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38 Mesenteric and peritoneal anatomy
bearing continuity in mind. At the intestinal margin of the conuence, the intestinal tract is seen to round the ex­ure, centered on the mesentery [10,16,20,21]. is complex of anatomic structures and pattern of attachment is again obscured from direct visualization by the right peritoneal and hepatocolic regions of the peritoneal reection.
Splenic exure
e reverse occurs at the splenic exure where the colon changes from being mobile (transverse colon) to attached (descending colon) to the posterior abdominal wall. e mes­enteric component of the conuence changes from having a mobile component at the mesenteric border of the intestinal tract to fully attaching to the posterior abdominal wall across the transverse extent of the lemesocolon. is anatomic arrangement is obscured from direct view by the splenocolic and le peritoneal regions of the peritoneal reection.
Colosigmoid and rectosigmoid exures
At the junction between the descending and sigmoid colon, the intestinal margin of the mesentery separates from the posterior abdominal wall and elongates considerably in tan­dem with the colon itself. At the exure between the rectum and sigmoid, the reverse occurs as the intestinal margin of the mesentery tapers distally, becomes adherent, and continues caudally as the mesorectum.
MESENTERIC CONFORMATION INGENERAL
All anatomic structures have limits (i.e., a beginning and an end). e mesentery distal to the duodenum emerges from the superior mesenteric root region. e anatomic end, or termination, occurs at the distal end of the mesorectum. In between these, the mesentery is best understood as fan­ning out and adopting a spiral conformation. In addition, it undergoes an extensive elongation at the intestinal margin. is combination, in conjunction with plication, ensures that despite extensive elongation at the intestinal margin, the entire gastromesenteric complex can be neatly packaged into a limited intraperitoneal space.
FUTURE DIRECTIONS
An improved understanding of mesenteric and peritoneal anatomy enables one to propose areas for future research, of which there are many. To the surgeon involved in excising the rectum, it remains unclear as to how, structurally, vessels and nerves interface with the mesorectum. Some propose the pres­ence of lateral ligaments or the “zone of adhesion,” while oth­ers demonstrate that the mesorectum remains separate from surrounding structures, throughout its full circumference.
Splanchnic nerves supply postganglionic parasympa-
thetic and sympathetic nerve bers to the intestinal tract.
Although the distribution of these nerves is of enormous clinical relevance, little is known regarding their trajectory within the mesenteric organ [21]. Most descriptions men­tion three ganglia from which postganglionic bers arise, but none accurately delineate the course of these, once they leave the ganglia. It is not unreasonable to suggest that the mesentery provides a crucial platform in this regard and that the lack of comprehensive descriptions of enteric neuroanatomy stems from the inaccurate appraisals of mes­enteric anatomy in the rst instance. It will be important to redress this in future studies and accurately determine the course of postganglionic nerves once these have le the celiac, superior or inferior mesenteric ganglia.
SUMMARY
e mesenteric organ, distal to the duodenojejunal ex­ure, fans out from its origin at the superior mesenteric artery to span the intestinal tract from DJ exure to ano­rectal junction[2]. It is continuous, as are the associated intestinal tract and peritoneal reections. Anatomic conti­nuity has implications for all related basic and clinical dis­ciplines. Clarication of mesenteric, peritoneal and fascial anatomy now enables a systematic (i.e., scientic) study of each [21].
REFERENCES
1. Treves, F., Lectures on the anatomy of the intesti­nal canal and peritoneum in man. Br Med J, 1885. 1(1264): 580–583.
2. Standring, S., Gray’s Anatomy: The Anatomical Basis of Clinical Practice. Elsevier Health Sciences, London, U. K ., 2015, pp. 10 9 8 –1111, 1124–1160.
3. Blackburn, S.C. and M.P. Stanton, Anatomy and physiology of the peritoneum. Semin Pediatr Surg,
2014. 23(6): 326–330.
4. Sinnatamby, C.S., Last’s Anatomy: Regional and Applied. Elsevier Health Sciences, London, U.K., 2011, pp. 234–238, 247–259.
5. Snell, R.S., Clinical Anatomy by Regions. Lippincott Williams & Wilkins, 2008, pp. 216, 226, 228, 233, 237,
240.
6. Schoenwolf, G.C. etal., Larsen’s Human Embryology. Elsevier Health Sciences, Philadelphia, PA, 2014, pp.341–375.
7. Cochard, L.R., Netter’s Atlas of Human Embryology: Updated Edition. Elsevier Health Sciences, London, U.K., 2012, pp. 133–134.
8. Sadler, T.W., Langman’s Medical Embryology. Wolters Kluwer Health, Philadelphia, PA, 2011, pp.208–232.
9. Sehgal, R. and J.C. Coffey, The development of consensus for complete mesocolic excision (CME) should commence with standardisation of anatomy and related terminology. Int J Colorectal Dis, 2014. 29(6): 763–764.
References 39
10. Culligan, K. etal., The mesocolon: A prospective
observational study. Colorectal Dis, 2012. 14(4): 421–428; discussion 428–430.
11. Coffey, J.C., Surgical anatomy and ana­tomic surgery—Clinical and scientic mutualism. Surgeon, 2013. 11(4): 177–182.
12. Treves, F., Discussion on the subsequent course and later history of cases of appendicitis after operation. Med Chir Trans, 1905. 88: 429– 610.
13. Standring, S., Gray’s Anatomy: The Anatomical Basis of Clinical Practice. Churchill Livingstone/ Elsevier, Edinburgh, Scotland, 2008, pp. 1069–1083, 10 9 9 –1111, 1125 –1163.
14. Moore, K.L., A.F. Dalley, and A.M.R. Agur, Clinically Oriented Anatomy. Wolters Kluwer Health, Philadelphia, PA, 2013, pp. 219–221, 239–263.
15. McConnell, A.A. and T.H. Garratt, Abnormalities of x­ation of the ascending colon: The relation of symptoms to anatomical ndings. Br J Surg, 1923. 10: 532–557.
16. Culligan, K. etal., The mesocolon: A histological and electron microscopic characterization of the mesen­teric attachment of the colon prior to and after surgi­cal mobilization. Ann Surg, 2014. 260(6): 1048–1056.
17. Coffey, J.C. etal., An appraisal of the computed axial tomographic appearance of the human mesentery based on mesenteric contiguity from the duodenojejunal exure to the mesorectal level. EurRadiol, 2016. 26(3): 714–721.
18. Coffey, J.C. etal., Mesenteric-based surgery exploits gastrointestinal, peritoneal, mesenteric and fascial continuity from duodenojejunal exure to the anorectal junction—A review. Dig Surg, 2015. 32(4): 291–300.
19. Culligan, K. etal., Review of nomenclature in colonic surgery—Proposal of a standardised nomencla­ture based on mesocolic anatomy. Surgeon, 2013. 11(1):1–5.
20. Coffey, J.C. etal., Terminology and nomencla­ture in colonic surgery: Universal application of a rule-based approach derived from updates on mesenteric anatomy. Tech Coloproctol, 2014. 18(9): 789–794.
21. Coffey, J.C. and D.P. O’Leary, The mesentery: structure, function, and role in disease. Lancet Gastroenterol Hepatol, 2016. 1(3): 2 3 8 –247.
Embryologic development of the mesentery, peritoneal reection, and Toldt’s fascia
J. CALVIN COFFEY, RISHABH SEHGAL, AND JOEP KNOL
3
Aim 41 Introduction 41 Part I: Classic teaching on mesenteric anatomy 41
Current teaching on mesenteric anatomy 42 Mesenteric rotation, anchorage, andelongation 42 Attachment: The peritoneal reection 42 Attachment: Toldt’s fascia 43 Summary 43
Part II: Reverse engineering gastromesenteric
embryology 43
Mesentery: Reverse engineered 43
Peritoneal reection 43 Reverse engineering the reection: Detachment 43 Hindgut: Shortening and detachment ofthe
mesorectum 44
Being entirely honest with oneself is a good exercise.
Sigmund Freud
AIM
e current interpretation of mesenteric anatomy means that the embryologic development of mesentery and associ­ated structures must be reappraised. e aim of this chapter is to conduct such an appraisal and attempt to reconcile cur­rent anatomic thinking with a plausible series of embryo­logic events.
INTRODUCTION
e embryologic development of the mesentery is a truly astonishing biologic process. e relative lack of pub­lished reports on mesenteric abnormalities indicates that it is highly conserved in man. One of the earliest identi­able forms of mesentery is the dorsal mesentery, and this undergoes a remarkable series of changes in size, shape, and
Reverse engineering the mesorectum 44 Hindgut: Shortening and detachment ofthe
mesosigmoid 44 Reverse engineering the mesosigmoid 44 Hindgut: Shortening and detachment ofthe
leftmesocolon 44 Reverse engineering the left mesocolon 44 Midgut 44 Reverse engineering mesentery between
theduodenojejunal and splenic exure 44
Summary 44
Future directions 45 Summary 45 References 45
orientation during development in and ex utero. e fol- lowing chapter is divided into two parts. In the rst, classic teaching and the current one are contrasted and the pro­cesses involved in the development of mesentery discussed. e second component attempts to develop an explanation as to how the dorsal mesentery might develop into the shape observed in the adult human.
PART I: CLASSIC TEACHING ON MESENTERIC ANATOMY
Classic teaching holds that the mesentery elongates con­siderably and then rotates counterclockwise around the superior mesenteric artery. Rotation is a concept used to explain how the right colon and associated mesentery ulti­mately reach their position in the adult, in the right ank. It is also used to explain how, in the adult, the small intes­tine and associated mesentery end up centrally positioned. When rotation does not occur (i.e., mal or nonrotation), the duodenum and small bowel continue vertically down the right ank. e lecolon lies in the le ank, and the right colon (and mesentery) takes up a position betweenboth.
41
42 Embryologic development of the mesentery, peritoneal reection, and Toldt’s fascia
As it is not known why rotation occurs in the rst instance, the cause of nonrotation is similarly obscure. Interestingly, a study of porcine mesentery demonstrates a similar gas­tromesenteric conformation to that observed in humans with mal or nonrotation. In the porcine context, the small intestinal gastromesenteric context continues vertically downward in the right side of the abdominal cavity. e lecolon and mesocolon are positioned on the leside of the abdomen, and the right colon and mesentery are cen­trally positioned between both [1,2].
Following rotation, the gastromesenteric complex
returns to the peritoneal cavity where components attach to the posterior abdominal wall. Attachment is dened as attening of the mesentery against the posterior abdomi­nal wall. It does not mean “insertion” of the mesentery into the posterior abdominal wall. Attachment is mediated by the development of Toldt’s fascia and the peritoneal reec­tion. In this manner, the intestine and mesentery take up the adult conformation (see earlier discussion). According to classic teaching, the small intestinal, transverse, and sig­moid mesentery persists into adulthood. However, the right and lemesocolon regress and the mesentery becomes dis­continuous. Although the sliding and regression theories were developed to explain right and lemesenteric disap­pearance, neither gained broad acceptance [3–6].
Current teaching on mesenteric anatomy
is is detailed in Chapter 2; however, a brief summary will be provided here. Current teaching holds that the mesentery dis­tal to the duodenojejunal exure is continuous to the meso­rectal level [7–9]. It fans out from the root region to span the intestinal tract from jejunum to anorectal junction. Broadly speaking, it has attached (see denition given above) and nonattached regions. e conformation of the attached region resembles a question mark. e curve of the question mark is made up by zones of attachment of the right, transverse, and lemesocolon. e stem of the question mark is formed by attachment of the lemesocolon, mesosigmoid, and meso­rectum. Nonattached mesentery elongates considerably at the intestinal (nonattached) margin. Together, the intestinal and mesenteric components of the gastromesenteric complex are compactly plicated into the peritoneal cavity.
Crucially, this relatively simplistic shape forms a new structural end point toward which embryologists must now work [9].
Mesenteric rotation, anchorage, andelongation
Relatively little is known regarding the cellular events involved in mesenteric development. Is it reasonable to suggest, however, that rotation, anchorage, elongation, and attachment are important and interrelated processes. ese will be briey discussed as follows.
During development, the mesorectum and rectum are anchored within the developing bony pelvis. is is
crucially important as it means that the developing hindgut is positioned to the leof the center. e vitello-intestinal duct anchors the intestine (and by denition the mesentery) in the midline (i.e., center) anteriorly. e superior and infe­rior mesenteric arteries anchor the mesentery (but not the intestine) in the midline (center) posteriorly. Anchorage of the hindgut to the leand the midgut in the center means that these are slightly oset in anatomic terms. e eects of anchorage at both ends means elongation between these leads to the adoption of a spiral conformation. It is illus­trated by taking a tube, xing it at both ends, then elongat­ing at between. e tube will curl in a spiral conformation. When this curling is viewed front on (i.e., anteriorly), it leads to the impression of “rotation.” As a result of elonga­tion between points of anchorage, the right colomesenteric (distal midgut) complex takes up position on the right side of the peritoneal cavity, the small bowel and associated mes­entery (proximal midgut) are centrally positioned, and the le colomesenteric (hindgut) complex remains on the le.
Elongation of the mid and hindgut occurs at dierent
rates. is means that the adult midgut remnant (i.e., small intestine, right colon, and proximal transverse) is consider­ably longer than the hindgut remnant (distal transverse and le side of large bowel).
Abnormalities in the process of elongation and the adop­tion of a spiral conformation are referred to as mal and non-rotation and can have devastating consequences (see
Chapter 7).
Attachment: The peritoneal reection
A further process must occur once the gastromesenteric complex has assumed its position within the abdominal cavity, that is, attachment. At present, this is understood to comprise two major events, the development of the perito­neal reection and Toldt’s fascia.
When the mesentery returns to its nal adult position a gap is present between the surface of the mesentery and the abdominal wall. In the adult, this gap is bridged by an extensive peritoneal reection, a layer of mesothelium that extends from the mesenteric surface (or intestinal surface) to the abdominal wall. e peritoneal reection occurs at the base of the small intestinal region of mesentery, where it attaches to the posterior abdominal wall. It continues around the ileocecal junction and back up around the right colon as the right peritoneal reection. From there, it continues around the upper surface of the hepatic exure. Itthen continues across the upper surface of the transverse colon, between this and the greater omentum. At the splenic exure, it continues as the splenocolic reection, which in turn continues along the lateral aspect of the lecolon as the leperitoneal reection. From here, the reection con­tinues into the pelvis as the lepararectal pelvic reection. A peritoneal reection also occurs on the right side of the mesosigmoid and continues into the pelvis as the right pararectal reection. e right and leperitoneal reection coalesce as the anterior reection in the pouch of Douglas.
Part II: Reverse engineering gastromesenteric embryology 43
Little is known regarding the embryologic development
of the peritoneal reection. ere are no reports in the liter­ature, of congenital absence of the reection. Even in nonro­tation, it is still present (albeit in a dierent conformation). Given the lack of valid data related to reection develop­ment, any statements made on the topic are highly specu­lative. It seems reasonable, however, to suggest how the reection develops once the gastromesenteric complex has adopted a nal intraperitoneal position and that it serves to secure the gastromesenteric complex in position. A sec­ondary and perhaps coincidental benet is that it limits the spread of future disease.
Attachment: Toldt’s fascia
Where the mesocolon and mesentery are attached to the retroperitoneum a fascial layer arises (i.e., Toldt’s fascia). e cellular and molecular basis of this process have not been determined although histologic composition provides important clues to both [10]. e fascia extends beneath the colon where this is apposed to the retroperitoneum (i.e., on the le and right side). It occupies a potential space between the mesentery and retroperitoneum. e peritoneal reec­tion is the anatomic limit of this space and the fascia [8].
Whenever two mesothelial surfaces come into direct
and prolonged contact, Toldt’s fascia develops between and bridges both. Movement between peritoneal surfaces (or lack of) may be a key determinant in the process and may explain the lack of fascial adhesion between pleural mesothelial surfaces and between intestinal mesothelial surfaces (outwith the context of previous abdominal sur­gery or trauma).
Pronounced similarities occur between Toldt’s fascia and intra-abdominal adhesions and may point to overlapping cellular and molecular processes. Adhesions can be areolar in appearance. Toldt’s fascia is similarly areolar and imsy within the pelvis. Alternatively, adhesions can be lmy, thus closely resembling Toldt’s fascia underlying the right and lemesocolon. Finally, adhesions can be dense and vascu­lar in nature, as can Toldt’s fascia in disease settings such as Crohn’s disease [7]. us, it is feasible that similar develop­mental processes underpin adhesion and fascia formation and that prolonged direct contact between two mesothelial surfaces is an important determinant.
Summary
A question also arises as to why the gastromesenteric com­plex attaches to the retroperitoneum in the rst place. Impaired attachment (coupled with other factors) predis­poses to volvulus formation. Volvulus is where the intes­tine and associated mesentery twist, possibly resulting in occlusion of their blood supply, obstruction, or perfora­tion. In nonrotation, attachment of the right mesocolon is considerably less than that seen in normality. As a result, the small intestine (and associated mesentery) is prone to twist around the superior mesenteric artery. us, it appears
that attachment via the root region alone is potentially life threatening and that additional mechanisms of attachment are necessary in general.
A further point is also important in relation to attach-
ment. If attachment did not occur, then mesenteric elonga­tion would likely follow intestinal elongation. is would occur across the radial breadth of the mesentery and not just at the intestinal margin. An enormous gastromesenteric tissue mass would result [9–11]. It would be hard to see this mass as functional, as if one were to stand upright the non­attached mass would collapse into the pelvis and peristalsis impaired. us, it is feasible that the attached mesenteric conformation observed in humans aids in upright mobi­lization, while a nonrotated and nonattached mesenteric conformation is sucient for species on all fours.
PART II: REVERSE ENGINEERING GASTROMESENTERIC EMBRYOLOGY
Current theory on mesenteric anatomy diers markedly from classic teaching. Mesenteric continuity presents a far more simplistic shape as well as a new structural end point toward to which embryologists must now work. In the fol­lowing, we aim to adopt the processes described in part one, in identifying a sequence of mesenteric-based events that could lead to the adult shape. Although it is highly specula­tive, it represents a rst step in reconciling current anatomic and embryologic thinking.
It would be possible, albeit extremely dicult, to concep­tualize mesenteric development from start to nish. Given that we already know the shape of the nished product (i.e.,the adult mesentery), then it may be easier to work backward from the adult to the embryologic shape.
e following is experimental in so far as we will com­mence with the adult mesentery and work backward to the embryologic dorsal mesentery. Events will be described in terms of the processes explained above, i.e., anchorage, elongation, rotation/curling, and attachment.
Mesentery: Reverse engineered
PERITONEAL REFLECTION
e peritoneal reection bridges the space between mesen­teric surfaces and the posterior abdominal wall. e reection continues laterally between the colon and lateral abdominal wall. e reection is also present between the colon and omentum. In the pelvis, it attaches the rectum to the lateral pelvic side wall and terminates at the anterior reection.
REVERSE ENGINEERING THE REFLECTION: DETACHMENT
e peritoneal reection regresses. is means that perito­neal connections between the surface of the mesentery and the posterior abdominal wall are lost. e reection between the right and lecolon and lateral abdominal wall is lost. e reection between the omentum and transverse colon is also lost. A similar phenomenon occurs in the pelvis.
44 Embryologic development of the mesentery, peritoneal reection, and Toldt’s fascia
e net eect is to release the mesentery, colon, and
rectum from the anatomic mainframe provided by the posterior wall of the abdomen. At this point, the mesen­tery remains attached via (1) Toldt’s fascia and (2)vascular points of suspension.
HINDGUT: SHORTENING AND DETACHMENT OFTHE MESORECTUM
e adult mesorectum is a downward extension of the meso­sigmoid. e major vessel of the mesorectum is the superior rectal artery (a caudal extension of the inferior mesenteric artery once the latter has given othe le colic artery). Atthis point, the mesorectum is attached to the pelvis via the mesorectal fascia (i.e., Toldt’s fascia). Waldeyer’s fascia is a distal condensation of Toldt’s fascia.
REVERSE ENGINEERING THE MESORECTUM
Dissolution of Waldeyer’s and Toldt’s fascia allows complete detachment of the mesorectum from its bony surroundings. e mesorectum and rectum then contract back to meso­sigmoid level. ey continue to occupy a central position in the midline.
HINDGUT: SHORTENING AND DETACHMENT OFTHE MESOSIGMOID
e mesosigmoid is continuous proximally and distally with the lemesocolon and mesorectum, respectively. Intrans­verse section, it is attached to the retroperitoneum medially, while laterally it is nonattached and mobile. Atthis point, in the overall sequence, Toldt’s fascia maintains attach­ment. e major vessel is the superior rectal and associated sigmoidal branches.
REVERSE ENGINEERING THE MESOSIGMOID
Toldt’s fascia beneath the mesosigmoid dissolves allowing complete detachment of the mesosigmoid from the retro­peritoneum. e intestinal margin of the sigmoid mesentery shortens in tandem with the sigmoid itself. e mesosig­moid contracts along vertical and transverse axes and the now mobile mesosigmoid rotates medially to adopt a central position in line with the mesorectum. e inferior mesen­teric artery and vein remain in position as the nal point of attachment of the hindgut.
retroperitoneum. e lemesocolon and colon can now rotate medially to take up a central position in line with the mesosigmoid and mesorectum.
At this point, the hindgut has been fully returned to a central and midline position, entirely detached from sur­rounding structures with exception of at points of vascular suspension.
MIDGUT
e midgut gastromesenteric complex will be considered as a single unit. e root region of the mesentery, where the superior mesentery artery emerges from the pancreas, is a crucially important anatomic landmark. e concept that the mesentery fans out from its root region to span the intes­tinal tract is also important. Finally, the central position of the middle colic and inferior mesenteric artery are relevant as they provide xed points of anchorage. e le side of the mesentery (from splenic exure to the mesorectum) has these, while the right does not.
We propose that during normal mesenteric development, the mesentery fans out from its root region. As it does so, it adopts a spiral conformation centered on the root region. e overall impression (when viewed from in front) is that of rotation. Rotation stops where the middle colic and infe­rior mesenteric vessels exert a break-like eect (i.e., at the level of the splenic exure).
REVERSE ENGINEERING MESENTERY BETWEEN THEDUODENOJEJUNAL AND SPLENIC FLEXURE
Toldt’s fascia, attaching the midgut to the posterior abdomi­nal wall, dissolves. is means the midgut is now free to unwind. If viewed from the front on, unwinding gives the impression of straightening in a clockwise direction. Unwinding occurs as the midgut shortens (i.e., reverse of elongation). It unwinds in a clockwise direction because of the mechanical eect of the middle colic and inferior mes­enteric vessels. As it unwinds, the vitello-intestinal duct reforms and connects the small intestine with the umbilicus in the midline. e process of unwinding clockwise means that the gastromesenteric complex of the midgut takes up a central position, above (but continuous with) the hindgut.
Summary
HINDGUT: SHORTENING AND DETACHMENT OFTHE LEFT MESOCOLON
is is continuous proximally and distally with the trans­verse and sigmoid mesocolon, respectively. e major vessel is the lecolic, a side branch of the inferior mesenteric artery. e lemesocolon is attached throughout its full extent, to the retroperitoneum via Toldt’s fascia.
REVERSE ENGINEERING THE LEFT MESOCOLON
Dissolution of Toldt’s fascia beneath the lemesocolon allows detachment of the mesocolon and colon from the
e following is the sequence of events which, if reversed, explain how the laminar dorsal mesentery transforms into the adult shape:
1. e peritoneal reection regresses.
2. Toldt’s fascia regresses beneath the hindgut mesentery
leaving it entirely detached with the exception of at
vascular points of suspension.
3. e hindgut then shortens and adopts a central position
in the midline.
4. Toldt’s fascia regresses beneath the small intestinal
mesentery and right mesocolon, thereby fully freeing
References 45
the midgut mesentery with exception of at points of vascular suspension (i.e., the superior mesenteric root region).
5. e midgut mesentery and intestine shortens.
6. Shortening causes to the midgut to appear to unwind
clockwise under the mechanical inuence of the hindgut.
7. e midgut mesentery and intestine take up a position
above the hindgut mesentery and intestine.
8. e embryonic dorsal mesentery has reformed.
e reverse of this process may occur as follows:
1. e laminar dorsal mesentery is positioned in the
midline, connected anteriorly via the vitello-intestinal duct and posteriorly via the superior and inferior mesentery vessels.
2. e hindgut mesentery and intestine elongate and then attaches to the le of the midline.
3. e midgut elongates and because of the le- sided position of the hindgut, it takes up a spiral conformation.
4. e midgut adopts a position whereby the right colon and mesentery are located on the right side, with the small bowel and mesentery centrally positioned.
5. Toldt’s fascia attaches the mesentery to the posterior abdominal wall.
6. e peritoneal reection develops to bridge any space between the intestine, mesentery, and abdominal wall (including also the greater omentum).
FUTURE DIRECTIONS
e mesentery and associated intestine are integrated at anatomical and histological levels. e cellular and extracel­lular developmental events that underpin such integration must also be tightly coordinated at multiple levels. is is supported by ndings related to neural crest cells, as well as by vascular, lymphatic, and connective tissue contiguity. As mentioned earlier, the incidence of embryologic mesenteric abnormalities is extremely low, indicating the process is highly conserved. It is remarkable to think that at the intesti­nal margin of the mesentery, a multilayered intestine devel­ops involving as of yet uncharacterized interactions between intestinal endoderm and mesenteric mesoderm. Future studies should aim to better characterize the embryologic development of the intestine, as well as the manner in which this is integrated with that of the mesentery in general.
Future studies should determine the conformation of the
gastromesenteric complex in other species. In particular, com­parisons should be made between species that mobilize on all fours with those that stand upright for prolonged periods.
Finally, online archives of images that depict human embry­ologic development should be reviewed in the context of the current understanding of mesenteric and peritoneal anatomy.
SUMMARY
Classic embryologic appraisals of mesenteric development were based on the concept of it being fragmented and complex, in the adult. As the mesenteric organ distal to the third part of the duodenum is now regarded as continu­ous, the embryologic development of this and associated structures (i.e., peritoneum and intestinal tract) must be reappraised.
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3. Cochard, L.R., Netter’s Atlas of Human Embryology:
Updated Edition. Elsevier Health Sciences, London, U.K., 2012, pp. 131–140.
4. Beck, D.E. etal., The ASCRS Manual of Colon and
Rectal Surgery. Springer, New York, 2014, pp. 1–27.
5. Moore, K.L., T.V.N. Persaud, and M.G. Torchia, The
Developing Human: Clinically Oriented Embryology. Elsevier Health Sciences, London, U.K., 2015, pp.221–233.
6. Schoenwolf, G.C. etal., Larsen’s Human
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Histology of the mesentery

J. CALVIN COFFEY, MIRANDA KIERNAN, AND LEON G. WALSH
4
Aim 47 Introduction 47 Mesenteric mesothelium and Toldt’s fascia: The
mesofascial plane 47 Nomenclature: The visceral andparietal peritoneum 49 Immunohistochemical analysis ofmesenteric
mesothelium 50
Order is the shape upon which beauty depends.
Pearl Buck
AIM
e aim of this chapter is to demonstrate recent ndings related to the histology of the mesentery and associated peritoneum. A secondary aim is to highlight the clinical relevance of these properties.
INTRODUCTION
Remarkably, little data are available on the histology of the mesentery and associated peritoneal reection. Still less is known regarding the clinical relevance of mesen­teric and peritoneal histology. is probably relates to the erroneous concept that the mesentery functions solely as a scaold for nerves and vessels and as a result has attracted little attention [1–3]. Recent investigation has increas­ingly focused on omental and visceral fat leading to the identication of numerous immunologic functions for both [4–8]. Data are emerging that supports endocrine and metabolic roles for adipose tissue in general [9–16]. Similar ndings are emerging in relation to the mesen­tery in particular, with direct links identied between the mesenteric production of C-reactive protein and systemic levels of this in disease states [17–22]. Increasing data indicates that the mesentery is highly active in coordinat­ing local and systemic physiologic events and is far from the inert bystander previously thought. is, coupled with recent clarications of macroscopic anatomy, prompt a
Mesenteric connective tissue lattice 51 Mesothelial cell plasticity 53 Future directions 54 Summary 54 References 54
reappraisal of mesenteric histology and its relevance to clinical practice [23,24].
In order to redress the aforementioned decit, our group
conducted a broad evaluation of mesenteric histology [25]. Full thickness mesenteric biopsies were examined at all mesenteric regions from duodenojejunal exure to the anorectal junction (Figure 4.1). A cadaveric approach was adopted as a similar study in living patients would endanger retroperitoneal organs such as the ureters, gonadal vessels, and duodenum. Samples were examined using a variety of histologic, immunohistochemical, and electron micro­scopic approaches to generate a comprehensive appraisal. is chapter focuses largely on the ndings of this study and their implications across multiple clinical and nonclinical sciences.
MESENTERIC MESOTHELIUM AND TOLDT’S FASCIA: THE MESOFASCIAL PLANE
Mesenteric histologic appearances are remarkably consistent from duodenojejunal to anorectal level (Figure 4.1a through
c). e small intestinal mesentery and mobile component of
the mesosigmoid have medial and lateral surfaces. In keep­ing with the above, both surfaces are covered by mesothe­lium (Figure 4.2). e transverse mesocolon has an upper and lower surface that is lined with mesothelium (Figure
4.3). While the right and lemesocolon have an upper sur-
face that is readily apparent, the undersurface of each is attached to Toldt’s fascia and the underlying retroperito­neum. Again, both surfaces of the right and lemesocolon are lined with mesothelium (Figure 4.4).
47