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What is the Function of the Human Vermiform Appendix

What is the Function of the Human Vermiform Appendix

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Published by MagicWand2
Evolution-Based Surgery: A New Perspective in the Darwinian Year 2009
Evolution-Based Surgery: A New Perspective in the Darwinian Year 2009

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Published by: MagicWand2 on Apr 05, 2011
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Fax +41 61 306 12 34E-Mail karger@karger.chwww.karger.com
Eur Surg Res 2009;43:67–71DOI:10.1159/000219236
What Is the Function of the HumanVermiform Appendix?
Evolution-Based Surgery: A New Perspective in the Darwinian Year 2009
L. Ansaloni F. Catena A.D. Pinna
Unit of General, Emergency and Transplant Surgery, St. Orsola-Malpighi University Hospital, Bologna, Italy
a more important physiological role in past ancestorsthan in present descendants. Independent of evolution-ary theory, a vestige can also be defined typologically asa reduced and rudimentary structure when compared tohomologous structures in other organisms, one that lacksthe complex functions usually found for that structure inother organisms. Typical examples of vestiges are thewings of the ostrich and the eyes of blind cavefish. Al-though these vestigial structures may currently serve ei-ther some evident or obscure purpose, rudimentary os-trich wings are useless as normal wings for flying just asrudimentary cavefish eyes are useless as normal eyes forseeing. The vermiform appendix is a developmental deriva-tive and evolutionary vestige of a much larger herbivo-rous caecum: in most vertebrates, the caecum is a largecomplex gastrointestinal organ, enriched in mucosallymphatic tissue. The caecum varies in dimension amongspecies, but in general the size of the caecum is directly relative to the quantity of plant matter in a given organ-ism’s diet. It is largest in obligate herbivores, animalswhose diets consist entirely of plant stuff, because thecaecum is essential for digestion of cellulose, a key mol-ecule found in plants. Since the caecum houses special-ized and symbiotic bacteria that secrete cellulase (an en-zyme that digests cellulose), without this specializedfunction, it is impossible for mammals to digest cellulose[7]. In vertebrate comparative anatomy, it has long beenknown that the human appendix and the end of the mam- The vermiform appendix has been known as an organsince the late fifteenth century; it was clearly depicted inLeonardo da Vinci’s anatomical drawings in 1492, firstdescribed in detail by Berengario da Carpi in 1521, andfinally, the worm-like organ was named the vermiformappendix in 1530 by Vido Vidius (Guido Guidi). The laterecognition of the appendix in the scientific community is probably due to the fact that early anatomical studieswere typically done on animal species possessing no suchorgan [1]. Although the first known surgical removal of the appendix occurred in December 1735 by ClaudiusAmyand, who operated on an 11-year-old boy with alongstanding scrotal hernia and a fecal fistula of the thighat St. George’s Hospital in London [2], it was not until thelate nineteenth century that it was acknowledged thatmost inflammatory diseases of the lower-right quadrantoriginate in the appendix. Due largely to the works of thephysician Reginald Fitz and the surgeon Charles McBur-ney in the United States, the clinical features of appendi-citis were clearly described and, most importantly, early surgical removal of the appendix became a commonly recommended procedure [3, 4]. A few years before this surgical revelation, CharlesDarwin and other proponents of the evolutionary theory (like the Italian naturalist, Giovanni Canestrini) listedthe vermiform appendix among the rudimentary organsof the human species, stressing its vestigial nature as evi-dence of human evolutionary history [5, 6]. Evolutionary vestiges are, in principle, diminished structures that had
Published online: May 19, 2009
Dr. Luca Ansaloni, MDUnit of General, Emergency and Transplant Surgery St. Orsola-Malpighi University Hospital, Via Massarenti 9IT–40138 Bologna (Italy)Tel. +39 051 636 3584, Fax +39 051 636 4745, E-Mail luca.ansaloni@aosp.bo.it© 2009 S. Karger AG, Basel0014–312X/09/0432–0067$26.00/0Accessible online at:www.karger.com/esr
Eur Surg Res 2009;43:67–71
malian caecum are structurally homologous by usualsystematic criteria. Within the gastrointestinal tract of many mammals, particularly primates, the end of thecaecum and the vermiform appendix share the same rel-ative position; both have a similar structure and form,both are blind sacs enriched with lymphatic tissue, andboth share a common developmental origin [7, 8]: a con-clusion that was further confirmed by cladistic system-atic analysis [9]. A vermiform appendix is not unique tohumans: it exists in all the hominoid apes, includingchimpanzees, gorillas, orangutans, and gibbons, and it isfound to varying degrees in several species of both NewWorld and Old World monkeys [10]. A few other mam-mals appear to have an organ similar to the hominoidvermiform appendix, including the wombat and SouthAmerican opossum (both marsupials), some rodents,and the rabbit. However, extensive comparative analysishas shown that the caecal appendixes of humans andthese other mammals were derived from the caecum in-dependently, and accordingly, these anatomic structuresare not homologous in origin [11]. Contrastingly, creationist authors and advocates of in-telligent design have argued that the appendix is not ves-tigial in an evolutionary sense, because, being functional,it appears to have a creative design and organization as if formed according to a plan for a specific purpose. In thisway, creationists contend that vermiform appendix util-ity acts as evidence against evolutionary theory [12, 13]. Although the precise function of the human vermi-form appendix is still uncertain, many hypotheses havebeen made. Because the appendix is associated with sub-stantial lymphatic tissue, it was once thought to play arole in immune function. Although it was suggested by some authors that the appendix itself could be the site of B-lymphocyte induction (a bursa of Fabricius equivalent)[14], recent conjecture favours the localization of this bio-logical programming in the bone marrow. The appendixmay still play a role in this highly significant task, but notby itself. Its lymphoid tissue is conclusively known to beinvolved in antibody production (the function of B-typelymphocytes), especially the IgA type immunoglobulinsfor secretion or mucosal surface immunity as well as partof the gut-associated lymphoid tissue (GALT) [15]. Thissuggests that the appendix plays a specific role in im-mune function, a theory further supported by the factthat the appendix is a priming site in the development of ulcerative colitis [16, 17], and so appendectomies seem tohave a protective role in patients who undergo them be-fore 20 years of age [18]. Some other authors have pro-posed that the human appendix is well suited as a ‘safehouse’ for commensal bacteria, providing support forbacterial growth and potentially facilitating re-inocula-tion of the colon in the event that the contents of the in-testinal tract are purged following exposure to a patho-gen [19]. This could explain the observation that appen-dectomies have been reported as a significant underlyingrisk factor in functional gastrointestinal disorders, likeirritable bowel syndrome [20, 21]. However, if the vermiform appendix does indeed havea specific function in humans (even as part of a largermulti-organ system), it would be an example of an organthat is not adapted, but rather exapted. Exaptation is awell-known and accurately described mechanism of evo-lution that must be properly distinguished from adapta-tion. Adaptation, a central concept in evolutionary biol-ogy, describes a trait that evolved by natural selection be-cause it served a particular function, while exaptationinstead refers to shifts in the function of a trait duringevolution. The idea that the function of a trait might shiftand reconfigure itself during its evolutionary pathway was originally proposed by Charles Darwin, who cited asevidence the swimbladder in fishes versus the lungs of other vertebrate animals [22], as well as the various fertil-izing contrivances of orchids [23]. Unfortunately, formany years the phenomenon was inappropriately labeled‘pre-adaptation’, a term suggesting planning and fore-thought, contradictory concepts to the basic principles of natural selection. Later, in 1982, the prominent paleon-tologists and evolutionary biologists Stephen J. Gouldand Elisabeth S. Vrba defined the idea of ‘exaptation’ as‘features that now enhance fitness, but were not designedby natural selection for their current role’. This term,which subsequently found major applications in the fieldof evolutionary biology, underscores the universal role of ‘redesign’ in the evolution of complex biological mecha-nisms and includes 2 categories: (1) a character trait, pre-viously shaped by natural selection for a particular func-tion (an adaptation), is co-opted for a new use; (2) a char-acter trait whose origin cannot be ascribed to the directaction of natural selection (a non-aptation), is co-optedfor a current use [24]. A multi-stage example of exapta-tion involves human hands, which evolved to facilitatetool use and are an exaptation of primate hands that wereused for grasping tree branches. Those primate hands, inturn, were an exaptation of front legs that were used forlocomotion on the ground, and those legs were an exap-tation of the fins of fish, which were used for locomotionin the water. As this lineage exploited different ecologicalniches (water, land, trees, and eventual ground-level tooluse), natural selection progressively reshaped its limbs.
What Is the Function of the HumanVermiform Appendix?
Eur Surg Res 2009;43:67–71
 This exaptation mechanism is crucial for resolvingone of the challenges to Darwin’s theory of evolution:how complex structures could evolve gradually, giventhat their incipient forms may have been inadequate toserve any function. As Mivart (a critic of Darwin) point-ed out, 5% of a bird wing would not be functional. Hence,the incipient form of complex traits would not have sur-vived long enough to evolve into a useful form. As Dar-win elaborated in
On the Origin of Species
 , many complextraits evolved from earlier traits that had once served dif-ferent functions [22]. By trapping air, primitive wings en-abled birds to efficiently regulate their temperature, inpart by lifting up their feathers to dissipate heat when toowarm. Individual members of this species that more ef-ficiently exploited this trait had better chances of surviv-al; therefore, bearing more offspring, resulting in thespread of this trait. Eventually, feathers became suffi-ciently large that they enabled some individuals to glide.These individuals in turn would bear more offspring, re-sulting in the further spread and promotion of this traitwithin the gene pool due to its beneficial, secondary lo-comotive function. Hence, the evolution of bird wingscan be explained by a shifting in function from the regu-lation of temperature to flight. Furthermore, regarding the concept of exaptation,Darwin’s theory explains how the traits of living organ-isms are often well-designed for their environment whilesimultaneously conceding that many traits can be imper-fectly designed. They appear to have been made fromavailable pre-existing structures and material, that is,they are a product of improvised design. Understandingexaptation involves scrutinizing the subtleties in the pro-gressive adaptation process. For instance, understandingthat feathers evolved initially for thermal regulation may help explain many of their features unrelated to flight[25]. Hence, the current functions of the human vermiformappendix appear to stem from the exaptation of the pri-mordial caecum, which was originally large and enrichedin mucosal lymphatic tissue, a structure essential for thedigestion of cellulose in herbivores. Additionally, the hu-man appendix seems to be suboptimally designed, as it isnotorious for the life-threatening complications it cancause. Acute appendicitis is the most common abdomi-nal emergency, and accounts for more than 40,000 hos-pital admissions in England every year [26]; the overalllifetime risk is 8.6% for males and 6.7% for females in theUnited States [27]. Before modern twentieth-century sur-gical techniques were available, a case of acute appendi-citis was usually fatal and even today, with all modernadvancements in medicine and technology, appendicitisfatalities remain significant [28]. The small entrance tothis dead-end pocket makes the appendix difficult toclean out and prone to physical blockage, a detrimentwhich is ultimately the cause of appendicitis. This pecu-liar structural layout is quite beneficial for a larger cellu-lose-fermenting caecum, but it is unclear why gut lym-phoid tissue would need to be housed in a remote, dead-end tube with negligible surface area. In fact, 60% of appendicitis cases are due to lymphoid hyperplasia lead-ing to occlusion of the interior of the appendix, indicatingthat the appendix is unusually prone to abnormal prolif-eration of its lymphoid tissue [29]. Such an occurrencewould be much less problematic if the interior of the ap-pendix were not so small, confined, and inaccessible fromthe rest of the gut. In many other primates and mammals,the GALT lymphoid tissue appears to function withoutdifficulty in a much more open, bulbous caecum withample surface area. Furthermore, as mentioned previ-ously, there is mounting evidence that removing the ap-pendix helps prevent ulcerative colitis, a nasty inflamma-tory disease of the colon [18]. This evidence suggests thatthe appendix is actually maladaptive, and that the lym-phoid tissue contained in the appendix is prone to chron-ic pathological inflammatory states. Unless the appendixdoes in fact have an important function that we have yetto discover, it is a leading candidate for the most poorly designed organ in the human body. Any biological struc-ture that supposedly safeguards our livelihood, yet para-doxically kills a percentage of its bearers prematurely, isindeed poorly designed. The vermiform appendix, in itspresent form, illustrates a delicate balance between exap-tive functional benefits and disease-causing detriments.It can aid a given individual by acting as a part of GALTfor the intestinal immune system or by re-inoculating thecolonic flora through bacterial growth, yet it can just aseasily trigger life-threatening conditions. This balance of exaptive benefits and vestigial detriments is directly re-lated to an individual’s ability to survive and reproducemore offspring of our species. Further complicating thisbalance is the fact that selective pressures have beenknown to change, thereby shifting this delicate equilib-rium. The vermiform appendix was an important evolu-tionary asset before the benefits of civilization (clean wa-ter, cooked food, etc.) eliminated some of the commonthreats to individual survival, rendering many of the or-gan’s primordial functions obsolete. Shortly thereafter,the detrimental properties of the appendix became moreprominent. However, civilization can further overridecertain evolutionary mechanisms with medicine, facili-

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