Enzymes for starch digestion quimica,multi strain probiotics side effects 10mg,probiotics in pet food redakce,yakult probiotics benefits - PDF 2016

Minerals, vitamins and water are already small enough to be absorbed by the body without being broken down, so they are not digested.
This website uses essential cookies without which it will not work, along with other harmless cookies aimed at improving your use of our website. Large food molecules (for example, proteins, lipids, nucleic acids, and starches) must be broken down into subunits that are small enough to be absorbed by the lining of the alimentary canal. In the small intestine, pancreatic amylase does the ‘heavy lifting’ for starch and carbohydrate digestion ([link]). The digestion of protein starts in the stomach, where HCl and pepsin break proteins into smaller polypeptides, which then travel to the small intestine ([link]). The three lipases responsible for lipid digestion are lingual lipase, gastric lipase, and pancreatic lipase. The mechanical and digestive processes have one goal: to convert food into molecules small enough to be absorbed by the epithelial cells of the intestinal villi. Absorption can occur through five mechanisms: (1) active transport, (2) passive diffusion, (3) facilitated diffusion, (4) co-transport (or secondary active transport), and (5) endocytosis. Because the cell’s plasma membrane is made up of hydrophobic phospholipids, water-soluble nutrients must use transport molecules embedded in the membrane to enter cells. In contrast to the water-soluble nutrients, lipid-soluble nutrients can diffuse through the plasma membrane. Active transport mechanisms, primarily in the duodenum and jejunum, absorb most proteins as their breakdown products, amino acids. The large and hydrophobic long-chain fatty acids and monoacylglycerides are not so easily suspended in the watery intestinal chyme. The free fatty acids and monoacylglycerides that enter the epithelial cells are reincorporated into triglycerides.
The products of nucleic acid digestion—pentose sugars, nitrogenous bases, and phosphate ions—are transported by carriers across the villus epithelium via active transport. The electrolytes absorbed by the small intestine are from both GI secretions and ingested foods. In general, all minerals that enter the intestine are absorbed, whether you need them or not.
Iron—The ionic iron needed for the production of hemoglobin is absorbed into mucosal cells via active transport.
Bile salts and lecithin can emulsify large lipid globules because they are amphipathic; they have a nonpolar (hydrophobic) region that attaches to the large fat molecules as well as a polar (hydrophilic) region that interacts with the watery chime in the intestine. Intrinsic factor secreted in the stomach binds to the large B12 compound, creating a combination that can bind to mucosal receptors in the ileum. The growth of knowledge in the biochemistry of insect digestion had a bright start during the first decades of the last century, but slowed down after the development of synthetic chemical insecticides in the 1940s. Applebaum (1985), in his review on the biochemistry of digestion, described the beginning of the renewed growth of the field. The aim of this topic is to review the recent and spectacular progress in the study of insect digestive biochemistry. Digestion is the process by which food molecules are broken down into smaller molecules that are absorbed by cells in the gut tissue. Enzyme kinetic parameters are meaningless unless assays are performed in conditions in which enzymes are stable.
If enzyme characterization is performed as part of a digestive physiology study, emphasis should be given to enzyme compartmentalization, substrate specificity, and substrate preference, in order to discover the sequential action of enzymes during the digestive process. Knowledge of the effect of pH on enzyme activity is useful in evaluating enzyme action in gut compartments (Figure 1) with different pH values. Complete enzymological characterization requires purification to homogeneity, and sequencing.
Cloning cDNA sequences encoding digestive enzymes enables the expression of large amounts of recombinant enzymes that may be crystallized or used for the production of antibodies.
However, detailed 3D structures are necessary to understand enzyme mechanisms and the binding of inhibitors to enzyme molecules.
Lipids are a large and heterogeneous group of substances that are relatively insoluble in water but readily soluble in apolar solvents.
Most food molecules to be digested are polymers, such as proteins and starch, and are digested sequentially in three phases. Any description of the spatial organization of digestion in an insect must relate the midgut compartments (cell, ecto-, and endoperitrophic spaces) to each phase of digestion, and hence to the corresponding enzymes. Despite the fact that digestive enzymes of some insects are thought to be derived from the microbiota, there are relatively few studies that show an unambiguous contribution of microbial hydrolases.
Microorganisms play a limited role in digestion, but they may enable phytophagous insects to overcome biochemical barriers to herbivory — for example, detoxifying flavonoid alkaloids and the phenolic aglycones of plant glycosides.
The pH of the contents of the midgut is one of the important internal environmental properties that affect digestive enzymes. A pH in the ectoperitrophic space lower than in the midgut lumen was reported in some lepidopteran larvae. The high alkanity of lepidopteran midgut contents is thought to allow these insects to feed on plant material rich in tannins, which bind to proteins at lower pH, reducing the efficiency of digestion (Berenbaum, 1980). A high midgut pH may also be of importance, in addition to its role in preventing tannin binding to proteins, in freeing hemicelluloses from plant cell walls ingested by insects.
The acid region in the cyclorrhaphous Diptera mid-gut is assumed to be involved in the process of killing and digesting bacteria, which may be an important food for maggots. Redox conditions in the midgut are regulated and may be the result of phylogeny, although data are scarce. Although several allelochemicals other than phenols may be present in the insect gut lumen, including alkaloids, terpene aldehydes, saponins, and hydroxamic acids (Appel, 1994), data on their effect on digestion are lacking. Figure 4 Diagrammatic representation of ion movements, proposed as being responsible for maintenance of pH in the larval midgut contents of Musca domestica. Action: hydrolysis of ester bonds between the glycerol molecules and the fatty acid chains.
The lacteals (green) that receive the lipoproteins before transporting them to the circulatory system. This is only a start to the process of digestion, as chewed pieces of food are still too large to be absorbed by the body. If you chew a piece of bread for long enough, the starch it contains is digested to sugar, and it begins to taste sweet. Glucose, galactose, and fructose are the three monosaccharides that are commonly consumed and are readily absorbed. After amylases break down starch into smaller fragments, the brush border enzyme ?-dextrinase starts working on ?-dextrin, breaking off one glucose unit at a time. Chemical digestion in the small intestine is continued by pancreatic enzymes, including chymotrypsin and trypsin, each of which act on specific bonds in amino acid sequences. The most common dietary lipids are triglycerides, which are made up of a glycerol molecule bound to three fatty acid chains.
However, because the pancreas is the only consequential source of lipase, virtually all lipid digestion occurs in the small intestine. Two types of pancreatic nuclease are responsible for their digestion: deoxyribonuclease, which digests DNA, and ribonuclease, which digests RNA. As you will recall from Chapter 3, active transport refers to the movement of a substance across a cell membrane going from an area of lower concentration to an area of higher concentration (up the concentration gradient). Moreover, substances cannot pass between the epithelial cells of the intestinal mucosa because these cells are bound together by tight junctions.
Once inside the cell, they are packaged for transport via the base of the cell and then enter the lacteals of the villi to be transported by lymphatic vessels to the systemic circulation via the thoracic duct. The small intestine is highly efficient at this, absorbing monosaccharides at an estimated rate of 120 grams per hour. Bile salts not only speed up lipid digestion, they are also essential to the absorption of the end products of lipid digestion. However, bile salts and lecithin resolve this issue by enclosing them in a micelle, which is a tiny sphere with polar (hydrophilic) ends facing the watery environment and hydrophobic tails turned to the interior, creating a receptive environment for the long-chain fatty acids. The triglycerides are mixed with phospholipids and cholesterol, and surrounded with a protein coat. Since electrolytes dissociate into ions in water, most are absorbed via active transport throughout the entire small intestine.
Once inside mucosal cells, ionic iron binds to the protein ferritin, creating iron-ferritin complexes that store iron until needed. When blood levels of ionic calcium drop, parathyroid hormone (PTH) secreted by the parathyroid glands stimulates the release of calcium ions from bone matrices and increases the reabsorption of calcium by the kidneys. Fat-soluble vitamins (A, D, E, and K) are absorbed along with dietary lipids in micelles via simple diffusion. Chemical digestion breaks large food molecules down into their chemical building blocks, which can then be absorbed through the intestinal wall and into the general circulation.
Later on, with the environmental problems caused by chemical insecticides, new approaches for insect control were investigated. He discussed contemporary research showing that most insect digestive enzymes are similar to their mammalian counterparts, but that insect exotic diets require specific enzymes.
To provide a broad coverage while keeping the topic within reasonable size limits, only a brief account with key references is given for work done prior to 2000. This process is controlled by digestive enzymes, and is dependent on their localization in the insect gut.


If researchers adopt uniform parameters and methods, comparisons among similar and different insect species will be more meaningful.
Finally, the determination of the molecular masses of digestive enzymes, associated with the ability of enzymes to pass through the peri-trophic membrane, allows estimation of the pore sizes of the peritrophic membrane. Furthermore,details of the catalytic mechanisms, including involvement of amino acid residues in catalysis and substrate specificity, should be determined. Antibodies are used in Western blots to identify a specific enzyme in protein mixtures, or to localize the enzyme in tissue sections in a light or electron microscope. Glycocalyx: the carbohydrate moiety of intrinsic proteins and glycolipids occurring in the luminal face of microvillar membranes. Alternatively, cDNA may be amenable to site-directed mutagenesis for structure—function studies. The enzyme classification and numbering system used here is that recommended by the Nomenclature Committee of the International Union of Biochemistry and Molecular Biology (Enzyme Commission).
They include the enzymes that cleave internal bonds in polysaccharides and are usually named from their substrates – for example, amylase, cel-lulase, pectinase, and chitinase.
Some contain fatty acids (fats, phospholipids, glycolipids, and waxes), while others lack them (terpenes, steroids, and carotenoids). Primary digestion is the dispersion and reduction in molecular size of the polymers, and results in oligomers.
To accomplish this, enzyme determinations must be performed in each midgut luminal compartment and in the corresponding tissue.
Exceptions to this rule, and the procedures for studying the organization of the digestive process, will be detailed below.
Microorganisms might be symbiotic or fortuitous contaminants from the external environment. Best examples are found among wood- and humus-feeding insects like termites, tipulid fly larvae, and scarabid beetle larvae. Although midgut pH is hypothesized to result from adaptation of an ancestral insect to a particular diet, its descendants may diverge, feeding on different diets, while still retaining the ancestral midgut pH condition. This is an artefact caused by a halt in alkaline secretion by the isolated midgut tissue (Grigorten et al., 1993). This explanation may also hold for scarab beetles and for detritus-feeding nematoceran Diptera larvae that usually feed on refractory materials such as humus. Tannic acid is frequently oxidized in the midgut lumen, generating peroxides, including hydrogen peroxide, which readily diffuses across cell membranes and is a powerful cytotoxin. Hemicelluloses are usually extracted in alkaline solutions for analytical purposes (Blake et al., 1971), and insects such as the caterpillar Erinnyis ello are able to digest hemicelluloses efficiently without affecting the cellulose from the leaves they ingest to any degree (Terra, 1988). This region is retained in Muscidae that have not diverged from the putative ancestral bacteria-feeding habit, as well as in the flesh-feeding Calli-phoridae and the fruit-feeding Tephritidae (Terra and Ferreira, 1994).
Dow (1992) described a carbonate secretion system, which may be responsible for the high pH found in Lepidoptera midguts (Figure 3). Carbonic anhydrase (CA) produces carbonic acid that dissociates into bicarbonate and a proton. Reducing conditions are observed in clothes moth, sphinx moths, owlet moths, and dermestid beetles (Appel and Martin, 1990), and in Hemiptera (Silva and Terra, 1994). Carbonic anhydrase (CA) in cup-shaped oxyntic cells in the middle of the midgut (A) produces carbonic acid which dissociates into bicarbonate and a proton. Food has to be broken down chemically into really small particles before it can be absorbed. Digestion of proteins in the stomach is helped by stomach acid, which is strong hydrochloric acid. Chemical digestion, on the other hand, is a complex process that reduces food into its chemical building blocks, which are then absorbed to nourish the cells of the body ([link]). At the same time, the cells of the brush border secrete enzymes such as aminopeptidase and dipeptidase, which further break down peptide chains. Pancreatic lipase breaks down each triglyceride into two free fatty acids and a monoglyceride. The nucleotides produced by this digestion are further broken down by two intestinal brush border enzymes (nucleosidase and phosphatase) into pentoses, phosphates, and nitrogenous bases, which can be absorbed through the alimentary canal wall. Each day, the alimentary canal processes up to 10 liters of food, liquids, and GI secretions, yet less than one liter enters the large intestine. In this type of transport, proteins within the cell membrane act as “pumps,” using cellular energy (ATP) to move the substance. Thus, substances can only enter blood capillaries by passing through the apical surfaces of epithelial cells and into the interstitial fluid.
The absorption of most nutrients through the mucosa of the intestinal villi requires active transport fueled by ATP.
All normally digested dietary carbohydrates are absorbed; indigestible fibers are eliminated in the feces. Short-chain fatty acids are relatively water soluble and can enter the absorptive cells (enterocytes) directly.
During absorption, co-transport mechanisms result in the accumulation of sodium ions inside the cells, whereas anti-port mechanisms reduce the potassium ion concentration inside the cells. When the body has enough iron, most of the stored iron is lost when worn-out epithelial cells slough off.
PTH also upregulates the activation of vitamin D in the kidney, which then facilitates intestinal calcium ion absorption. This is why you are advised to eat some fatty foods when you take fat-soluble vitamin supplements. Intestinal brush border enzymes and pancreatic enzymes are responsible for the majority of chemical digestion.
With the help of bile salts and lecithin, the dietary fats are emulsified to form micelles, which can carry the fat particles to the surface of the enterocytes. Midgut studies were particularly stimulated after the realization that the gut is a very large and relatively unprotected interface between the insect and its environment.
In the next decade it became apparent that even enzymes similar to those of mammals have distinct characteristics, because each insect taxon deals with food in a special way (Terra and Ferreira, 1994). Papers after 2000 have been selected from those richer in molecular details, and, when they were too numerous, representative papers were chosen, especially when abundant in references to other papers. A rectilinear plot of product formation (or substrate disappearance) versus time will ensure that enzymes are stable in a given condition. Molecular masses determined in non-denaturing conditions are preferred, since in these conditions the enzymes should maintain their in vivo aggregation states (not only their quaternary structures if present). This permits the classification of insect digestive enzymes into catalytic families, and discloses the structural basis of substrate specificities; it will also enable us to establish evolutionary relationships with enzymes from other organisms.
Enzyme crystals used for resolving three-dimensional (3D) structures (via X-ray diffraction or nuclear magnetic resonance, NMR) need amounts of purified enzymes that frequently are difficult to isolate from insects by conventional separation procedures. Site-directed mutagenesis tests the role of individual amino acid residues in enzyme function or structure. During intermediate digestion, these undergo a further reduction in molecular size to dimers; in final digestion, they become monomers. Techniques of sampling enzymes from midgut luminal compartments and enzymes trapped in cell glycocalyx have been reviewed elsewhere (Terra and Ferreira, 1994). They are found in the lumen, adhering to the peritrophic membrane, attached to the midgut surface, or within cells. Nevertheless, the pH in the immediate neighborhood of the negatively-charged microvillar glycocalyx is expected to be lower than in the bulk solution, because of proton retention (Quina et al., 1980). Nevertheless, mechanisms other than high gut pH must account for the resistance to tannin displayed by some locusts (Bernays et al., 1981) and beetles (Fox and Macauley, 1977). This explanation is better than the previous one in accounting for the very high pH observed in several insects, since a pH of about 8 is sufficient to prevent tannin binding to proteins (Terra, 1988). Phosphorus NMR microscopy has been used to show that valinomycin leads to a loss of alkalinization in the midgut of Spodoptera litura (Skibbe et al., 1996). The proton is pumped by a V-ATPase into the goblet cell cavity, from where it is removed in exchange with K+ that eventually diffuses into lumen. Bicarbonate is transported into the hemolymph, whereas the proton is actively translocated into the midgut lumen acidifying its contents to pH 3.2. In this section, you will look more closely at the processes of chemical digestion and absorption. Your bodies do not produce enzymes that can break down most fibrous polysaccharides, such as cellulose.
The fatty acids include both short-chain (less than 10 to 12 carbons) and long-chain fatty acids.
Almost all ingested food, 80 percent of electrolytes, and 90 percent of water are absorbed in the small intestine. Passive diffusion refers to the movement of substances from an area of higher concentration to an area of lower concentration, while facilitated diffusion refers to the movement of substances from an area of higher to an area of lower concentration using a carrier protein in the cell membrane. Water-soluble nutrients enter the capillary blood in the villi and travel to the liver via the hepatic portal vein. The monosaccharides glucose and galactose are transported into the epithelial cells by common protein carriers via secondary active transport (that is, co-transport with sodium ions).
Despite being hydrophobic, the small size of short-chain fatty acids enables them to be absorbed by enterocytes via simple diffusion, and then take the same path as monosaccharides and amino acids into the blood capillary of a villus.
Without micelles, lipids would sit on the surface of chyme and never come in contact with the absorptive surfaces of the epithelial cells.


After being processed by the Golgi apparatus, chylomicrons are released from the cell ([link]). To restore the sodium-potassium gradient across the cell membrane, a sodium-potassium pump requiring ATP pumps sodium out and potassium in. When the body needs iron because, for example, it is lost during acute or chronic bleeding, there is increased uptake of iron from the intestine and accelerated release of iron into the bloodstream. Most water-soluble vitamins (including most B vitamins and vitamin C) also are absorbed by simple diffusion. Water absorption is driven by the concentration gradient of the water: The concentration of water is higher in chyme than it is in epithelial cells. Hence, an understanding of gut function was thought to be essential when developing methods of control that act through the gut, such as the use of transgenic plants to control phytophagous insects. Since then, the field of digestive physiology and biochemistry has progressed dramatically at the molecular level (Terra and Ferreira, 2005). Throughout, the focus is on providing a coherent picture of phenomena and highlighting further research areas. Finally, the fourth part (section 11.10) discusses data on digestive enzyme secretion mechanisms.
Such knowledge is a prerequisite in developing new biotechno-logical approaches to control insects via the gut. Oligosaccharidases and disaccharidases are usually named based on the monosaccharide that gives its reducing group to the glycosidic bond, and on the configuration (a or P) of this bond (Figure 2).
Digestion usually occurs under the action of digestive enzymes from the midgut, with little or no participation of salivary enzymes. Intracellular bacteria are usually found in special cells, mycetocytes, which may be organized in groups, mycetomes. In fact, midgut pH correlates well with insect phylogeny (Terra and Ferreira, 1994; Clark, 1999). Thus, midgut contents are acidic in the anterior midgut and nearly neutral or alkaline in the posterior midgut in Dictyoptera, Orthoptera, and most families of Coleop-tera.
One possibility is the effect of surfactants such as lysolecithin, which is formed in insect fluids due to the action of phospholipase A on cell membranes (Figure 2), and which occurs widely in insect digestive fluids (De Veau and Schultz, 1992). Dihydroxy phenolics in an alkaline medium are converted to quinones that react with lysine E-amino groups. As valinomycin is known to transport K+ down its concentration gradient, this result gives further support to the model described in Figure 3. Bicarbonate is secreted in exchange with chloride and loses a proton due to the intense field near the membrane, forming carbonate and raising the gut pH.
In spite of this, the artificial lowering of in vivo redox potentials did not significantly impact digestive efficiency of the herbivore Helicoverpa zea, although the reducing agent used (dithiothreitol) inhibited some proteinases in vitro (Johnson and Felton, 2000). While indigestible polysaccharides do not provide any nutritional value, they do provide dietary fiber, which helps propel food through the alimentary canal. Although the entire small intestine is involved in the absorption of water and lipids, most absorption of carbohydrates and proteins occurs in the jejunum. Co-transport uses the movement of one molecule through the membrane from higher to lower concentration to power the movement of another from lower to higher. The monosaccharides leave these cells via facilitated diffusion and enter the capillaries through intercellular clefts. Short chains of two amino acids (dipeptides) or three amino acids (tripeptides) are also transported actively. Too big to pass through the basement membranes of blood capillaries, chylomicrons instead enter the large pores of lacteals. Since women experience significant iron loss during menstruation, they have around four times as many iron transport proteins in their intestinal epithelial cells as do men.
The fats are then reassembled into triglycerides and mixed with other lipids and proteins into chylomicrons that can pass into lacteals. Amino acid residues are denoted by the one-letter code, if in peptides, for the sake of brevity. According to the International Union of Biochemistry and Molecular Biology, the assay temperature should be 30°C, except when the enzyme is unstable at this temperature or altered for specific purposes. Finally, interference RNA may be used to suppress the expression of one enzyme, in order to test hypotheses regarding its physiological role.
Microorganisms produce and secrete their own hydrolases, and cell death will result in the release of enzymes into the intestinal milieu. Cyclorrhaphan Diptera midguts have nearly neutral contents in the anterior and posterior regions, whereas in middle midgut the contents are very acid (Terra and Ferreira, 1994).
Surfactants are known to prevent the precipitation of proteins by tannins even at pH as low as 6.5 (Martin and Martin, 1984).
Midgut alkalinization in nematoceran Diptera occurs by mechanisms similar to those of lepidopteran larvae (Okech et al., 2008), whereas no data are available for scarab beetles. Midgut antioxidant enzymes in Spodoptera litto-ralis are upregulated in response to increased oxidative stress caused by oxidizable allelochemicals (Krishnan and Kodrik, 2006). Finally, endocytosis is a transportation process in which the cell membrane engulfs material. The monosaccharide fructose (which is in fruit) is absorbed and transported by facilitated diffusion alone. However, after they enter the absorptive epithelial cells, they are broken down into their amino acids before leaving the cell and entering the capillary blood via diffusion.
Intrinsic factor secreted in the stomach binds to vitamin B12, preventing its digestion and creating a complex that binds to mucosal receptors in the terminal ileum, where it is taken up by endocytosis. Other absorbed monomers travel from blood capillaries in the villus to the hepatic portal vein and then to the liver.
When mentioned in text with a position number, amino acid residues are denoted by the three-letter code to avoid ambiguity. Owing to partial inactivation, the optimum temperature is not a true property of enzymes, and therefore should not be included in the characterization. Any consideration of the spectrum of hydrolase activity in the midgut must include the possibility that some of the activity may derive from microorganisms. Present knowledge is not sufficient to relate midgut detergency to diet or phylogeny, or to both. Terra and Regel (1995) determined pH values and concentrations of ammonia, chloride, and phosphate in the presence or absence of ouabain and vanadate in Musca domestica midguts.
While you will be able to view the content of this page in your current browser, you will not be able to get the full visual experience. By the time chyme passes from the ileum into the large intestine, it is essentially indigestible food residue (mainly plant fibers like cellulose), some water, and millions of bacteria ([link]).
The monosaccharides combine with the transport proteins immediately after the disaccharides are broken down.
The chylomicrons are transported in the lymphatic vessels and empty through the thoracic duct into the subclavian vein of the circulatory system. For consistency, traditional abbreviations, like BAPA for benzoyl-arginine p-nitroan-ilide, have been changed, in the example to B-R-pNA, because the one-letter code for arginine is R.
Optimum enzyme pH should be determined using different buffers to discount the effects of chemical constituents of the buffers and their ionic strength on enzyme activity. These phenomena are inhibited in larvae of several lepidopteran species by secreting glycine into the midgut lumen. From the results, they proposed that middle midgut acidification is accomplished by a proton pump of mammalian-like oxyntic cells, whereas the neutralization of posterior midgut contents depends on ammonia secretion (Figure 4). Inside the cells, NH4+ forms NH3, which diffuses into midgut lumen, and proton that is transferred into the hemolymph. Please consider upgrading your browser software or enabling style sheets (CSS) if you are able to do so. Once in the bloodstream, the enzyme lipoprotein lipase breaks down the triglycerides of the chylomicrons into free fatty acids and glycerol.
The number of molecular forms of a given enzyme should be evaluated by submitting the enzyme preparation to a separation process (gel permeation, ion-exchange chromatography, hydrophobic chromatography, electrophoresis, gradient ultracentrifugation, etc.), followed by assays of the resulting fractions.
These breakdown products then pass through capillary walls to be used for energy by cells or stored in adipose tissue as fat. In some insects, tannins reduce the overall efficiency of conversion of ingested matter to body mass by an unknown mechanism. Liver cells combine the remaining chylomicron remnants with proteins, forming lipoproteins that transport cholesterol in the blood. Substrate preference expressed as the percentage activity towards a given substrate in relation to the activity upon a reference substrate may be misleading because, in this condition, enzyme activities are determined at different substrate saturations. Nevertheless, the performance of these insects remains unchanged, because of compensatory feeding (Barbehenn et al., 2009). The isoelectric points of many enzymes can be determined after staining with specific substrates following the separation of the native enzymes on isoelectrofocusing gels.




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