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Based on the work of Atwater, it became common practice to calculate energy content of foods using 4 kcal/g for carbohydrates and proteins and 9 kcal/g for lipids. [2] The system was later improved by Annabel Merrill and Bernice Watt of the USDA, who derived a system whereby specific calorie conversion factors for different foods were proposed. [3]
The unit is most commonly used to express food energy, namely the specific energy (energy per mass) of metabolizing different types of food. For example, fat (triglyceride lipids) contains 9 kilocalories per gram (kcal/g), while carbohydrates (sugar and starch) and protein contain approximately 4 kcal/g. [29] Alcohol in food contains 7 kcal/g. [30]
Fatty acids, stored as triglycerides in an organism, are a concentrated source of energy because they contain little oxygen and are anhydrous. The energy yield from a gram of fatty acids is approximately 9 kcal (37 kJ), much higher than the 4 kcal (17 kJ) for carbohydrates.
Fat has a food energy content of 38 kilojoules per gram (9 kilocalories per gram) proteins and carbohydrates 17 kJ/g (4 kcal/g). [2] Water makes up a large proportion of the total mass ingested as part of a normal diet but it does not provide any nutritional value.
It can take up to 20 hours of little physical output (e.g., walking) to "burn off" 17,000 kJ (4,000 kcal) [17] more than a body would otherwise consume. For reference, each kilogram of body fat is roughly equivalent to 32,300 kilojoules of food energy (i.e., 3,500 kilocalories per pound or 7,700 kilocalories per kilogram). [18]
Carbohydrates and proteins provide 17 kJ approximately (4 kcal) of energy per gram, while fats provide 37 kJ (9 kcal) per gram. [8] However, the net energy derived from the macronutrients depends on such factors as absorption and digestive effort, which vary substantially from instance to instance.
In humans and many animals, fats serve both as energy sources and as stores for energy in excess of what the body needs immediately. Each gram of fat when burned or metabolized releases about nine food calories (37 kJ = 8.8 kcal). [5] Fats are also sources of essential fatty acids, an important dietary requirement.
Plant thylakoid membranes have the largest lipid component of a non-bilayer forming monogalactosyl diglyceride (MGDG), and little phospholipids; despite this unique lipid composition, chloroplast thylakoid membranes have been shown to contain a dynamic lipid-bilayer matrix as revealed by magnetic resonance and electron microscope studies. [59]