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Digestion


Digestion is the mechanical and chemical breaking down of food into smaller components that can be absorbed into a blood stream, for instance. Digestion is a form of catabolism: a break-down of larger 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. 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 is great for enzymes. After some time (typically an hour or two in humans, 4–6 hours in dogs, somewhat shorter duration in house cats, ...), the results go through the small intestine, through the large intestine, and are excreted during defecation.[1]

Other organisms use different mechanisms to digest food.

Contents

[edit] Digestive systems

Digestive systems take 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 breakdown 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.

[edit] Secretion systems

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

[edit] 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). 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]

[edit] Molecular syringe

A 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]

[edit] Conjugation machinery

Schematic drawing of bacterial conjugation. Conjugation diagram 1- Donor cell produces pilus. 2- Pilus attaches to recipient cell, brings the two cells together. 3- The mobile plasmid is nicked and a single strand of DNA is then transferred to the recipient cell. 4- Both cells recircularize their plasmids, synthesize second strands, and reproduce pili; both cells are now viable donors.

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.

[edit] 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]

[edit] 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.[10]

Trophozoites of Entamoeba histolytica with ingested erythrocytes

[edit] 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.

Aboral end
Oral end
Mouth
Oral end
Aboral end
    Exoderm
    Gastroderm
    Mesoglea
    Digestive cavity
Medusa (left) and polyp (right)[11]

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.[12]

Venus Flytrap (Dionaea muscipula) leaf

[edit] Specialized organs and behaviors

Catalina Macaw exhibits its seed shearing beak.
Squid beak and ruler for size comparison.
Teeth of a Carcharodon megalodon.
Rough illustration of a ruminant digestive system.

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

[edit] 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[13] 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.[14] The beak is the only indigestible part of the squid.

[edit] 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 molecules 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.

[edit] 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.

[edit] 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.[15]

Certain insects may have a crop or enlarged oesophagus.

[edit] 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.

[edit] Specialized behaviors

A flesh fly "blowing a bubble". One explanation of this behaviour is that the fly regurgitates its food into a bubble in order to increase the concentration of its food by evaporating excessive water content

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.[16].

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.

[edit] 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 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.

[edit] 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:

  1. Ingestion: placing food into the mouth (entry of food in the digestive system),
  2. 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,
  3. Absorption: of nutrients from the digestive system to the circulatory and lymphatic capillaries through osmosis, active transport, and diffusion, and
  4. 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]

[edit] Human digestion process

Salivary glands Parotid gland Submandibular gland Sublingual gland Pharynx Tongue Esophagus Pancreas Pancreatic duct Stomach Ileum Anus Rectum Vermiform appendix Cecum Descending colon Ascending colon Transverse colon Bile duct Duodenum Gallbladder Liver Oral cavity
Upper and Lower human gastrointestinal tract

The whole digestive system is around 9 meters long. In a healthy human adult this process can take between 24 and 72 hours.

[edit] 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 distention of the stomach, presence of food in stomach and decrease in pH. Distention activates long and myentric 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 lowers the pH of the stomach to around pH 1-3. Inhibition of gastrin and HCl secretion is lifted. This triggers G cells to release gastrin, which in turn stimulates parietal cells to secrete HCl. HCl 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.

[edit] Oral cavity

In humans, digestion begins in the oral 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. There are two types of saliva. One is a thin, watery secretion, and its purpose is to wet the food. The other is a thick, mucous secretion, and it acts as a lubricant and causes food particles to stick together and form a bolus. The saliva serves to clean the oral cavity and moisten 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 mucous, a glycoprotein which helps soften the food into a bolus.

Swallowing transports the chewed food into the oesophagus, 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.

[edit] Pharynx

The pharynx is the part of the neck and throat situated immediately posterior to (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.

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.

[edit] Esophagus

The esophagus is a narrow muscular tube about 20-30 centimeters long which starts at 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 open 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 no digestion takes place.

[edit] Stomach

The stomach is a small,'J'-shaped pouch with walls made of thick, elastic muscles, 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 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. 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.

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 the it develops from the mesoderm rather than the endoderm (thus making a total of 5 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.

[edit] Small intestine

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:

As the pH level changes in the small intestines and gradually becomes basic, more enzymes are activated further that chemically break down various nutrients into smaller molecules to allow absorption into the circulatory or lymphatic systems. 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.

[edit] 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 resorption 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.

[edit] Skye digestion

The presence of fat in the small intestine produces hormones which stimulate the release of lipase from the pancreas, largely to the liver for further processing, or to fat tissue for storage.

[edit] Digestive hormones

Action of the major 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.

[edit] 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 not function in a low-pH environment like the stomach. A pH below 7 indicates an acid, while a pH above 7 indicates a base; the concentration of the acid or base, however, does also play a role.

The pH of the stomach is very low (highly acidic) which inhibits the breakdown of carbohydrates while there. The strong acid content of the stomach provides two benefits; it serves to denature proteins for further digestion in the small intestines, and provides non-specific immunity, retarding 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 neutralise the acidic conditions from the stomach. Also 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.

[edit] 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.
  • 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 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 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]

[edit] See also

[edit] References

  1. ^ a b Maton, Anthea; Jean Hopkins, Charles William McLaughlin, Susan Johnson, Maryanna Quon Warner, David LaHart, Jill D. Wright (1993). Human Biology and Health. Englewood Cliffs, New Jersey, USA: Prentice Hall. ISBN 0-13-981176-1. OCLC 32308337. 
  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.
  4. ^ Wooldridge K (editor) (2009). Bacterial Secreted Proteins: Secretory Mechanisms and Role in Pathogenesis. Caister Academic Press. ISBN 978-1-904455-42-4. 
  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
  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. 
  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. 
  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)
  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)
  10. ^ Boettner DR, Huston CD, Linford AS, et al. (January 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. PMID 18208324. PMC 2211552. http://www.plospathogens.org/article/info:doi/10.1371/journal.ppat.0040008. 
  11. ^ Ruppert, E.E., Fox, R.S., and Barnes, R.D. (2004). Invertebrate Zoology (7 ed.). Brooks / Cole. pp. 76–97. ISBN 0030259827. 
  12. ^ Leege, Lissa. "How does the Venus flytrap digest flies?". Scientific American. http://www.sciam.com/article.cfm?id=how-does-the-venus-flytra. Retrieved 2008-08-20. 
  13. ^ Clarke, M.R. (1986). A Handbook for the Identification of Cephalopod Beaks. Oxford: Clarendon Press. ISBN 0-19-857603-X. 
  14. ^ Miserez, A; Li, Y; Waite, H; Zok, F (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. 
  15. ^ Gordon John Larkman Ramel (2008-09-29). "The Alimentary Canal in Birds". http://www.earthlife.net/birds/digestion.html. Retrieved 2008-12-16. 
  16. ^ Levi, Wendell (1977). The Pigeon. Sumter, S.C.: Levi Publishing Co, Inc. ISBN 0853900132. 
  17. ^ "World's oldest condom". Ananova. 2008. http://www.ananova.com/news/story/sm_1870958.html?menu=news.quirkies.sexlife. Retrieved 2008-04-11. 

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