The Evolution and Development of the Gut. Dr Mike Wride School of Natural Sciences Zoology Department

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1 The Evolution and Development of the Gut Dr Mike Wride School of Natural Sciences Zoology Department

2 The gut? Gut Function and Regulation (Dr. Alan Tuffery) Absorption of nutrients (the digestive system) Structure of the gut: Anatomy Embryological Origin Development Comparative anatomy/development

3 Topics to be covered in today s lecture Early Development - gastrulation (a reminder) Endoderm Gut development and genetics Gut nervous innervation: Hirschsprung s disease Gut evolution

4 Gut anatomy Wikipedia: Anatomy_and_Physiology_of_Animals/The_Gut_and_Digestion

5 Gut ultrastructure: Small intestine Villi

6 Useful texts for gut structure, physiology and development Developmental Biology 8th Ed. Scott S Gilbert. Sinauer. Chapter 15 pages Vertebrates, Comparative Anatomy, Function and Evolution. Kenneth V. Kardong. McGraw Hill 4th International Ed. Chapter 13, pages Animal Physiology. Hill, Wyse, Anderson. 2nd Ed. Chapter 5, pages

7 Gastrulation: a reminder Gastrulation begins between days 14 and 16 of human development and at about E6.5 in the mouse Gastrulation results in formation of the 3 germ layers: mesoderm (e.g. muscle, bone, kidneys), endoderm (e.g. gut) and ectoderm (e.g. nervous system/epidermis)

8 Gastrulation cont... Figure Gilbert Chapter 11, page 354 Endoderm forms gut is derived from endoderm

9 Endoderm 1: Induces several mesodermal tissues (notochord, heart, blood vessels, germ layer) II: Constructs linings of 2 tubes of vertebrate body plan: Digestive tube and respiratory tube i.e. both derived from primitive gut Buds from digestive tube form: liver, gallbladder, pancreas Region of digestive tube anterior to respiratory tube: called the pharynx

10 Formation of the Human Digestive System at (A) 16 days (B) 18 days

11 Formation of the Human Digestive System at (C) 22 days (D) 28 days

12 Development of the Pharynx

13 The digestive tube and its derivatives Regional specification of the gut endoderm and its derivatives Involves reciprocal interactions between endoderm and mesoderm does this ring a bell?

14 Regional specification of gut Digestive tube meets different kinds of mesenchyme (splanchnic lateral plate mesoderm) at different levels (rostral-caudal; head-tail) Digestive tube differentiates into different kinds of structures: oesophagus, stomach, small intestine, large intestine and colon There is an emerging consensus on the steps leading to regional specification of gut tube

15

16 The role of Barx1 in gut development Kim BM, Buchner G, Miletich I, Sharpe PT, Shivdasani RA. The stomach mesenchymal transcription factor Barx1 specifies gastric epithelial identity through inhibition of transient Wnt signaling. Dev Cell Apr;8(4): /- Barx1 knockout mouse

17 BMP signaling influences development of muscle types Theodosiou NA, Tabin CJ. Sox9 and Nkx2.5 determine the pyloric sphincter epithelium under the control of BMP signaling. Dev Biol Mar 15;279(2):

18 Liver pancreas and gall bladder

19 Nervous innervation of the gut - Hirschsprung s disease Enteric nervous system (ENS): provides intrinsic innervation of the gastrointestinal tract Controls essential functions such as motility, secretion and blood flow Comprises a vast number of neurons and glial cells organized into complex networks of interconnected ganglia distributed throughout the entire length of the gut wall Hirschsprung s disease (congenital aganglionic megacolon): 1in 5000 babies - enlargement of colon caused by bowel obstruction resulting from an aganglionic section of bowel (the normal enteric nerves are absent)

20 Arrangement of the ENS Farlie et al Birth Defects Research (Part C) 72: (2004)

21 The Neural Crest Enteric neurons and glia - derived from neural crest cells (undergo extensive migration, proliferation, differentiation and survival to form a functional ENS).

22 Migration paths of neural crest cells to colonize the gut Farlie et al (2004)

23 Farlie et al (2004) Holschneider and Puri (2008)

24 Comparative Gut Anatomy/Evolution Symbiosis with heterotrophic microbes - provide metabolic capabilities that the animal lacks Include anaerobic bacteria, protists, yeasts, fungi carrying out fermentation Development of specialised enlarged/ dilated gut chambers during evolution

25 Foregut fermenters: ruminant mammals 1. fermentation - of non digestible compounds 2. Synthesis of B vitamins and essential amino acids 3. Recycling of waste nitrogen (urea) to make proteins

26 Hindgut fermenters Similar to foregut fermenters in terms of fermentation but vitamin synthesis and nitrogen recycling do not occur to the same extent

27 Variations in the stomach and intestines of lower vertebrates and birds

28 Variations in the stomach and intestines of mammals

29 Stomachs of Various Vertebrates

30 Summary of Today s Lecture Early Development - gastrulation (a reminder) Endoderm Gut development and genetics gut nervous innervation: Hirschsprung s disease Gut evolution

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