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a specific isoform of the Na-H antiporter NHE3 as described in 6 and shown in Figure 9 2 This transporter is the major means not only of hydrogen ion secretion but also of sodium uptake from the proximal tubule lumen Second, a primary active H-ATPase exists in all the hydrogen ion secreting distal tubular segments The Type A intercalated cells of the collecting-duct system, in addition to their primary active H-ATPase, possess a primary active H-K-ATPase, which simultaneously moves hydrogen ions into the lumen and potassium into the cell, both actively (Figure 9 3) Note that, as described in 8, the luminal membrane H-K-ATPase also mediates active potassium reabsorption by these cells and contributes to potassium homeostasis The basolateral membrane exit step for bicarbonate is via Cl-HCO3 antiporters or Na-HCO3 symporters (Figures 9 2 and 9 3), depending on the tubular segment In both cases, the movement of bicarbonate is down its electrochemical gradient (ie, the transport is passive) Symport with sodium is the dominant means of extruding bicarbonate in the proximal tubule and is particularly interesting because efflux of sodium is up its electrochemical gradient This is a rare case of a secondary active transporter that does not use the sodium gradient as the energy source, but actually moves sodium up its electrochemical gradient Through its secretion of hydrogen ions, the proximal tubule reabsorbs 80 90% of the filtered bicarbonate The thick ascending limb of Henle s loop reabsorbs another 10%, and almost all the remaining bicarbonate is normally reabsorbed by the distal convoluted tubule and collecting-duct system (except for alkalotic individuals, who will excrete some of the bicarbonate; see later discussion) Throughout the tubule, intracellular carbonic anhydrase is involved in the reactions generating hydrogen ion and bicarbonate In the proximal tubule, carbonic anhydrase is also located in the lumen-facing surface of apical cell membranes, and this carbonic anhydrase catalyzes the intraluminal generation of CO2 and water from the large quantities of secreted hydrogen ions combining with filtered bicarbonate.

Development (3+ mos)

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Sometimes my father would work the team into a near-frenzy before a critical game Bill Forrester, a linebacker on Lombardi s early Packer teams, says he sustained his worst injury of Lombardi s first season in Green Bay when he jumped up and smashed his elbow against his locker after listening to my father s pep talk before a game against the Bears Another player once begged my father to Stop, stop! in the middle of his pre-game pep talk It was too much, too much But he was just as capable of pushing the team in the other direction As he boarded the bus that would take the Packers to the first-ever superbowl game against the Kansas City Chiefs, Lombardi stepped into the middle of the aisle and got the team s attention Then, to everyone s amazement, he did a soft-shoe dance in the aisle and sat down What was that all about, an astounded assistant asked him They were too tight, my father responded matter-of-factly

If all the filtered bicarbonate is reabsorbed, there is no acid-base consequence for the body of filtering a large amount of bicarbonate; it is as though none had been filtered in the first place When base has been added to the body fluids, the effect is to increase the plasma concentration of bicarbonate Whether this increased bicarbonate comes from metabolism of protein that contains many anionic amino acids or from ingestion of baking soda (sodium bicarbonate), the result is the same: The body fluids contain more bicarbonate The renal handling of such base loads is relatively straightforward: We excrete enough bicarbonate in the urine to match the input If the addition of base to the body is 30 mEq and the kidneys put out 30 mmol of bicarbonate, the kidneys have achieved their goal: balance The kidneys do this in 2 ways: (1) allow some filtered bicarbonate to pass through to the urine and (2) secrete bicarbonate via Type B intercalated cells The Type B intercalated cells, which are found only in the cortical collecting duct, do indeed secrete bicarbonate In essence, the Type B intercalated cell is a flipped-around Type A

LOMBARDI RULE #4

QA (1+ mos)

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Before a game against the arch-rival Bears in 1967, he skipped the high-volume exhortation that many of the Packers must have been expecting Instead, he told a joke, keying off the fact that the area around Green Bay had been settled many years earlier by Belgian immigrants Did you ever hear why Belgians are so strong

mechanisms in Type A intercalated cells for the secretion of hydrogen ions that result in the formation of titratable acidity The apical membrane contains H-ATPases, which transport hydrogen ions alone or in exchange for potassium B, The Type B intercalated cell secretes bicarbonate and simultaneously transports hydrogen ions into the interstitium The difference between this cell type and the Type A cell and those in the proximal tubule is that the location of the transporters for hydrogen ions and bicarbonate are switched between apical and basolateral membranes ATP, adenosine triphosphate

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