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An Overview of Metabolism
Metabolism
Metabolism is all the chemical reactions that occur in an organism Cellular metabolism
Cells break down excess carbohydrates first, then lipids, finally amino acids if energy needs are not met by carbohydrates and fat Nutrients not used for energy are used to build up structure, are stored, or they are excreted 40% of the energy released in catabolism is captured in ATP, the rest is released as heat
Anabolism
Performance of structural maintenance and repairs Support of growth Production of secretions Building of nutrient reserves
Catabolism
Breakdown of nutrients to provide energy (in the form of ATP) for body processes
Cells provide small organic molecules to mitochondria Mitochondria produce ATP used to perform cellular functions
Metabolism of Carbohydrates
Carbohydrate Metabolism
Primarily glucose
Glucose uptake from blood to cells usually mediated by insulin and transporters
Fates of Glucose
Fed state
Storage as glycogen
Storage as lipids
Fasted state
Muscle
Glycogen
Glucose absorbed
Adipose Cells Glucose absorbed
Glucose absorbed
Glucose Metabolism
Four major metabolic pathways: Immediate source of energy Pentophosphate pathway Glycogen synthesis in liver/muscle Precursor for triacylglycerol synthesis
Energy status of body regulates which pathway gets energy Same in ruminants and non-ruminants
Oxidized to ATP
Only excess glucose Stored as triglycerides in adipose
Glucose Utilization
Adipose
Energy Stores
Glycogen
Glucose
Pentose Phosphate Pathway Glycolysis
Ribose-5-phosphate
Pyruvate
Glucose Utilization
Adipose
Energy Stores
Glycogen
Glucose
Pentose Phosphate Pathway Glycolysis
Ribose-5-phosphate
Pyruvate
Glycolysis
Glucose Pyruvate
Acetyl-CoA (TCA cycle)
Glycolysis
Glucose + 2 ADP + 2 Pi 2 Lactate + 2 ATP + 2 H2O
rechargeable battery
ADP + P + Energy
ATP
Substrate-level phosphorylation
Substrate transfers a phosphate group directly Requires enzymes Phosphocreatine + ADP Creatine + ATP
Oxidative phosphorylation
Method by which most ATP formed Substrate transfers electrons to transporter which enters the electron transport chain
Traps glucose in cells (irreversible in muscle cells) Hexokinase is used by most plants, animals, and microbes to phosphorylate glucose Glucokinase in hepatic tissue Cows do not have hexokinase
Glycolysis - Summary
Glucose
2 ATP 4 ADP 2 ADP 4 ATP 2 NAD 2 NADH + H
2 Pyruvate
Pyruvate Metabolism
Conversion to lactate (anaerobic) Conversion to alanine (amino acid) Entry into the TCA cycle via pyruvate dehydrogenase pathway
Pyruvate Metabolism
Conversion to lactate (anaerobic) Conversion to alanine (amino acid) Entry into the TCA cycle via pyruvate dehydrogenase pathway
Problem:
During glycolysis, NADH is formed from NAD+ Without O2, NADH cannot be oxidized to NAD+ No more NAD+
Solution:
NAD+ COO HC OH
Lactate dehydrogenase
CH3 Lactate
(reduced)
ATP yield
Two ATPs (net) are produced in the anaerobic breakdown of one glucose
Pyruvate
NADH NAD+
Lactate
Cori Cycle
Lactate is converted to pyruvate in the liver
Pyruvate Metabolism
Conversion to lactate (anaerobic) Conversion to alanine (amino acid) Entry into the TCA cycle via pyruvate dehydrogenase pathway
Pyruvate metabolism
Keto acid
Amino acid
Pyruvate Metabolism
Conversion to lactate (anaerobic) Conversion to alanine (amino acid) Entry into the TCA cycle via pyruvate dehydrogenase pathway
Aerobic Conditions
Electron Transport Chain TCA Cycle
PDH - Summary
Pyruvate
2 NAD
2 NADH + H
CO2
Acetyl CoA
TCA Cycle
In aerobic conditions TCA cycle links pyruvate to oxidative phosphorylation Occurs in mitochondria Generates 90% of energy released from feed Metabolizes carbohydrate, protein, and fat Strategy is to oxidize acetyl-CoA to CO2 and capture energy as NADH(FADH2) and ATP
1 ATP 1 ADP
3 NAD 3 NADH + H
1 FAD 1 FADH2
Oxidative Phosphorylation
Requires coenzymes (NAD and FAD) as H+ carriers and consumes oxygen Key reactions take place in the electron transport system (ETS)
H+ flow through ETC and eventually attach to O2 forming water 3 ATP 2 ATP
NADH + H+ FADH2
Produced by bacteria in the fermentation of pyruvate Provides important energy sources for animals, particularly ruminants or those that rely on cellulose as a key carbohydrate in their diet Three major VFAs
Acetate
Propionate
Energy source and for fatty acid synthesis Used to make glucose through gluconeogenesis Energy source and for fatty acid synthesis Some use and metabolism (alterations) by rumen wall and liver before being available to other tissues
Butyrate
Acetic acid
Yields 10 ATP Yields 18 ATP Yields 27 ATP Little butyrate enters blood
Propionic acid
Butyric acid
Propionate Butyrate
Energy Precursor of glucose
Energy Carbon source for fatty acids - mammary
Oxidative Phosphorylation
A BRIEF INTERLUDE
Working animals
Horses, dogs, dairy cattle, hummingbirds! Increased gut size relative to body size Increased feed intake Increased digestive enzyme production Increased liver size and blood flow to liver
Increased ability to deliver oxygen to tissues and get rid of carbon dioxide
Lung size and efficiency increases Heart size increases and cardiac output increases Increase capillary density
Increased numbers of mitochondria in cells Locate mitochondria closer to cell walls (oxygen is lipid-soluble)
Hummingbirds
Lung oxygen diffusing ability 8.5 times greater than mammals of similar body size Heart is 2 times larger than predicted for body size Cardiac output is 5 times the body mass per minute Capillary density up to 6 times greater than expected
Substrate-level phosphorylation
Anaerobic glycolysis
Fatty Acid
Glucose Glucose
Acetate
Anaerobic glycolysis
Pyruvate Pyruvate
Substrate-level phosphorylation
Phosphocreatine + ADP
Creatine + ATP
Glucose Utilization
Adipose
Energy Stores
Glycogen
Glucose
Pentose Phosphate Pathway Glycolysis
Ribose-5-phosphate
Pyruvate
Produces NADPH
Similar to NADH Required for fatty acid synthesis Ribose Used for synthesis of nucleic acids
D-Ribulose5-phosphate
RNA or DNA
D-Ribose5-phosphate
Glucose Utilization
Adipose
Energy Stores
Glycogen
Glucose
Pentose Phosphate Pathway Glycolysis
Ribose-5-phosphate
Pyruvate
Energy Storage
Energy from excess carbohydrates (glucose) stored as lipids in adipose tissue Acetyl-CoA (from TCA cycle) shunted to fatty acid synthesis in times of energy excess
High ATP, acetyl-CoA goes to fatty acid synthesis Low ATP, acetyl CoA enters TCA cycle to generate MORE ATP
Glucose Utilization
Adipose
Energy Stores
Glycogen
Glycogenesis
Glucose
Pentose Phosphate Pathway Glycolysis
Ribose-5-phosphate
Pyruvate
Glycogenesis
Liver
710% of wet weight Use glycogen to export glucose to the bloodstream when blood sugar is low Glycogen stores are depleted after approximately 24 hrs of fasting (in humans) De novo synthesis of glucose for glycogen
Glycogenesis
Skeletal muscle
1% of wet weight
Use glycogen (i.e., glucose) for energy only (no export of glucose to blood) Use already-made glucose for synthesis of glycogen
Fates of Glucose
Fed state
Storage as glycogen
Storage as lipids
Fasted state
Gluconeogenesis
Necessary process
Not simply a reversal of glycolysis Insulin and glucagon are primary regulators
Gluconeogenesis
Little glucose absorbed from small intestine VFA can not fuel CNS and RBC
Feline species
Lipolysis
Mobilization of fat stores stimulated by glucagon and epinephrine Triglyceride = glycerol + 3 free fatty acids Glycerol can be used as a glucose precursor
Proteolysis
The breakdown of muscle protein with release of amino acids Alanine can be used as a glucose precursor
Muscle
Proteins Broken Down
Glycogen
Glucose released
Adipose Cells
Glycerol, fatty acids released
In a fasted state, substrates for glucose synthesis (gluconeogenesis) are released from storage
Gluconeogenesis
Glycerol Amino acids Lactate Supply carbon skeleton Pyruvate Propionate There is no glucose synthesis from fatty acids
Occurs primarily in liver, but can also occur in kidneys and small intestine
Carbohydrate Comparison
Carbohydrate Comparison
Importance of gluconeogenesis
MOST monogastrics = less important Ruminant = very important MOST monogastrics = abnormal situation Ruminant = normal situation
Ketone production