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Digestion is the mechanical and chemical breakdown of food into smaller components that are more easily absorbed into a blood stream, for instance. Digestion is a form of catabolism: a breakdown of large food molecules to smaller ones.

In mammals, food enters the mouth, being chewed by teeth, with chemical processing beginning with chemicals in the saliva from the salivary glands. This is called mastication. Then it travels down the esophagus into the stomach, where hydrochloric acid kills most contaminating microorganisms and begins mechanical break down of some food (e.g., denaturation of protein), and chemical alteration of some. The hydrochloric acid also has a low pH, which allows enzymes to work more efficiently. After some time (typically an hour or two in humans, 4–6 hours in dogs, somewhat shorter duration in house cats, ...) {{citation}}: Empty citation (help), the resulting thick liquid is called chyme. Chyme will go through the small intestine, where 95% of absorption of nutrients occurs, through the large intestine with waste material eventually being eliminated during defecation.^[1]

Other organisms use different mechanisms to digest food.

Digestive systems

Digestive systems take many, many forms. There is a fundamental distinction between internal and external digestion. External digestion was the first to evolve, and most fungi still rely on it.^[2] In this process, enzymes are secreted into the environment surrounding the organism, where they break down an organic material, and some of the products diffuse back to the organism. Later, animals evolved by rolling into a tube and acquiring internal digestion, which is more efficient because more of the broken down products can be captured, and the chemical environment can be more efficiently controlled.^[3]

Some organisms, including nearly all spiders, simply secrete biotoxins and digestive chemicals (e.g., enzymes) into the extracellular environment prior to ingestion of the consequent "soup". In others, once potential nutrients or food is inside the organism, digestion can be conducted to a vesicle or a sac-like structure, through a tube, or through several specialized organs aimed at making the absorption of nutrients more efficient.

Secretion systems

Bacteria use several systems to obtain nutrients from other organisms in the environments.

Channel transport system

In a channel transport system several proteins form a contiguous channel traversing the inner and outer membranes of the bacteria. It is a simple system, which consists of only three protein subunits: the ABC protein, membrane fusion protein (MFP), and outer membrane protein (OMP)^[specify]. This secretion system transports various molecules, from ions, drugs, to proteins of various sizes (20 - 900 kDa). The molecules secreted vary in size from the small Escherichia coli peptide colicin V, (10 kDa) to the Pseudomonas fluorescens cell adhesion protein LapA of 900 kDa.^[4]

Molecular syringe

One molecular syringe is used through which a bacterium (e.g. certain types of Salmonella, Shigella, Yersinia) can inject proteins into eukaryotic cells. One such mechanism was first discovered in Y. pestis and showed that toxins could be injected directly from the bacterial cytoplasm into the cytoplasm of its host's cells rather than simply be secreted into the extracellular medium.^[5]

Conjugation machinery

The conjugation machinery of some bacteria (and archaeal flagella) is capable of transporting both DNA and proteins. It was discovered in Agrobacterium tumefaciens, which uses this system to introduce the Ti plasmid and proteins into the host, which develops the crown gall (tumor).^[6] The VirB complex of Agrobacterium tumefaciens is the prototypic system.^[7]

The nitrogen fixing Rhizobia are an interesting case, wherein conjugative elements naturally engage in inter-kingdom conjugation. Such elements as the Agrobacterium Ti or Ri plasmids contain elements that can transfer to plant cells. Transferred genes enter the plant cell nucleus and effectively transform the plant cells into factories for the production of opines, which the bacteria use as carbon and energy sources. Infected plant cells form crown gall or root tumors. The Ti and Ri plasmids are thus endosymbionts of the bacteria, which are in turn endosymbionts (or parasites) of the infected plant.

The Ti and Ri plasmids are themselves conjugative. Ti and Ri transfer between bacteria uses an independent system (the tra, or transfer, operon) from that for inter-kingdom transfer (the vir, or virulence, operon). Such transfer creates virulent strains from previously avirulent Agrobacteria.

Release of outer membrane vesicles

In addition to the use of the multiprotein complexes listed above, Gram-negative bacteria possess another method for release of material: the formation of outer membrane vesicles.^[8] Portions of the outer membrane pinch off, forming spherical structures made of a lipid bilayer enclosing periplasmic materials. Vesicles from a number of bacterial species have been found to contain virulence factors, some have immunomodulatory effects, and some can directly adhere to and intoxicate host cells. While release of vesicles has been demonstrated as a general response to stress conditions, the process of loading cargo proteins seems to be selective.^[9]

Gastrovascular cavity

The gastrovascular cavity functions as a stomach in both digestion and the distribution of nutrients to all parts of the body. Extracellular digestion takes place within this central cavity, which is lined with the gastrodermis, the internal layer of epithelium. This cavity has only one opening to the outside that functions as both a mouth and an anus: waste and undigested matter is excreted through the mouth/anus, which can be described as an incomplete gut.

In a plant such as the Venus Flytrap that can make its own food through photosynthesis, it does not eat and digest its prey for the traditional objectives of harvesting energy and carbon, but mines prey primarily for essential nutrients (nitrogen and phosphorus in particular) that are in short supply in its boggy, acidic habitat.^[10]

Phagosome

A phagosome is a vacuole formed around a particle absorbed by phagocytosis. The vacuole is formed by the fusion of the cell membrane around the particle. A phagosome is a cellular compartment in which pathogenic microorganisms can be killed and digested. Phagosomes fuse with lysosomes in their maturation process, forming phagolysosomes. In humans, Entamoeba histolytica can phagocytose red blood cells.^[11]

Specialized organs and behaviors

To aid in the digestion of their food animals evolved organs such as beaks, tongues, teeth, a crop, gizzard, and others.

A Catalina Macaw's seed-shearing beak

Squid beak with ruler for size comparison

Beaks

Macaws primarily eat seeds, nuts, and fruit, using their impressive beaks to open even the toughest seed. First they scratch a thin line with the sharp point of the beak, then they shear the seed open with the sides of the beak.

The mouth of the squid is equipped with a sharp horny beak mainly made of chitin^[12] and cross-linked proteins. It is used to kill and tear prey into manageable pieces. The beak is very robust, but does not contain any minerals, unlike the teeth and jaws of many other organisms, including marine species.^[13] The beak is the only indigestible part of the squid.

Tongue

The tongue is skeletal muscle on the floor of the mouth that manipulates food for chewing (mastication) and swallowing (deglutition). It is sensitive and kept moist by saliva. The underside of the tongue is covered with a smooth mucous membrane. The tongue is utilised to roll food particles into a bolus before being transported down the esophagus through the use of peristalsis. The sublingual region underneath the front of the tongue is a location where the oral mucosa is very thin, and underlain by a plexus of veins. This is an ideal location for introducing certain medications to the body. The sublingual route takes advantage of the highly vascular quality of the oral cavity, and allows for the speedy application of medication into the cardiovascular system, bypassing the gastrointestinal tract.

Teeth

Teeth (singular, tooth) are small whitish structures found in the jaws (or mouths) of many vertebrates that are used to tear, scrape, milk and chew food. Teeth are not made of bone, but rather of tissues of varying density and hardness. The shape of an animal's teeth is related to its diet. For example, plant matter is hard to digest, so herbivores have many molars for chewing.

The teeth of carnivores are shaped to kill and tear meat, using specially shaped canine teeth. Herbivores' teeth are made for grinding food materials, in this case, plant parts.

Crop

A crop, or croup, is a thin-walled expanded portion of the alimentary tract used for the storage of food prior to digestion. In some birds it is an expanded, muscular pouch near the gullet or throat. In adult doves and pigeons, the crop can produce crop milk to feed newly hatched birds.^[14]

Certain insects may have a crop or enlarged esophagus.

Abomasum

Herbivores have evolved cecums (or an abomasum in the case of ruminants). Ruminants have a fore-stomach with four chambers. These are the rumen, reticulum, omasum, and abomasum. In the first two chambers, the rumen and the reticulum, the food is mixed with saliva and separates into layers of solid and liquid material. Solids clump together to form the cud (or bolus). The cud is then regurgitated, chewed slowly to completely mix it with saliva and to break down the particle size.

Fiber, especially cellulose and hemi-cellulose, is primarily broken down into the volatile fatty acids, acetic acid, propionic acid and butyric acid in these chambers (the reticulo-rumen) by microbes: (bacteria, protozoa, and fungi). In the omasum water and many of the inorganic mineral elements are absorbed into the blood stream.

The abomasum is the fourth and final stomach compartment in ruminants. It is a close equivalent of a monogastric stomach (e.g., those in humans or pigs), and digesta is processed here in much the same way. It serves primarily as a site for acid hydrolysis of microbial and dietary protein, preparing these protein sources for further digestion and absorption in the small intestine. Digesta is finally moved into the small intestine, where the digestion and absorption of nutrients occurs. Microbes produced in the reticulo-rumen are also digested in the small intestine.

Specialized behaviors

Regurgitation has been mentioned above under abomasum and crop, referring to crop milk, a secretion from the lining of the crop of pigeons and doves with which the parents feed their young by regurgitation.^[15]

Many sharks have the ability to turn their stomachs inside out and evert it out of their mouths in order to get rid of unwanted contents (perhaps developed as a way to reduce exposure to toxins).

Other animals, such as rabbits and rodents, practice coprophagia behaviors - eating specialized feces in order to re-digest food, especially in the case of roughage. Capybara, rabbits, hamsters and other related species do not have a complex digestive system as do, for example, ruminants. Instead they extract more nutrition from grass by giving their food a second pass through the gut. Soft fecal pellets of partially digested food are excreted and generally consumed immediately. They also produce normal droppings, which are not eaten.

Young elephants, pandas, koalas, and hippos eat the feces of their mother, probably to obtain the bacteria required to properly digest vegetation. When they are born, their intestines do not contain these bacteria (they are completely sterile). Without them, they would be unable to get any nutritional value from many plant components.

In earthworms

An earthworm's digestive system consists of a mouth, pharynx, esophagus, crop, gizzard, and intestine. The mouth is surrounded by strong lips, which act like a hand to grab pieces of dead grass, leaves, and weeds, with bits of soil to help chew. The lips break the food down into smaller pieces. In the pharynx the food is lubricated by mucus secretions for easier passage. The esophagus adds calcium carbonate to neutralize the acids formed by food matter decay. Temporary storage occurs in the crop where food and calcium carbonate are mixed. The powerful muscles of the gizzard churn and mix the mass of food and dirt. When the churning is complete, the glands in the walls of the gizzard add enzymes to the thick paste, which helps chemically breakdown the organic matter. By peristalsis the mixture is sent to the intestine where friendly bacteria continue chemical breakdown. This releases carbohydrates, protein, fat, and various vitamins and minerals for absorption into the body.

Overview of vertebrate digestion

In most vertebrates, digestion is a multi-stage process in the digestive system, starting from ingestion of raw materials, most often other organisms. Ingestion usually involves some type of mechanical and chemical processing. Digestion is separated into four steps:

Ingestion: placing food into the mouth (entry of food in the digestive system),
Mechanical and chemical breakdown: mastication and the mixing of the resulting bolus with water, acids, bile and enzymes in the stomach and intestine to break down complex molecules into simple structures,
Absorption: of nutrients from the digestive system to the circulatory and lymphatic capillaries through osmosis, active transport, and diffusion, and
Egestion (Excretion): Removal of undigested materials from the digestive tract through defecation.

Underlying the process is muscle movement throughout the system through swallowing and peristalsis. Each step in digestion requires energy, and thus imposes an "overhead charge" on the energy made available from absorbed substances. Differences in that overhead cost are important influences on lifestyle, behavior, and even physical structures. Examples may be seen in humans, who differ considerably from other hominids (lack of hair, smaller jaws and musculature, different dentition, length of intestines, cooking, etc.).

The major part of digestion takes place in the small intestine. The large intestine primarily serves as a site for fermentation of indigestible matter by gut bacteria and for resorption of water from digesta before excretion.

In mammals, preparation for digestion begins with the cephalic phase in which saliva is produced in the mouth and digestive enzymes are produced in the stomach. Mechanical and chemical digestion begin in the mouth where food is chewed, and mixed with saliva to begin enzymatic processing of starches. The stomach continues to break food down mechanically and chemically through churning and mixing with both acids and enzymes. Absorption occurs in the stomach and gastrointestinal tract, and the process finishes with defecation.^[1]

Human digestion process

The whole digestive system is around 9 meters long. In a healthy human adult this process can take between 24 and 72 hours. Food digestion physiology varies between individuals and upon other factors such as the characteristics of the food and size of the meal.^[16]

Phases of gastric secretion

Cephalic phase - This phase occurs before food enters the stomach and involves preparation of the body for eating and digestion. Sight and thought stimulate the cerebral cortex. Taste and smell stimulus is sent to the hypothalamus and medulla oblongata. After this it is routed through the vagus nerve and release of acetylcholine. Gastric secretion at this phase rises to 40% of maximum rate. Acidity in the stomach is not buffered by food at this point and thus acts to inhibit parietal (secretes acid) and G cell (secretes gastrin) activity via D cell secretion of somatostatin.
Gastric phase - This phase takes 3 to 4 hours. It is stimulated by distension of the stomach, presence of food in stomach and decrease in pH. Distention activates long and myenteric reflexes. This activates the release of acetylcholine, which stimulates the release of more gastric juices. As protein enters the stomach, it binds to hydrogen ions, which raises the pH of the stomach. Inhibition of gastrin and gastric acid secretion is lifted. This triggers G cells to release gastrin, which in turn stimulates parietal cells to secrete gastric acid. Gastric acid is about 0.5% hydrochloric acid (HCl), which lowers the pH to the desired pH of 1-3. Acid release is also triggered by acetylcholine and histamine.
Intestinal phase - This phase has 2 parts, the excitatory and the inhibitory. Partially digested food fills the duodenum. This triggers intestinal gastrin to be released. Enterogastric reflex inhibits vagal nuclei, activating sympathetic fibers causing the pyloric sphincter to tighten to prevent more food from entering, and inhibits local reflexes.

Oral cavity

In humans, digestion begins in the oral cavity, otherwise known as the "Buccal Cavity", where food is chewed. Saliva is secreted in large amounts (1-1.5 litres/day) by three pairs of exocrine salivary glands (parotid, submandibular, and sublingual) in the oral cavity, and is mixed with the chewed food by the tongue. Saliva cleans the oral cavity, moistens the food, and contains digestive enzymes such as salivary amylase, which aids in the chemical breakdown of polysaccharides such as starch into disaccharides such as maltose. It also contains mucus, a glycoprotein that helps soften the food and form it into a bolus. An additional enzyme, lingual lipase, hydrolyzes long-chain triglycerides into partial glycerides and free fatty acids.

Swallowing transports the chewed food into the esophagus, passing through the oropharynx and hypopharynx. The mechanism for swallowing is coordinated by the swallowing center in the medulla oblongata and pons. The reflex is initiated by touch receptors in the pharynx as the bolus of food is pushed to the back of the mouth.

Pharynx

The pharynx is the part of the neck and throat situated immediately behind the mouth and nasal cavity, and cranial, or superior, to the esophagus. It is part of the digestive system and respiratory system. Because both food and air pass through the pharynx, a flap of connective tissue, the epiglottis closes over the trachea when food is swallowed to prevent choking or asphyxiation.

The oropharynx is that part of the pharynx behind the oral cavity. It is lined with stratified squamous epithelium. The nasopharynx lies behind the nasal cavity and like the nasal passages is lined with ciliated columnar pseudostratified epithelium.

Like the oropharynx above it the hypopharynx (laryngopharynx) serves as a passageway for food and air and is lined with a stratified squamous epithelium. It lies inferior to the upright epiglottis and extends to the larynx, where the respiratory and digestive pathways diverge. At that point, the laryngopharynx is continuous with the esophagus. During swallowing, food has the "right of way", and air passage temporarily stops.

Esophagus

The esophagus is a narrow muscular tube about 20-30 centimeters long, which starts at the pharynx at the back of the mouth, passes through the thoracic diaphragm, and ends at the cardiac orifice of the stomach. The wall of the esophagus is made up of two layers of smooth muscles, which form a continuous layer from the esophagus to the colon and contract slowly, over long periods of time. The inner layer of muscles is arranged circularly in a series of descending rings, while the outer layer is arranged longitudinally. At the top of the esophagus, is a flap of tissue called the epiglottis that closes during swallowing to prevent food from entering the trachea (windpipe). The chewed food is pushed down the esophagus to the stomach through peristaltic contraction of these muscles. It takes only about seven seconds for food to pass through the esophagus and now digestion takes place.

Stomach

The stomach is a small, 'J'-shaped pouch with walls made of thick, elastic muscles, which stores and helps break down food. Food reduced to very small particles is more likely to be fully digested in the small intestine, and stomach churning has the effect of assisting the physical disassembly begun in the mouth. Ruminants, who are able to digest fibrous material (primarily cellulose), use fore-stomachs and repeated chewing to further the disassembly. Rabbits and some other animals pass some material through their entire digestive systems twice. Most birds ingest small stones to assist in mechanical processing in gizzards.

Food enters the stomach through the cardiac orifice where it is further broken apart and thoroughly mixed with gastric acid, pepsin and other digestive enzymes to break down proteins. The enzymes in the stomach also have an optimum, meaning that they work at a specific pH and temperature better than any others. The acid itself does not break down food molecules, rather it provides an optimum pH for the reaction of the enzyme pepsin and kills many microorganisms that are ingested with the food. It can also denature proteins. This is the process of reducing polypeptide bonds and disrupting salt bridges, which in turn causes a loss of secondary, tertiary, or quaternary protein structure. The parietal cells of the stomach also secrete a glycoprotein called intrinsic factor, which enables the absorption of vitamin B-12. Mucus neck cells are present in the gastric glands of the stomach. They secrete mucus, which along with gastric juice plays and important role in lubrication and protection of the mucosal epithelium from excoriation by the highly concentrated hydrochloric acid. Other small molecules such as alcohol are absorbed in the stomach, passing through the membrane of the stomach and entering the circulatory system directly. Food in the stomach is in semi-liquid form, which upon completion is known as chyme.

After consumption of food, digestive "tonic" and peristaltic contractions begin, which helps break down the food and move it through.^[16] When the chyme reaches the opening to the duodenum known as the pylorus, contractions "squirt" the food back into the stomach through a process called retropulsion, which exerts additional force and further grinds down food into smaller particles.^[16] Gastric emptying is the release of food from the stomach into the duodenum; the process is tightly controlled with liquids being emptied much more quickly than solids.^[16] Gastric emptying has attracted medical interest as rapid gastric emptying is related to obesity and delayed gastric emptying syndrome is associated with diabetes mellitus, aging, and gastroesophageal reflux.^[16]

The transverse section of the alimentary canal reveals four (or five, see description under mucosa) distinct and well developed layers within the stomach:

Serous membrane, a thin layer of mesothelial cells that is the outermost wall of the stomach.
Muscular coat, a well-developed layer of muscles used to mix ingested food, composed of three sets running in three different alignments. The outermost layer runs parallel to the vertical axis of the stomach (from top to bottom), the middle is concentric to the axis (horizontally circling the stomach cavity) and the innermost oblique layer, which is responsible for mixing and breaking down ingested food, runs diagonal to the longitudinal axis. The inner layer is unique to the stomach, all other parts of the digestive tract have only the first two layers.
Submucosa, composed of connective tissue that links the inner muscular layer to the mucosa and contains the nerves, blood and lymph vessels.
Mucosa is the extensively folded innermost layer. It can be divided into the epithelium, lamina propria, and the muscularis mucosae, though some consider the outermost muscularis mucosae to be a distinct layer, as it develops from the mesoderm rather than the endoderm (thus making a total of five layers). The epithelium and lamina are filled with connective tissue and covered in gastric glands that may be simple or branched tubular, and secrete mucus, hydrochloric acid, pepsinogen and rennin. The mucus lubricates the food and also prevents hydrochloric acid from acting on the walls of the stomach.

Small intestine

It has three parts Duodenum, Jejunum, and Ileum.

After being processed in the stomach, food is passed to the small intestine via the pyloric sphincter. The majority of digestion and absorption occurs here after the milky chyme enters the duodenum. Here it is further mixed with three different liquids:

Bile, which emulsifies fats to allow absorption, neutralizes the chyme and is used to excrete waste products such as bilin and bile acids. Bile is produced by the liver and then stored in the gallbladder. The bile in the gallbladder is much more concentrated.
Pancreatic juice made by the pancreas.
Intestinal enzymes of the alkaline mucosal membranes. The enzymes include maltase, lactase and sucrase (all three of which process only sugars), trypsin and chymotrypsin.

The pH level increases in the small intestine. A more basic environment causes more helpful enzymes to activate and begin to help in the breakdown of molecules such as fat globules. Small, finger-like structures called villi, each of which is covered with even smaller hair-like structures called microvilli improve the absorption of nutrients by increasing the surface area of the intestine and enhancing speed at which nutrients are absorbed. Blood containing the absorbed nutrients is carried away from the small intestine via the hepatic portal vein and goes to the liver for filtering, removal of toxins, and nutrient processing.

The small intestine and remainder of the digestive tract undergoes peristalsis to transport food from the stomach to the rectum and allow food to be mixed with the digestive juices and absorbed. The circular muscles and longitudinal muscles are antagonistic muscles, with one contracting as the other relaxes. When the circular muscles contract, the lumen becomes narrower and longer and the food is squeezed and pushed forward. When the longitudinal muscles contract, the circular muscles relax and the gut dilates to become wider and shorter to allow food to enter.

Large intestine

After the food has been passed through the small intestine, the food enters the large intestine. Within it, digestion is retained long enough to allow fermentation due to the action of gut bacteria, which breaks down some of the substances that remain after processing in the small intestine; some of the breakdown products are absorbed. In humans, these include most complex saccharides (at most three disaccharides are digestible in humans). In addition, in many vertebrates, the large intestine reabsorbs fluid; in a few, with desert lifestyles, this reabsorbtion makes continued existence possible.

In humans, the large intestine is roughly 1.5 meters long, with three parts: the cecum at the junction with the small intestine, the colon, and the rectum. The colon itself has four parts: the ascending colon, the transverse colon, the descending colon, and the sigmoid colon. The large intestine absorbs water from the bolus and stores feces until it can be egested. Food products that cannot go through the villi, such as cellulose (dietary fiber), are mixed with other waste products from the body and become hard and concentrated feces. The feces is stored in the rectum for a certain period and then the stored feces is eliminated from the body due to the contraction and relaxation through the anus. The exit of this waste material is regulated by the anal sphincter.

Breakdown into nutrients

Protein digestion

Protein digestion occurs in the stomach and duodenum in which 3 enzymes: pepsin secreted by the stomach and trypsin and chymotrypsin secreted by the pancreas breakdown food proteins into polypeptides that are then broken down by the enzyme erepsin into amino acids.

Fat digestion

Digestion of fat begins in the mouth where lingual lipase breaks down some lipids into diglycerides. The presence of fat in the small intestine produces hormones that stimulate the release of pancreatic lipase from the pancreas and bile from the liver for breakdown of fats into fatty acids.

Carbohydrate digestion

Starches are broken down into sugars (glucose and fructose) by amylase and hydrochloric acid in the stomach.

DNA and RNA digestion

DNA and RNA are broken down into mononucleotides by the nucleases deoxyribonuclease and ribonuclease (DNase and RNase) from the pancreas.

Digestive hormones

There are at least five hormones that aid and regulate the digestive system in mammals. There are variations across the vertebrates, as for instance in birds. Arrangements are complex and additional details are regularly discovered. For instance, more connections to metabolic control (largely the glucose-insulin system) have been uncovered in recent years.

Gastrin - is in the stomach and stimulates the gastric glands to secrete pepsinogen (an inactive form of the enzyme pepsin) and hydrochloric acid. Secretion of gastrin is stimulated by food arriving in stomach. The secretion is inhibited by low pH .
Secretin - is in the duodenum and signals the secretion of sodium bicarbonate in the pancreas and it stimulates the bile secretion in the liver. This hormone responds to the acidity of the chyme.
Cholecystokinin (CCK) - is in the duodenum and stimulates the release of digestive enzymes in the pancreas and stimulates the emptying of bile in the gall bladder. This hormone is secreted in response to fat in chyme.
Gastric inhibitory peptide (GIP) - is in the duodenum and decreases the stomach churning in turn slowing the emptying in the stomach. Another function is to induce insulin secretion.
Motilin - is in the duodenum and increases the migrating myoelectric complex component of gastrointestinal motility and stimulates the production of pepsin.

Significance of pH in digestion

Digestion is a complex process controlled by several factors. pH plays a crucial role in a normally functioning digestive tract. In the mouth, pharynx, and esophagus, pH is typically about 6.8, very weakly acidic. Saliva controls pH in this region of the digestive tract. Salivary amylase is contained in saliva and starts the breakdown of carbohydrates into monosaccharides. Most digestive enzymes are sensitive to pH and will denature in a high or low pH environment.

The stomach's high acidity inhibits the breakdown of carbohydrates within it. This acidity confers two benefits: it denatures proteins for further digestion in the small intestines, and provides non-specific immunity, damaging or eliminating various pathogens.^{[citation needed]}

In the small intestines, the duodenum provides critical pH balancing to activate digestive enzymes. The liver secretes bile into the duodenum to neutralize the acidic conditions from the stomach, and the pancreatic duct empties into the duodenum, adding bicarbonate to neutralize the acidic chyme, thus creating a neutral environment. The mucosal tissue of the small intestines is alkaline with a pH of about 8.5.^{[citation needed]}

Uses of animal gut by humans

The stomachs of calves have commonly been used as a source of rennet for making cheese.
The use of animal gut strings by musicians can be traced back to the third dynasty of Egypt. In the recent past, strings were made out of lamb gut. With the advent of the modern era, musicians have tended to use strings made of silk, or synthetic materials such as nylon or steel. Some instrumentalists, however, still use gut strings in order to evoke the older tone quality. Although such strings were commonly referred to as "catgut" strings, cats were never used as a source for gut strings^{[citation needed]}.
Sheep gut was the original source for natural gut string used in racquets, such as for tennis. Today, synthetic strings are much more common, but the best gut strings are now made out of cow gut.
Gut cord has also been used to produce strings for the snares that provide a snare drum's characteristic buzzing timbre. While the modern snare drum almost always uses metal wire rather than gut cord, the North African bendir frame drum still uses gut for this purpose.
"Natural" sausage hulls (or casings) are made of animal gut, especially hog, beef, and lamb. Similarly, Haggis is traditionally boiled in, and served in, a sheep stomach.
Chitterlings, a kind of food, consist of thoroughly washed pig's gut.
Animal gut was used to make the cord lines in longcase clocks and for fusee movements in bracket clocks, but may be replaced by metal wire.
The oldest known condoms, from 1640 AD, were made from animal intestine.^[17]

References

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^ Gordon John Larkman Ramel (2008-09-29). "The Alimentary Canal in Birds". Retrieved 2008-12-16.
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^ "World's oldest condom". Ananova. 2008. Retrieved 2008-04-11.

External links

[Maton-1] Maton, Anthea (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall. ISBN 0-13-981176-1. OCLC 32308337. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)

[2] Dusenbery, David B. (1996). “Life at Small Scale”, pp. 113-115. Scientific American Library, New York. ISBN 0-7167-5060-0.

[3] Dusenbery, David B. (2009). Living at Micro Scale, p. 280. Harvard University Press, Cambridge, Mass. ISBN 978-0-674-03116-6.

[Wooldridge-4] Wooldridge K (editor) (2009). Bacterial Secreted Proteins: Secretory Mechanisms and Role in Pathogenesis. Caister Academic Press. ISBN 978-1-904455-42-4. {{cite book}}: |author= has generic name (help)

[Salyers-5] Salyers, A. A. & Whitt, D. D. (2002). Bacterial Pathogenesis: A Molecular Approach, 2nd ed., Washington, D.C.: ASM Press. ISBN 1-55581-171-X

[Cascales-6] Cascales E & Christie P.J. (2003). "The versatile Type IV secretion systems". Nat Rev Microbiol. 1 (2): 137–149. doi:10.1038/nrmicro753. PMID 15035043.

[Christie-7] Christie PJ, Atmakuri K, Jabubowski S, Krishnamoorthy V & Cascales E. (2005). "Biogenesis, architecture, and function of bacterial Type IV secretion systems". Ann Rev Microbiol. 59: 451–485. doi:10.1146/annurev.micro.58.030603.123630. PMID 16153176.{{cite journal}}: CS1 maint: multiple names: authors list (link)

[Chatterjee-8] Chatterjee, SN and J Das. "Electron microscopic observations on the excretion of cell wall material by Vibrio cholerae." "J.Gen.Microbiol." "49" : 1-11 (1967) ; Kuehn, MJ and NC Kesty. "Bacterial outer membrane vesicles and the host-pathogen interaction." Genes Dev.and then the 19(22):2645-55 (2005)

[McBrrom-9] McBroom, AJ and MJ Kuehn Release of outer membrane vesicles by Gram-negative bacteria is a novel envelope stress response. Mol. Microbiol. 63(2):545-58 (2007)

[Leege-10] Leege, Lissa. "How does the Venus flytrap digest flies?". Scientific American. Retrieved 2008-08-20.

[Boettner-11] Boettner DR, Huston CD, Linford AS; et al. (2008). "Entamoeba histolytica phagocytosis of human erythrocytes involves PATMK, a member of the transmembrane kinase family". PLoS Pathog. 4 (1): e8. doi:10.1371/journal.ppat.0040008. PMC 2211552. PMID 18208324. {{cite journal}}: Explicit use of et al. in: |author= (help); Unknown parameter |month= ignored (help)CS1 maint: multiple names: authors list (link) CS1 maint: unflagged free DOI (link)

[Clarke-12] Clarke, M.R. (1986). A Handbook for the Identification of Cephalopod Beaks. Oxford: Clarendon Press. ISBN 0-19-857603-X. {{cite book}}: Cite has empty unknown parameter: |coauthors= (help)

[Miserez-13] Miserez, A (2007). "Jumbo squid beaks: Inspiration for design of robust organic composites". Acta Biomaterialia. 3 (1): 139–149. doi:10.1016/j.actbio.2006.09.004. PMID 17113369. {{cite journal}}: Cite has empty unknown parameter: |month= (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)

[Gordon-14] Gordon John Larkman Ramel (2008-09-29). "The Alimentary Canal in Birds". Retrieved 2008-12-16.

[Levi-15] Levi, Wendell (1977). The Pigeon. Sumter, S.C.: Levi Publishing Co, Inc. ISBN 0853900132.

[KongSingh2008-16] Kong F, Singh RP (2008). "Disintegration of solid foods in human stomach". J. Food Sci. 73 (5): R67–80. doi:10.1111/j.1750-3841.2008.00766.x. PMID 18577009. {{cite journal}}: Unknown parameter |month= ignored (help)Free full-text

[17] "World's oldest condom". Ananova. 2008. Retrieved 2008-04-11.

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@@ Line 30: / Line 30: @@
 ====Conjugation machinery====
-The [[Bacterial conjugation|conjugation]] machinery of some bacteria (and archaeal flagella) is capable of transporting both DNA and proteins. It was discovered in ''Agrobacterium tumefaciens'', which uses this system to introduce the Ti plasmid and proteins into the host which develops the crown gall (tumor).<ref name=Cascales>{{cite journal |author=Cascales E & Christie P.J. |title=The versatile Type IV secretion systems |journal=Nat Rev Microbiol |volume=1 |issue=2 |pages=137–149 |year=2003 |doi=10.1038/nrmicro753 |pmid=15035043}}</ref> The VirB complex of ''Agrobacterium tumefaciens'' is the prototypic system.<ref name=Christie>{{cite journal |author=Christie PJ, Atmakuri K, Jabubowski S, Krishnamoorthy V & Cascales E. |title=Biogenesis, architecture, and function of bacterial Type IV secretion systems |journal=Ann Rev Microbiol |volume=59 |pages=451–485 |year=2005 |doi=10.1146/annurev.micro.58.030603.123630 |pmid=16153176}}</ref>
+The [[Bacterial conjugation|conjugation]] machinery of some bacteria (and archaeal flagella) is capable of transporting both DNA and proteins. It was discovered in ''Agrobacterium tumefaciens'', which uses this system to introduce the Ti plasmid and proteins into the host, which develops the crown gall (tumor).<ref name=Cascales>{{cite journal |author=Cascales E & Christie P.J. |title=The versatile Type IV secretion systems |journal=Nat Rev Microbiol |volume=1 |issue=2 |pages=137–149 |year=2003 |doi=10.1038/nrmicro753 |pmid=15035043}}</ref> The VirB complex of ''Agrobacterium tumefaciens'' is the prototypic system.<ref name=Christie>{{cite journal |author=Christie PJ, Atmakuri K, Jabubowski S, Krishnamoorthy V & Cascales E. |title=Biogenesis, architecture, and function of bacterial Type IV secretion systems |journal=Ann Rev Microbiol |volume=59 |pages=451–485 |year=2005 |doi=10.1146/annurev.micro.58.030603.123630 |pmid=16153176}}</ref>
 The [[diazotroph|nitrogen fixing]] ''[[Rhizobia]]'' are an interesting case, wherein conjugative elements naturally engage in inter-[[Kingdom (biology)|kingdom]] conjugation. Such elements as the ''[[Agrobacterium]]'' Ti or Ri plasmids contain elements that can transfer to plant cells. Transferred genes enter the plant cell nucleus and effectively transform the plant cells into factories for the production of [[opines]], which the bacteria use as carbon and energy sources. Infected plant cells form [[Agrobacterium tumefaciens|crown gall]] or [[Agrobacterium rhizogenes|root tumors]]. The Ti and Ri plasmids are thus [[endosymbiont]]s of the bacteria, which are in turn endosymbionts (or parasites) of the infected plant.
@@ Line 42: / Line 42: @@
 [[Image:VFT ne1.JPG|thumb|right|125px|Venus Flytrap (''Dionaea muscipula'') leaf]]
 ===Gastrovascular cavity===
-The [[gastrovascular cavity]] functions as a stomach in both digestion and the distribution of nutrients to all parts of the body.  Extracellular digestion takes place within this central cavity which is lined with the gastrodermis, the internal layer of [[epithelium]]. This cavity has only one opening to the outside that functions as both a mouth and an anus: waste and undigested matter is excreted through the mouth/anus, which can be described as an incomplete gut.
+The [[gastrovascular cavity]] functions as a stomach in both digestion and the distribution of nutrients to all parts of the body.  Extracellular digestion takes place within this central cavity, which is lined with the gastrodermis, the internal layer of [[epithelium]]. This cavity has only one opening to the outside that functions as both a mouth and an anus: waste and undigested matter is excreted through the mouth/anus, which can be described as an incomplete gut.
 In a plant such as the [[Venus Flytrap]] that can make its own food through photosynthesis, it does not eat and digest its prey for the traditional objectives of harvesting energy and carbon, but mines prey primarily for essential nutrients (nitrogen and phosphorus in particular) that are in short supply in its boggy, acidic habitat.<ref name=Leege>{{cite web | url = http://www.sciam.com/article.cfm?id=how-does-the-venus-flytra | work = Scientific American | author = Leege, Lissa | title = How does the Venus flytrap digest flies? | accessdate= 2008-08-20 }}</ref>
@@ Line 109: / Line 109: @@
 ===In earthworms===
-An [[earthworm]]'s digestive system consists of a [[mouth]], [[pharynx]], [[esophagus]], [[Crop (anatomy)|crop]], [[gizzard]], and [[intestine]]. The mouth is surrounded by strong lips which act like a hand to grab pieces of dead grass, leaves, and weeds, with bits of soil to help chew. The lips break the food down into smaller pieces. In the pharynx the food is lubricated by mucus secretions for easier passage. The esophagus adds calcium carbonate to neutralize the acids formed by food matter decay. Temporary storage occurs in the crop where food and calcium carbonate are mixed. The powerful muscles of the gizzard churn and mix the mass of food and dirt. When the churning is complete, the glands in the walls of the gizzard add enzymes to the thick paste which aid in the chemical breakdown of the organic matter. By [[peristalsis]] the mixture is sent to the intestine where friendly bacteria continue chemical breakdown. This releases carbohydrates, protein, fat, and various vitamins and minerals for absorption into the body.
+An [[earthworm]]'s digestive system consists of a [[mouth]], [[pharynx]], [[esophagus]], [[Crop (anatomy)|crop]], [[gizzard]], and [[intestine]]. The mouth is surrounded by strong lips, which act like a hand to grab pieces of dead grass, leaves, and weeds, with bits of soil to help chew. The lips break the food down into smaller pieces. In the pharynx the food is lubricated by mucus secretions for easier passage. The esophagus adds calcium carbonate to neutralize the acids formed by food matter decay. Temporary storage occurs in the crop where food and calcium carbonate are mixed. The powerful muscles of the gizzard churn and mix the mass of food and dirt. When the churning is complete, the glands in the walls of the gizzard add enzymes to the thick paste, which helps chemically breakdown the organic matter. By [[peristalsis]] the mixture is sent to the intestine where friendly bacteria continue chemical breakdown. This releases carbohydrates, protein, fat, and various vitamins and minerals for absorption into the body.
 ==Overview of vertebrate digestion==
@@ Line 134: / Line 134: @@
 *[[Cephalic phase]] -  This phase occurs before food enters the stomach and involves preparation of the body for eating and digestion. Sight and thought stimulate the [[cerebral cortex]]. Taste and smell stimulus is sent to the [[hypothalamus]] and [[medulla oblongata]]. After this it is routed through the [[vagus nerve]] and release of acetylcholine. Gastric secretion at this phase rises to 40% of maximum rate. Acidity in the stomach is not buffered by food at this point and thus acts to inhibit parietal (secretes acid) and [[G cell]] (secretes gastrin) activity via [[D cell (biology)|D cell]] secretion of [[somatostatin]].
-*Gastric phase - This phase takes 3 to 4 hours. It is stimulated by [[Gastric distension|distension]] of the stomach, presence of food in stomach and decrease in [[pH]]. Distention activates long and myenteric reflexes. This activates the release of [[acetylcholine]] which stimulates the release of more [[gastric juice]]s. As protein enters the stomach, it binds to [[hydrogen]] ions, which raises the [[pH]] of the [[stomach]]. Inhibition of gastrin and [[gastric acid]] secretion is lifted. This triggers [[G cell]]s to release [[gastrin]], which in turn stimulates [[parietal cell]]s to secrete gastric acid.  Gastric acid is about 0.5% [[hydrochloric acid]] (HCl), which lowers the pH to the desired pH of 1-3.  Acid release is also triggered by [[acetylcholine]] and [[histamine]].
+*Gastric phase - This phase takes 3 to 4 hours. It is stimulated by [[Gastric distension|distension]] of the stomach, presence of food in stomach and decrease in [[pH]]. Distention activates long and myenteric reflexes. This activates the release of [[acetylcholine]], which stimulates the release of more [[gastric juice]]s. As protein enters the stomach, it binds to [[hydrogen]] ions, which raises the [[pH]] of the [[stomach]]. Inhibition of gastrin and [[gastric acid]] secretion is lifted. This triggers [[G cell]]s to release [[gastrin]], which in turn stimulates [[parietal cell]]s to secrete gastric acid.  Gastric acid is about 0.5% [[hydrochloric acid]] (HCl), which lowers the pH to the desired pH of 1-3.  Acid release is also triggered by [[acetylcholine]] and [[histamine]].
 *Intestinal phase - This phase has 2 parts, the excitatory and the inhibitory. Partially digested food fills the [[duodenum]]. This triggers intestinal gastrin to be released. Enterogastric reflex inhibits vagal nuclei, activating [[Sympathetic nervous system|sympathetic fibers]] causing the [[Pylorus|pyloric sphincter]] to tighten to prevent more food from entering, and inhibits local reflexes.
@@ Line 140: / Line 140: @@
 {{Main|Mouth (human)}}
-In humans, digestion begins in the [[Mouth (human)|oral cavity]], otherwise known as the "Buccal Cavity", where food is [[Mastication|chewed]]. [[Saliva]] is secreted in large amounts (1-1.5 litres/day) by three pairs of exocrine salivary glands (parotid, submandibular, and sublingual) in the oral cavity, and is mixed with the chewed food by the tongue. The saliva serves to clean the oral cavity and moisten the food, and contains digestive [[enzyme]]s such as salivary [[amylase]], which aids in the [[Hydrolysis|chemical breakdown]] of [[polysaccharide]]s such as [[starch]] into [[disaccharide]]s such as [[maltose]].  It also contains mucus, a [[glycoprotein]] which helps soften the food and form it into a [[Bolus (digestion)|bolus]]. An additional enzyme, [[lingual lipase]], hydrolyzes long-chain triglycerides into partial glycerides and free fatty acids.
+In humans, digestion begins in the [[Mouth (human)|oral cavity]], otherwise known as the "Buccal Cavity", where food is [[Mastication|chewed]]. [[Saliva]] is secreted in large amounts (1-1.5 litres/day) by three pairs of exocrine salivary glands (parotid, submandibular, and sublingual) in the oral cavity, and is mixed with the chewed food by the tongue. Saliva cleans the oral cavity, moistens the food, and contains digestive [[enzyme]]s such as salivary [[amylase]], which aids in the [[Hydrolysis|chemical breakdown]] of [[polysaccharide]]s such as [[starch]] into [[disaccharide]]s such as [[maltose]].  It also contains mucus, a [[glycoprotein]] that helps soften the food and form it into a [[Bolus (digestion)|bolus]]. An additional enzyme, [[lingual lipase]], hydrolyzes long-chain triglycerides into partial glycerides and free fatty acids.
 [[Swallowing]] transports the chewed food into the [[esophagus]], passing through the [[oropharynx]] and [[hypopharynx]]. The mechanism for swallowing is coordinated by the swallowing center in the [[medulla oblongata]] and [[pons]]. The reflex is initiated by touch receptors in the pharynx as the bolus of food is pushed to the back of the mouth.
@@ Line 149: / Line 149: @@
 The pharynx is the part of the neck and throat situated immediately behind the mouth and nasal cavity, and cranial, or superior, to the esophagus. It is part of the [[digestive system]] and [[respiratory system]]. Because both food and air pass through the pharynx, a flap of connective tissue, the [[epiglottis]] closes over the trachea when food is swallowed to prevent choking or asphyxiation.
-The [[oropharynx]] is that part of the pharynx which lies behind the oral cavity and is lined by [[stratified squamous epithelium]]. The [[nasopharynx]] lies behind the nasal cavity and like the nasal passages is lined with ciliated columnar [[pseudostratified epithelium]].
+The [[oropharynx]] is that part of the pharynx behind the oral cavity. It is lined with [[stratified squamous epithelium]]. The [[nasopharynx]] lies behind the nasal cavity and like the nasal passages is lined with ciliated columnar [[pseudostratified epithelium]].
 Like the oropharynx above it the hypopharynx ([[laryngopharynx]]) serves as a passageway for food and air and is lined with a stratified squamous epithelium. It lies inferior to the upright epiglottis and extends to the larynx, where the respiratory and digestive pathways diverge. At that point, the laryngopharynx is continuous with the esophagus. During swallowing, food has the "right of way", and air passage temporarily stops.
@@ Line 156: / Line 156: @@
 {{Main|esophagus}}
-The esophagus is a narrow muscular tube about 20-30 centimeters long which starts at the [[pharynx]] at the back of the mouth, passes through the [[thoracic diaphragm]], and ends at the [[Cardiac|cardiac orifice]] of the [[stomach]].  The wall of the esophagus is made up of two layers of [[smooth muscle]]s, which form a continuous layer from the esophagus to the colon and contract slowly, over long periods of time. The inner layer of muscles is arranged circularly in a series of descending rings, while the outer layer is arranged longitudinally.  At the top of the esophagus, is a flap of tissue called the [[epiglottis]] that closes during swallowing to prevent food from entering the [[Vertebrate trachea|trachea]] (windpipe).  The chewed food is pushed down the esophagus to the stomach through [[Peristalsis|peristaltic]] contraction of these muscles.  It takes only about seven seconds for food to pass through the esophagus and now digestion takes place.
+The esophagus is a narrow muscular tube about 20-30 centimeters long, which starts at the [[pharynx]] at the back of the mouth, passes through the [[thoracic diaphragm]], and ends at the [[Cardiac|cardiac orifice]] of the [[stomach]].  The wall of the esophagus is made up of two layers of [[smooth muscle]]s, which form a continuous layer from the esophagus to the colon and contract slowly, over long periods of time. The inner layer of muscles is arranged circularly in a series of descending rings, while the outer layer is arranged longitudinally.  At the top of the esophagus, is a flap of tissue called the [[epiglottis]] that closes during swallowing to prevent food from entering the [[Vertebrate trachea|trachea]] (windpipe).  The chewed food is pushed down the esophagus to the stomach through [[Peristalsis|peristaltic]] contraction of these muscles.  It takes only about seven seconds for food to pass through the esophagus and now digestion takes place.
 ===Stomach===
 {{Main|Stomach}}
-The stomach is a small, 'J'-shaped pouch with walls made of thick, elastic [[muscle]]s, which stores and helps break down food. Food which has been reduced to very small particles is more likely to be fully digested in the small intestine, and stomach churning has the effect of assisting the physical disassembly begun in the mouth. Ruminants, who are able to digest fibrous material (primarily [[cellulose]]), use fore-stomachs and repeated chewing to further the disassembly. Rabbits and some other animals pass [[Cecotrope|some material]] through their entire digestive systems twice. Most birds ingest small stones to assist in mechanical processing in gizzards.
+The stomach is a small, 'J'-shaped pouch with walls made of thick, elastic [[muscle]]s, which stores and helps break down food. Food reduced to very small particles is more likely to be fully digested in the small intestine, and stomach churning has the effect of assisting the physical disassembly begun in the mouth. Ruminants, who are able to digest fibrous material (primarily [[cellulose]]), use fore-stomachs and repeated chewing to further the disassembly. Rabbits and some other animals pass [[Cecotrope|some material]] through their entire digestive systems twice. Most birds ingest small stones to assist in mechanical processing in gizzards.
-Food enters the stomach through the cardiac orifice where it is further broken apart and thoroughly mixed with [[gastric acid]], [[pepsin]] and other [[digestive enzyme]]s to break down proteins. The enzymes in the stomach also have an optimum, meaning that they work at a specific pH and temperature better than any others. The acid itself does not break down food molecules, rather it provides an optimum pH for the reaction of the enzyme [[pepsin]] and kills many microorganisms that are ingested with the food. It can also denature proteins. This is the process of reducing polypeptide bonds and disrupting salt bridges which in turn causes a loss of secondary, tertiary or quaternary protein structure.  The [[parietal cell]]s of the stomach also secrete a [[glycoprotein]] called [[intrinsic factor]] which enables the absorption of [[Cyanocobalamin|vitamin B-12]].  Mucus neck cells are present in the gastric glands of the stomach. They secrete mucus which along with gastric juice plays and important role in lubrication and protection of the mucosal epithelium from excoriation by the highly concentrated hydrochloric acid. Other small molecules such as [[alcohol]] are [[Absorption (digestive)|absorbed]] in the stomach, passing through the membrane of the stomach and entering the [[circulatory system]] directly. Food in the stomach is in semi-liquid form, which upon completion is known as [[chyme]].
+Food enters the stomach through the cardiac orifice where it is further broken apart and thoroughly mixed with [[gastric acid]], [[pepsin]] and other [[digestive enzyme]]s to break down proteins. The enzymes in the stomach also have an optimum, meaning that they work at a specific pH and temperature better than any others. The acid itself does not break down food molecules, rather it provides an optimum pH for the reaction of the enzyme [[pepsin]] and kills many microorganisms that are ingested with the food. It can also denature proteins. This is the process of reducing polypeptide bonds and disrupting salt bridges, which in turn causes a loss of secondary, tertiary, or quaternary protein structure.  The [[parietal cell]]s of the stomach also secrete a [[glycoprotein]] called [[intrinsic factor]], which enables the absorption of [[Cyanocobalamin|vitamin B-12]].  Mucus neck cells are present in the gastric glands of the stomach. They secrete mucus, which along with gastric juice plays and important role in lubrication and protection of the mucosal epithelium from excoriation by the highly concentrated hydrochloric acid. Other small molecules such as [[alcohol]] are [[Absorption (digestive)|absorbed]] in the stomach, passing through the membrane of the stomach and entering the [[circulatory system]] directly. Food in the stomach is in semi-liquid form, which upon completion is known as [[chyme]].
-After consumption of food, digestive "tonic" and [[peristalsis|peristaltic]] contractions begin which help to break down the food and move it through.<ref name=KongSingh2008/> When the chyme reaches the opening to the duodenum known as the [[pylorus]], contractions "squirt" the food back into the stomach through a process called retropulsion, which exerts additional force and further grinds down food into smaller particles.<ref name=KongSingh2008/> Gastric emptying is the release of food from the stomach into the duodenum; the process is tightly controlled with liquids being emptied much more quickly than solids.<ref name=KongSingh2008/> Gastric emptying has attracted medical interest as [[gastric dumping syndrome|rapid gastric emptying]] is related to obesity and [[Gastroparesis|delayed gastric emptying syndrome]] is associated with diabetes mellitus, aging, and gastroesophageal reflux.<ref name=KongSingh2008/>
+After consumption of food, digestive "tonic" and [[peristalsis|peristaltic]] contractions begin, which helps break down the food and move it through.<ref name=KongSingh2008/> When the chyme reaches the opening to the duodenum known as the [[pylorus]], contractions "squirt" the food back into the stomach through a process called retropulsion, which exerts additional force and further grinds down food into smaller particles.<ref name=KongSingh2008/> Gastric emptying is the release of food from the stomach into the duodenum; the process is tightly controlled with liquids being emptied much more quickly than solids.<ref name=KongSingh2008/> Gastric emptying has attracted medical interest as [[gastric dumping syndrome|rapid gastric emptying]] is related to obesity and [[Gastroparesis|delayed gastric emptying syndrome]] is associated with diabetes mellitus, aging, and gastroesophageal reflux.<ref name=KongSingh2008/>
 The transverse section of the alimentary canal reveals four (or five, see description under mucosa) distinct and well developed layers within the stomach:
@@ Line 189: / Line 189: @@
 {{Main|Large intestine}}
-After the food has been passed through the small intestine, the food enters the [[large intestine]]. Within it, digestion is retained long enough to allow fermentation due to the action of gut bacteria, which breaks down some of the substances which remain after processing in the small intestine; some of the breakdown products are absorbed. In humans, these include most complex saccharides (at most three disaccharides are digestible in humans). In addition, in many vertebrates, the large intestine reabsorbs fluid; in a few, with desert lifestyles, this reabsorbtion makes continued existence possible.
+After the food has been passed through the small intestine, the food enters the [[large intestine]]. Within it, digestion is retained long enough to allow fermentation due to the action of gut bacteria, which breaks down some of the substances that remain after processing in the small intestine; some of the breakdown products are absorbed. In humans, these include most complex saccharides (at most three disaccharides are digestible in humans). In addition, in many vertebrates, the large intestine reabsorbs fluid; in a few, with desert lifestyles, this reabsorbtion makes continued existence possible.
 In humans, the large intestine is roughly 1.5 meters long, with three parts: the [[cecum]] at the junction with the [[small intestine]], the [[Colon (anatomy)|colon]], and the [[rectum]]. The colon itself has four parts:  the [[ascending colon]], the [[transverse colon]], the [[descending colon]], and the [[sigmoid colon]].  The large intestine absorbs water from the [[bolus (digestion)|bolus]] and stores [[feces]] until it can be [[Defecation|egested]].  Food products that cannot go through the [[Intestinal villus|villi]], such as [[cellulose]] ([[dietary fiber]]), are mixed with other waste products from the body and become hard and concentrated [[feces]]. The feces is stored in the [[rectum]] for a certain period and then the stored feces is eliminated from the body due to the contraction and relaxation through the [[anus]]. The exit of this waste material is regulated by the [[anal sphincter]].
@@ Line 197: / Line 197: @@
 ===Protein digestion===
-Protein digestion occurs in the stomach and [[duodenum]] in which 3 enzymes: [[pepsin]] secreted by the stomach and [[trypsin]] and [[chymotrypsin]] secreted by the pancreas breakdown food proteins into [[polypeptides]] which are then broken down by the enzyme [[erepsin]] into [[amino acids]].
+Protein digestion occurs in the stomach and [[duodenum]] in which 3 enzymes: [[pepsin]] secreted by the stomach and [[trypsin]] and [[chymotrypsin]] secreted by the pancreas breakdown food proteins into [[polypeptides]] that are then broken down by the enzyme [[erepsin]] into [[amino acids]].
 ===Fat digestion=== <!--Fat digestion redirects here-->
-Digestion of fat begins in the mouth where [[lingual lipase]] breaks down some lipids into [[diglyceride]]s.  The presence of fat in the small intestine produces hormones which stimulate the release of [[pancreatic lipase]] from the pancreas and [[Bile acid|bile]] from the liver for breakdown of fats into [[fatty acids]].
+Digestion of fat begins in the mouth where [[lingual lipase]] breaks down some lipids into [[diglyceride]]s.  The presence of fat in the small intestine produces hormones that stimulate the release of [[pancreatic lipase]] from the pancreas and [[Bile acid|bile]] from the liver for breakdown of fats into [[fatty acids]].
 ===Carbohydrate digestion===
@@ Line 220: / Line 220: @@
 ==Significance of pH in digestion==
-Digestion is a complex process which is controlled by several factors. [[pH]] plays a crucial role in a normally functioning digestive tract. In the mouth, pharynx, and esophagus, pH is typically about 6.8, very weakly acidic. [[Saliva]] controls pH in this region of the digestive tract. [[Salivary amylase]] is contained in saliva and starts the breakdown of carbohydrates into [[monosaccharides]]. Most digestive enzymes are sensitive to pH and will denature in a high or low pH environment.
+Digestion is a complex process controlled by several factors. [[pH]] plays a crucial role in a normally functioning digestive tract. In the mouth, pharynx, and esophagus, pH is typically about 6.8, very weakly acidic. [[Saliva]] controls pH in this region of the digestive tract. [[Salivary amylase]] is contained in saliva and starts the breakdown of carbohydrates into [[monosaccharides]]. Most digestive enzymes are sensitive to pH and will denature in a high or low pH environment.
-The stomach's high acidity inhibits the breakdown of [[carbohydrates]] within it. This acidity confers two benefits: it serves to [[Denaturation (biochemistry)|denature]] proteins for further digestion in the small intestines, and provides [[Innate immune system|non-specific immunity]], damaging or eliminating various [[pathogens]].{{Citation needed|date=November 2009}}
+The stomach's high acidity inhibits the breakdown of [[carbohydrates]] within it. This acidity confers two benefits: it [[Denaturation (biochemistry)|denatures]] proteins for further digestion in the small intestines, and provides [[Innate immune system|non-specific immunity]], damaging or eliminating various [[pathogens]].{{Citation needed|date=November 2009}}
 In the small intestines, the duodenum provides critical pH balancing to activate digestive enzymes. The liver secretes bile into the duodenum to neutralize the acidic conditions from the stomach, and the [[pancreatic duct]] empties into the duodenum, adding [[bicarbonate]] to neutralize the acidic [[chyme]], thus creating a neutral environment. The mucosal tissue of the small intestines is alkaline with a pH of about 8.5.{{Citation needed|date=May 2011}}
@@ Line 230: / Line 230: @@
 * The use of animal gut [[Strings (music)|strings]] by musicians can be traced back to the [[third dynasty of Egypt]].  In the recent past, strings were made out of [[domestic sheep|lamb]] gut.  With the advent of the modern era, musicians have tended to use strings made of [[silk]], or synthetic materials such as [[nylon]] or [[steel]].  Some instrumentalists, however, still use gut strings in order to evoke the older tone quality.  Although such strings were commonly referred to as "[[catgut]]" strings, [[cat]]s were never used as a source for gut strings{{Citation needed|date=October 2010}}.
 * Sheep gut was the original source for natural gut string used in [[racquet]]s, such as for [[tennis]].  Today, synthetic strings are much more common, but the best gut strings are now made out of [[cow]] gut.
-* Gut cord has also been used to produce strings for the snares which provide the [[snare drum]]'s characteristic buzzing timbre.  While the snare drum currently almost always uses metal wire rather than gut cord, the [[North Africa]]n [[bendir]] frame drum still uses gut for this purpose.
+* Gut cord has also been used to produce strings for the snares that provide a [[snare drum]]'s characteristic buzzing timbre.  While the modern snare drum almost always uses metal wire rather than gut cord, the [[North Africa]]n [[bendir]] frame drum still uses gut for this purpose.
 * "Natural" sausage hulls (or [[Casing (sausage)|casings]]) are made of animal gut, especially hog, beef, and lamb.  Similarly, [[Haggis]] is traditionally boiled in, and served in, a sheep stomach.
 * [[Chitterlings]], a kind of food, consist of thoroughly washed [[pig]]'s gut.