PMC:6194691 / 115757-123744 JSONTXT

{"target":"http://pubannotation.org/docs/sourcedb/PMC/sourceid/6194691","sourcedb":"PMC","sourceid":"6194691","source_url":"https://www.ncbi.nlm.nih.gov/pmc/6194691","text":"Observed fluxes of amino acids\nQuantitative measurements of fluxes of amino acids have been either of influx or net flux. Influx is measured by adding a tracer to the blood perfusing the brain and measuring the amount that enters the brain over a short period. Net flux of an amino acid is calculated as10 net flux=A-V difference×blood flowby using measurements of the blood flow and the A − V difference equal to the difference between the concentrations in arterial blood entering and venous blood leaving the brain. Direct measurements of efflux have proved difficult. In practice efflux into the blood has been calculated as the difference between influx and net flux from the blood.\nInflux of amino acids into brain parenchyma across the blood–brain barrier has been studied in rats. In a highly influential early study, rates were compared to that for water using 14C-labeled amino acids and 3HOH added together as a single bolus arterial injection. The results were reported as the brain uptake index (BUI), defined as a ratio of ratios ((uptake of 14C-aa)/[14C-aa])/((uptake of 3HOH)/[3HOH]) [300]. When added one at a time, the influxes of the amino acids varied greatly, with BUI for phenylalanine or leucine found to be more than 50% (i.e. each enters about half as easily as water) while at the other extreme influxes of proline, glutamate, asparagine and glycine were below the background limit of detection by the technique, BUI \u003c ~ 3%. Influx of each of the essential amino acids (those not able to be formed within the brain) was easily measurable.\nAll of the influxes that were clearly above baseline were inhibited when the radiolabeled amino acids were added using serum rather than a simple buffer suggesting competition for transport with the amino acids present in serum. Competition was investigated further and confirmed by measuring uptake of tracer in the presence of an excess of individual unlabelled amino acids [300].18 Quantitative estimates of the influxes of various amino acids in rats when plasma concentrations of tracers were held constant by controlled infusions [369] or during perfusion of isolated brains [43, 370, 371] have confirmed the pattern seen using BUI measurements [300, 372] (see Table 4).\nTable 4 Influx and net flux of amino acids in the indicated species\nRat1 Rat2 Rat3 Rat4 Rat5 Rat6 Dog7 Dog7 Cat8 Sheep9 Human10 Human11 Human12 Human13 Human14\nInflux* Influx Influx Influx Influx Net Net* Net* Net Net Net Net Net Net Net\nPhenylalanine 6.5 7 8 13.2 − 0.6 0.12 0.01 2.2 0.06 0.3 − 0.5\nLeucine 6.2 15 14.5 9.7 1.56 1.28 1.9 6 5.2 3.2 2.2 0.6\nIsoleucine 1.8 4 5.7 0.78 0.46 1.5 3.1 1.2 1.4 0.7 0.3\nValine 1.3 1.8 2.9 1.26 0.37 2.7 11.0 1.7 0.8 − 1.7\nTyrosine 5.3 7 4.1 − 1.1 0 0.06 1.6 0.1 − 0.1 − 0.1\nMethionine 1.6 1.7 − 1.1 0 − 0.12 1.0 0.6 0.2 0.2\nTryptophan − 1.1\nHistidine 2.5 2.5 − 1.7 − 0.23 − 0.7 0.7 − 0.3 − 1.3\nthreonine 1.2 0.8 − 3.4 0.62 − 0.1 5.0 2.6 0.1\nArginine 1 − 7.4 − 0.2 − 0.05 2 0.7 − 0.5 − 0.1\nLysine 6.2 9 − 1.1 0.58 − 1.6 2.5 − 0.3 − 0.9\nGlutamate 1.7 − 0.24 − 0.18 − 0.05 1.4 − 0.5 − 6\nGlutaminea 11.6 1 − 15.4 − 6.6 − 4.3 − 1.94 − 7.4 11.0 − 20 − 20.4\nnerv. stim.b glutamate − 1.26\nnerv. stim.b glutamine − 6.69\nValues are stated in nmol g−1 min−1. Influxes have been measured only in rats. The only available measurements of net fluxes in rats, shown in the italicized column, failed to reach statistical significance\n* Data used by Pardridge [379] in his comparison of influxes and net fluxes\naIn some columns the values are for glutamine + asparagine as the assay used detected both\nbRelease measured during nervous stimulation\n1Banos et al. [369]\n2Hawkins et al. [583]\n3Mans et al. [584]\n4Ennis et al. [415]\n5Smith and Stoll [43]\n6Brosnan et al. [378]\n7Betz and Gilboe [365]\n8Abdul-Ghani et al. [394]\n9Pell and Bergman [585]\n10Felig et al. [586]\n11Lying-Tunell et al. [587]\n12Eriksson et al. [588]\n13Grill et al. [589]\n14Strauss et al. [590]\nFrom the patterns of competition between amino acids for influx across the blood–brain barrier it appeared that there were four separate systems of transport (see e.g. [43, 44]).System L primarily for neutral amino acids, which can be inhibited by 2-aminobicyclo-(2,2,1)-heptane-2-carboxylic acid (BCH);\nSystem ASC primarily for neutral amino acids, which is not inhibited by BCH;\nSystem y+ (sometimes called system Lys+) primarily for basic amino acids;\nSystem N primarily for the nitrogen-rich amino acids glutamine, histidine and asparagine.\nA number of amino acids fit into more than one of these groups. Most of the amino acids with large BUI values are substrates for system L.\nStudies with isolated brain microvessels, which provide access to the abluminal membranes of the endothelial cells, identified two more systems.A Na+-linked transport system for small neutral amino acids (system A, with identifying substrate N-methyl-a-aminoisobutyric acid, MeAIB) [373].\nAnother system for glutamate [374].\nThe ability to prepare vesicles enriched in membranes from either the luminal or abluminal membranes of the endothelial cells [375] allowed localization of transport activities to the separate membranes with the generalization (since revised, see Sect. 5.5.6) that transporters in the luminal membrane are not Na+-linked and hence bidirectional while those in the abluminal membrane are Na+-linked favouring transport from ISF into the endothelial cells [44]. There are now known to be many more types of transporter present at the blood–brain barrier than initially suggested by identification of these systems (see Sect. 5.5.6).\nLarge rates of efflux of amino acids from CSF to blood were detected in cats [376] and rabbits [377] undergoing ventriculo-cisternal or ventriculo-cortical subarachnoid space perfusions. However, it was not possible in these studies to determine how much of the efflux was going via the choroid plexuses and how much via the parenchyma and the blood–brain barrier. Evidence that the latter route is important derives from the observation that transfer was much more rapid in ventriculo-subarachnoid infusion than in ventriculo-cisternal infusion. Both types of perfusion expose the infused fluid to the choroid plexuses, but in the former a much larger surface area of parenchyma is exposed to the fluid [376].\nThe net flux into a region can be calculated if the blood flow to that region and the concentrations of the solute in arterial blood and the venous outflow can be measured (see Sect. 5.5.6). (Equating net flux out of blood with net flux into the brain ignores possible metabolism within the endothelial cells, see the end of Sect. 5.5.6). Net flux measurements have been attempted using rats [378], but all except one of the A − V differences were not statistically significant. The rest of the net flux data in Table 4 are for larger species.\nPardridge [379] compared the influx data for rats obtained by Banos et al. [369] with the net flux data for dogs obtained by Betz et al. [365] (see Table 4) and noted that the net fluxes are much smaller than the unidirectional influxes. With the assumption that the fluxes are similar in various species, this comparison implies that there must be large effluxes, comparable in size to the influxes. Measurements of net fluxes in dogs, sheep, and humans have produced data broadly comparable with each other (see Table 4) favouring the assumption that when expressed per gram of tissue the fluxes are the same in all species.19\nAt present there are strong indications that the net flux of glutamine is outwards. This was seen in five out of six studies. There is also indication that the combined net flux of the branched chain amino acids, leucine + isoleucine + valine, is inwards. This was seen in six out of seven studies. But as described in the next section there is no evidence for a sufficiently large inwards net flux of neutral amino acids to provide for all of the transamination invoked in the explanations of glutamate turnover, at least in 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