Animal Nutrition
Nutrient classes, non-ruminant and ruminant digestion, feed additives, and lipid metabolism (ANSC 223, Lectures 1–5)
Classes of Nutrients
Definitions
- Nutrition: series of processes by which an organism takes in and assimilates food for growth, tissue maintenance, and production (milk, eggs, wool)
- Nutrient: any chemical element or compound in the diet that supports growth, reproduction, work, lactation, or maintenance of life processes
- Essential nutrient: removing it from the diet causes an abnormality; adding it back eliminates the abnormality (e.g. Ca deficiency → milk fever; Ca/P/Vitamin D deficiency → rickets)
- Feed = food for farm animals; Diet = mixture of feedstuffs supplying nutrients; Ration = supply of feed at a feeding or daily
Six Main Classes of Nutrients
Water
- Animals need ~2× as much water as dry feed intake; can survive much longer without food than without water
- Makes up ~60% of body; muscle tissue ~75% water, fat tissue ~15% water
- Sources: drinking water, water in food, metabolic water (from C₆H₁₂O₆ → H₂O + CO₂ + ATP)
- Water loss: urine (~60%), feces, skin and lungs (15–60%)
- Intake increased by: lactation (dairy cow needs ~0.9 kg water per kg milk), dietary salt, heat stress
Carbohydrates
- Primary source of energy; made of repeating CH₂O units; derived from plants (except lactose)
- Structural carbs: cellulose, hemicellulose (high fiber, not digestible by monogastrics without hindgut fermenters or rumen)
- Non-structural carbs: sugars, starch (corn = 85% carbohydrate)
Lipids
- Soluble in organic solvents; provide 2.25× more energy than carbs/proteins (9.4 kcal/g vs ~4 kcal/g)
- Fats: long-chain, saturated, solid at room temp (e.g. tallow)
- Oils: long-chain, unsaturated, liquid at room temp (e.g. soy oil)
- Functions: energy, solvent for fat-soluble vitamins (A, D, E, K), source of essential fatty acids, palatability
Protein
Most expensive nutrient added to diets. Contains C, O, H, and N (nitrogen). Source of essential amino acids. Molecular makeup involves chains of amino acids linked by peptide bonds.
Vitamins
Fat-Soluble (A, D, E, K)
Can be stored in the animal. Fortification/enrichment needed (e.g. milk fortified with Vitamin D).
Water-Soluble (8 B vitamins + Vitamin C)
Cannot be stored — must be provided daily. B vitamins synthesized by rumen microbes in ruminants.
Fortification = adding nutrients not already present. Enrichment = replenishing nutrients lost during processing.
Minerals
Macrominerals (large amounts in body)
Calcium, Sodium, Chloride, Potassium, Magnesium, Sulfur
Microminerals / Trace Minerals
Selenium, Copper, Iron, Zinc, Cobalt
Minerals must be fed in proper balance — over or underfeeding causes serious problems. Cannot be decomposed or synthesized by the body.
Forages vs Concentrates
| Component | Forage (Roughage) | Concentrate (Grain) |
|---|---|---|
| Energy | Low | High |
| Protein | Low | Low–moderate |
| Fiber | High (stimulates rumination) | Low |
| Ca & K | High | Low |
| Fed to | Cattle, horses, sheep, exotics | Poultry, pigs, dogs, cats, fish |
Feed Efficiency
Ability of an animal to convert a unit of feed into a unit of body mass. Major determinant of cost.
| Animal | Gain (kg/d) | Feed (kg/d) | Feed:Gain | Gain:Feed |
|---|---|---|---|---|
| Chicken | 0.9 | 1.1 | 1.2 | 0.82 |
| Young pig | 0.3 | 0.5 | 1.7 | 0.60 |
| Finishing pig | 1.0 | 2.4 | 2.4 | 0.42 |
| Beef steer | 1.6 | 10 | 6.3 | 0.16 |
Factors affecting efficiency: diet quality, age (mature animals less efficient), composition of gain (fat vs lean), endocrine status, environment, genetics.
Non-ruminant Digestion
GI Tract Types
Simple Stomach
Dogs, cats, pigs, humans. One stomach compartment.
Hindgut Fermenter
Horses, rabbits, guinea pigs, rhinoceros, elephant. Large cecum for fiber fermentation.
Avian
Chickens, turkeys. Crop → proventriculus → gizzard → short small intestine → ceca → cloaca.
Three forces acting on food in the GIT: mechanical (chewing, peristalsis), chemical (HCl, bile), enzymatic (amylase, lipase, proteases).
Stomach — Three Regions
- Cardiac region: mucus secretion
- Fundic region: HCl (acidifies), zymogens (pepsinogen, prorennin), mucus
- Pyloric region: mucus, gastrin (hormone controlling gastric juice flow)
- Zymogens are inactive enzyme precursors activated by hydrolysis (e.g. pepsinogen → pepsin)
- Rennin: complex of enzymes in stomach of young pre-ruminants; curdles milk by coagulating casein into curds → retains milk longer for digestion
- Lingual lipase: secreted by tongue glands; initiates fat digestion in neonates
Small Intestine
- Three sections: Duodenum (digestion), Jejunum (digestion + absorption), Ileum (mostly absorption)
- Villi increase surface area for absorption. Malnutrition causes villus atrophy.
- Secretin (triggered by low pH) and Cholecystokinin (CCK) (triggered by lipid + peptides) secreted by duodenum
| Source | Enzymes / Secretions | Substrate |
|---|---|---|
| Pancreas | Trypsin/Chymotrypsin/Carboxypeptidase (as zymogens), Pancreatic amylase, Pancreatic lipase + colipase | Protein, Starch, Lipid |
| Small intestine | Enteropeptidase (activates trypsinogen), Maltase, Sucrase, Lactase, Peptidases | Trypsinogen, Maltose, Sucrose, Lactose, Peptides |
| Liver/Gallbladder | Bile (bile salts) | Lipid emulsification → ↑ surface area for lipase |
Absorption Mechanisms
Simple Diffusion
High → low concentration. No energy needed.
Active Transport
Against concentration gradient. Requires ATP. Important for glucose and some amino acids.
Protein-Mediated Transport
Facilitated diffusion via carrier proteins.
Hindgut Fermenters (Horses)
- Large cecum (blind pouch) + large colon provide microbial fermentation of fiber
- Produces short-chain VFAs (Volatile Fatty Acids), water-soluble vitamins, and protein
- Only VFAs and water-soluble vitamins are absorbed from the cecum/colon
- VFAs can provide over 70% of horse's energy requirement
Avian GI Tract
- Crop: food storage and moistening
- Proventriculus: glandular stomach (HCl + enzymes)
- Gizzard: muscular stomach; mechanical breakdown using grit (sand and small stones). No sphincter separating it from duodenum.
- Ceca (2): microbial fermentation, water absorption
- Cloaca: common chamber for GI tract, urinary tract, and egg laying
- Phytate-phosphorus (main plant P storage form) requires phytase enzyme for release
Ruminant Digestion
Ruminant Animals
True ruminants (Bovidae, Cervidae, Giraffidae): cattle, sheep, goats, bison, moose, reindeer, giraffe, wildebeest, antelope. Pseudo-ruminants (Tylopoda): camels, llamas.
Key characteristic: consume large amounts of fibrous material quickly, then rest and ruminate (chew cud). Microbial fermentation in a multi-chambered foregut makes otherwise indigestible plant cellulose available — animals cannot make cellulase, but rumen microbes can.
4-Compartment Stomach
Reticulum (Honeycomb)
Origin of contractions, rumination (regurgitation), and eructation. Traps foreign objects. No enzymes secreted.
Rumen
Large fermentation vat (~100L). Papillae for absorption. pH 6–7 maintained by saliva. Contains 500,000 billion bacteria, 50 billion protozoa. Gas produced must be released via eructation — failure causes bloat (diaphragm compression → asphyxia).
Omasum
Filters digesta flow from rumen. Laminae reduce particle size. VFAs and water absorbed. No enzymes secreted.
Abomasum (True Stomach)
Only glandular compartment. Secretes HCl, pepsin, and mucus. Lysozyme in mucus breaks down bacterial cell walls. Displaced abomasum occurs when diet is too low in forage.
Rumen Microbes & Fermentation
- Strictly anaerobic environment. >60 bacterial species and 20 protozoal species are normal inhabitants.
- Three niches: sugar fermenters (~3% of typical diets), fiber digesters (cellulose/hemicellulose), starch fermenters (grain diets)
- Protozoa: prey on bacteria, engulf starch granules, important in N recycling
- Heat of fermentation: rumen is warm; up to 10% of total energy lost as heat. Asset in cold stress, liability in heat stress.
- Methane: 5–10% of ingested energy lost as CH₄. Highest with forage, lowest with grain diets. Cattle in US contribute ~20% of all atmospheric CH₄.
VFA Production & Use
Fiber (forage) diet → high acetate
Acetate (C2): lipogenesis, energy. Propionate (C3): small proportion. Butyrate (C4): energy.
Starch (grain) diet → high propionate
Acetate (C2): reduced. Propionate (C3): 70–90% goes to liver → gluconeogenesis. Butyrate (C4): energy.
VFAs absorbed through rumen papillae → portal vein → liver. Propionate is primary gluconeogenic substrate in ruminants.
Protein Digestion in Rumen
- Bacteria break down soluble protein to peptides and ammonia (enzymes on bacterial surface)
- Bacteria synthesize microbial protein from the peptides and ammonia
- Microbial protein digested in the abomasum (true stomach)
- Excess ammonia → liver → urea → recycled in saliva or excreted in urine
Acidosis
- Caused by: accidental grain overload, too-rapid diet change to high grain
- Lactic acid accumulates (10× more acidic than VFAs) → rapid pH drop
- Sub-acute ruminal acidosis (SARA): pH 5.0–5.5; Acute: pH ≤ 4.5
- Below pH 6: cellulolytic and methanogenic bacteria decrease, amylolytic bacteria increase, Lactobacillus spp. increase
- Prevention: ≥10% roughage in finishing rations, gradual diet transitions, buffers (NaHCO₃), ionophores
Neonate Ruminants
Born with non-functional rumen. Cleaned by mother. Rumen nearly functional at ~60 days. Suckling causes esophageal (reticular) groove to close → milk bypasses rumen directly to omasum. Diet drives rumen development: grain promotes papillae growth more than hay alone.
Feed & Food Additives
Definitions
- Non-nutritive additives: any compound added to the diet for reasons other than nutrient supply
- Drugs: substances intended for treatment, prevention, or diagnosis of disease — require FDA approval
- GRAS (Generally Recognized As Safe): substances with published safety/utility information; efficacy must still be proven for each new product
Non-regulated Additives
Regulated Additives
Antibiotics
- Bacteriostatic (prevent growth) or bactericidal (kill bacteria)
- Used as growth promotants: control sub-clinical disease, nutrient-sparing effect, metabolic effect (more ATP available)
- Problem: antibiotic resistance and potential transfer to human pathogens
- FDA Guidance 209/213: strict withdrawal times before market (monitored by APHIS)
- Common: Oxytetracycline, Chlortetracycline, Tylosin (Tylan), Carbadox (swine)
Ionophores (Ruminants)
- Allow ions to move through cell membranes of gram-positive bacteria → selectively inhibit acetate-producing and lactic acid-producing bacteria
- ↑ propionate, ↓ acetate in rumen → improved feed efficiency
- Also reduce bloat, acidosis, coccidiosis risk, and methane production
- Used in ~95% of feedlot cattle diets (fed at 10–13 g/ton)
- Monensin (Rumensin): 80% market share. ↑ feed efficiency 5–10%, ↑ weight gain 2–7%
- Lasalocid (Bovatec): 15% market share
- Warning: Horses DO absorb ionophores — very toxic, no treatment available
Beta-Agonists (Ractopamine)
- β-adrenergic agonist — shifts energy partitioning from fat to lean muscle tissue
- Not an antibiotic, not a hormone. No withdrawal period.
- Paylean (pigs): last 28 days before market, 4.5–9 g/ton. ↑ carcass wt 10.6 lbs, ↑ feed efficiency 7–15%, ↑ daily gains 6–20%
- Optaflexx (cattle): 200–400 mg/head/day for 28–42 days. ↑ carcass wt 15–20 lbs, ↑ feed efficiency 10–15%
Buffers, Probiotics & Enzymes
- Buffers: NaHCO₃ at 0.75% DM most common for dairy cattle and feedlot cattle during diet adjustment. MgO also used. Prevent rumen acidosis on high-grain diets.
- Organic acids: Citric acid most common for young pigs and chicks. Lowers stomach pH → improved protein digestion, reduced pathogen incidence.
- Phytase: most common added enzyme. From Aspergillus or E. coli. Releases phytate-bound phosphorus from plant feeds; ↑ P absorption 5–7%, ↓ P excretion 50–75%. Primarily for non-ruminants.
- Probiotics (direct-fed microbials): live Lactobacillus, Streptococcus, Bacillus, yeast cultures. Most effective in stressed, newly weaned, or relocated animals.
- Prebiotics: non-digestible ingredients (food for good bacteria); promote lactic-acid bacteria, suppress E. coli. Primarily in pet foods.
- Yeast (S. cerevisiae): source of B vitamins and mannan-oligosaccharides. In dairy cattle: higher rumen pH, ↑ cellulolytic bacteria, ↑ fiber digestion, ↑ microbial protein synthesis.
Bloat
Frothy (pasture) bloat: caused by soluble proteins and saponins from legumes (clover, alfalfa) → insufficient saliva → slime production → foam traps gas. Sudden death risk. Prevention: poloxalene (Bloat Guard) = non-ionic surfactant that consolidates gas bubbles → eructation. Mineral oil also used.
Lipids
Classification of Lipids
Simple Lipids (glycerol-based)
Triacylglycerols (TAGs): 3 fatty acids ester-bonded to glycerol. >90% of dietary lipids.
Sterols: cholesterol (zoosterol), phytosterols (plants). Precursor of steroid hormones, bile acids, Vitamin D.
Compound Lipids
Phospholipids (Lecithin): major cell membrane component; amphiphilic → emulsification; body store of choline (deficiency → fatty liver).
Glycolipids: CHO + lipid; major lipids in forage leaves.
Lipoproteins: transport lipids in blood (chylomicrons, VLDL, LDL, HDL).
Non-glycerol Lipids
Waxes (surface lipids), sphingomyelins, cerebrosides, terpenes, prostaglandins.
Fatty Acids by Carbon Chain Length
Named by convention: C[length]:[double bonds], ω-[methyl-end position]. Melting point ↑ with chain length; ↓ with unsaturation (double bonds).
| Formula | Name | Class | Source / Origin | Biological Role | Animal Notes |
|---|---|---|---|---|---|
| Short-Chain (C2–C6) — Volatile Fatty Acids (VFAs), water-soluble, absorbed directly into portal blood | |||||
| C2:0 | Acetic acid (acetate) | Sat VFA | Rumen fermentation of fiber (forage diet); vinegar | Energy (enters TCA as acetyl-CoA); substrate for lipogenesis and milk fat synthesis | 60–70% of rumen VFAs on forage; ↓ on grain diets; ionophores reduce acetate production |
| C3:0 | Propionic acid (propionate) | Sat VFA | Rumen starch fermentation (grain diet); silage | Primary gluconeogenic substrate in ruminants; enters TCA as succinyl-CoA | 70–90% goes to liver on grain diet; ↑ with ionophores; critical for blood glucose maintenance in dairy cows |
| C4:0 | Butyric acid (butyrate) | Sat VFA | Rumen fermentation; butter (3–4% of milk fat) | Energy for rumen epithelial cells and colonocytes; drives rumen papillae development | ~15% of VFAs; ↑ grain diet; critical for neonatal rumen development (grain feeding initiates papillae growth) |
| C6:0 | Caproic acid | Sat | Milk fat; minor rumen product | Minor energy; direct portal absorption without chylomicrons | Antimicrobial against gram-positive bacteria; rapid absorption in neonates |
| Medium-Chain (C8–C14) — MCTs; absorbed via portal vein, bypass lymph and chylomicron packaging | |||||
| C8:0 | Caprylic acid | Sat MCT | Coconut oil, palm kernel oil | Rapid oxidation to ketones; immediate energy; antifungal/antimicrobial | MCT oil supplement; used in pet foods for cognitive support; antimicrobial in poultry |
| C10:0 | Capric acid | Sat MCT | Coconut oil, palm kernel oil | Rapid energy; antifungal (disrupts lipid membranes of pathogens) | Reduced incidence of pathogenic E. coli in piglets when added to diets |
| C12:0 | Lauric acid | Sat MCT | Coconut oil (45–55% of fat); palm kernel; some milk fat | Potent antimicrobial (gram-positive bacteria); disrupt viral lipid envelopes | Raises LDL and HDL; monolaurin derived from it is antiviral; added to nursery pig diets; soap/personal care industry |
| C14:0 | Myristic acid | Sat | Coconut/palm kernel, dairy fat, nutmeg | Protein myristoylation (signal transduction); structural membrane FA | Raises LDL cholesterol; significant component (~10%) of ruminant milk fat |
| Long-Chain Saturated (C16–C18:0) — Solid at room temp; main storage fats in animals; absorbed via lymph as chylomicrons | |||||
| C16:0 | Palmitic acid | Sat | Palm oil; beef/pork tallow; de novo lipogenesis end product; ~25% of mammalian adipose | Structural FA in membranes and stored fat; primary product of fatty acid synthase (FAS) | ↑ LDL in excess; excess accumulates in liver (fatty liver); most abundant saturated FA in animal body; ↑ on high-grain dairy diets |
| C18:0 | Stearic acid | Sat | Beef tallow, cocoa butter; end product of rumen biohydrogenation (MP +70°C) | Desaturated to oleic (C18:1) by Δ9-desaturase in tissues; structural membrane FA | Uniquely does NOT raise LDL (converted to oleic in body); high in ruminant fat post-biohydrogenation; solid at room temp |
| Long-Chain Unsaturated (C16:1–C18:3) — Liquid at room temp; one or more double bonds | |||||
| C16:1, ω-7 | Palmitoleic acid | MUFA | Macadamia nuts; fish; ruminant adipose | Acts as a lipokine (adipose-secreted signaling molecule); anti-inflammatory | Biomarker for metabolic syndrome risk; produced by Δ9-desaturase from palmitic acid |
| C18:1, ω-9 | Oleic acid | MUFA | Olive/canola oil; lard; tallow; most abundant MUFA in animal tissues | ↓ LDL without ↓ HDL; structural membrane FA; neutral/beneficial cardiovascular effect | Dominant FA in poultry and pork fat; high-oleic pork has improved shelf stability; product of Δ9-desaturation of stearic acid |
| C18:2, ω-6 | Linoleic acidEssential | PUFA ω-6 | Soybean, sunflower, corn oil; most plant oils | Skin integrity, RBC membrane structure, fertility, normal growth; precursor to arachidonic acid (C20:4, ω-6) via Δ-6 desaturase | Dogs: ≥1% DM required. Cats: ≥1% DM required. Swine/poultry most commonly deficient; soybean oil supplemented. EFA deficiency → scaly dermatitis, poor coat, hair loss |
| C18:3, ω-3 | α-Linolenic acid (ALA)Essential | PUFA ω-3 | Flaxseed oil (55%), chia, hemp; green leaves; canola (10%) | Eye and CNS structural lipid; precursor to EPA and DHA; competes with ω-6 for Δ-6 desaturase | Flaxseed feeding to hens → ω-3 enriched eggs; feeding to dairy cows → ω-3 enriched milk; high conversion rate to EPA/DHA in ruminants vs. monogastrics |
| Very Long-Chain PUFA (C20–C22) — EFA derivatives; precursors to eicosanoids and critical for neural/retinal function | |||||
| C20:4, ω-6 | Arachidonic acid (AA)Semi-essential | PUFA ω-6 | Animal tissues, egg yolk; synthesized from linoleic via Δ-6 desaturase → elongase → Δ-5 desaturase | Membrane phospholipid; primary eicosanoid precursor: prostaglandins (PG), thromboxanes (TX), leukotrienes (LT), prostacyclins | Cats are essential: low Δ-6 desaturase activity → cannot convert linoleic to AA fast enough; require ≥0.02% DM dietary AA (from animal-origin food). Eicosanoids regulate inflammation, platelet aggregation, vasodilation/constriction |
| C20:5, ω-3 | EPA (Eicosapentaenoic acid) | PUFA ω-3 | Fish oil; algae; elongation/desaturation of ALA | Competes with AA for COX/LOX enzymes → produces less inflammatory eicosanoids; reduces platelet aggregation; ↓ TG levels | Fish oil supplement in breeding mares and dairy cows → improved fertility and reduced inflammation; anti-inflammatory eicosanoids (PGE3, TXA3); omega-3 enriched products |
| C22:6, ω-3 | DHA (Docosahexaenoic acid) | PUFA ω-3 | Fish oil; algae; brain (40% of brain PUFA); retina (60% of retinal PUFA) | Critical for neural membrane fluidity, retinal photoreceptor function, fetal brain development | Essential for neonatal brain/vision; omega-3 enriched eggs via algae meal feeding to hens; farmed salmon fed algae-based DHA; critical in gestation for fetal CNS |
Melting point rule: ↑ chain length → ↑ MP; ↑ double bonds → ↓ MP (unsaturation dominates). Cis bonds = natural (bent chain, low MP); trans = biohydrogenation or industrial processing (straighter chain, behaves like saturated FA).
Biohydrogenation: Rumen microbes add H atoms to unsaturated FAs → saturated FAs. Net result: dietary C18:2/C18:3 largely converted to C18:0 (stearic) before small intestine. Ruminant body fat is therefore more saturated than monogastric fat on the same diet.
Eicosanoids from PUFA
From Arachidonic Acid (ω-6) — Pro-inflammatory series
- PGE2, PGF2α (Prostaglandins): inflammation, fever, uterine contraction, ovulation
- TXA2 (Thromboxanes): platelet aggregation, vasoconstriction
- LTB4 (Leukotrienes): neutrophil chemotaxis, bronchoconstriction
From EPA (ω-3) — Anti-inflammatory / weaker series
- PGE3, PGF3α (Prostaglandins): weaker pro-inflammatory effect
- TXA3 (Thromboxanes): weak platelet aggregation
- LTB5 (Leukotrienes): weak chemotaxis
ω-3 and ω-6 compete for the same COX and LOX enzymes. A higher dietary ω-3:ω-6 ratio shifts production toward less inflammatory eicosanoids.
Lipid Digestion & Absorption
- Gastric/lingual lipase begins lipid digestion in stomach (primary in neonates)
- In duodenum: bile salts emulsify dietary lipids (↑ surface area)
- Pancreatic lipase + colipase hydrolyze TAGs → monoacylglycerol + 2 free fatty acids
- Products form micelles → absorbed across intestinal epithelium
- Re-esterified into TAGs → packaged into chylomicrons → absorbed into lymph via lacteals → enter bloodstream
- Lipoprotein lipase removes fatty acids from chylomicrons at capillaries → deposited in adipose or muscle. Chylomicron remnant → liver.
Short- and medium-chain FAs (C2–C12) bypass chylomicron packaging and are absorbed directly into the portal vein → liver.
Lipoproteins in Blood
| Lipoprotein | % Protein | % TAG | % Cholesterol | Role |
|---|---|---|---|---|
| Chylomicron | 1–2 | 85–95 | 3–7 | Transport dietary fat from gut → tissues |
| VLDL | 6–10 | 50–65 | 20–30 | Transport liver-synthesized TAG → tissues |
| LDL (bad) | 18–22 | 4–8 | 51–58 | Deliver cholesterol to tissues (atherosclerosis risk) |
| HDL (good) | 45–55 | 2–7 | 18–25 | Reverse cholesterol transport (tissue → liver) |
Ruminant Lipid Considerations
- Biohydrogenation: rumen microbes convert dietary unsaturated FAs → saturated FAs (adds H atoms). End product: mostly C18:0 (stearic). Produces trans FAs as intermediates. After rumen, small intestine functions like monogastric.
- Fatty liver disease: common in high-producing dairy cows post-partum and overweight non-ruminants. High free fatty acids from adipose overwhelm liver oxidation and VLDL synthesis capacity → hepatocytes fill with TAG → reduced liver function.
- Ruminants have more bile acids than monogastrics, compensating for the more saturated FA profile entering the small intestine.