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Normal root elongation requires arginine produced by argininosuccinate lyase in rice. The sum of the molar content of arginine 0. Then, it follows that arginine or proline intake is likely inadequate during periods of the NICU stay as this level of protein intake for preterm infants cannot be met with human milk until full feeds are established and fully fortified. These observations suggest that the low arginine documented in preterm infants 7 relates to inadequate intake of proline and arginine.
Whether this is clinically important with either short- or long-term effects is unknown. The enteral turnover measurements of arginine and proline in our study are the first to be published for healthy growing preterm infants; therefore, it is necessary to discuss our results with reference to adult human studies and piglet studies as a model for the preterm infant.
However in the study by Castillo et al. The higher rate in premature neonates compared with adults can be explained by the higher rate of endogenous protein turnover in newborns, which may be as much as 4—6 times greater than that of adults 25 , 26 and the increased requirement for arginine to meet the metabolic demands of growth. There are no comparable data in either human infants or adults.
The only available data in humans is i. This similarity suggests that the pattern of metabolism of the two amino acids, i. The similarity in this ratio for arginine and proline suggests that the splanchnic metabolism of proline in humans may be much higher than previously believed and deserves further investigation to elucidate the proline requirements for the splanchnic organs and whether it is required for protein synthesis alone, amino acid precursor, or other metabolic roles.
Of note, in our estimates of enteral turnover, for both arginine and proline, is the considerable variation among subjects, which is greater for arginine than proline reflecting the multiple metabolic roles for arginine, notably creatine and NO synthesis. The source of this variation is likely multifactorial. It is well recognized that protein turnover rates in preterm infants are much higher than adults and show considerable variation among individuals.
For example, Denne et al. Protein synthesis is the largest component of arginine flux, and therefore, this likely explains the variation observed in our study. In addition, and by necessity, it was not possible to standardize feeds in this study beyond feed volume, and therefore, variation in nutrient intake will have contributed to the overall variation. The milk of choice is expressed breast milk to which fortifier is added when preterm infants reach full feeds.
The amount of protein and arginine in breast milk also displays considerable variation. Montagne et al.
There are no human data with which to compare the fractional synthesis rates of ornithine and citrulline; however, these are similar to the synthesis rates reported for the piglet We did not measure any glutamate tracer in the urine glutamate pool. Recognizing that this was a possibility because of high dilution from dietary and endogenous glutamate, we dosed more on a per kg body weight basis than the other tracers.
Because glutamate has a relatively low solubility, we estimated that this was the maximum dose of glutamate that could be given enterally to the infants without risk of inducing feed intolerance. It is possible that absorption of free intraluminal glutamate may have been incomplete; however, this is an unlikely explanation as enrichment of the arginine and proline tracers was significant, and in piglets, it has been demonstrated that almost all enteral glutamate is absorbed and subsequently metabolized by the small intestine A further possible reason for the lack of enrichment of the glutamate pool is that of proton substitution.
This happens when a labeled deuterium atom is replaced by an unlabeled proton. This is an unlikely explanation in this case because the isomer used was [2,4,4,D 3 ] glutamate, and these deuterium atoms are unlikely to be spontaneously replaced because of their location on the molecule.
For example, studying deuterium exchange in arginine, Huang et al. By studying leucine metabolism using two tracers, one carbon the other deuterium labeled, Hoerr et al. Despite the lack of evidence that deuterium substitution may be occurring, the use of a carbon-labeled tracer would circumvent the issue and also allow measurement of oxidation.
Notwithstanding these issues, we conclude that as there was no measurable enrichment of glutamate, it is clear that enterally supplied glutamate is entirely metabolized by the splanchnic organs on first pass.
This is in keeping with the findings of others that there is a substantial metabolism of glutamate and glutamine by the gut 30 , 34 , Therefore, although it is clear that P5C synthase is present in piglet enterocytes 18 and that arginine may be synthesized from glutamine in isolated enterocytes 18 , there is negligible, or undetectably low, synthesis of arginine from dietary glutamate in healthy growing preterm infants in vivo , despite the significant metabolism of glutamate by the splanchnic organs.
We hypothesize that in the neonatal enterocyte in vivo that the carbon chain of glutamate is used primarily as an energy source and is rapidly oxidized 36 with the nitrogen atom either excreted or used through transamination. The main finding in our study is that the dietary precursor for arginine is proline, with no synthesis from glutamate. This is consistent with our work using the neonatal pig in which we have demonstrated, using a number of dietary approaches 14 , 15 , 37 , 38 , that arginine can only be synthesized from proline through citrulline and that there is no in vivo synthesis from enteral glutamate.
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