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54         Small Animal Clinical Nutrition



                                                                      and flatulence). However, because methane production is
        VetBooks.ir                                                   considered to be negligible in dogs and cats (McKay and
                                                                      Eastwood, 1984), ME can be defined in terms of DE and uri-
                                                                      nary energy losses.
                                                                        Animals continuously produce heat as a result of basal
                                                                      metabolism and physical work. Heat production increases after
                                                                      a meal. This increase in heat due to food ingestion is called the
                                                                      heat increment of food (HI). HI consists of the heat of intes-
                                                                      tinal microbial fermentation and heat produced in intermediary
                                                                      metabolism as a result of using nutrients. A study in people
                                                                      showed that 5 to 10% of the energy consumed was lost as heat.
                                                                      The HI during the postprandial period was 60% greater for the
                                                                      protein-consuming group when compared to the isocaloric car-
                                                                      bohydrate-consuming group. This increase in HI was attrib-
                                                                      uted to the increased protein turnover observed by the protein-
                                                                      consuming group. The metabolic cost for protein turnover was
                                                                      approximately 36 and 68% for carbohydrate and protein feed-
                                                                      ing, respectively (Robinson et al, 2000). The energy of HI is
                                                                      normally wasted except when the environmental temperature is
                                                                      below an animal’s critical temperature (i.e., shivering). In this
                                                                      situation, the HI is used to keep the body warm.
                                                                        Subtracting HI from the ME gives the net energy (NE) of
                                                                      food. NE can also be partitioned into the amount used for
                                                                      maintenance (NE ) and the amount used for production
                                                                                     m
                                                                      (NE : growth, pregnancy, lactation, exercise). NE values of
                                                                          p
                                                                      foods and ingredients are typically used when discussing live-
                                                                      stock nutrition (beef cattle, dairy cattle, swine), whereas DE
                                                                      and ME are more typically used in canine and feline nutrition.
                                                                      Although not commonly measured and used, the NE princi-
                  Figure 5-5. Schematic of how animals obtain and use energy.
                                                                      ples of partitioning energy for maintenance and production
                  Plants use solar energy to produce energy-containing nutrients
                                                                      separately hold true for dogs and cats.
                  (i.e., proteins, fats and carbohydrates) via photosynthesis. Animals
                  eat the energy-containing plant nutrients and other animals. Once
                  eaten, energy-containing nutrients are digested, absorbed and  Energy Use
                  metabolized by body cells to release energy that fuels the process-  The initial biochemical reactions by which energy is derived
                  es that sustain life.                               from carbohydrates, fats and amino acids are different.
                                                                      However, all three nutrients eventually go through a final
                                                                      common pathway for energy generation (i.e., TCA cycle).
                  to as being  “bulk limited.” Low-energy, bulk-limited foods  Glucose derived from dietary carbohydrates is first oxidized
                  designed for weight loss can be formulated to provide adequate  through the glycolysis pathway to yield pyruvate and then
                  intake of non-energy nutrients (Chapter 27).        acetyl-CoA. Acetyl-CoA is oxidized in the TCA cycle pro-
                                                                      ducing carbon dioxide, and electrons, which are captured by
                  Energy Metabolism                                   important heme-containing compounds called cytochromes
                  Digestion, Absorption and Excretion                 (Figure 5-7). Electrons produced in the TCA cycle are shut-
                  Digestion and absorption of the energy-supplying nutrients  tled by nicotinamide-adenine dinucleotide (NAD) and flavin
                  (protein, carbohydrate and fat) are discussed in other sections of  adenine dinucleotide (FAD) to the electron transport chain
                  this chapter. The total amount of potential energy in food is  where the cytochromes participate in electron transfer through
                  termed gross energy (GE). Burning the food and measuring  valence changes in their heme iron (Figure 5-4). NAD and
                  the heat produced in a bomb calorimeter determine GE in  FAD are synthesized from the vitamins niacin and riboflavin,
                  food. Animals are unable to use 100% of the GE in foods  respectively. The electrons are passed between successive oxi-
                  because some of the food energy is lost in the form of solid, liq-  dation/reduction reactions to the end of the chain where oxy-
                  uid and gaseous excretions as well as radiant heat.  gen accepts the final electrons and is converted to water. ATP
                    Nutritionists have partitioned dietary energy based on the  is formed as the electrons are passed down the chain (oxidative
                  losses that occur (Figure 5-6). Digestible energy (DE) refers  phosphorylation). A net of 36 ATP is generated for each mol-
                  to the GE content of food minus energy lost in feces (FE).  ecule of glucose that is oxidized to carbon dioxide and water.
                  Typically, ME is defined as DE minus energy lost in urine  Fatty acids and glycerol from dietary fats are initially oxidized
                  and as intestinal gaseous products of digestion (i.e., eructation  to acetyl-CoA by the  β-oxidation pathway (Figure 5-8).
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