
| Protein-Energy Interactions (International Dietary Energy Consultative Group - IDECG, 1991, 437 pages) |
| (introductory text...) |
| Introduction |
| Some basic aspects of protein-energy interrelationships |
![]() | (introductory text...) |
![]() | 1. Introduction |
![]() | 2. Energy dependency of protein and amino acid metabolism |
![]() | 2.1. Qualitative aspects |
![]() | 2.2. Quantitative aspects |
![]() | 2.3. Correlations between energy and protein metabolism |
![]() | 3. Summary and conclusions |
![]() | References |
| Amino acid oxidation and food intake |
![]() | (introductory text...) |
![]() | 1. Introduction |
![]() | 2. Nitrogen balance and amino acid oxidation |
![]() | 3. Amino acid oxidation during periods of positive or negative energy balance |
![]() | 4. Interactions between energy and protein metabolism |
![]() | 5. Amino acid degradation and gluconeogenesis |
![]() | 6. Summary |
![]() | References |
| The metabolic basis of amino acid requirements |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction: The nature of the problem |
![]() | 2. Nutrient requirement models |
![]() | 3. The Millward & Rivers requirement model: Qualitative aspects |
![]() | 4. The variable extrinsic component of the maintenance requirement |
![]() | (introductory text...) |
![]() | 4.1. Indispensable amino acids as toxic metabolites |
![]() | 4.2. Diurnal cycling |
![]() | 5. The anabolic drive |
![]() | 6. Hormonal components of the anabolic drive |
![]() | 7. Protein requirements: Formal statement |
![]() | 8. The issue of protein quality |
![]() | (introductory text...) |
![]() | 8.1. Accretion: Both net and transient |
![]() | 8.2. Minimum obligatory needs: Theoretical predictions |
![]() | 9. Stable isotope studies |
![]() | 10. Practical experience of biological values of dietary protein |
![]() | 11. Urea salvage |
![]() | 12. Indispensable amino acid requirements for the anabolic drive |
![]() | 13. Conclusions |
![]() | References |
| Commentary on paper by D.J. Millward |
![]() | (introductory text...) |
![]() | References |
| Critique of protein-energy interactions in vivo: Urea kinetics |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. General considerations |
![]() | 2.1. Functional metabolic demand |
![]() | 2.2. Carbon flux and nitrogen flux |
![]() | 2.3. Functional metabolic mass of protein |
![]() | 2.4. Specific limiting nutrients |
![]() | 2.5. Limitations imposed by protein quality |
![]() | 2.6. Amino acids: Essential, non-essential and conditionally essential |
![]() | 3. The Millward model |
![]() | (introductory text...) |
![]() | 3.1. Present perception of nitrogen disposal |
![]() | 3.2. Urea production |
![]() | 3.3. Urea excretion |
![]() | 3.4. Salvaged urea nitrogen |
![]() | 3.5. The 'effective dietary intake' of nitrogen |
![]() | 3.6. Limits of adaptation to low-protein diets |
![]() | 3.7. Implications of salvaged urea nitrogen |
![]() | 4. Conclusions |
![]() | References |
| The effects of different levels of energy intake on protein metabolism and of different levels of protein intake on energy metabolism: A statistical evaluation from the published literature |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. The effects of different levels of energy intake on protein metabolism |
![]() | 2.1. Adults |
![]() | 2.2. Children |
![]() | 3. The effects of different levels of protein intake on energy metabolism |
![]() | 4. Protein/energy ratios |
![]() | 5. Summary and conclusions |
![]() | References |
| Effect of different levels of carbohydrate, fat and protein intake on protein metabolism and thermogenesis |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Influence of nutrient intake on nutrient oxidation |
![]() | 3. Effect of energy intake on nitrogen retention |
![]() | 3.1. Fasting and very low caloric intake |
![]() | 3.2. Moderately hypocaloric diets |
![]() | 3.3. Maintenance diets |
![]() | 3.4. Energy intake in excess of maintenance |
![]() | 4. Effect of protein intake on nitrogen retention |
![]() | 4.1. Normal and obese subjects |
![]() | 4.2. Severely depleted subjects |
![]() | 4.3. Moderately depleted subjects |
![]() | 5. The role of glucose and lipid in nitrogen sparing |
![]() | 5.1. Healthy young subjects |
![]() | 5.2. Patients receiving total parenteral nutrition |
![]() | 6. Mechanisms of the sparing effect of dietary carbohydrate and fat |
![]() | 6.1. Effect of dietary glucose on leucine oxidation |
![]() | 7. Effect of amino acid plasma levels on protein synthesis |
![]() | 8. Practical considerations: Role of the thermic effect of nutrients |
![]() | 9. Conclusions |
![]() | References |
| Respiratory quotients and substrate oxidation rates in the fasted and fed state in chronic energy deficiency |
![]() | (introductory text...) |
![]() | 1. Respiratory quotients in semi-starvation |
![]() | 2. Respiratory quotients in experimental semi-starvation |
![]() | 3. Respiratory quotients and substrate oxidation rates in chronically energy deficient subjects |
![]() | 4. Substrate oxidation rates during dietary thermogenesis in chronic energy deficiency |
![]() | 5. Effects of refeeding or supplementation on respiratory quotients and substrate oxidation rates of CED subjects |
![]() | Acknowledgements |
![]() | References |
| Effects of protein-energy interactions on growth |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Mechanisms for effects of protein and energy on growth |
![]() | (introductory text...) |
![]() | 2.1. Insulin and insulin-like growth factors |
![]() | 2.2. Growth hormone |
![]() | 2.3. Epidermal growth factor |
![]() | 2.4. Corticosteroids |
![]() | 3. The determinants of catch-up growth |
![]() | 4. Effect of the protein/energy ratio on growth of premature infants |
![]() | 5. Effect of protein and energy on growth of children with primary malnutrition |
![]() | 6. Effect of the P/E ratio on growth of children with malnutrition secondary to chronic renal insufficiency |
![]() | 7. Conclusions and speculations |
![]() | Acknowledgements |
![]() | References |
| Protein-energy interrelationships during rapid growth |
![]() | (introductory text...) |
![]() | 1. Efficiency of protein deposition |
![]() | 2. Protein turnover during rapid growth |
![]() | 3. Energy cost of protein synthesis |
![]() | References |
| Quantitative relationships between protein and energy metabolism: Influence of body composition |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Theoretical basis |
![]() | 3. Constancy of tissue mobilisation |
![]() | 4. Tissue mobilisation in the obese |
![]() | 5. Allometric analysis |
![]() | 6. Conclusions |
![]() | References |
| Protein-energy relationships in pregnancy and lactation |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Influence of gestational weight gain on pregnancy outcomes |
![]() | 2. Protein needs during pregnancy |
![]() | 2.1. Influence of gestational weight gain on protein needs |
![]() | 2.2. Efficiency of protein utilization during pregnancy |
![]() | 2.3. Influence of dietary energy on protein utilization |
![]() | 2.4. Summary of protein requirements during pregnancy |
![]() | 3. Energy requirements during pregnancy |
![]() | 3.1. Influence of gestational weight gains on energy needs |
![]() | 3.2. Physical activity and pregnancy |
![]() | 3.3. Summary of energy requirements during pregnancy |
![]() | 4. Protein needs during lactation |
![]() | 4.1. Estimation of protein needs |
![]() | 4.2. Influence of protein intake on milk composition |
![]() | 4.3. Studies of whole-body protein turnover |
![]() | 4.4. Effects of protein intake on milk production |
![]() | 4.5. Summary of protein needs during lactation |
![]() | 5. Energy needs during lactation |
![]() | (introductory text...) |
![]() | 5.1. Summary of energy needs for lactation |
![]() | 6. Conclusions |
![]() | References |
| Effects of physical activity on protein-energy interactions: Metabolic and nutritional considerations |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Energy metabolism in exercise |
![]() | 2. Are protein requirements affected by exercise when energy requirements are met? |
![]() | 3. Muscle protein breakdown and amino acid oxidation |
![]() | 4. Substrate metabolism in exercise |
![]() | 5. Effect of exercise on protein synthesis |
![]() | 6. Summary and dietary recommendations |
![]() | Acknowledgements |
![]() | References |
| Influence of physical activity on energy and protein metabolism |
![]() | (introductory text...) |
![]() | 1. Exercise and efficiency of dietary energy and protein utilization |
![]() | 2. Effects of reduced physical activity on energy and protein metabolism |
![]() | 3. Energy substrates and changes in exercise pattern |
![]() | References |
| Exercise, aging and protein metabolism |
![]() | (introductory text...) |
![]() | 1. Body composition changes with age and their consequences |
![]() | 2. Fuels used to meet various components of energy requirements |
![]() | 3. Age and dietary protein needs |
![]() | 4. Exercise-induced muscle damage and acute phase response |
![]() | 5. Exercise and protein metabolism |
![]() | 6. Summary |
![]() | References |
| Effect of starvation and very low calorie diets on protein-energy interrelationships in lean and obese subjects |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Early total starvation |
![]() | 2.1. Energy metabolism |
![]() | 2.2. Protein metabolism |
![]() | 2.3. Protein/energy ratios |
![]() | 3. Prolonged total starvation |
![]() | 3.1. Body fat |
![]() | 3.2. Implications of initial body weight and fat stores on protein-energy interrelationships |
![]() | 3.3. Evidence for the first postulate of the model: Survival time in relation to body composition |
![]() | 3.4. Evidence for second postulate of the model: During prolonged starvation the contribution of protein oxidation to energy expenditure is less in obese than lean subjects |
![]() | 3.5. Starvation in man and other species |
![]() | 4.1. Duration of dieting |
![]() | 4.2. Protein and energy intake |
![]() | 4.3. Body composition |
![]() | 4.4. Exercise |
![]() | 5. Some other issues, conclusions and recommendations |
![]() | References |
| Impact of gastrointestinal function on protein-energy interactions and nutritional needs |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Gastrointestinal function in protein-energy malnutrition |
![]() | 2. Diarrheal diseases |
![]() | 3. Nutritional recommendations |
![]() | References |
| Role of the gastrointestinal tract in energy and protein metabolism |
![]() | (introductory text...) |
![]() | 1. Introduction |
![]() | 2. Cell and protein turnover |
![]() | 3. Nutrient absorption |
![]() | 4. Protein synthesis |
![]() | 5. Restriction of energy and protein intake |
![]() | 6. Fat absorption and exocytosis |
![]() | 7. Chronic environmental enteropathy |
![]() | References |
| Effect of protein-energy interaction with reference to immune function and response to disease |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Outlining the issues |
![]() | 3. Host metabolism and host defense |
![]() | 4. The metabolic profile of the infected host |
![]() | 5. The role of cytokines |
![]() | 6. Cytokine regulation: Natural antagonists and biological modulators |
![]() | 7. The future |
![]() | References |
| Nutrition of immune cells: The implications for whole body metabolism |
![]() | (introductory text...) |
![]() | 1. Introduction |
![]() | (introductory text...) |
![]() | 1.1. Lymphocytes |
![]() | 1.2. Macrophages |
![]() | 2. An introduction to metabolic-control logic and its application to the structure of a biochemical pathway |
![]() | 2.1. Near-equilibrium and non-equilibrium reactions |
![]() | 2.2. The flux-generating reaction |
![]() | 3. Use of maximum activities of enzymes as quantitative indices of maximum flux through metabolic pathways |
![]() | 4. Enzyme activities as indication of the capacity of major energy providing pathways in immune cells |
![]() | 5. Glutamine and the immune cells |
![]() | 6. Glutamine - A link between muscle and the immune system |
![]() | (introductory text...) |
![]() | 6.1. Glutamine synthesis in skeletal muscle |
![]() | 6.2. The transport of glutamine across the muscle membrane: Glutamine uptake and release |
![]() | 7. Large decreases in the concentration of glutamine in plasma |
![]() | 8. The clinical significance of the role of glutamine in immune cells |
![]() | 9. The effects of glutamine provision for the patient |
![]() | 10. Branched-point sensitivity, substrate cycles and thermogenesis |
![]() | References |
| Metabolic and nutritional interrelationships between energy and protein in sepsis, trauma and depletion |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. History 1900-1960 |
![]() | 3. Indirect calorimetry and N balance in surgical patients |
![]() | 4. Nitrogen balance: The role of energy balance and N intake |
![]() | (introductory text...) |
![]() | 4.1. Normal subjects |
![]() | 4.2. Depleted patients |
![]() | 4.3. Injured patients |
![]() | 5. Summary |
![]() | References |
| Protein and energy requirements following burn injury |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Resting energy expenditure |
![]() | 2.1. Mechanism of hypermetabolism |
![]() | 2.2. Prediction of resting energy expenditure in burned patients |
![]() | 3. Relationship of total energy expenditure (TEE) to REE |
![]() | 4. Sources of energy |
![]() | 5. Protein requirements |
![]() | Acknowledgements |
![]() | References |
| Protein-energy relationships: Experience with parenteral nutrition |
![]() | (introductory text...) |
![]() | Abstract |
![]() | 1. Introduction |
![]() | 2. Metabolic response to starvation |
![]() | 3. Metabolic response to stress |
![]() | References |
| Modifications of parenteral nutrition support for critical surgical illness |
![]() | (introductory text...) |
![]() | References |
| Dietary protein/energy ratios for various ages and physiological states |
![]() | (introductory text...) |
![]() | 1. Definition, interpretation and uses |
![]() | 2. Calculation of recommended P/E ratios |
![]() | 3. Recommended P/E ratios |
![]() | 4. Food sources of energy and proteins |
![]() | References |
| Effects of disease on desirable protein/energy ratios |
![]() | (introductory text...) |
![]() | 1. Effects of infections on nutritional status |
![]() | (introductory text...) |
![]() | 1.1. Anorexia |
![]() | 1.2. Cultural and therapeutic practices |
![]() | 1.3. Malabsorption |
![]() | 1.4. Catabolic losses |
![]() | 1.5. Anabolic losses |
![]() | 1.6. Fever |
![]() | 1.7. Additional intestinal losses |
![]() | 2. Environmental ('tropical') enteritis |
![]() | 3. Other chronic infections |
![]() | 4. Energy vs protein requirements |
![]() | 5. Possible role of specific amino acids |
![]() | 5.1. Branched-chain amino acids |
![]() | 5.2. Glutamine |
![]() | 5.3. Cancer |
![]() | 6. Summary |
![]() | 7. Recommendations |
![]() | References |
| Amino acid scoring in health and disease |
![]() | (introductory text...) |
![]() | 1. Introduction |
![]() | 2. Amino acid scoring in health |
![]() | 2.1. Protein quality evaluation: The protein digestibility-corrected amino acid score method |
![]() | 2.2. Protein digestibility |
![]() | 2.3. Amino acid scoring patterns |
![]() | 3. Amino acid scoring in special cases and disease |
![]() | (introductory text...) |
![]() | 3.1. Amino acid essentiality |
![]() | 3.2. Glycine |
![]() | 3.3. Glutamine |
![]() | 3.4. Arginine |
![]() | 3.5. Cysteine/taurine |
![]() | 3.6. Branched-chain amino acids (BCAAs) |
![]() | References |
| Research needs |
![]() | (introductory text...) |
![]() | 1. Energy expenditure and metabolism |
![]() | 1.1. Energy expenditure of free-living populations |
![]() | 1.2. More measurements of activity patterns in free-living populations |
![]() | 1.3. Effects of carbohydrates in the diet on fat deposition |
![]() | 2. Protein metabolism and requirements |
![]() | 2.1. Amino acid oxidation |
![]() | 2.2. Amino acid requirements |
![]() | 2.3. Protein requirements during pregnancy and lactation |
![]() | 2.4. Control of urea recycling from the gut |
![]() | 2.5. Limits to the de novo synthesis of 'conditionally essential' amino acids |
![]() | 2.6. Special roles of particular amino acids |
![]() | 3. Body composition |
![]() | 3.1. Methods of measurement |
![]() | 3.2. Composition of lean body mass |
![]() | 3.3. Composition of weight gain during pregnancy |
![]() | 4. Weight gain in children |
![]() | 4.1. Variability of weight gain and its effect on protein requirements |
![]() | 4.2. Factors limiting protein deposition |
![]() | 4.3. Effects of frequent versus intermittent feeding on growth |
![]() | 4.4. Quantitative and qualitative requirements for catch-up growth |
![]() | 5. Linear growth |
![]() | 5.1. Potential causes of stunting |
![]() | 5.2. Reversibility of stunting |
![]() | 6. Physical activity |
![]() | 6.1. Effects of physical activity on metabolism and body composition |
![]() | 6.2. Energy intake and physical activity |
![]() | 6.3. Changes in life-style |
![]() | 7. Infection |
![]() | 7.1. Interactions between energy, protein and amino acid intakes and cytokine responses |
![]() | 7.2. Methods of quantifying losses imposed by infection |
![]() | 7.3. Development of field methods for assessing the severity and intensity of infection |
![]() | 7.4. Interaction of protein-energy status, immunizations and immune status |
![]() | 8. Functional consequences |
![]() | 9. Variation |
![]() | List of participants |
The foregoing has dealt with studies performed on adults. Brief consideration will now be given to protein-energy needs of children. First, however, an overview of assessment methods will be presented. During the first year of life, the protein content of the body increases rapidly. The average increase in body protein is about 3.5 g/d during the first 4 months of life and 3.1 g/d during the next 8 months. By 4 years of age, body protein concentration reaches the adult value of 18 to 19% of body weight. As the growth rate drops rapidly after the first year of life, the maintenance requirement represents a gradually increasing and major proportion of the total protein requirement (FAO/WHO/UNU, 1985).
For the first months of life, requirements are based on intake data because of the difficulties in estimating accurate allowances for growth and maturation. Infants breast-fed by healthy, well-nourished mothers can grow at a satisfactory rate for 4 to 6 months. Measurements of human milk consumption (FAO/WHO/UNU, 1985) demonstrated that protein intakes ranged from 2.43 g/kg/d in the first month to 1.51 g/kg/d in the fourth month. A rounded value of 2.00 g/kg/d was selected as an allowance for this age group, but it was recommended that protein requirements should be considered in relation to energy needs. The protein needs of an infant up to 4 months of age will be met if the energy needs are met, provided the food contains protein of quality and quantity equivalent to that of breast milk. This implies that at least 6°10 of the food-energy in the form of high-quality protein is desirable.
A modified factorial procedure was used to estimate needs for older children, and the FAO/WHO/UNU (1985) group also recognized the need for continuous provision of protein in anticipation of extra, unpredictable demands from the daily variations in growth rate, since protein provided on a day of no growth is probably not held in reserve for later growth. There were no data on which to base reliable estimates for the extra protein needed for this purpose, but a value 50°70 higher than the estimated daily nitrogen increment was considered realistic.
Tabulated values for daily protein and energy allowances for the various age and sex groupings are shown in Table 5 and are initially expressed as high-quality, highly digestible proteins such as meat, milk, fish and eggs. These would need to be corrected for amino acid composition and digestibility to be converted to protein as consumed in ordinary dietaries. Final recommended allowances for dietary protein, as provided by a diet typical of an industrialized country, would thus range from somewhat over 2 g/kg/d in the first year of life to approximately 1 g/kg/d for adolescents. The protein/energy ratios for the various age and sex recommendations will be discussed in a subsequent section.
Table 5. Recommended Allowances for Protein and Energy (NAS-NRC 1989) and calculated values for PCal%
|
Age and Sex years |
Weight kg |
Height cm |
Protein gm |
Energy kcal |
APCal%1 |
B |
|
Infants | ||||||
|
Breast Milk2 |
11 |
720 |
6.1 |
6.1 | ||
|
0 - 0.5 |
6 |
60 |
13 |
650 |
8.0 |
5.2 |
|
0.5 - 1.0 |
9 |
71 |
14 |
850 |
6.6 |
5.1 |
|
Children | ||||||
|
1 - 3 |
13 |
90 |
16 |
1300 |
4.9 |
4.0 |
|
4 - 6 |
20 |
112 |
24 |
1800 |
5.3 |
3.6 |
|
7 - 10 |
28 |
132 |
28 |
2000 |
5.6 |
4.5 |
|
Males | ||||||
|
11 - 14 |
45 |
157 |
45 |
2500 |
7.2 |
4.3 |
|
15 - 18 |
66 |
176 |
59 |
3000 |
7.9 |
5.3 |
|
19 - 24 |
72 |
177 |
58 |
2900 |
8.0 |
6.0 |
|
25 - 50 |
79 |
176 |
63 |
2900 |
8.7 |
6.5 |
|
51+ |
77 |
173 |
63 |
2300 |
11.0 |
8.0 |
|
Females | ||||||
|
11 - 14 |
46 |
157 |
46 |
2200 |
8.4 |
5.0 |
|
15 - 18 |
55 |
163 |
44 |
2200 |
8.0 |
6.0 |
|
19 - 24 |
58 |
164 |
46 |
2200 |
8.4 |
6.3 |
|
25 - 50 |
63 |
163 |
50 |
2200 |
9.1 |
6.9 |
|
51+ 65 |
160 |
50 |
1900 |
10.5 |
8.2 | |
|
Pregnancy3 |
60 |
2500 |
9.6 |
7.8 | ||
|
Lactation 1st |
6m |
65 |
2700 |
9.6 |
7.9 | |
2nd |
6m |
62 |
2700 |
9.2 |
7.5 | |
1
PCal% = Protein g per 100g x 400/kcal per
100g. Since energy allowances are averages and protein allowances are safe
levels (means + 2SD) the direct ratios of allowances (A) are misleading. The
ratios (B) are calculated using average protein needs and are, thus,
lower.
2 Mature milk composition.
3 Second and third
trimesters.
Protein-energy interactions in diets for children have been reviewed by ZLOTKIN (1986). Although recommended intakes of nutrients are listed according to population norms for age and size on an individual basis, protein and energy intakes are generally adjusted to meet specific outcome goals. Recommended intakes for protein and energy for the full-term infant are based on the composition and volume intake of mature human milk, defined as milk produced sometime after 30 days post partum. For infants born prior to term, oral nutrient intake recommendations have been established either from:
(a) analysis of body composition
(b) direct clinical studies of various feeding regimens, or
(c) analysis of the nutrient content of mature human milk.
The establishment of an appropriate outcome for the newborn infant, who is parenterally fed, is even more complicated, since there are potential differences in the body's handling of intravenously versus orally ingested nutrients. As reviewed by ZLOTKIN (1986), the literature describes intravenous formulations delivering 1.5 to 4 g/kg/d of amino acids and 50 to 140 kcal/kg/d for energy, depending on clinical circumstances and whether the infusion is via the central or peripheral vein.
For children, the energy intake necessary to minimize nitrogen loss associated with an amino acid free diet has not been established but is likely to be in the range of 70 kcal/kg/d (ZLOTKIN 1986). For infants who are receiving an energy intake that is adequate to maintain good health, daily nitrogen retention can be shown to increase with increasing nitrogen intake. Even at the highest nitrogen intake level (640 mg/kg/d), there was no decrease in the linear increase in nitrogen retention.
This response is in marked contrast to that seen in the adult where nitrogen retention is assumed, by current theory, to plateau at or slightly above the zero balance line, when excess dietary nitrogen is provided. It should be noted, however, that infants receiving the highest nitrogen intakes also had the highest plasma amino acid levels, with these levels reflecting the composition and quantity of the amino acid solutions infused. Thus, achieving even higher nitrogen retention in infants is possible, though probably undesirable, because of the concomitant metabolic consequences (ZLOTKIN, 1986).
When both nitrogen and energy are taken concurrently both interact to affect nitrogen retention. If energy intake is inadequate to meet maintenance requirements, then protein may be used as an energy source. Similarly, if protein intake is inadequate to meet maintenance needs, then increasing energy intakes will spare protein for protein synthesis. This holds true also for infants and it has been demonstrated (ZLOTKIN, 1986) that, for an energy intake just below maintenance requirements (50-60 non-protein kcal/kg/d), increasing nitrogen intake from 400 to 500 mg/kg/d results in a significant increase in nitrogen retention.
At this same low energy intake, however, a further increase in nitrogen intake to 655 mg/kg/d did not result in any further increase in nitrogen retention. The clinical consequences suggest that, when infants are maintained on a hypocaloric intake, nitrogen should be provided at no more than 480 mg/kg/d, since the administration of excess nitrogen will not contribute to nitrogen retention and may lead to increasing plasma urea and amino acid levels.
Milk formula will meet the protein needs of an infant up to about 4 months, if energy needs are met, since the protein and energy content of infant formulas, at least quantitatively, reflects mature human milk (ZLOTKIN, 1986). Therefore, once again, if the energy needs of the infant are met, so too will be the protein needs. It is generally accepted that the value of 1.5 g of crude protein per 100 kcal (Pcal% = 6) approaches the ideal protein/energy ratio for infant foods (FAO/WHO/UNU, 1985).
For the premature infant fed orally, the situation is much more complicated, since it is not universally accepted that mature human milk is the ideal food for the premature infant. The nutrient content of the milk produced by mothers giving birth to premature infants is different from mature human milk (ATKINSON et al., 1978). The estimated protein needs of the premature infant are 3 g/kg/d and energy needs at 120 kcal/kg/d. Thus, the protein/energy ratio for these infants would be 2.5 g of protein per 100 kcal or Pcal% = 10).
Although the actual requirements for growth and development are independent of the route of feeding, the latter has a significant effect on recommended intakes because bioavailability and absorption considerations are of less consequence and activity levels may differ. ZLOTKIN (1986) has calculated that a energy intake of 80 to 100 kcal/kg/d should be adequate to meet requirements of intravenously fed infants. This value is some 20% lower than the recommended oral intake. It has been estimated also that, for the pre-term infant, as long as adequate energy is provided, a nitrogen intake of about 480 mg/kg/d will safely achieve the goal of duplication of the rates of intrauterine nitrogen accretion. This corresponds to about 13% calories from protein.
The relative effect of glucose and lipids on whole-body protein-metabolism kinetics was assessed in seven infants undergoing parenteral feeding by BRESSON et al. (1991). Protein intake was kept constant, and non-protein energy was either provided as glucose alone or as an isoenergetic glucose-lipid mixture. Protein metabolism and energy-substrate utilization were assessed by a primed, constant L-[13C]leucine infusion, combined with indirect calorimetry. There was a significant difference in the pattern of energy-substrate utilization according to regime. Protein turnover, protein breakdown, and amino acid oxidation rates were higher for the glucose than the glucose-lipid treatment, whereas protein-synthesis rates did not significantly differ. The nature of energy substrates delivered to parenterally fed infants may thus affect protein metabolism.
For requirements of children living in developing countries, the studies that we have considered are those from Thailand, Guatemala and Jamaica. Protein-energy relationships have been reported for children in Thailand by TONTISIRIN et al. (1984b). Nine children, aged 9 to 36 months, weighing 8.1 to 11.1 kg and living in a metabolic unit were given normal local weaning diets at three levels of energy intake, varying from 87 to 118 kcal/kg/d. The diets consisted of rice, fish and banana, and the protein intake was fixed at the safe level of 1.7 g/kg/d. Each level of energy was fed for 7 days and vitamin and mineral supplements were given daily. At the lowest level of energy intake, 87 kcal/kg/d, apparent nitrogen retention and weight gain were quite low being 44 mg N/kg and 3.5 g/d, respectively. At the two higher levels of energy intake (100 and 118 kcal/kg), apparent nitrogen retention was greater than 60 mg/kg per day, and weight gain was 20 g per day or more. The results from this study suggested that at the safe level of protein intake, as recommended by FAO/WHO in 1973, the usual Thai weaning food provided adequate protein for the needs of young children, if energy intake was supplied at 100 kcal/kg/d or more. A longer-term study, lasting 4 months for six normal, healthy young male children, aged 8 to 12 months, that had been rehabilitated for 8 weeks or longer following protein-energy malnutrition and had reached normal weights for height, confirmed the earlier observations that 1.7 g/kg protein was adequate for one-year-old children, provided that the energy intake was 100 kcal/kg/d or more. This would represent Pcal%. = 6.8. Habitual Guatemalan diets were investigated for their capacity to allow for catch-up growth in children with mild to moderate malnutrition (TORUN et al., 1984). Children were fed diets mainly based on corn and beans, but with some animal foods such that protein contents averaged from 2.1 to 1.8 g/kg, depending on the stage of the recovery. The energy density of the diet was increased by adding oil to the beans and sugar to the beverages. It was observed that needs of children 2-4 years old could be satisfied with 85-90 kcal/kg (Pcal% = 8.5-9.5) while catch-up growth required 95-105 kcal/kg (Pcal% = 8).
Nitrogen balance and whole-body protein turnover were measured in children aged about one year by JACKSON et al. (1983). Diets provided 1.7 or 0.7 g milk protein/kg/d and were fed at three levels of metabolizable energy, 80, 90 and 100 kcal/kg/d. The Pcal% values of the diets thus ranged from 2.8 to 8.5%. All the children were in positive nitrogen balance at all levels of energy intake on 1.7 g protein/kg/d. Nitrogen equilibrium was maintained on 0.7 g protein/kg/d when the energy intake exceeded 90 kcal/kg/d, but on 80 kcal/kg/d nitrogen balance was negative. Whole-body protein turnover was measured from the enrichment in urinary ammonia, following a continuous infusion of 15N-glycine.
It was observed that the variation between individuals on the same diet was significantly greater than the variation within individuals at different levels of energy intake. For the group as a whole, protein synthesis on 1.7 g protein/kg/d was 0.74, 0.75 and 0.87 g N/kg/d on 100, 90 and 80 kcal/kg/d, respectively; whereas on 0.7 g protein/kg/d it was 0.37, 0.38 and 0. 40 g N/kg/d. These results show that, over this range of intakes, protein synthesis decreased as dietary protein fell, but tended to remain unchanged or even increase as energy intake fell.
Physical activity is also a factor in energy-protein interrelationships (YOUNG et al., 1983) and may be especially important in children. A study by TORUN et al. (1976) revealed the beneficial effects of continued mild exercise on the utilization of dietary protein in young children aged 2 to 4 years undergoing treatment for protein-energy malnutrition. These children were stimulated to be more physically active through a daily program of games that required a mild increase in energy expenditure. Measurements were made of growth and of body energy and nitrogen balances, and results were compared with those obtained in a similar group of children treated in the traditional manner at INCAP. The more active children were shown to grow better in height and in lean body mass. Thus, the efficiency of utilization of dietary protein-energy for growth was greater in the more active children. It could be concluded that moderate, systemic exercise has a growth-enhancing affect and a favorable impact on the utilization of dietary protein. Similar relationships for other age groups are described elsewhere in this volume.