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Clinical Biochemistry of Domestic Animals (Sixth Edition) - UMK ...

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414<br />

Chapter | 14 Gastrointestinal Function<br />

In those species that produce salivary amylase (e.g., pig<br />

and human), the secretory granules are the zymogen<br />

precursors <strong>of</strong> this enzyme. A third cell type is found lining<br />

the striated ducts. The striations along the basal borders<br />

<strong>of</strong> these cells are caused by vertical infoldings <strong>of</strong> the<br />

cell membrane, a characteristic <strong>of</strong> epithelial cells involved<br />

in rapid movement <strong>of</strong> water and electrolytes. The primary<br />

secretion <strong>of</strong> the acinar cells is modified by active transport<br />

processes <strong>of</strong> the ductal epithelium.<br />

The distribution <strong>of</strong> the different types <strong>of</strong> secretory<br />

cells in the salivary glands varies among species. The<br />

parotid glands <strong>of</strong> most animals are serous glands, which<br />

produce a secretion <strong>of</strong> low-specific gravity and osmolality<br />

containing electrolytes and proteins including certain<br />

hydrolytic enzymes. The mandibular (submaxillary) and<br />

sublingual glands are mixed salivary glands that contain<br />

both mucous and serous types <strong>of</strong> cells and produce a more<br />

viscous secretion that contains large amounts <strong>of</strong> mucus<br />

( Dukes, 1955 ).<br />

B . Composition <strong>of</strong> Saliva<br />

1 . Mucus<br />

Mucus is an aqueous mixture <strong>of</strong> proteoglycans and glycoproteins.<br />

One <strong>of</strong> the most completely studied glycoproteins<br />

is mucin. Salivary mucins are O-glycosylated and consist<br />

<strong>of</strong> peptides with many oligosaccharides linked covalently<br />

to the hydroxyamino acid serine or threonine. The carbohydrate<br />

portion <strong>of</strong> submaxillary mucin from sheep is a<br />

disaccharide <strong>of</strong> N-acetylneuraminic acid (sialic acid) and<br />

N-acetylgalactosamine ( Carlson et al., 1973 ). The enzymes<br />

that link protein with hexosamine have been purified from<br />

the mandibular glands <strong>of</strong> sheep ( Carlson et al., 1973 ) and<br />

swine ( Schachter et al., 1971 ).<br />

The physiological functions <strong>of</strong> mucin are related to its<br />

high viscosity. N-acetylneuraminic acid is the component<br />

responsible for the formation <strong>of</strong> viscous aqueous solutions<br />

and, at physiological pH, causes expansion and stiffening <strong>of</strong><br />

the mucin molecule. The resistance <strong>of</strong> mucin to enzymatic<br />

breakdown is also due to the presence <strong>of</strong> disaccharide residues.<br />

Removal <strong>of</strong> terminal N-acetylneuraminic acid residues<br />

by action <strong>of</strong> neuraminidase significantly increases the<br />

susceptibility <strong>of</strong> peptide bonds to trypsin.<br />

2 . Electrolytes<br />

The principal inorganic constituents <strong>of</strong> saliva are sodium,<br />

potassium, chloride, and bicarbonate, which, with the<br />

exception <strong>of</strong> bicarbonate, originate directly from the plasma.<br />

Rates <strong>of</strong> salivary flow vary depending on stimulation, and<br />

there are wide variations in electrolyte concentration. Saliva<br />

is formed by a process that initially requires uptake <strong>of</strong><br />

sodium and other electrolytes from the interstitium <strong>of</strong> the<br />

terminal structural unit <strong>of</strong> the salivary gland, the acinus or<br />

end piece. Water flows passively. This primary or precursor<br />

fluid has a sodium concentration similar to plasma, and<br />

the potassium concentration is similar to or slightly higher<br />

than plasma. As the primary fluid passes from the acinus<br />

along the duct system, the concentration <strong>of</strong> sodium, potassium,<br />

and other electrolytes changes. In most species, there<br />

is net sodium absorption and potassium secretion. Wide<br />

variations in electrolyte composition may occur depending<br />

on the flow rate ( Young and Schneyer, 1981 ), the salivary<br />

gland <strong>of</strong> origin, and the species ( Table 14-1 ).<br />

3 . Amylase<br />

The saliva <strong>of</strong> rodents contains the α -amylase, ptyalin,<br />

but this enzyme activity is absent in the saliva <strong>of</strong> dogs,<br />

cats, horses, cattle, and sheep ( Dukes, 1955 ; Young and<br />

Schneyer, 1981 ). Salivary amylase splits the α 1,4-glucosidic<br />

bonds <strong>of</strong> various polysaccharides. Salivary amylase is<br />

similar in major respects to pancreatic α -amylase, which is<br />

described in Section V.B. Salivary amylase initiates digestion<br />

<strong>of</strong> starch and glycogen in the mouths <strong>of</strong> those species<br />

that secrete the enzyme. The optimal pH for amylase activity<br />

is approximately 7, so that this activity ceases when the<br />

enzyme mixes with acidic gastric contents.<br />

TABLE 14-1 Electrolyte Concentration <strong>of</strong> Mandibular Gland and Parotid Gland Saliva Observed during<br />

Maximum Rates <strong>of</strong> Secretion (mmol/l)<br />

Mandibular Gland Parotid Gland<br />

Na K HCO 3 Na K HCO 3 <br />

Sheep 20 7 23 160–175 9–10 113–140<br />

Dog 70–100 12–15 10–30 80–110 6–14 50<br />

Cat 40–51 9–10 26 — — —<br />

Rabbit 50–100 10–40 25 110–140 10 12–30

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