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Macronutrient Caloric Density

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Carbohydrates and proteins supply about four to five kilocalories per gram, while fat supplies about nine, making fat the most energy-dense macronutrient.

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Animals seeking lipid-rich food gain more stored energy per gram eaten than from an equal mass of carbohydrate.

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Higher caloric density is not automatically harmful — it reflects efficient long-term energy storage, useful during famine or migration.