AM.EKIC:AN .JOURN.I\L OF PHYSI0I.OC.

Y
Vol. 226, No. 3, March 197 4. Printed in U.S..4.

Medullary collecting-duct function in antidiuretic

and in salt- or water-diuretic rats

H. SONNENBERG (With the Technical Assistance of A. Marsden-Potter and C. J. Potter)

Department of Physiology, University of Toronto, Toronto 5, Ontario, Canada

SONNENBERG, H. Medullary collecting-duct function in antidiuretic rats) received a glucose-saline mixture as drinking fluid
and in salt- or water-diuretic rats. Am. J. Physiol. 226(3): 501-506. (0.9 g NaCl + 5 g glucose/l00 ml), while a third group
1974.-Microcatheterization of the medullary collecting duct of drank a glucose solution in distilled water (5 g/l00 ml).
adult rats was used to study the contribution of water, sodium, and Animals were used between 1 and 3 wk after being placed
potassium transport in this nephron segment to urinary excretion. on the fluid regimen. Rats were anesthetized with Inactin
In antidiuresis medullary reabsorption of fluid and sodium was

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(10 mg/lOO g body wt) and maintained at a body tempera-
50/;, and 3y0 of filtered load, respectively. Animals chronically in-
gesting saline, followed by infusion at a rate comparable to prior
ture near 38°C on a heated operating table. The trachea
voluntary intake, showed an increase in the absolute amount of and a jugular vein were cannulated and an intravenous
water and Na delivered to the medullary duct system from the infusion was begun. Control rats received Ringer solution
cortex of the experimental kidney. The absence of any further re- (1 ml/l00 g body wt per h), while the saline- and water-
absorption of either substance resulted in diuresis and natriuresis. drinking rats received isotonic saline and glucose solution
In rats on high water intake with subsequent infusion, an in- (2 g/l00 ml>, respectively, at a rate of 12 ml/h. Since these
creased amount of water, but not sodium, was delivered to the two groups had an overnight fluid intake of 160 ml (range:
medullary collecting duct. Essentially complete inhibition of fur- 80-400 ml) and 190 ml (range : 100-275 ml), respectively,
ther water reabsorption, associated with Na reabsorption as great
the voluntary rate of fluid ingestion was equal to or ex-
as, or greater than, that during antidiuresis, resulted in excretion
ceeded the rate of infusion during the experiment. In
of a large volume of hypotonic urine. No net transport of potas-
sium was found in the collecting duct either in antidiuresis or
water-loaded rats the infusion did not cause hemolysis in
saline diuresis, -while in water diuresis net reabsorption was de- either plasma or urine, and no glucose was excreted. Follow-
tectable. It is concluded that the medullary collecting duct is a ing cannulation of a femoral artery for blood pressure
major site for the regulation of urinary excretion of both salt and measurements and sampling, the right ureter was cannu-
water. lated through a flank incision. Tubing (PE-50), pulled out
at the tip if necessary, was used for ureteral cannulations
renal Na and K transport; water reabsorption (2). The left kidney was mobilized through the same in-
cision, placed in a Lucite cup, and covered with Parafilm
and cotton wool to prevent drying and cooling of the sur-
face. The left ureter was carefully dissected to the pelvis
THE MAJOR FUNCTION of the mammalian collecting duct has
and cannulated. A small longitudinal slit was made in the
been thought to be the passive equilibration of tubular
upper surface of the ureteral wall as close to the body of
with medullary interstitial fluid. During antidiuresis the
the kidney as possible, and the incision was spread by ad-
high permeability of collecting-duct epithelium allows
vancing the ureteral cannula to the level of the cut. Usually
diffusion of water into the medulla, thereby concentrating
the tip of the papilla became visible in the incision; occa-
the final urine. In the absence of antidiuretic hormone the
sionally, however, a relatively short papilla remained in-
collecting duct is impermeable to water, preventing equi-
accessible even after careful repositioning of the kidney.
libration of hypotonic distal tubular fluid and resulting in
Such animals could not be used for an experiment. Urine
water diuresis (11, 18).
from the left, experimental kidney was collected by applying
Recent experimental evidence suggests, however, that the
continuous gentle suction to the ureteral catheter. Since the
collecting duct may play a more active role in the regulation
sides of the opened ureter formed a small leakproof “well,”
of urinary excretion, affecting both natriuresis (3, 14, 15)
loss of urine through the incision was not a problem. Aspira-
and water diuresis (5, 6). The microcatheterization tech-
tion of tissue fluid into the opened ureter was prevented by
nique (7) was, therefore, used to determine transport
careful prior dissection and by continuous removal of any
characteristics of the medullary collecting duct in anesthe-
free tissue fluid via suction. The similarly low osmotic pres-
tized rats during antidiuresis, during the natriuresis following
sure of urine collected from both kidneys during water
saline infusion, and during water diuresis.
diuresis (see Table 1) indicates that contamination of
experimental urine was precluded if these precautions were
METHODS
observed. Conversely, similar filtration rates in control and
Male Sprague-Dawley rats (weight range: 240-405 g) experimental kidneys of saline- and water-diuretic rats
were used. Control animals (11 rats) were maintained on indicate the absence of significant loss of urine. Since the
water and Purina laboratory chow. A second group (seven same techniques were used in all three series, it is concluded
502 H. SONNENBERG

that the suction method results in quantitative collection of technique was the advancement of the catheter through the
urine. Inulin-3H was added to the constant intravenous outer glass capillary guide which was fixed on a micro-
infusion 1 h prior to the beginning of urine collections to manipulator. Polyethylene catheters were pulled to an
deliver 125 &i over the course of the experiment. Total outer-tip diameter between 16 and 40 p (ID 8-18 cl), by
time of infusion before collections varied between 1.5 and means of a heated wire loop (7) and were mounted on glass
2.5 h; duration of the experiments was from 160 to 240 min. capillary tubes to leave 5-10 mm of length available for
Urine from the right, control kidney was collected continu- insertion. The catheters were filled with colored, heavy
ously at ZO-min intervals. Arterial blood samples were taken mineral oil and tested in a water bath to determine the
at the beginning and end of each collection period. Urine degree of suction required to just overcome resistance to
from the left, experimental kidney was taken at intervals to flow in the catheter. After insertion of the catheter into a
coincide with simultaneous sampling of collecting-duct papillary duct, the same level of suction was set by means of
fluid. Sodium and potassium concentrations in plasma and a mercury manometer and maintained throughout the
urine were determined by flame photometry; inulin-3H collection (5-20 min).
concentrations, by liquid scintillation counting in a toluene- Catheters were advanced into a duct, usually until ob-
based scintillant. Urinary excretion of sodium (U,,V) and struction was felt, and then pulled back slightly. In attempts
potassium (U,V) was calculated, as were glomerular to advance the catheter as far as possible toward the cortex,
filtration (GFR) rates. penetration into the tissue could occur. Samples were dis-
The microcatheterization technique of Jarausch and carded if red cells appeared in the collected fluid or if the
Ullrich (7) was used to obtain samples of tubular fluid at predetermined suction did not result in a steady rate of

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different levels in the collecting duct. A modification of this collection. On completion of the collection, the catheter

TABLE I. Comparison of average control and experimental kidney function in antidiuretic and in saline- and water-diuretic rats

V, pl/min per g KW UN&V, neq/min per g KW UKV, neq/min per g KW Uosm, mosM GFR, ml/min per g KW
Control Exptl Control Exptl Control Exptl Control Exptl Control Exptl

Antidiuresis
2.64 4.50 70 165 817 817 940 1.21 0.94
1.41 5.98 59 332 570 1,269 1.24 0.87
2.33 4.59 107 211 1,063 673 1,070 1.16 0.72
2.10 6.00 117 175 594 891 994 1.58 0.74
3.68 10.92 203 498 823 672 537 1.18 0.90
5.17 5.26 258 108 740 703 0.76 0.68
1.98 3.82 217 390 349 576 1,540 1,344 1.14 1.07
1.89 4.49 52 142 630 567 1,097 1.17 1.16
5.10 6.87 57 118 1,733 904 1,573 726 1.58 0.97
3.15 11.53 227 767 1,048 1,056 1,880 587 1.31 0.90
3.68 10.84 91 640 1,334 1,376 1,875 734 1.34 1.04

z 3.01 6.8Ot 133 322* 882 864 1,717 892t 1 .24 0.91t
SE ho.38 zt.87 &24 &68 Ml8 zt82 zt93 *88 A.07 334
Saline diuresis
32.4 58.8 5,500 6,919 1,248 1,225 496 345 1.15 0.99
18.1 47.8 4,514 6,345 396 624 323 295 1.02 0.95
49.0 52.9 6,841 6,770 546 657 752 351 1.06 1.19
53.6 58.8 8,293 8,347 916 1,005 425 358 1.32 1.30
51.5 59.5 6,897 8,689 883 930 409 311 1.28 1.23
27.2 62.5 4,571 8,725 540 768 531 357 1.09 1.14
42.3 46.3 7,288 6,988 805 1,025 450 412 1.29 1.28

2 39.2 55.2*5 6,272 7 95wi 762 891 484 347* 1.17 1.155
SE A5.1 AZ.4 zt544 +380 *110 rt82 It51 +14 zt.05 It.05
Water diuresis
60.9 45.8 15 346 153 139 73 72 1.39 1.04
89.3 77.1 123 237 740 567 51 51 1.26 1.31
91.5 93.5 167 385 544 576 41 38 1.47 1.35
77.4 78.7 62 168 573 668 49 47 1.20 1.18
56.4 64.1 43 166 371 491 61 52 1.28 1.19
73.7 104.8 63 307 430 1,052 51 46 1.35 1.29
67.2 66.6 58 164 504 522 43 44 1.05 1.11

z 73.8 75.85 76 253-f 474 575$ 53 50 1.29 1.215

SE zt5.1 zt7.4 *20 rt35 zt69 *lo2 &4 *4 zt.05 LO4
~~
gKW= gram kidney weight. *t Significant difference (P < 0.05, P < 0.01) between control and experimental kidney function
within groups. $8 Significant difference (P < 0.05, P < 0.01) between experimental kidney function of the control group and that of
either saline- or water-diuretic groups.
COLLECTING-DUCT FUNCTION IN DIFFERENT DIURETIC STATES 503

was removed from the duct and its distance of insertion was tained by dividing the tubular fluid-to-plasma concentra-
measured with an ocular micrometer. In each animal tion ratio for sodium and potassium by (TF/P)r,.
attempts were made to obtain a representative range of Orthogonal regression analysis (19) was used to obtain
sampling depths. An average of nine samples (range 2-14) lines of best fit. Paired and unpaired t-test analysis as well
was collected. At the end of the experiment a saggital sec- as correlation coefficients were used to obtain levels of
tion of the experimental kidney was taken, from which it statistical significance of differences in data (13).
was possible to determine the distance from the tip of a
papilla to the border between the outer and inner stripes of
RESULTS
the outer medulla. Since it has been found that the outer
stripe of the outer medulla comprised approximately one- A comparison of the average function of control and
third of the remaining distance to the kidney surface (un- experimental kidneys is given for individual animals in
published observations), this amount was added to the Table 1. In nondiuretic rats urine volume and sodium ex-
measurement to obtain the total medullary length. The cretion were uniformly higher in the experimental kidney,
different distances of insertion of catheters were then ex- despite relative reduction in the urinary solute concentra-
pressed as percents of total medullary length. tion and glomerular filtration rate. With the exception of
Sodium and potassium concentrations, in duplicate GFR, these distinctions tended to persist in saline diuresis;
aliquots (10 nl) of tubular fluid, were determined in an whereas during water diuresis, only sodium excretions re-
Aminco helium-glow photometer, Inulin-3H (30 nl), by mained significantly different. In saline-loaded animals
liquid scintillation counting. A Clifton nanoliter osmometer both renal fluid and sodium output were increased more

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was used to measure the total solute concentration in than lo-fold compared to control rats. Diuresis in the
tubular fluid samples and in urine of the control group. Pre- water-loaded group showed further enhancement; natri-
cision of the different measurements was as follows: for uresis, however, was not different from the control group.
inulin-3H, radioactivity of tubular fluid samples was always Both control and high-salt animals had similar plasma
greater than 5 X background (background = 8-10 counts/ sodium concentrations (142 rt 2 SE and 141 rt 1 meq/liter,
min). Two sets of ZO-min counts were obtained, resulting in respectively), although plasma potassium levels were sig-
an average deviation of 2.0 % =t 1.8 SD from the mean. nificantly (P < 0.01) reduced in the latter (4.5 rt 0.1 to
Duplicate determinations on sample concentrations of 3.7 =t 0.1 meq/liter). In contrast, high water intake with
sodium, potassium, and total solute resulted in average subsequent infusion led to significant reduction (P < 0.01)
deviationsof2.3% 2t 2.2 SD,4.5% rt 4.6SD, and 1.0% =t in plasma sodium (131 rt 2 meq/liter) and no change in
0.8 SD, respectively. Urine of diuretic animals was tested by plasma potassium (4.7 rt 0.1 meq/liter). Similar average
macro osmometry. Plasma inulin concentration was calcu- hematocrit values of 47.5 & 1.6, 46.5 rt 1.6, and 49.0 &
lated for the midpoint of each fluid-collection period and 0.8 % in control and in salt- and water-loaded groups, re-
the tubular fluid-to-plasma concentration ratio ((TF/P)r,) spectively, indicate that high fluid intake did not result in
was determined. In addition, the fractions of filtered sodium significant increase of circulating blood volume. In addi-
and potassium remaining at the collection site were ob- tion, hematocrit values in both diuretic series tended to rise
throughout the course of experiment, whereas plasma
CORTEX 1 MEDULLA I URINE sodium concentrations did not change significantly; this
1200
r0A 0
0

P
shows that urinary
balancing
excretion was more than capable
the large volume of fluid infused. Renal function
of

tended to remain relatively constant throughout an experi-

ment, although absolute rates of excretion differed widely
600
among groups.
Total solute concentration in tubular fluid was related to
400 medullary length in Figs. 1 and 2.1 Since in the control
i
E series there was a large variability in osmotic pressure of
2 corresponding urine samples, the points were divided arbi-
I
k 200 I I I I I f f-f trarily into two groups according to simultaneous urine
e -I- concentrations above (Fig. 1A) and below (Fig. 1B) the
average for control animals (see Table 1). In each case there
was a significant rise (P < 0.01) of osmotic pressure toward
the tip of the papilla with correlation coefficients (r) of
0.809 and 0.73 1 for high and low groups, respectively. In
contrast, tubular fluid concentration during saline diuresis
(Fig. 2A) did not change with medullary length and re-
2ooL I 1 I I 1 J U mained near isotonicity throughout. Tubular fluid samples
0 20 40 60 80 100
percent medullary length from water-diuretic rats (Fig. 2B) showed a significant
FIG. 1. Change of osmotic pressure in collecting-duct fluid with
medullary length in antidiuresis. A: data with simultaneous urine con- 1 For a table of individual values for concentrations of inulin, so-
centrations greater than 890 mosM. 23: data with simultaneous urine dium, potassium, and total solute in tubular fluid samples from differ-
concentrations less than 890 mosM. Regression lines of log osmolality ent medullary depths, order NAPS Document 02302 from Micro-
with length are indicated; open circles represent average osmotic fiche Publications, 305 East 46th St., New York, N. Y. 10017, remitting
pressure &SD of corresponding urine samples. $1.50 for microfiche or $5.00 for photocopies.
504 H. SONNENBERG

CORTEX 1 MEDULLA 1 URINE

absorption reduced this tubular load to a value less than
0.3 76 at the tip of the papilla. In saline-loaded rats (B),
initial sodium load was increased to 4.5 ‘%. No net transport
of the ion occurred in the collecting duct, which accounts
for the large natriuresis in this group. During water diuresis
(C), despite a tubular-volume flow rate greater than in the
high-salt group, fractional sodium reabsorption in the
medullary collecting duct was indistinguishable from that
in control animals.
A similar plot for potassium transport (Fig. 5) showed no
net secretion or reabsorption of the ion in the collecting duct
of either nondiuretic or saline-diuretic rats. In water-
diuretic animals there was significant (P < 0.01) reabsorp-
tion of K, although only 13 % of total variability could be
0 l!J accounted for by this mechanism (T = -0.361).
-20 0 20 40 60 80 100 Absolute tubular load of fluid, Na, and K, in the medul-
percent medullory length
lary collecting duct was calculated for the three different
FIG. 2. Change of osmotic pressure in collecting duct fluid with groups of rats as follows: mean glomerular filtration rate in a
mrdullary length in- A: saline diuresis; B: water diuresis. Linear re-
given series of animals was obtained by averaging the inulin
grcssion is indicated. Symbols as in Kg. 2.

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clearances that were measured simultaneously with the
individual tubular fluid collections. For Na and K this
CORTEX 1 MEDULLA I URINE
8 value was multiplied by average plasma concentrations to
0
200 obtain absolute filtered load of each ion. Filtered load was
0A 8
too then multiplied by the fraction remaining at beginning and
1 8
end of the collecting duct, using the calculated regression
50
t lines (Figs. 3-5). Results are given in Table 2. With the
20 exception of volume flow during saline diuresis, the calcu-
t I I I I I
IOL 1 1
A ,- CORTEX 1 MEDULLA 1 URINE

+ 5L I I I I I I 1

0.001 L 1 I I I I I
a

5L 1 I I I I I JI 1
-20 0 20 40 60 80 100
percent medullary length

FIG. 3. Change of collecting-duct fluid-to-plasma concentration

ratio of inulin with medullary length in--4: antidiurcsis; B: saline
diuresis; C: water diuresis. Average urinary values from experimental
kidneys of individual annuals in each group are given (open circles).
Regression lines of log of concentration ratio with length are indicated.

decline
-0.429,
of concentration
1’ < 0.01).
toward the papillary tip (r =
0.05
roc a a a
Individual tubule TF/P inulin ratios are depicted in
Fig. 3 for control (,4), saline-diuretic (B), and water-diuretic
(C) series. Significant increase with medullary length was
seen only in nondiuretic animals (r = 0.747, I’ < O.Ol),
although the TF/P inulin at the beginning of the medulla
was comparable for all three groups. Further reabsorption
of the similar fractional fluid load, therefore, did not occur
during either saline or water diuresis.
The fraction of filtered sodium remaining in the tubule
was calculated for each collection and related to medullary pwcmt mdullary length

distance in Fig. 4. During antidiuresis (A), approximately FIG. 4. Fraction of filtered sodium remaining at different levels of
3 YL of filtered Na entered the medullary duct. Further re- medullary collecting duct. Symbols and explanations as in Fig. 3.
COLLECTING-DUCT FUNCTION IN DIFFERENT DIURETIC STATES 505

CORTEX 1 MEDULLA 1 URINE TABLE 2. Tubular load and reabsorbtion of ions and water 1 J
l
in medullary collecting duct in antidiuresis
.
and 2n saline or water diuresis

V, pl/min per g KW Ka, neq/min per K, neq/min per

id-’ &SW
-e l
l
l

0.02L I I I I I I I
Antidiuresis 884 122
Saline diuresis 688 -142
Water diuresis 641 331*

ns;r 0B l l

g KW = gram kidney weight. Statistically significant reabsorption (P <

0.0 1) based on correlation coefficients.

l
0. I lm l l e As demonstrated by Ullrich (17), under antidiuretic con-
l 8
te I I 1 ditions there is significant reabsorption of both sodium and
0.05 t, I I
water in the medullary collecting duct, comprising in the
present experiments 3 % and 5 % of filtered load, respec-

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tively. During saline diuresis, there is no net fluid or Na
reabsorption, whereas medullary delivery is increased
(Table 2). Since osmolality in the medulla is reduced by
saline loading ( I), reduction of the usual gradient for water
reabsorption could account for the observed diuresis;
-20 0 20 40 60 80 too however, assuming a constant rate of sodium reabsorption,
percent medullary length
the increased Na load can account for only a part of the
FIG. 5. Fraction of filtered potassium remaining at different levels natriuretic response. Inhibition of collecting-duct transport
of medullary collecting duct. Symbols and explanations as in Fig. 3.
of sodium is, therefore, a demonstrable feature of the saline
natriuresis.
lated tubular load at the papillary tip was within 1 SD of Aldosterone has been shown to increase Na transport
the observed urinary excretion (Table 1) ; this provides a capacity in the collecting duct (16). In the present saline-
loaded animals, chronically decreased release of aldosterone
degree of validation for the microcatheterization technique.
In antidiuresis there was extensive reabsorption of both associated with the observed reduction in plasma K levels
water and sodium in the medullary collecting-duct system, (10) could account for both increased medullary delivery
and lack of collecting-duct reabsorption of sodium. Such
while net potassium transport was insignificant. In saline
dependence of medullary Na transport on aldosterone
diuresis both fluid and Na reabsorption were completely
inhibited, resulting in total excretion of the increased could also explain the apparent discrepancy between the
present results and those of Diezi et al. (3). Using micro-
tubular load entering the medulla. Again, no net K trans-
port was observed. In water diuresis, while fluid reabsorp- puncture techniques, these investigators found an increase
tion was inhibited, absolute sodium reabsorption was in- in fractional sodium reabsorption in the terminal collecting
ducts of young rats subjected to acute saline infusion. If the
creased compared to control conditions. A small but signifi-
acute treatment is less effective in altering plasma aldos-
cant rate of reabsorption of potassium was seen in this
terone levels, reduction of the hormone in chronic but not
group.
acute saline loading provides a satisfactory explanation for
the difference in transport characteristics. The data, how-
DISCUSSION
ever, do not exclude a mediation of the inhibition of collect-
The difference in excretion between control and experi- ing-duct transport by factors other than aldosterone (14).
mental kidneys could be due to at least two different mecha- During water diuresis, there is little or no fluid reabsorp-
nisms. First, the opening of the ureter may reduce hydro- tion in the collecting duct. Absolute sodium transport,
static pressure and increase flow rate in collecting ducts, however, is, if anything, increased compared to control
thus allowing less time for equilibration. The opposite effect conditions. Inhibition of release of antidiuretic hormone due
is believed responsible for exaggeration of transport mecha- to reduction in plasma osmolality (PosM = 272 & 2.5 SE
nisms in stop-flow experiments (9). Second, it has been mosM), combined with comparatively low medullary solute
shown that a change in solute concentration of the fluid concentration (8), can adequately explain the diuresis.
bathing an exposed papilla results in a corresponding Since, as a result of reduced plasma Na concentration in this
change of concentration in the terminal collecting-duct group, aldosterone levels may be enhanced (lo), maintained
fluid (12). Variable decreases in osmolality of the fluid sur- reabsorption of the ion is not unexpected. When paired col-
rounding the experimental papilla, depending on exposure lections from the same duct were used, an increase in net
and urine flow, could explain the difference in osmotic fluid reabsorption was seen in the terminal collecting duct of
pressure of urine from control and experimental kidneys young water-loaded rats with hereditary diabetes insipidus,
(Table l), as well as the variability in antidiuresis (Fig. 1). compared to antidiuresis in the same rats (5). The larger
506 H. SONNENBERG

variability in the present experiments, due to summation of exchange of the two ions in the isolated cortical collecting
collections from different ducts and animals, allows no duct of the rabbit (4). It should be emphasized, however,
conclusion regarding water reabsorption in the final l-2 mm. that the scatter in the potassium data precludes any but
It is evident, however, that such increased reabsorption, if tentative conclusions about K transport.
present, does not extend over the whole medullary duct (see It is concluded that tubular reabsorption of sodium in the
Fig. 3C and Table 2). On the other hand, the data do sup- medullary collecting duct, possibly under the influence of
port the finding of urinary dilution (6) in the collecting aldosterone, contributes critically to the regulation of
system of water-diuretic rats (see Fig. 2B). urinary excretion of both salt and water. Inhibition of
In corroboration with findings of Ullrich (17), and Diezi collecting-duct transport of sodium has been demonstrated
et al. (3), no evidence for net potassium secretion into as a major mechanism of saline natriuresis. On the other
medullary collecting-duct fluid is seen in any of the present hand, under the present experimental conditions, regulation
experimental groups. On the contrary, in water diuresis a of potassium excretion seems to occur upstream from the
small but statistically significant net reabsorption of the medullary collecting system.
ion is evident. Net reabsorption of potassium was also
found after potassium depletion in rats (3). The dissociation This study was supported by the Medical Research Council of
between Na and reverse K transport in the medullary col- Canada, Grant MA-4043.
lecting duct is in sharp contrast to the carrier-mediated Received for publication 29 August 1973.

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5. JAMISON, R. L., J. BUERKERT, AND F. LACY. A micropuncture 15. SONNENBERG, H. Proximal and distal tubular function in salt-
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6 JAMISON, R L., AND F. B. LACY. Evidence for urinary dilution by 16. UHLICH, E., C. A. BALDAMUS, AND K. J. ULLRICH. Einfluss von
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