Académique Documents
Professionnel Documents
Culture Documents
Joelal ACHMADI; Laboratory of Nutrirional Biochemistry; Faculty of Animal Agriculture; Diponegoro University.
www.fp.undip.ac.id; jachmadi@undip.ac.id; joelal.achmadi@yahoo.com
Nutrition
The interrelated steps by which a living organism assimilates food and uses it for growth, tissue repair and replacement, or elaboration of products. It encompasses all forms of life, including plants and animals. It requires the application of chemistry, physics, and mathematics as well as the integration of advances in soil science, plant science, animal science, biochemistry, engineering, production systems, and other disciplines.
jachmadi@undip.ac.id
Feed. An edible material that provides nutrients. Diet. A mixture of feedstuffs used to supply
jachmadi@undip.ac.id
nutrients to an animal.
A simplified schematic chart of element and compounds that may be present to feed
1. Protein 1. Nitrogen Containing 2. Nonprotein 1. Essential fatty acid 2. Sterols 3. Terpenoids 4. Waxes 5. Phospholipids 6. Miscellaneous 1. 2. 3. 4. 5. Monosaccharides Disaccharides Oligosaccharides Nonfibrous polysccharides Fibrous saccharides
3. Carbohydrates
jachmadi@undip.ac.id
2. Micro
Inorganic
jachmadi@undip.ac.id
a.1.2. Non protein Peptides, amides, amines, nucelic acids, nitrates, urea, many nonprotein amino acids, and hundreds of other compounds containing N
a.2. Lipids
a.2.1. Essential fatty acids: linoleic, linolenic a.2.2. Sterols: cholesterol, vit. D, many other related compounds a.2.3. Terpenoids: carotene, xantophylls, others a.2.4. Waxes: cutin, others a.2.5. Phospholipids: lecithin, others a.2.6. Miscellaneous: free fatty acids, others
jachmadi@undip.ac.id
a.3. Carbohydrates
a.3.1. Monosaccharides: simple pentose or hexose a.3.2. Disaccharides: sugars with two molecules of simple sugars a.3.3. Oligosaccharides: sugars with more than two simple sugars but still relatively small molecules a.3.4. Nonfibrous polysaccharides: dextrins, starches, pectins a.3.5. Fibrous polysaccharides: hemicelluloses, celluloses, xylans
a.4. Miscellaneous
jachmadi@undip.ac.id
Lignin; organic acids; compound contributing to color, flavor and odor; toxins or inhibitors of various types; animal and plant hormones
b.1.2. Micro elements: Co, Cr, Cu, F, Fe, I, Mn, Mo, Ni, Se, Si, Sn, V, Zn
b.3. Nonessential elements: Ag, Al, Au, Bi, Ge, Hg, Pb, Rb, Sb, Ti
jachmadi@undip.ac.id
b.4. Often toxic: As, Cd, Cu, F, Hg, Mo, Pb, Se, SI
Non Ruminant
Require specific amino acid Unsaturated fatty acids pass through G.I. tract wihtout hydrogenation Ration is limited by dietary crude fiber [cellulose] Most essential vitamins must be supplied Most dietary P [from plant] could not be utilized
Ruminant
Do not require specific amino acid Rumen microbes undergo hydrogenation on unsaturated fatty acids Rumen microbes could degrade beta-linkage of several glucose molecules Rumen microbes could synthesize water soluble vitamins and vitamin K Rumen microbes could break phytate
Vitamins Minerals
DIGESTIVE PHYSIOLOGY
Digestion
Preparation of feed for absorption Reduction in food particle size mechanical - chewing chemical - HCl, bile enzymatic - lipase microbial
Monogastric (Simple stomach): humans, swine, poultry Monogastric with a functional cecum: horses, elephant, rabbit Ruminant (polygastric - 4 compartments): cattle, sheep, goats
10
jachmadi@undip.ac.id
11
Non Ruminant
Mechanical - chewing Physical breakdown Chemical digestion
Ruminant
Mechanical - chewing Physical breakdown Chemical digestion Microbial digestion Enzimatic digestion
Enzimatic digestion
jachmadi@undip.ac.id
Microbial digestion
Microbial digestion
12
DM
jachmadi@undip.ac.id
VS
DM DM basis
[concentrated nutrients]
13
Nutrient utilization to determine the proportion of nutrients in a feed or diet that are absorbed from gastrointestinal tract of animal [a biological trial].
14
Dry at 105oC
MOISTURE-FREE SAMPLE
ETHER EXTRACTION
ETHER EXTRACT
CRUDE PROTEIN
FAT-FREE SAMPLE
BURN IN FURNACE
CRUDE FIBER
ASH
15
SAMPLE OF ROUGHAGE
[Cell content]
N D S
[Cell wall]
N D F
[Lignocellulose]
jachmadi@undip.ac.id
A D F
A D S
[Cellulose]
ACID SOLUBLE
ACID INSOLUBLE
[Lignin]
16
17
Nutrient Digestibility [100%]: Nutrient intake - Nutrient in feces Nutrient intake X 100
18
Animals are fed a diet of known composition over a time period, during which the feces are collected and later analyzed for the components of interest. Maintaining a constant daily feed intake is advisable to minimize day-today variation in fecal excretion. Time required for feed residues to travese the GI tract 1 or 2 days for nonruminant, and 4 to 7 days for ruminant preliminary period of 3 days [nonruminant] and 10 days is needed to void the GI tract of residues of pretest feed and to allow adaptation of the animal to the diet.
jachmadi@undip.ac.id
A collection period of 4 days [nonruminant] and 10 days [for ruminant] follows the adjustment period. Values can be obtained for apparent digestibility of any desired nutrients, but meaningless for vitamin or minerals that are present in extremely small amounts.
19
20
Nutrient Utilization of Feed: Indicator Methods of Digestion Trials Often a choice when it is impossible or inconvenient to measure total feed intake or to collect total feces. It uses reference substance which should be indigestible, nonabsrobable, nontoxic, and easily analyzed in feed and feces.
jachmadi@undip.ac.id
This methods provides an estimate of digestibility of any or all nutrients without need to know either the total feed consumption or the excretion of feces.
21
Internal Indicator
Extensively uses for ruminant. Internal markers that are present in feed naturally, such as lignin that is digested to a negligable degree though it is incomplete recovery. The use of silica have a problem presumably because of contamination of soil.
External Indicator
Extensively uses for both nonruminant and ruminant.
jachmadi@undip.ac.id
External markers or indicators such as chromic oxide or rare earth elements that are either added to the feed or given to animal orally to animal [or administered into the rumen or cannula]
22
100
{ 100 X
% ind.feed % ind.feces
% nutr.feces % nutr.feed
Amount of feed consumption may be esimated using indicator methods: DMfc [units/day] = X [amount. ind. per unit dry feces] ind.fd/unit DMfd
jachmadi@undip.ac.id
If 1% of marker Cr2O3 was added to feed, and the marker Cr2O3 in feces were 3%; 3,1%; 2,9%; and 2,7% Then estimate the coefficient of feed digestibility!
23
jachmadi@undip.ac.id
24
It is extensively used for nonruminant, but there is possibility for ruminant. 1. Feeding a nitrogen-free diet and determining the amount of nitrogen excreted in feces. 2. Feeding a completely digestible nitrogen.
jachmadi@undip.ac.id
3. Feeding several levels of nitrogen and calculating the fecal level by regression analysis to a zero intake [use specific steps of calculation].
25
Item Daily N intake, g Daily fecal N, g Apparent N absorption, g [line 1 line 2] Apparent dig.,% [3/1x100] Metabolic fecal N, g Unabsorbed dietary N, g [line2 line5] True N absorption, g [line1 - line6] [1-6] True N dig., % [7/1 x 100]
7 8
To determine true digestibility of a protein, proceed sequenttially through the 8 steps as indicated in the column labeled Line No.. Note that true digestibility is not changed by level of protein intake, eventhough apparent digestibility increased by a high protein intake. [Pond et al., 1995].
26
jachmadi@undip.ac.id
27
Carbohydrates:
Cellulose 1,2,3,4--glucan Arabinoxylan Galactomannan -galactoside
Polypeptides
Pepsin
Trypsin Chymotrypsin Elastase Carboxy peptidase A& B
Oligosacchar ide Maltose Maltr iose Dextrins
Polypeptides Oligopeptides
Polypeptides Oligopeptides
jachmadi@undip.ac.id
Glucose
Glucoamylase; Disaccharidase
Amino acids
Monoglycerides
Glucose
Protein
Lipids
Lactose
Saccharose
Starch
28
jachmadi@undip.ac.id
Schematic representation of rumnal digestion of carbohydrate. Dg = degradable; Ug = undegradable; Hc, Hcell = hemicellulose; C, Cell = cellulose; St = starch; WSC = water soluble carbohydrate; ATP = adenosin triphosphate; VFA = volatile fatty acids; M, Micr = microbes, microbial; Duod = duodenal. (from : Beever, 1993).
29
Carbohydrate Metabolism
Digestion
In the mouth of nonruminants, amylase from saliva degrades starch to oligosaccharide units. The amylase saliva randomly catalyses the breakdown of 1,4-glicosidic bonds in starch. The action of amylase saliva is inhibited by a low pH of the stomach. In the small intestine, the digestion of starch is continued by the action of amylase from pancreas [amylopsin]. The oligosaccahrides are digested to monosccharides by the action of disaccharidases. In ruminants, some bacteria secrete cellulase, pentonase, and other carbohydases. These intracelluler enzymes catalyse the breakdown of dietary carbohydrates. Monosaccahrides and disaccharides are rapidly fermented to volatile fatty acids [VFA], and hemicellulose and cellulose are fermented more slowly. Acetic, propionic, and butiric acids are main products of carbohydrate digestion in rumen. The proportion of this VFA depends on the ratio of dietary roughage to concentrate.
comparative animal nutrition 30
jachmadi@undip.ac.id
jachmadi@undip.ac.id
31
Non carbohydrate compounds are also converted to glucose via gluconeogenesis pathway in liver. Glucogenic amino acids and short chain fatty acid could be converted to glucose via gluconeogenesis.
32
The storage of glycogen is limited, when ingested carbohydrate is in excessive amount over the requirement for glycogen formation, then glucose will be converted to the fat body. This process is accomplished by breakdown of glucose to pyruvate, and therefore it is ready for fat synthesis. The conversion of glucose to pyruvate is continued by lactate formation that occurs in anaerobic condition in muscle tissue [recall pathway of glycolisis]. Other route for glucose catabolism via oxidative pathway of phosphogluconate [pentose-phosphate pathway; oxidative shunt; pentose shunt]. Some enzymes that drive glucose utilization are found in liver, adipose tissue, cell of mammary gland. The pathway of pentose-phosphate is impportant: 1. to allow synthesis of ribose-5-phosphate, the main component of nucleic acid, 2. to allow by passing process for some tissues in glycolitic pathway, 3. to produce nicotine adenin dinucleotide phosphate [NADPH], an essential reductor in synthesis of fatty acids and reaction of hydroxylation.
jachmadi@undip.ac.id
33
In ruminants, oxidation of acetate mainly occurs in adipose and muscle tissues. Propionate is mainly for glucose synthesis. Propionate plays an important role for stabilization of blood glucose, especially in ruminant fed on high roughage, thus only small amount of glucose is absorbed from GI tract. Some studies reported that more than 30% of propionate from rumen is utilized for glucose synthesis. After transportation to blood circulation, VFA enters the tricarboxylic acid [TCA] cycle, and is used to meet the energy requirement of some body processes:
Acetic acid + 2 ATP Acetyl coenzyme A TCA cycle
10 mol ATP + CO2 + H2O Succinyl coA TCA cycle
jachmadi@undip.ac.id
Propionic acid
Methylmalonyl coA
Butyric acid
Butyryl coA
Acetyl coA
34
Feed
NFE
VFA
SIMPLE SUGAR
Blood
NFE Rumen
VFA
Small Intestine
NFE
VFA
FIBRE COMPONENTS
Tissue
jachmadi@undip.ac.id
MICROBIAL CARBOHYDRATE
Feces
35
Lipids Metabolism
Digestion The ingested lipids react with bile salts in small intestine, and then are digested by pancreatic lipase digest. The bile salts decrease the surface tension of the ingested lipids mass [ as a chyme], this causes an emulsification of fatty materials. The pancreatic lipase then forms emulsion from lipids particle, this may significantly result in increased surface area of the water insoluble material. And this materials are more reactively to pancreatic lipase. Pancreatic lipase are secreted in an inactive form in the lumen of small intestine. The presence of Ca2+ may activate the pancreatic lipase. This enzyme is very reactive to emulsified substrates. The main action of this enzyme is on an ester group of hydroxyl [in the position of 1 and 3 of glycerol]. The end product of pancreatic lipase action in small intestine is 1,2diglycerides.
jachmadi@undip.ac.id
36
Rumen microbes hydrolyze triglycerides and some lipids. Generally, lipid from plants could be hydrolized completely compared to lipid and glycerol from animal origin. The product of lipid hydrolyzation is triglycerides, then are fermented to be propionic acid by rumen microbes. In most lipids of plant, unsaturated fatty acids have cis configuration, but most fatty acids of ruminant body have trans configuration. This fact showed that the fermentation of ingested lipids in rumen produces some fatty acids with configuration of trans-isomer. When ruminants are fed on free fat diet then the fatty acids of C14, C15, C16 and C18 will be synthesized in the rumen. Fatty acids in the body of ruminants could be synthesized from ingested carbohydrate, protein and lipid. And the most interesting thing, methionine could stimulate the synthesis of LCFA from glucose and acetic acid.
jachmadi@undip.ac.id
37
Schematic Representation of lipid digestion, absorption, and resynthesis. The heavy arrows indicate the more important pathway [Pond et. al. 1995]
jachmadi@undip.ac.id
38
Triglyceride
1,2-diglyceride
Pancreatic lipase
jachmadi@undip.ac.id
2-monoglyceride
39
Hydrogenation isomerase [+ H ]
2+
Isomeration
Hydrogenation [+ H ]
40
Schematic diagram of major conversions that occur in transport lipids accross the intestinal mucosal cell during absorption [Pond et al., 1995]
jachmadi@undip.ac.id
41
jachmadi@undip.ac.id
42
After entering the epithelial cell, LCFA are converted to derivatives of coenzyme A in the presence of ATP. This fatty acid-coenzyme A complex [fatty acyl coenzyme A] reacts with monoglycerides within cell to form di and then triglycerides. The triglycerides thus fromed contain only FA of C12 or greater chain length because shorter chain FA are absorbed directly into portal system. Before leaving the mucosal cell, the mixed lipid droplets become coated with a thin layer of protein absorbed to the surface. These protein coated lipid droplets are called chylomicrons and consist mainly of triglycerides with small quantities of phospholipids, cholesterol esters, and protein. The chylomicrons leave the mucosal cell by reverse pinocytosis and enter the lacteals via intercelluler spaces. Lacteal lead to the lymphatic system, which carries the chylomicrons to the blood via the thoracic duct. Although mammals absorb most of these LCFA into the lymphatic system, the chicken apparently absorbs its dietary lipids into the portal blood, which carries them dierctly to the liver. Nevertheless, the processof reesterification of FA to triglycerides in the mucosal cell is similar in birds and mammals.
jachmadi@undip.ac.id
43
Fatty acids are absorbed via active transportation within rumen because there is a gradation of energy concentration between fatty acid in the rumen and in the blood circulation. This fact showed that each two molecules of absorbed fatty acid may loss one molecule of CO2 and forming one molecule of bicarbonate (HCO3). After the absorption of VFA from rumen wall: The concentration of -hydroxy butyric acid in blood vena is higher than in blood artery, because butyric acid is oxidised to -hydroxy butyric acid; The cells of rumen wall do not convert acetic acid; The propionic acid do not change significantly. The fatty acids in blood circulation become precursor for LCFA, and along with glycerol forming milk or body fat. Some factors affecting the synthesis of milk fat: The ration of acetate : propionat, the more higher this ratio value more higher the synthesis of milk fat, The more lower rumen pH [to normal limit] more higher the synthesis milk fat; The -hydroxy butyric acid become precursor of milk fat, because 8-10% of hydroxy butyric acid supplies the carbon skeleton of milk fat; The more higher glucogenic precursor [glucose] more higher milk fat, but the precursor of non glucogenic decreases LCFA.
comparative animal nutrition 44
jachmadi@undip.ac.id
Mobilization and Deposition of Fat The Free fatty acid [non esterified fatty acid] is transported as a complex with albumin. Chylomicrons are removed rapidly toward liver, fat storage, and other tissues. The body tissues store triglycerides, the adipose tissue is an important fat storage. The adipose tissues have capability in synthesis and mobilisation of fat from carbohydrate source and oxidized fatty acid. The synthesis and mobilisation of triglicerides in adipose tissue occur continuously, because triglyceride storage is readily as the energy source. When the energy intake exceeds the requirement, triglycerides will be stored, and vice versa. The triglyceride storage tends to be specifically according to the animal species. In non ruminants, the feed lipids reflects on the fat storage. In ruminants, the fat storage in body is less sensitive to the the feed lipids because of lipid metabolism in the rumen, though there is a little effect of feed lipids on body fat storage. When the action of rumen microbes is inhibited, the body fat storage is influenced by the feed lipids. The protection of feed lipid may inhibit the action of rumen microbes.
comparative animal nutrition 45
jachmadi@undip.ac.id
jachmadi@undip.ac.id
The body fat storage of ruminant could be manipulated by adding more readily available carbohydrate. Normally, the body fat storage in ruminants is characterized by odd and branhced carbon chain as the ruminal VFA derivatives. Also the body fat storage in ruminants is characterized by configuration of trans isomer as the results of metabolism of feed unsaturated fatty acid. The fat body of ruminants apparently on a dynamic of metabolic status. The turnover rate of triglyceride is very rapidly in the storage of ruminant body. The lipid metabolism of adult ruminants is apparently different to nonruminant: Feed lipid [triglycerides, phospholipids, and galactolipids] is degraded by the rumen microbes, and the occurence of microbial lipid synthesis. The hydrogenation of fatty acid, the unsaturated fatty acid is saturated to saturated fatty acid. Lysolecithin did not act as micell membrane formation, and monoglycerides do not act as stabilizator in micell formation. The resynthesis of triglycerides in epithelial intestine uses the pathway of -glyceroacetic, and does not use the monoglycerides pathway.
46
Protein Metabolism
Digestion
Digestion of ingested protein or hydrolysis of peptide bonds begins in stomach with the presence of acidic condition [pH 2-3], it is an optimum pH for pepsin proteolytic. The product of pepsin digestion, oligopeptides and polypeptides, then react with pancreatic proteases, trypsin, chymotripsin, and carboxypeptidase. The product of these protease digestions are free amino acid and oligopeptides. In young ruminants, rennin of stomach coagulates ingested casein rapidly.
jachmadi@undip.ac.id
In adult ruminants, ingested protein consist of pure protein and non protein nitrogen. The protein that are bypassed from reticulo rumen fermentation then will be degraded in the small intestine [just like in the monogastric].
47
In adult ruminants, amino acid is hydrolyzed for : a] synthesis of protein microbes, or b] deamination to form organic acids, ammonia and CO2. The ammonia then is combined with keto acids to form new amino acids for synthesis of protein microbes, and are absorbed to protal circulation liver urea. Most of urea are filtered by kidney and then are excreted via urine. A part of urea may be recycled to rumen via saliva or passthrough the rumen wall [via blood circulation]. In rumen, urea is converted to ammonia and CO2 by the action of urease. In the synthesis of protein non nitrogen, rumen microbes need a large amount of organic acids that are originated from ingested starch.
jachmadi@undip.ac.id
Ingested protein that enters to abomasum [true stomach] : bypassed protein from reticulorumen degradation, protein saliva from degestion process, microbe protein, amino acids originated from resynthesis of deamination.
48
jachmadi@undip.ac.id
Mode of Nitrogen Transaction in the Rumen. The ovals delinetae the microbial cell wall and numbers adjacent to arrows refer to individual pathways as follows: 1 proteolysis by bacterial, protozoal and fungal proteases; 2 carrier-mediated peptide uptake across microbial cell walls; 3 peptidolysis; 4 amination/demination; 5 protein synthesis; 6 microbial assimilation/excretion or equilibration of amino acids and ammonia; 7 protein not hydrolyzed before efflux from rumen [UDP]; 8 microbial protein efflux; 9 efflux of extracellular peptides and amino acids; 10 efflux of extracellular ammonia; 11 absorption of ammonia through rumen wall; 12 movement of endogenous urea through the rumen wall; 13 N compounds excreted by living cells and debris of lysed cells; 14 engulfment of proteinaceous particle by protozoa [from: Nolan, 1993]
49
The digestion of ingested protein in small intestine is mainly by the action of pancreatic proteases, i.e. trypsinogen, chymotrypsinogen, and procarboxypeptidase]. Upon a proteolitic cleveage, these pancreatic proproteases are then activated. Trypsinogen is activated by trypsin itself or by enterokinase that is secreted from small intestine mucose. Activation of trypsinogen consist of transfer of simple hydrolytic on one hexapeptide, Val-Asp4-Lys, from end terminal of amine group. The conversion of chymotripsinogen to chymotripsin occurs in three different steps. The combination of these pancreatic proteases degrade protein molecule to free amino acids and small amount of peptides. As in carbohydrate digestion, the hydrolysis is ended by hydrolase action from mucose of small intestine. Some hydrolysis action occur in the lumen of small intestine, but most of hydrolysis occur as peptides enter and contact to brush border epithel or in mucose cells. Leucine aminopeptidase, a non-specific exopeptidase, is responsible for most of the peptidase activities. Though some peptidases remain to be elucidated for their roles in the end of digestion process.
comparative animal nutrition 50
jachmadi@undip.ac.id
Absorption The intact protein or small chain of polypeptides is unable to be absrobed by small intestine of adult animal. The absorption of free amino acids occur in the small intestine using an active transportation system. The rate of absorption depends on type of amino acid, though there is no spesific transportation system for each type of amino acid, there is a common transportation system. The absorbed amino acids from small intestine then enter portal vena, thus they are preceded a process in the liver to be available for body tissue metabolism. The absorption of amino acid by an active transportation system. The amino acids that enter membrane cell of small intestine move againts gradient of concentration, this uses energy from cellular metabolism. L-amino acids are more absorbable than D-amino acid. Each neutral amino acid may compete for each transportation. For example, a high amount of leucine in the diet may increase the need of isoleucine.
jachmadi@undip.ac.id
51
Removing a carboxyl group by forming an ester, removing the electric charge of amino group with acetylation, or introducing the electric charge on side chain of some amino acids may destroy the system of active transport. This caused by a high natural specifity of the carrier system. The basic amino acids, ornithine, arginine, and lysine have a similar transportation system along with cystine. Arginine, cystine and ornithine inhibit the transportation of lysine. Some neutral amino acids inhibit the transportation system of some basic amino acids. For example, methionine inhibits the transportation of lysine.
jachmadi@undip.ac.id
The basic amino acids is unable to inhibit the transportation of neutral amino acids apparently. Proline and hydroxyproline have a simlar transportation system to sarcosine and betaine, and have a high affinity for the transportation of neutral amino acid.
52
Feed
PROTEIN PROTEIN
Rumen, Reticulum
AMMONIA
Liver
jachmadi@undip.ac.id
Urine
Feces
Tissue
53
Tissue protein
Amino acids
Deamination
Relationship between ingested proteins and tissue proteins. Amino acids from both sources form a common metabolic pool which can provide amino acids for resynthesis of proteins or which can be metabolized to other end products.
54
References cited: Forbes, J.M. and J. France [eds.]. 1993. Quantitatice Aspetcs of Ruminant Digestion and Metabolism. CAB international, the University press, Cambridge. Pond, W.G., D.C. Church, and K.R. Pond. 1995. Basic Animal Nutrition and Feeding. John Wiley & Sons. New York. Preston, T.R. And R.A. Leng. 1987.Matching Ruminant Production Systems with Available Resources in the Tropics and Sub-Tropics. Preambul Books, Armidale.
55