A lymphatic gland is a lymph node; a Lieberkühnian gland is an intestinal crypt; a Brunner gland is a predominantly mucous duodenal gland. The first is a lymphoid organ with traversing lymphatic channels; the other two are epithelial glands opening towards the intestinal lumen. Similar historical terminology must not erase that difference.
A lymph node has a fibrous connective tissue capsule sending trabeculae into the organ and an internal reticular framework supporting mobile cells. Its outer cortex contains lymphoid follicles, whose appearance changes with immune activity; activated secondary follicles have paler germinal centres surrounded by more densely packed lymphocytes. The deeper cortex or paracortex lies between follicular cortex and medulla. The medulla contains branching cellular cords separated by medullary sinuses. A hilum provides the entry and exit of the main blood vessels and the exit of efferent lymphatics.
Afferent lymphatic vessels enter at the convex surface and deliver lymph to the subcapsular sinus. Flow then passes through cortical or trabecular channels into medullary sinuses and out through efferent vessels near the hilum. Macrophages and the reticular arrangement bring carried material into contact with the cellular apparatus. The node's blood circulation is distinct from that lymphatic route. In modern terms, follicles are predominantly B cell territories, the paracortex is rich in T cells, and medullary cords contain many antibody-producing plasma cells. Those cellular classifications go beyond an ordinary 1865 microscopic description.[ref]
Histology, Lymph Nodes (StatPearls). Structure; Function; Histochemistry and Cytochemistry: cortex, paracortex and medulla
Thin stained sections show the overall architecture, while a silver stain highlights the supporting reticular fibres rather than every lymphocyte. The fibres include type III collagen; the thicker capsule has a different connective-tissue organization. One must not interpret the meshwork itself as an epithelial secretory tube, or assume a section missing the hilum proves that the node has no efferent vessel.[ref]
Lymphatic Tissues: Lymph Nodes by Columbia University. Slides 22 and 31: silver-stained reticulum, sinuses, capsule, hilum and postcapillary venules
The intestinal crypts are simple tubular epithelial invaginations opening between the intestinal villi in the small intestine. Their bases approach the muscularis mucosae, remaining within the mucosa and surrounded by lamina propria. Each gland has a lumen continuous with the intestinal surface and a lining supported by a basement membrane. It contains columnar epithelial cells at different stages of differentiation, goblet cells producing mucus, endocrine cells and, in small-intestinal crypt bases, conspicuous granular Paneth cells. A proliferative compartment supplies replacement cells that move towards the surface; this modern cell-renewal account explains why dividing cells are common in the crypts. Crypts also occur in the colon, whose mucosa lacks ordinary small-intestinal villi. One should therefore specify the region rather than make villi part of every crypt's definition.
A Brunner gland lies chiefly in the submucosa of the proximal duodenum, below the muscularis mucosae. It is a branched, compound tubular gland with pale mucus-rich secretory cells, basal nuclei and small lumina. Its ducts pass through the muscularis mucosae and commonly join overlying intestinal crypts before opening into the lumen. Its alkaline, mucus-rich secretion helps protect the mucosa from acidic gastric contents. The position below the muscularis mucosae distinguishes it particularly well from a Lieberkühn gland in the mucosa.[ref]
GI Tract Notes by Duke University. Small Intestine; regional differences: crypts, cell types and submucosal Brunner glands with ducts joining crypts
The most useful comparison is consequently one of compartments: a reticular lymphoid organ containing sinuses, a mucosal epithelial tube, and a branched submucosal secretory system. Recognition of the capsule, epithelial lining, muscularis mucosae and duct connections establishes those distinctions more securely than a name alone.
The principal intestinal Entozoa to describe are roundworms, threadworms, whipworms and tapeworms; the chief visceral larval form is the hydatid. “In this country” refers to the British setting of the paper. It is not a claim that all human helminths were common or acquired locally. A contemporary account already distinguished Nematoda, Cestoda and Trematoda, and recognised that an adult worm and a larval cyst could be different phases of one life cycle.[ref]
Entozoa, an Introduction to the Study of Helminthology by Thomas Spencer Cobbold. 1864; systematic treatments of human nematodes, cestodes and trematodes
The life cycles below use present knowledge to explain transmission; their inclusion does not imply that every migratory stage or intermediate host was established in 1865.Ascaris lumbricoides lives as a large, cylindrical, unsegmented, tapering roundworm in the small intestine; adults are cream-coloured or pinkish and roughly 15–35 cm long, with the male generally smaller and its tail curved. Eggs leave in faeces and become infective after development in the environment. After ingestion, larvae cross the intestinal wall, pass through the liver and lungs, ascend the airway, are swallowed and mature in the small intestine. Adults can wander into the bile ducts. The pulmonary migration belongs to the larval history, not to the usual location of the adult.[ref]
DPDx: Ascariasis. Life cycle; adult morphology
Enterobius vermicularis, the threadworm, is a small white nematode of the caecal and adjacent colonic region. The female is about 8–13 mm long and has a conspicuous pointed tail; the male is shorter. The gravid female migrates to the perianal skin and deposits eggs, often at night, so itching and transfer by contaminated fingers favour renewed infection. Eggs become infective rapidly; a pulmonary migration is not part of its usual life cycle. This explains why a perianal adhesive sample can be more useful than an ordinary stool examination.[ref]
DPDx: Enterobiasis. Life cycle; adult worms; diagnostic findings
Trichuris trichiura, historically Trichocephalus dispar, inhabits the caecum and colon. The adult is about 3–5 cm long, with a very slender anterior portion embedded in the mucous membrane and a thicker posterior portion projecting into the lumen: the outline resembles a whip. Eggs pass in faeces, mature in soil and infect after ingestion. The adult shape and the characteristic eggs with polar plugs distinguish it from the preceding worms.[ref]
DPDx: Trichuriasis. Life cycle and morphology
Adult Taenia are flat, ribbon-like tapeworms, several metres long, with a small attachment head or scolex, a narrow neck and a chain of proglottids. Taenia solium has four suckers and a hooked rostellum; Taenia saginata, also called Taenia mediocanellata in period literature, has four suckers but lacks that crown of hooks. Their larval cysticerci occur in pig and cattle tissues respectively; ingestion of inadequately cooked infected meat permits an adult to develop in the human small intestine. Eggs and segments pass in faeces. Egg appearance alone does not reliably distinguish these two species.[ref]
For Taenia solium, swallowing eggs is a different event from swallowing pork cysticerci: eggs can establish tissue cysticercosis in humans, including abdominal tissues, whereas pork larvae cause intestinal taeniasis. A cysticercus is a small fluid-filled bladder with an invaginated scolex, not a miniature chain of segments.[ref]
DPDx: Cysticercosis. Life cycle and tissue cyst morphology
The hydatid of Echinococcus granulosus is particularly important in the liver. Humans acquire eggs from contamination associated with infected dogs and act as intermediate hosts. Larvae reach viscera and form a fluid-filled cyst with a laminated wall and an inner germinal layer; brood capsules, protoscolices and sometimes daughter cysts develop inside. The tiny adult tapeworm belongs in the dog's intestine, not as the normal human liver worm. Dogs complete the cycle by eating infected viscera from an intermediate host.[ref]
Less usual examples deserve proportionate mention. Diphyllobothrium latum is a broad, segmented fish tapeworm with two longitudinal attachment grooves rather than four suckers; aquatic intermediate hosts and infected fish link it to humans. It should not be presented as a uniformly common British infection.[ref]
DPDx: Diphyllobothriasis. Life cycle; adult morphology
Fasciola hepatica, a leaf-shaped fluke roughly 2–3 cm long with oral and ventral suckers, can inhabit the bile ducts; its cycle requires a freshwater snail and infective stages on aquatic vegetation. Human disease was an unusual counterpart of an important infection of grazing animals.[ref]DPDx: Fascioliasis. Life cycle and adult morphology
Trichinella spiralis has minute intestinal adults after ingestion of infected meat, but its enduring encysted larvae occupy skeletal muscle; those muscle cysts should not be misdescribed as the typical abdominal hydatid.[ref]Gross form and location guide identification, but microscopy of eggs, larvae or segments and knowledge of the life cycle supply the confirmation.
The two pneumogastric nerves are the right and left vagus nerves, the tenth cranial nerves: mixed sensory, branchial motor and parasympathetic pathways from the medulla oblongata to the neck, thorax and abdomen. A useful description separates their apparent origin at the surface of the brain from the origins and terminations of their component fibres.
Rootlets emerge in the groove behind the inferior olive, between the glossopharyngeal nerve above and the accessory rootlets below. Branchial motor fibres arise chiefly in the nucleus ambiguus; preganglionic parasympathetic fibres arise in the dorsal motor nucleus of the vagus and, particularly for cardiac control, the nucleus ambiguus. Visceral sensory fibres have their cell bodies chiefly in the inferior vagal ganglion and terminate centrally in the nucleus of the solitary tract; somatic sensory fibres associated with the superior vagal ganglion project to the spinal trigeminal nucleus. Consequently a sensory fibre is not accurately described as originating in a motor nucleus simply because its peripheral nerve emerges nearby.[ref]
Neuroanatomy, Nucleus Ambiguus (StatPearls). Introduction and Structure and Function: branchial motor output and cardiac vagal control
Each vagus nerve leaves the skull through the jugular foramen, with the glossopharyngeal nerve and accessory nerve. It bears the small superior vagal ganglion at the foramen and the larger inferior vagal ganglion below it. Connections include communications with adjacent cranial nerves and the cervical sympathetic nervous system. Traditional descriptions assign fibres joining the vagus nerve from the cranial part of the accessory nerve to the latter peripheral nerve; their modern classification is more nuanced and does not change the peripheral branches being described. In the neck the vagus nerve descends in the carotid sheath, behind and between the internal jugular vein and the internal, then common, carotid artery.
Its meningeal and auricular branches supply small sensory territories of the cranial covering and external ear. Pharyngeal branches contribute to the pharyngeal plexus, supplying much of the soft palate and pharynx; the stylopharyngeus muscle belongs to the glossopharyngeal nerve and the tensor veli palatini muscle to the mandibular division of the trigeminal nerve. The internal branch of the superior laryngeal nerve supplies sensation above the vocal folds; its external branch supplies the cricothyroid muscle. The recurrent laryngeal nerve supplies the other intrinsic laryngeal muscles and sensation below the vocal folds. The right recurrent laryngeal nerve loops beneath the right subclavian artery; the left loops under the aortic arch beside the ligamentum arteriosum. Both return towards the larynx near the groove between trachea and oesophagus.[ref]
Neuroanatomy, Cranial Nerve 10 (Vagus Nerve) (StatPearls). Structure and Function; Vagus Nerve Branches; Muscles
In the thorax, both vagus nerves pass behind the roots of the lungs, contrasting with the anterior course of the phrenic nerves. Cardiac branches join the cardiac plexus; pulmonary branches join the pulmonary plexus. The trunks break up into the oesophageal plexus, then collect principally as an anterior trunk derived from the left vagus nerve and a posterior trunk derived from the right. Through the oesophageal hiatus they reach the stomach; hepatic and coeliac branches distribute to abdominal viscera. Their parasympathetic territory includes foregut and midgut derivatives, extending into the proximal colon, rather than every abdominal and pelvic organ.[ref]
Neuroanatomy, Cranial Nerve 10 (Vagus Nerve) (StatPearls). Structure and Function: thoracic course and oesophageal plexus
This extensive distribution explains why the level of experimental section is indispensable information.The main consequences of section can be organized by the interrupted functions:
- Heart: cutting both cervical vagus nerves removes tonic inhibition and usually increases heart rate. The heart continues beating because its rhythm is intrinsic. Electrical stimulation of the peripheral cut end can slow the heart, whereas stimulation of the central end tests reflex effects through the brainstem. Cutting a pathway and stimulating its intact output are different experiments.
- Breathing: bilateral cervical section commonly produces slower, deeper breaths in experimental mammals. Sensory information from the lungs has been interrupted; the respiratory diaphragm is not directly paralysed, because its motor peripheral nerves are the phrenic nerves. Persistence of breathing establishes that the vagus nerves modify the respiratory rhythm rather than being its sole generator. The subsequently named Hering-Breuer inflation reflex provides a later explanation of one component of this feedback; its formal description belongs to 1868, after this examination.[ref]Reflexes from the lungs and airways: historical perspective. Discussion of the 1868 lung inflation and deflation experiments
- Larynx and lungs: section proximal to the recurrent laryngeal nerves can impair opening and closure of the vocal folds, causing altered voice, obstruction and impaired airway protection. Classical bilateral cervical experiments sometimes ended in death with pulmonary congestion, oedema or aspiration changes. Such outcomes must be distinguished from an immediate necessary arrest of breathing: species, age, lesion level and airway management affect them.
- Stomach: bilateral section can reduce vagally driven gastric acid secretion and disturb gastric motor coordination and emptying. Nevertheless digestion and nutrient absorption are not abolished as chemical possibilities: enteric nervous system activity, local stimuli and hormones remain. Likewise, interruption of gastric sensory fibres does not establish that the entire experience of hunger originates in them.
A discriminating historical experiment would record respiratory movements and heart rate before and after section, distinguish unilateral from bilateral section, and compare an intact airway with an airway bypass. A tracheal cannula can separate obstruction at the glottis from changes in the respiratory rhythm, while the central-versus-peripheral stimulation comparison separates incoming reflex traffic from outgoing motor effects. Later separate-lung recordings confirmed that a change following unilateral section could affect both lungs, supporting a central reflex interpretation rather than simple local paralysis.[ref]
A Study of the Hering-Breuer Reflex. Summary and Conclusions, points 1–4; separate recordings of the two lungs, 1927
The gastric distinction is also borne out by measurements of persistent acidity after surgical vagotomy, despite major reductions in stimulated gastric acid secretion.[ref]Twenty-four hour gastric acidity after vagotomy. Abstract: hourly intragastric sampling, seven patients, four years after operation