator der Kommentierten Referate. Prof. Dr. Stefan Müller-Lissner Park-Klinik Weißensee Schönstraße 80, 13086 Berlin Tel. + 49 30 96 28 36 00, Fax 36 05, [email protected]
References 01 Bassotti G, de Roberto G, Castellani D et al. Normal aspects of colorectal motility and abnormalities in slow transit constipation. World J Gastroenterol 2005; 11: 2691 – 2696 02 Brading AF, Ramalingam T. Mechanisms controlling normal defecation and the potential effects of spinal cord injury. Prog Brain Res 2006; 152: 345 – 358 03 Bittorf B, Ringler R, Forster C et al. Cerebral representation of the anorectum using functional magnetic resonance imaging. Br J Surg 2006; 93: 1251 – 1257 04 Bajwa A, Emmanuel A. The physiology of continence and evacuation. Best Pract Res Clin Gastroenterol 2009; 23: 477 – 485 05 Kiddoo DA. Toilet training children: when to start and how to train. CMAJ 2012; 184: 511 – 512 06 Kyrklund K, Koivusalo A, Rintala RJ et al. Evaluation of bowel function and fecal continence in 594 Finnish individuals aged 4 to 26 years. Dis Colon Rectum 2012; 55: 671 – 676 07 Rose L, Marshall F. Meat eating, hominid sociality, and home bases revisited. Curr Anthropol 1996; 37: 307 – 338 08 Hall JR, Cuthill IC, Baddeley R et al. Camouflage, detection and identification of moving targets. Proc Biol Sci 2013; 280 20130064 09 Liberles SD. Trace amine-associated receptors are olfactory receptors in vertebrates. Ann N Y Acad Sci 2009; 1170: 168 – 172 10 Hughes NK, Price CJ, Banks PB. Predators are attracted to the olfactory signals of prey. PLoS One 2010; 5: e13114 11 Hein AM, McKinley SA. Sensing and decisionmaking in random search. Proc Natl Acad Sci USA 2012; 109: 12070 – 12074 12 Ferrero DM, Liberles SD. The secret codes of mammalian scents. Wiley Interdiscip Rev Syst Biol Med 2010; 2: 23 – 33 13 Moore JG, Krotoszynski BK, O'Neill HJ. Fecal odorgrams. A method for partial reconstruction of ancient and modern diets. Dig Dis Sci 1984; 29: 907 – 911 14 Moore JG, Jessop LD, Osborne DN. Gas-chromatographic and mass-spectrometric analysis of the odor of human feces. Gastroenterology 1987; 93: 1321 – 1329
15 Charles-Dominique P. Urine marking and territoriality in Galago alleni (Waterhouse, 1837 – Lorisoidea, Primates) – a field study by radio-telemetry. Z Tierpsychol 1977; 43: 113 – 138 16 Brashares JS, Arcese P. Scent marking in a territorial African antelope: II. The economics of marking with faeces. Anim Behav 1999; 57: 11 – 17 17 Bekoff M. Observations of scent-marking and discriminating self from others by a domestic dog (Canis familiaris): tales of displaced yellow snow. Behav Processes 2001; 55: 75 – 79 18 Dröscher I, Kappeler PM. Maintenance of familiarity and social bonding via communal latrine use in a solitary primate (Lepilemur leucopus). Behav Ecol Sociobiol 2014; 68: 2043 – 2058 19 Brain CK. New finds at the Swartkrans australopithecine site. Nature 1970; 225: 1112 – 1119 20 Lee-Thorp J, Thackeray JF, van der Merwe N. The hunters and the hunted revisited. J Hum Evol 2000; 39: 565 – 576 21 Pickering TR. Taphonomy of the Swartkrans hominid postcrania and its bearing on issues of meat-eating and fire management. In Stanford CB, Bunn HT eds. Meat-eating and human evolution. Oxford: Oxford University; 2001: 33 – 51 22 Blumenschine RJ, Stanistreet IG, Njau JK et al. Environments and hominin activities across the FLK Peninsula during Zinjanthropus times (1.84 Ma), Olduvai Gorge, Tanzania. J Hum Evol 2012; 63: 364 – 383 23 Eppinger M, Baar C, Linz B et al. Who ate whom? Adaptive Helicobacter genomic changes that accompanied a host jump from early humans to large felines. PLoS Genet 2006; 2: e120 24 Zu J, Takeuchi Y. Adaptive evolution of antipredator ability promotes the diversity of prey species: critical function analysis. Biosystems 2012; 109: 192 – 202 25 Shultz S, Dunbar RI. Chimpanzee and felid diet composition is influenced by prey brain size. Biol Lett 2006; 2: 505 – 508 26 Walter M. Defence of bipedalism. Human Evol 2004; 19: 19 – 44 27 Ferrero DM, Lemon JK, Fluegge D et al. Detection and avoidance of a carnivore odor by prey. Proc Natl Acad Sci USA 2011; 108: 11235 – 11240 28 Anson JR, Dickman CR. Behavioral responses of native prey to disparate predators: naiveté and predator recognition. Oecologia 2013; 171: 367 – 377 29 Apfelbach R, Blanchard CD, Blanchard RJ et al. The effects of predator odors in mammalian prey species: a review of field and laboratory studies. Neurosci Biobehav Rev 2005; 29: 1123 – 1144 30 Parsons MH, Blumstein DT. Familiarity breeds contempt: kangaroos persistently avoid areas with experimentally deployed dingo scents. PLoS One 2010; 5: e10403 31 Takahashi LK, Nakashima BR, Hong H et al. The smell of danger: a behavioral and neural analysis of predator odor-induced fear. Neurosci Biobehav Rev 2005; 29: 1157 – 1167 32 Takahashi LK, Chan MM, Pilar ML. Predator odor fear conditioning: current perspectives and new directions. Neurosci Biobehav Rev 2008; 32: 1218 – 1227
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
461
Downloaded by: Collections and Technical Services Department. Copyrighted material.
Kommentiertes Referat
Kommentiertes Referat 33 Bassotti G, Villanacci V. The control of defecation in humans: an evolutionary advantage? Tech Coloproctol 2013; 17: 623 – 624 34 Porges RF, Porges JC, Blinick G. Mechanisms of uterine support and the pathogenesis of uterine prolapse. Obstet Gynecol 1960; 15: 711 – 726 35 Dubrovsky B, Filipini D. Neurobiological aspects of the pelvic floor muscles involved in defecation. Neurosci Biobehav Rev 1990; 14: 157 – 168 36 Rosenberg K, Trevathan W. Birth, obstetrics and human evolution. Br J Obstet Gynaecol 2002; 109: 1199 – 1206 37 Altomare DF, Rinaldi M, Veglia A et al. Contribution of posture to the maintenance of anal
38
39
40
41
continence. Int J Colorectal Dis 2001; 16: 51 – 54 Schimpf M, Tulikangas P. Evolution of the female pelvis and relationships to pelvic organ prolapse. Int Urogynecol J 2005; 16: 315 – 320 Abitbol MM. Evolution of the ischial spine and of the pelvic floor in the Hominoidea. Am J Phys Anthropol 1988; 75: 53 – 67 Somel M, Liu X, Khaitovich P. Human brain evolution: transcripts, metabolites and their regulators. Nat Rev Neurosci 2013; 14: 112 – 127 Whiten A, Erdal D. The human socio-cognitive niche and its evolutionary origins. Phi-
los Trans R Soc Lond B Biol Sci 2012; 367: 2119 – 2129
Prof. Dr. Gabrio Bassotti Department of Clinical & Experimental Medicine, Gastroenterology & Hepatology Section Via Enrico Dal Pozzo 06100 Perugia Italy Tel.: ++ 39/075/5784423 Fax.: ++ 39/0 75/5 84/75 70 [email protected]
Downloaded by: Collections and Technical Services Department. Copyrighted material.
462
Bassotti G, Müller-Lissner S. Controlling defecation: to … Z Gastroenterol 2015; 53: 460–462 · DOI http://dx.doi.org/10.1055/s-0034-1399242
