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Upon completion of this chapter, you should be able to answer the following questions:
What constitutes the functional anatomy of salivary glands, including their secretory elements?
What are the cephalic and oral phases (what, why, how it happens) of the response to a meal?
What are the general principles of secretion along the gastrointestinal (GI) tract (where do secretions come from, what are the components)?
How do the components of secretion vary with the gland or region of the GI tract?
What is the correlation between the composition and functions of salivary secretion?
How are primary and secondary secretions within salivary glands generated and regulated?
What is the sequence of events in swallowing?
What are the stimulus and neural pathways generating primary and secondary esophageal peristalsis?
What changes in gastric motility take place during swallowing, and what is the significance?
What are the major functions of the esophagus and associated structures in terms of protection and propulsion?
This chapter will describe the processes that occur in the gastrointestinal (GI) tract in the early stages of the integrated response to a meal. There are changes in GI tract physiology (1) before food is ingested (the cephalic phase), (2) when ingested food is in the mouth (the oral phase), and (3) when food is transferred from the mouth to the esophagus (the esophageal phase). The responses of the GI tract to the presence of food are mainly associated with preparing the GI tract for digestion and absorption.
The main feature of the cephalic phase is activation of the GI tract in readiness for the meal. The stimuli involved are cognitive and include anticipation or thinking about the consumption of food, olfactory input, visual input (seeing or smelling appetizing food when hungry), and auditory input. The latter may be an unexpected link but was clearly demonstrated in the classic conditioning experiments of Pavlov, in which he paired an auditory stimulus to the presentation of food to dogs; eventually the auditory stimulus alone could stimulate secretion. A real-life analogy is presumably being told that dinner is ready. All these stimuli result in an increase in excitatory parasympathetic neural outflow to the gut. Sensory input (e.g., smell) stimulates sensory nerves that activate parasympathetic outflow from the brainstem. Higher brain sites (e.g., limbic system, hypothalamus, cortex) are also involved in the cognitive components of this response. The response can be both positive and negative; thus, anticipation of palatable food and a person's psychological status, such as anxiety, can alter the cognitive response to a meal. However, the final common pathway is activation of the dorsal motor nucleus in the brainstem, the region where the cell bodies of the vagal preganglionic neurons arise. Activation of the nucleus leads to increased activity in efferent fibers passing to the GI tract in the vagus nerve. In turn the efferent fibers activate the postganglionic motor neurons (referred to as motor because their activation results in change of function of an effector cell). Increased parasympathetic outflow enhances salivary secretion, gastric acid secretion, pancreatic enzyme secretion, gallbladder contraction, and relaxation of the sphincter of Oddi (the sphincter between the common bile duct and duodenum). All these responses enhance the ability of the GI tract to receive and digest the incoming food. The salivary response is mediated via the ninth cranial nerve; the remaining responses are mediated via the vagus nerve.
Many of the features of the oral phase are indistinguishable from the cephalic phase. The only difference is that food is in contact with the surface of the GI tract. Thus, there are additional stimuli generated from the mouth, both mechanical and chemical (taste). However, many of the responses initiated by the presence of food in the oral cavity are identical to those initiated in the cephalic phase, because the efferent pathway is the same. The responses specifically initiated in the mouth, which consist mainly of the stimulation of salivary secretion, will be discussed next.
The mouth is important for the mechanical disruption of food and for initiation of digestion. Chewing subdivides and mixes the food with the enzymes salivary amylase and lingual lipase and with the glycoprotein mucin, which lubricates food for chewing and swallowing. Minimal absorption occurs in the mouth, although alcohol and some drugs are absorbed from the oral cavity, and this can be clinically important. However, as with the cephalic phase, it is important to realize that stimulation of the oral cavity initiates responses in the more distal GI tract, including increased gastric acid secretion, increased pancreatic enzyme secretion, gallbladder contraction, and relaxation of the sphincter of Oddi, mediated via the efferent vagal pathway.
Secretions in the GI tract come from glands associated with the tract (salivary glands, pancreas, and liver), from glands formed by the gut wall itself (e.g., submucosal glands in esophagus and duodenum), and from the intestinal mucosa itself. The exact nature of the secretory products can vary tremendously, depending on the function of that region of the GI tract. However, these secretions have several characteristics in common. Secretions from the GI tract and associated glands include water, electrolytes, protein, and humoral agents. Water is essential for generating an aqueous environment for efficient enzyme action. Secretion of electrolytes is important for generation of osmotic gradients to drive the movement of water. Digestive enzymes in secreted fluid catalyze the breakdown of macronutrients in ingested food. Moreover, many additional proteins secreted along the GI tract have specialized functions, some of which are fairly well understood, such as those of mucin and immunoglobulins, and others that are only just beginning to be understood, such as those of trefoil peptides.
Secretion is initiated by multiple signals associated with the meal, including chemical, osmotic, and mechanical components. Secretion is elicited by the action of specific effector substances called secretagogues acting on secretory cells. Secretagogues work in one of the three ways that have already been described in Chapter 27 —endocrine, paracrine, and neural.
Inorganic secretory components are region or gland specific, depending on the particular conditions required in that part of the GI tract. The inorganic components are electrolytes, including H + and HCO 3 − . Two examples of different secretions include acid (HCl) in the stomach, which is important to activate pepsin and start protein digestion, and HCO 3 − in the duodenum, which neutralizes gastric acid and provides optimal conditions for the action of digestive enzymes in the small intestine.
Organic secretory components are also gland or organ specific and depend on the function of that region of the gut. The organic constituents are enzymes (for digestion), mucin (for lubrication and mucosal protection), and other factors such as growth factors, immunoglobulins, bile acids, and absorptive factors.
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