Editorial
Eur Surg Res 2009;43:67–71
DOI: 10.1159/000219236
What Is the Function of the Human
Vermiform 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
The vermiform appendix has been known as an organ
since the late fifteenth century; it was clearly depicted in
Leonardo da Vinci’s anatomical drawings in 1492, first
described in detail by Berengario da Carpi in 1521, and
finally, the worm-like organ was named the vermiform
appendix in 1530 by Vido Vidius (Guido Guidi). The late
recognition of the appendix in the scientific community
is probably due to the fact that early anatomical studies
were typically done on animal species possessing no such
organ [1]. Although the first known surgical removal of
the appendix occurred in December 1735 by Claudius
Amyand, who operated on an 11-year-old boy with a
longstanding scrotal hernia and a fecal fistula of the thigh
at St. George’s Hospital in London [2], it was not until the
late nineteenth century that it was acknowledged that
most inflammatory diseases of the lower-right quadrant
originate in the appendix. Due largely to the works of the
physician 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, Charles
Darwin and other proponents of the evolutionary theory
(like the Italian naturalist, Giovanni Canestrini) listed
the vermiform appendix among the rudimentary organs
of 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
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a more important physiological role in past ancestors
than in present descendants. Independent of evolution-
ary theory, a vestige can also be defined typologically as
a reduced and rudimentary structure when compared to
homologous structures in other organisms, one that lacks
the complex functions usually found for that structure in
other organisms. Typical examples of vestiges are the
wings 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 as
rudimentary cavefish eyes are useless as normal eyes for
seeing.
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 large
complex gastrointestinal organ, enriched in mucosal
lymphatic tissue. The caecum varies in dimension among
species, 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, animals
whose diets consist entirely of plant stuff, because the
caecum 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 specialized
function, it is impossible for mammals to digest cellulose
[7]. In vertebrate comparative anatomy, it has long been
known that the human appendix and the end of the mam-
Dr. Luca Ansaloni, MD
Unit of General, Emergency and Transplant Surgery
St. Orsola-Malpighi University Hospital, Via Massarenti 9
IT–40138 Bologna (Italy)
Tel. +39 051 636 3584, Fax +39 051 636 4745, E-Mail luca.ansaloni@aosp.bo.it
malian caecum are structurally homologous by usual
systematic criteria. Within the gastrointestinal tract of
many mammals, particularly primates, the end of the
caecum 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, and
both 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 to
humans: it exists in all the hominoid apes, including
chimpanzees, gorillas, orangutans, and gibbons, and it is
found to varying degrees in several species of both New
World and Old World monkeys [10]. A few other mam-
mals appear to have an organ similar to the hominoid
vermiform appendix, including the wombat and South
American opossum (both marsupials), some rodents,
and the rabbit. However, extensive comparative analysis
has shown that the caecal appendixes of humans and
these other mammals were derived from the caecum in-
dependently, and accordingly, these anatomic structures
are 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 this
way, 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 have
been made. Because the appendix is associated with sub-
stantial lymphatic tissue, it was once thought to play a
role 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 appendix
may still play a role in this highly significant task, but not
by itself. Its lymphoid tissue is conclusively known to be
involved in antibody production (the function of B-type
lymphocytes), especially the IgA type immunoglobulins
for secretion or mucosal surface immunity as well as part
of the gut-associated lymphoid tissue (GALT) [15]. This
suggests that the appendix plays a specific role in im-
mune function, a theory further supported by the fact
that the appendix is a priming site in the development of
ulcerative colitis [16, 17], and so appendectomies seem to
have 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 ‘safe
68 Eur Surg Res 2009;43:67–71
house’ for commensal bacteria, providing support for
bacterial 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 underlying
risk factor in functional gastrointestinal disorders, like
irritable bowel syndrome [20, 21].
However, if the vermiform appendix does indeed have
a specific function in humans (even as part of a larger
multi-organ system), it would be an example of an organ
that is not adapted, but rather exapted. Exaptation is a
well-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 exaptation
instead refers to shifts in the function of a trait during
evolution. The idea that the function of a trait might shift
and reconfigure itself during its evolutionary pathway
was originally proposed by Charles Darwin, who cited as
evidence the swimbladder in fishes versus the lungs of
other vertebrate animals [22], as well as the various fertil-
izing contrivances of orchids [23]. Unfortunately, for
many 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. Gould
and Elisabeth S. Vrba defined the idea of ‘exaptation’ as
‘features that now enhance fitness, but were not designed
by natural selection for their current role’. This term,
which subsequently found major applications in the field
of 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 direct
action of natural selection (a non-aptation), is co-opted
for a current use [24]. A multi-stage example of exapta-
tion involves human hands, which evolved to facilitate
tool use and are an exaptation of primate hands that were
used for grasping tree branches. Those primate hands, in
turn, were an exaptation of front legs that were used for
locomotion on the ground, and those legs were an exap-
tation of the fins of fish, which were used for locomotion
in the water. As this lineage exploited different ecological
niches (water, land, trees, and eventual ground-level tool
use), natural selection progressively reshaped its limbs.
Ansaloni/Catena/Pinna
 This exaptation mechanism is crucial for resolving
one of the challenges to Darwin’s theory of evolution:
how complex structures could evolve gradually, given
that their incipient forms may have been inadequate to
serve 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 complex
traits 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, in
part by lifting up their feathers to dissipate heat when too
warm. Individual members of this species that more ef-
ficiently exploited this trait had better chances of surviv-
al; therefore, bearing more offspring, resulting in the
spread 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 trait
within the gene pool due to its beneficial, secondary lo-
comotive function. Hence, the evolution of bird wings
can 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 while
simultaneously conceding that many traits can be imper-
fectly designed. They appear to have been made from
available pre-existing structures and material, that is,
they are a product of improvised design. Understanding
exaptation involves scrutinizing the subtleties in the pro-
gressive adaptation process. For instance, understanding
that feathers evolved initially for thermal regulation may
help explain many of their features unrelated to flight
[25].
Hence, the current functions of the human vermiform
appendix appear to stem from the exaptation of the pri-
mordial caecum, which was originally large and enriched
in mucosal lymphatic tissue, a structure essential for the
digestion of cellulose in herbivores. Additionally, the hu-
man appendix seems to be suboptimally designed, as it is
notorious for the life-threatening complications it can
cause. 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 overall
lifetime risk is 8.6% for males and 6.7% for females in the
United 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 modern
What Is the Function of the Human
Vermiform Appendix?
advancements in medicine and technology, appendicitis
fatalities remain significant [28]. The small entrance to
this dead-end pocket makes the appendix difficult to
clean out and prone to physical blockage, a detriment
which 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, indicating
that the appendix is unusually prone to abnormal prolif-
eration of its lymphoid tissue [29]. Such an occurrence
would be much less problematic if the interior of the ap-
pendix were not so small, confined, and inaccessible from
the rest of the gut. In many other primates and mammals,
the GALT lymphoid tissue appears to function without
difficulty in a much more open, bulbous caecum with
ample 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 that
the appendix is actually maladaptive, and that the lym-
phoid tissue contained in the appendix is prone to chron-
ic pathological inflammatory states. Unless the appendix
does in fact have an important function that we have yet
to 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, is
indeed poorly designed. The vermiform appendix, in its
present 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 GALT
for the intestinal immune system or by re-inoculating the
colonic flora through bacterial growth, yet it can just as
easily trigger life-threatening conditions. This balance of
exaptive benefits and vestigial detriments is directly re-
lated to an individual’s ability to survive and reproduce
more offspring of our species. Further complicating this
balance is the fact that selective pressures have been
known 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 common
threats to individual survival, rendering many of the or-
gan’s primordial functions obsolete. Shortly thereafter,
the detrimental properties of the appendix became more
prominent. However, civilization can further override
certain evolutionary mechanisms with medicine, facili-
Eur Surg Res 2009;43:67–71 69
tating the survival and procreation of those individuals
who would not have survived alone in nature (e.g. diabe-
tes). Regarding the appendix, modern surgical interven-
tion has saved many lives, enabling affected individuals,
who would have died in an earlier era when the surgery
was not available, to survive and reproduce. Therefore,
the human appendix is a prime example of how evolu-
tionary history can explain the onset of a common surgi-
cal condition; this interpretation of acute appendicitis
ethiopathogenesis is a perfect example of Darwinian
medicine.
Darwinian medicine (or evolution-based medicine) is
the study of evolutionary selective forces that have result-
ed in human vulnerability to disease [30, 31]. As a term,
it was used in the early 1990s, most notably by the evolu-
tionary biologist George C. Williams and the physician
Randolph Nesse, to apply the principles of evolutionary
biology to medicine [32]. Since ‘… nothing in biology
makes any sense except in the light of evolution’ [33],
Darwinian Medicine tends to stray slightly from the tra-
ditional approach to medicine, focusing instead on the
proximate causes of disease, asking questions such as
‘what?’ and ‘how?’, with an approach that emphasizes the
ultimate, or evolutionary, origin of the disease, ‘why?’.
Evolutionary explanations of illness fall into a number of
categories, such as conflicts with other organisms, evolu-
tionary constraints, environmental factors, trade-offs,
and evolved defences. This adaptationist approach can
benefit our efforts to combat disease and improve our ef-
fectiveness as clinicians, researchers and educators [34,
35].
Even in the field of surgery, as the example of the ver-
miform appendix demonstrates, the evolution-based ap-
proach allows us to better understand the remote causes
of disease. In particular, an evolutionary perspective is
crucial in acknowledging that the human being is not a
perfectly designed machine, but rather the imperfect and
poorly designed result of evolutionary history. This un-
derstanding is pivotal for surgery, where surgical inter-
ventions are often required to remedy those biological
shortcomings induced by evolutionary happenstance
[36]. Other common surgical conditions can be interpret-
ed through evolution-based surgery. For example, the
predisposition of male H. sapiens for inguinal hernias is
ultimately attributable to the evolutionary shift from an
aquatic fish body to that of a land-dwelling mammal. Fish
have gonads that extend toward their chest, approaching
the heart, yet this structure is not possible in mammals
where sperm cells require a specific range of tempera-
tures to develop correctly for their 3 months of life: too
70 Eur Surg Res 2009;43:67–71
hot, and the sperm are malformed; too cold, and they die.
Male mammals have an ingenuitive mechanism for con-
trolling the temperature of the sperm-making apparatus:
the scrotum, in which the gonads are positioned outside
of the body. The disadvantage of this structure is that the
biological plumbing that carries sperm to the penis is cir-
cuitous. Sperm travel from the testes in the scrotum
through the sperm cord that leaves the scrotum, travels
up towards the waist, loops over the pelvis, then goes
through the pelvis to travel along the penis. The reason
for this absurdly complex route lies in our developmental
and evolutionary history: in the human male, testes de-
velop initially within the abdomen and later during ges-
tation, they migrate through the abdominal wall into the
scrotum. This causes 2 weak points in the abdominal wall
where hernias can later form. This weakness in the body
wall means that guts can escape the body cavity, squeez-
ing together and lying alongside the spermatic cord.
Men’s tendency to develop hernias illustrates a trade-off
between our fish ancestry and our current mammalian
physiology. Prior to modern surgical techniques, compli-
cations from hernias, including intestinal blockage, gan-
grene, etc., usually resulted in death [37].
The year 2009 sees the bicentenary of Darwin’s birth
and the 150th anniversary of the publication of his semi-
nal work, On the Origin of Species. Perhaps, to commem-
orate these historic anniversaries, we could all consider
the significance of evolution-based surgery in modern
medicine.
Dr. Luca Ansaloni, MD,
Dr. Fausto Catena, MD,
Members of the editorial board, together with
Prof. Antonio Daniele Pinna, MD, FACS
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