BULLETINOF

MATHEMATICALBIOLOGY
VOLUME 36, 19"/4

F L O W OF A N E W T O N I A N F L U I D T H R O U G H A
P E R M E A B L E T U B E : T H E A P P L I C A T I O N TO T H E
PROXIMAL RENAL TUBULE

[] E. A. iVlXaS~AT,TJand E. A. TROWBRIDGE
Department of Applied Mathematics and Computing Science,
University of Sheffield,
England

Creeping :flowof a Newtonian fluid through a rigid permeable tube is considered and the
transmural seepage is assumed to obey Darcy's law. Closed-formsolutions for the pres-
sure and velocity fields are presented and equations describing the axial variation of the
mean cross-sectional pressure, the axial volumetric flow and the transmural fluid flux are
derived. Approximate solutions for small seepage rates are given and are applied to the
flow in the proximal renal tubule. Probable values for the epithelium permeability and
the intra|uminal hydrostatic pressure drop are obtained.

1. Introduction. In certain physical and physiological processes filtration and

mass transfer occur as a fluid flows through a permeable tube. This pheno-
menon is encountered in reverse osmotic desalination; in extracorporeal cir-
cuitry for body fluid processing, such as the haemodialyzer; in lymphatic flow,
and in flow in the nephron tubules. The velocity and pressure fields in these
situations differ from simple Poiseuille flow in an impermeable tube since the
fluid in contact with the wall has a normal velocity component.
Laminar flow in a pipe with constant leakage or injection has been investi-
gated by Yuan and Finkelstein (1956) who obtained perturbation solutions for
"'small" and "large" transmural seepages. The situation has been considered
as a model for the flow in the proximal renal tubule (Macey, 1963, 1965) and
creeping flow solutions were obtained for leakage rates which vary linearly or
457
458 E. A. MARSHALL AND E. A. T R O W B R I D G E

decrease exponentially along the length of the tube. Macey also gave a formal
solution for the case where the leakage-rate has an arbitrarily prescribed axial
variation. Macey's solution for the latter cases has been extended by Kozinski
et al. (1970) who added solutions for the flow between porous parallel plates.
The problem has also been considered in connection with the proximal tubule by
Friedlander and Walser (1965) who used an approximation similar to that used
in lubrication theory to derive a cross-sectional velocity profile for small
leakage in which the variation with axial distance is arbitrary.
In the above investigations no consideration has been given to the physical
conditions existing at the permeable boundary when leakage occurs, although
Kozinski et al. noted that it was expected that the wall permeability would have
influence on the leakage-rate. A relationship frequently employed in calcula-
tions on flow through inorganic porous materials is Darcy's law (see, for example
Streeter, 1961). This states that the leakage-rate is directly proportional to the
transmural pressure difference. For permeable organic membranes the depen-
dence of seepage-rates on the transmural pressure difference was first postulated
by Ludwig (1861) (cf., Landis and Pappenheimer, 1963) and has subsequently
been employed extensively in theoretical investigations (e.g. Deen et al.,
1972, 1973).
Here Darcy's law is expressed as follows. If u, A denote respectively the
normal component of fluid velocity at the boundary and the difference between
the transmural pressures then
KA
u = - -~t ' (I)

where t is the thickness of the boundary, ~ is the viscosity of the seepage fluid
and K is the permeability of the boundary, commonly expressed in units of cm 2
or, alternatively, dareys (centipoise cm2/atm see). It may be verified that the
hydrodynamieal solutions obtained in the investigations referred to above are
not compatible with (1). The purpose of the present paper is to submit solu-
tions for the flow of a Newtonian fluid through a permeable tube which satisfy
this basic relationship.
This paper is arranged as follows. The low Reynolds number hydrodynami-
cal equations and the boundary conditions pertaining to the present problem
are set out in Section 2 and an exact, closed-form solution to these is presented.
For a particular value of a permeability parameter, graphs showing the cross-
sectional variation of pressure and velocity are constructed from this solution.
In Section 3 equations describing the axial variation of the wall leakage flux q,
the mean cross-sectional pressure P, and the axial volumetric flow Q are derived
from the exact solution and their properties are discussed. It is shown that
 FLUID THROUGH A PERMEABLE TUBE 459
P and Q are related by an expression which is analogous to Poiseuille's law.
This result finds an application elsewhere (Marshall and Trowbridge, 1974) in
the approximate analysis of a more complicated flow problem involving trans-
mural flnid leakage. Corresponding relationships between P and Q for the
cases of constant leakage and leakage with exponential attenuation with dis-
tance were obtained by Macey (loc. cir.). In Section 4 approximate solutions
valid for small seepage-rates are derived from the exact solution. The form of
these approximate solutions is consistent with the results obtained by the earlier
investigators referred to above. Finally, Section 5 is concerned with the
application of the solutions presented here to the problem of flow in the proxi-
mal renal tubule. Available experimental data is used together with the re-
sults obtained in this paper to estimate probable values for the permeability of
the tubular epithelium to isotonic glomerular filtrate and the intraluminal
hydrostatic pressure drop along this nephron segment.

2. Solution of the Hydro-Mechanical Equations. The steady flow of an incom-

pressible, homogeneous Newtonian fluid of viscosity/~ through an indefinitely
long circularly-cylindrical permeable tube of radius a is considered relative to a
system of cylindrical coordinates (r, 0, z) in which the z axis coincides with the
axis of the tube. The tube wall is considered to be rigid and it is assumed that
the fluid input is such that no free liquid surface is formed.
This investigation will be concerned with flow situations which are symmetri-
cal with respect to the tube axis and in which the magnitude of the Reynolds
number is such that the inertial terms in the Navier-Stokes equations are
negligibly small compared with those associated with the viscous and hydro-
static stresses. This quasi-statical fluid flow is thus described by the creeping-
flow equations (Happel and Brenner, 1965)

1 ~p ~2vT 1 ~vr Vr ~2v~

= + r r +
(2)
1 ~p ~2vz 1 ~vz ~2vz

and the continuity equation for an incompressible fluid

8-'7- + = o, (a)

where vr(r, z), v~(r, z) are the components of the particle velocity in the radial
and axial directions, and p(r, z) is the hydrostatic fluid pressure. A physical
requirement is that vT, vz and p are bounded at all points in the flow field.
460 E. A. MARSHALL AND E. A. T R O W B R I D G E

Also for the no-slip condition at the tube wall to be satisfied,

v (a, z) = o. (4)

With the form of solution presented here it is not possible to prescribe arbitra-
rily values of p and v~ over a typical cross-section, for example z = 0 and it is
assumed that the form of the entry cross-sectional distributions of pressure and
velocity is compatible with that developed at large downstream distances. To
evaluate integration constants occurring in the solution the conditions

Qo -- 2rr f ~ rvz(r, O) dr, (5)

-Po = ~ rp(r, O) dr, (6)

where Qo is the rate of fluid flow through the cross-section z = 0 and P0 is the
mean pressure applied to the fluid over the cross-section z = O, are introduced.
The application of the solutions to be given here to the problem of a finite tube
with an arbitrary distribution of pressure and velocity over the entry plane and
an arbitrary distribution of pressure over the exit plane will, in general, involve
some error. I t is expected that this error will be negligible at distances from
the tube ends greater than a small multiple of the diameter. This is completely
analogous to the "entry length" problem in conventional pipe flow.
The permeability condition (1) is used to evaluate an otherwise undetermined
parameter which appears in the solutions and for this purpose the non-dimen-
sional permeability
~* =- K/at (7)

is introduced. The solutions to be presented are strictly applicable only to the

leaking flow of a single-component fluid for which the transmural pressure
difference A is p(a, z) - Pb, the hydrostatic pressure pb of the fluid in contact
with the external surface of the tube being considered constant. However,
the effects of dissolved solutes m a y be taken into account, approximately, by
considering A to be composed of the differences of the hydrostatic fluid pressures
and the differences of the solution osmotic pressures. Thus

A = p(a, z) - r z) - pb + (s)

where r z), tb are the osmotic pressures of the fluids on adjacent sides of the
tube wall. For brevity we write

PT = Pb r z) -- tb, (9)
 FLUID THROUGH A PERMEABLE TUBE 461

where Pr may be visualized as an effective back-pressure opposing fluid leakage.

The combination of (1), (7), (8) and (9) then gives, since u = vr(a, z),

vr(a, z) =
aT*[p(a, z) - PT] , (lo)

as the condition which must be satisfied for the solutions to be compatible with
the Darey/Ludwig law [1856].
In the present investigation PT is considered to be constant. This would
apply to a single component fluid seeping into a constant pressure environment
such as atmospheric surroundings or to the leakage of a solution provided that
the osmotic pressure difference and the external hydrostatic pressure are con-
stant.
Variables separable solutions for p - Pr, v , v~ are sought, and by using
elementary analytical techniques (see Appendix) it is found that solutions of
this form which satisfy (2), (3), (4), (5) and (6) and which are bounded at r = 0
are

P - Pr 2Jo(2r/a) f 2z l ~}
-Po - P'c = 2j1(,~ ) cosh a C sinh ,

( 2 r _ [2r\ [AJl(2)

(-~Jl(-~) "1(2)7 (~)} (Csinh

where J0 and J1 are Bessel functions of zero and first order, )l ()l > 0) is a positivet
parameter which arises from separation of variables and

v o = Qo (12)
7ra 2

is the mean axial velocity at the plane z = 0. The coefficient C is given by

a(Po- PT)[J0()0 J1(2) _~)
C = ~V~ ~ J - - ~ + Jo(2~ " (13)

It remains to determine 2. By evaluating (11)1 and (11)2 at r = a and

The c a s e ~ ---- 0 leads to a solution in which t h e t r a n s m u r a l leakage is c o n s t a n t or has linear
variation w i t h axial distance. I t m a y be verified t h a t this situation, which was considered b y
Macey (1963 loc. cir.), is incompatible with (1) a n d so is excluded from the p r e s e n t study.
462 E.A. MARSHALL AND E. A. TROWBRIDGE
substituting the resulting expressions together with (13) into (10) it is found
that A is given inversely as a function of 7* by
Jl(~) Jl(~)

The function A(7*) has been computed and is shown in Figure I from which it is
seen that 0 < A < jo, 1 for 0 < 7* < ~ where Jo, 1 is the first zero of Jo(x),
i.e. Jo, 1 ~ 2.4048.

3"0

2"5 Asympr X (oo)~ 2.404

,,<

2-0
2(u
E
o
o
o.
f
I'C
o

0"5
(D

I { r [ ~ l I I f I f
0 I 2 3 4 5 6 7 8 9 I0 II

Permeabilify c o e f f i c i e n f , y*

Figure I. Graph showing separation parameter A as

function of the permeability coefficient ~*

The distributions of pressure and radial and axial velocity components in any
plane cross-section z = constant are conveniently expressed in the form

P - Pr A _/;~r\

izvr 1 (2tr /2tr\ [,~Jl(2t) ] (-~)t (15)

7*a(P(z) - PT) = 47*Jl(2t) t a J ~ + [ Jo(~t) 2 J1 '

vz 1 Xj-~v o~a - a J1 '

l?(z) = 4r,Jo(~ )
where

P(z) = 2 f:rp(r,z)dz,
(16)
2 f : rvz(r, z) dz = Q(z)
V(z) = -~ ~a 2.
 FLUID THROUGH A PERMEABLE TUBE 463
Graphs of the pressure and velocity distributions have been constructed b y
using (15) for the ease ~* = 0.09 ()~ = I). These are shown in Figure 2 to-
gether with distributions obtained from approximate equations obtained in a
following section of this paper. I t is seen that, for this value of ~,*, there is a
small discrepancy between approximate and exact solutions. The percentage
difference in these is greatest in the solutions for Vr at r = a, the approximate

2'r - - Exact solution

.... Approximate solution

Vz

1.6

P-PT
~(zI-p T
1.2

Q.
/,
O'B
"G /
/ / ~ /~Vr
:o
Q~
E

V I I I I \
0 0"2 0'4 0.6 0"8 1.0

rla

Figure 2. Cross-sectional distributions of pressure,

radial velocity and axial velocity ~* = 0.09 (h = 1)

solution exceeding that of (15)2 b y approximately 22 per cent. It is expected

that in many practical eases ~* could be much less than the above value and
the difference between the approximate and exact solutions would then be
insignificant. For example, in the application to the proximal tubule con-
sidered in Section 5 it is dedueed that y* _ 1.62 x 10 -7. In this case the eross-
sectional pressure and velocity distributions obtained from the approximate and
exact solutions are indistinguishable on plotting.

3. Equations for Mean Pressure and Flow. The distributions of the mean
464 E.A. M A R S H A L L A N D E. A. T R O W B R I D G E

cross-sectional pressure P(z), the axial flow Q(z), a n d the wall leakage flux q(z)
are readily determined using (11) and (16) together with the result t h a t
dQ(z)
q(z) = 21ravr(a, z) = dz (17)

I t is easily shown t h a t

P ( z ) - P r -- cosh Az
- ~1 sinh --,
)lz (18)
a a
Pc - Pr

Q(z) = cosh --~z _ C sinh _~z (19)

Qo a a
and

)tQ~
a q(z) = C cosh -~Za
- - sinh AZa, (20)

where C is given b y (13).

The right-hand sides of (18), (19) a n d (20) reveal t h a t the behavior of Q,
a n d q is strongly influenced b y the value of C. This behavior is conveniently
discussed b y considering the properties of the funtion Q(z) t h r o u g h o u t the
spectrum of values of C for positive z. For - c ~ < C < 0 it is seen from (19)
t h a t Q increases monotonically from Q = Q0 at z = 0 a n d Q --~ + co as z --> oo,
a n d at large z, Q is approximated to b y Q = ~Qo (1 + IC{)exp ()~z/a). The
particular case C = 0 also exhibits these properties since in this case Q =
Qo cosh )tz/a. F o r 0 < C < 1 it is seen t h a t Q decreases initially from the value
Q = Qo at z --- 0 to a m i n i m u m value of Qmin = Qo( 1 - c2) 112 when

a +Cl
z = ~log \1 - C]

a n d thereafter increases monotonically and Q --> + co as z --> co. I n this case,

for large z, Q ~ ~Q0(1 - C) exp ()tz/a). W h e n C = 1, Q = Qo exp ( - )tz/a) and
so Q decreases monotonically a n d Q-->o as z-->oo. For 1 < C < c~, Q
decreases monotonically from Q = Qo at z = o and becomes zero at
a [ c + It
z log

i.e. a stagnation point in the flow is predicted. F o r larger values of z, - Q

increases monotonically a n d Q - + - o o as z--> co being approximated b y
Q = - 1 Q o ( C - 1) exp (Az/a) for large z. Reverse flow is thus predicted in this
case. I n certain physical situations reverse flow m a y be inadmissible. In such
cases the solution in the z, C domain where Q < 0 would be inapplicable. A
 FLUID THROUGH A PERMEABLE TUBE 465

possible inference is that termination of flow may occur. Similar discussions

may be carried out for the behavior of q and P. In the case of the latter
quantity care is required in the consideration of the special case C = 0. In
the context of the present investigation C = 0 implies that Po = PT, and in
this case
ay*Jo(A)
t~Voj~(a ) [P(z) - Pr] = - sinh )lZa

and P(z) --> - m as z --> m.

Graphs of Q/Qo, [P(z) - TT][[Po -- PT] and [aq/)~Qo] for typical values of C
which illustrate these properties are shown in Figures 3-5. Similar curves have
been obtained, using an approximate theory, by Makiya et al. (1972) for the ease
of pure water transport. From the above discussion and from inspection of

1"0

0"8
dx .
0 O'E
Reverse lea.kag2.
o

:o
I ] ".i [ "-4 i
0 0"3 0"6 0"9"- 1"2 15 '~" ~ 1"8
Dimensionless axial distance \\ ~
Reverse f l o ~
-0"2 -- XZ \
a ",,Reverse flow

Figure 3. Variation of dimensionless volumetric axial flow

Q/Qo with dimensionless axial distance t~z/a

Figures 3-5 it is deduced that for the case of positive axial flow in a finite length
tube with nett positive leakage flux and non-stagnation flow, the value of C must be
approximately unity.
An interesting result is obtained from (18) and (19), since a consideration of
these equations shows that
dP
d--~ = -flQ' (21)

where
;~ Jl(;~)
fl = 7ra4y , j0(h )- (22)
466 E . A . MARSHALL AND E. A. TROWBRIDGE

~. ~ . o ~

' ~ B}-- \'%~'~. Increasing pressure

9 I \~-"-----...~ I~ _---'<dr~ with revers~
o~ I \ ~ flow
,,,
~

E 0"2 0"4 0"6 0"8 I'0 1"2 1"4 ~ 16 1'8

P Xz "-
~- -0"
Dimensionless axial distance,
a Reverse ""
pressure drop
and leakage

Figure 4. Variation of dimensionless mean cross-sectional pressure

(P - pr/Po Pr) with dimensionless axial distance ,~z/a
-

1 " 4 .~J/

I'2 /

oo
o" 0"~
o

C O'E

E
o
0"4
= .

o 0'3 0'6 0'9 1"2 "-~1- 5 IS

Dimensionless oxi01 disfonce, ),._~z
a Re.verse~..
-0.~ - le0kage

Figure 5. Variation of dimensionless wall leakage aqptQo with

dimensionless axial distance ~tz/a

E q u a t i o n (21) is analogous to Poiseuille's law relating axial flow and pressure

gradient in the classical case of laminar flow through a pipe with impermeable
walls. In t h a t case, however,
dP
dz = - flPQ' (23)
 FLUID THROUGH A PERMEABLE TUBE 467

where

~ = sg
~a 4
(24)

As a consequence of (23) and (24) an obvious rearrangement to (21) and (22)

is made giving
dP
d--~ = - r#.Q, (25)

where

F = )~ Jz()~)
8y* J0(~)' (26)

The q u a n t i t y F is shown plotted as a function of the wall permeability parameter

V* in Figure 6. I t is seen that, as expected, F --~ 1 as ~* -+ 0 a n d F -+ 0 as
y * --> oC).

[.-, ,0I
0'~

0'6
u

"O
0"4

m
0"2

1 2 3 4 5 @ 7 8 9 I0 II

Perrne~biH f y c o e f f i c i e n t , y*

Figure 6. Graph showing pressure drop parameter F as a

function of the permeability coefficient ~*

Equation (21) m a y be alternatively expressed in the form

P(z) - Po = -~,r~(z), (27)

z

where

Q(z) = z if Q(z*) dz*, (2s)

468 E. A. MARSHALL AND E. A. TROWBRIDGE

is the mean axial flow between plane cross-sections at the origin and some
generic value of the axial coordinate. Equation (27) is similar to a result
obtained by Maeey (1965, loc. cit.) except that there F = 1 and so this earlier
result is, as noted by Macey, only approximately true. In the context of the
present investigation the approximation is valid for vanishingly small wall
permeabilities.

4. P r e s s u r e a n d Velocity S o l u t i o n s f o r T u b e s w i t h S m a l l W a l l P e r m e a b i l i t i e s . In
developing solutions for small values of y* we note that J0()0, Jl()l) have the
series expansions
(-
go(~) = n=o (n!) 2 '
(29)
( - 1)n(l~) 2n+1
Jl(~) =
n=o (n + 1)!n! '

which are uniformly convergent in any finite domain of the complex plane.
The substitution of (29) into (14) and the expansion of the resulting expressions
in a series of ascending powers of 2 shows that

V$ ~- "16~'
( 1~2y 1 +~11~2 ..~ 0(~4)}, (30)

so that for vanishingly small )~

-- 4v~v*. (31)

Similar procedures show that, for small y* (i.e. small ~), (11) reduces to the
limiting form

YPo 2--G PT = 1+2v* _ 4V*a c o s h 4 ~ y * ~ - - - CLsinh4~V~y*a ,

v--Lr = 2%/ 2 -- C L cosh 4 ~ r ~* z - sinh , (32)

Vo a

= 2 1 - cosh ~%/~ - C~, sinh 4 V U * a ,

V--~m

where Cz is the limiting form taken by C and is given by

c,. = a(Po2 ~ v~)v~v*

o
- (33)
 F L U I D THROUGH A PERMEABLE T U B E 469

Likewise we find that results corresponding to (15) for the cross-sectional dis-
tribution of pressure and velocity in this limiting case are

P --PT = 1 -- 2~* 2 -- 1 ,
P(z) - Pr

(34)
~,*a[P(z) - Pr]

Equations (34)2 and (34)3 are of the same form as those derived by Friedlander
and Walser (loc. cir.) using the lubrication approximation, by Macey (1965,
loc. cir.), and by Kozinski et al. (loc. cir.). Distributions of cross-sectional
pressure and velocity constructed from (34) are shown in Figure 2.
Finally, from (26), (29) and (31), it is found that
F ~- 1 - la~* + O(y*2). (35)

5. Application to the Flow in the Proximal Renal Tubule. Micropuncture

studies of the proximal tubule of the rat (Giebisch et al., 1964; Gottschalk, 1961,
1962; Lassiter et al., 1961; Malnic et al., 1966; Ullrich et al., 1963; Windhager,
1968) have shown that the transfer of water and solutes such as sodium, glucose
and urea is such that the fluid osmolality remains constant. There is no evi-
dence to suggest that the renal interstitial hydrostatic pressure changes
significantly in distances along the proximal tubule which are a small multiple
of the tubule diameter. It therefore appears that the case considered in Section
2 where Pr is constant could apply to the proximal tubule and that the solutions
obtained above could provide a model for the reabsorption process in this
nephron segment.
In applying these solutions the data given by Gottschalk and Mylle (1956)
and used by Macey (1963, loc. cit.) are employed. Thus Q0 = 3 x 10 -7
cma/sec, a = 10 -3 cm and/x = 7 10 -3 dynes sec/cm 2 are taken as repre-
sentative parameter values for the rat kidney. It is noted that here Qo is the
single nephron glomerular filtration rate (SNGFR). A simple calculation using
this data shows that the diameter based Reynolds number _NRefor the flow is
approximately 3 x 10-2. For flows where 2YRois of this magnitude the inertial
forces are negligible and the use of the creeping flow equations (2) is justified.
Further, the total length of the proximal tubule (which includes both straight
and convoluted segments) is taken as 1 = 1 era. As assumed by Macey, in the
present investigation it is also considered that, in normal nephron function,
approximately 80 per cent of the glomerular filtrate is transferred through the
470 E. A. MARSHALL AND E. A. TROWBRIDGE
tubular epithelium. Thus z = 1 denotes the distal end of the proximal tubule
and Q(1) denotes the filtrate entering the descending limb of Henle's loop and we
have Q(t)/Qo = 0.2.
In order to test the accuracy with which the solutions presented here model
the phenomena in the proximal tubule it is necessary for the values of Po,/or
and ~* introduced in Section 2 to be known. Provided that these quantities,
and other parameters occurring in (33), are specified then the distributions of
mean pressure, axial flow and leakage flux can be constructed using (18), (19)
and (20) irrespective of the magnitudes of the/oarameters.
Pressure measurements performed recently by Allison et al. (1972) suggest
that, in the rat, the mean hydrostatic pressure at the entrance to the proximal
tubule is D 0 z 13 mm Hg (1.74 104 dynes/cm2). The q u a n t i t y p r is defined
in (8) and is seen to involve/oh the hydrostatic fluid pressure adjacent to the
exterior of the proximal tubule (i.e. the "interstitial" or "tissue" pressure) and
the osmotic pressures r z) and r on adjacent sides of the epithelium. Landis
and Pappenheimer (loc. cit.) quote an interstitial hydrostatic pressure of 10
m m Hg in the rat. However, Allison et al. remark that the value of this quan-
tity is uncertain and, moreover, that no satisfactory method exists, at present,
for its measurement. According to Pitts (1963) the osmotic concentrations of
the fluid on adjacent sides of the epithelium are equal and have a magnitude of
300 mOsm/1. For these concentrations r z) = 5100 m m Hg = r In view
of the large values for these osmotic pressures it is readily seen that very slight
differences of concentrations (e.g. 1 mOsm/1 across the epithelium could give
values of r z) - r which exceed or are comparable to, Pb. At present there-
fore exact values for PT seem to be uncertain. In fact it might be expected that/or
(which is considered to be uniform along the tube)might change in value depend-
ing on the physiological conditions. Values of the non-dimensional permeabi-
lity ~* for the tubular epithelium do not appear to be available.
I n view of lack of precise data, the available information has been used,
together with the theory presented here, to estimate probable values for ~* and
PT. I t is assumed a priori that ~* is sufficiently small for the approximate
solutions given in Section 4 to be applicable. Then, by substituting (31) and
(33) into (19) and evaluating the resulting expression at z = l, it is seen that
Q(1) cosh 4 t / ~ --/ a(Po - PT) ~/~-~'sinh 4%/~ 1 (36)
Qo = a 2,V~ a"
When the data given earlier in this section is substituted into (36) and D o - Pr
is expressed in mm Hg the transcendental equation
eosh 4 0 0 0 ~ / ~ - 1000(P o -/Or)V~-~* sinh 4000~/~ " - 0.2 = 0, (37)
is obtained for the determination of y* for a given value of D 0 - PT.
 FLUID THROUGH A PERMEABLE TUBE 471

By using (37), values of F* corresponding to various physiologically possible

assumed values of Po - Pr have been computed. The results of these calcula-
tions are displayed in Table I.

TABLE I
Values of y* Calculated from (37) for Various Assumed Values of/~o - Pr

Assumed Value of Po - Pr Corresponding Value

(ram Hg) o f ~*

1 0.99 x 10 - 6
2 2.45 x 10 -7
3 1.26 10 - 7
4 8.0 10 - 8
5 5.6 10 - 8
6 4.6 10 - 8
7 3.8 10 - 8
8 3.2 10 - 8
9 2.7 10 - s
10 2.4 10 - s
11 2.2 10 - s

An inspection of Table I shows that the values calculated for y* are sensitive
to the value of -P0 - Pr assumed for their calculation. However, in all the
cases considered here ~* is sufficiently small for the approximate solution of
Section 4 to be applicable. Some of the properties of the function Q(z) given by
(19) and used in the calculation of ~* have already been discussed in Section 3
where reasons were advanced for supposing that C -~ 1.
For definiteness it is assumed that, in normal nephron function, C = 1. The
flow profile resulting from this assumption is then of the same form as that
proposed empirically by Kelman (1962) and used by Macey (1965 loc. cir.). This
profile is the exponential distribution

Q = Qo exp ( - 4~/y-~ z).

This is a biologically important result since it provides a simple explanation, on

a mechanistic basis, for the possible occurrence of exponentially decreasing flow
and oftubulax balance. By substituting 0 = 1 in (19), evaluating the resulting
expression at z = t and using (33) and the above numerical data, it is found that
F* = 1.62 x 10-L For this value of F*, the pressure drop parameter F
[equations (26) and (35)] is such that 1 - F < 10 -s. From (33) the associated
472 E . A . MARSHALL AND E. A. T R O W B R I D G E

value of Po - PT is 2.48 mm Hg. This value is consistent with the data quoted
earlier in this section.
The variation of the transmural mean pressure difference along the tubule is
found from (18). B y inserting (33) together with the relevant data into this
expression it is found that

P(z) - PT = cosh z _ 1 sillh 4Y~F* (38)

Po - pT a l o o o ( P o - p )v y* a"
Again assuming that, in normal nephron function, C -- 1 so that Po - ~Dy
2.48 mm Hg and ~* = 1.62 x 10 .7 we have

P(z)-pr = 2.48exp - 1 . 6 1 x 10 - a ~ , (39)

and hence at z = l, P(l) - PT ---- 0.496 mm Hg and it follows that Po - P(1) --

1.984 mm Hg.
It has not been overlooked that the sensitivity of (19), which describes the
axial variation of the volumetric flow in the tubule, to the value of C m a y shed
some light on the nature of the delicate mechanism controlling the process of
fluid transfer across the epithelium. A consideration of (33) shows that, with
values prescribed for the other parameters, C increases due to an increase in
Po - Pr or ~,* or both. Thus from (19) (see also Figure 3) it is seen that an
increase in Po - / ~ f or ~* reduces the volume of filtrate entering the descending
portion of Henle's loop and can actually lead to the prediction of the termination
of the flow in this segment. Conversely a reduction of the value of -P0 - PT or
~,* causes the filtrate flow-rate into the descending portion of Henle's loop to
increase b y reducing the fluid transfer across the tubular epithelium. This
situation m a y be identified as a diuretic state. I t is not appropriate at this
juncture to speculate on the nature of a physiological mechanism by which
changes in Po - Pr or $* are initiated. I t should be emphasized that the
present investigation is concerned only with the process of fluid transfer across
the proximal tubule epithelium. The mechanism of the associated reabsorp-
tion process into the peritubular capillaries is a matter for further consideration
a simplified treatment of which has recently been attempted b y Deen et al.
(1973).

We are pleased to record our thanks to Dr. I. W. Henderson and his colleagues
in the Department of Zoology, University of Sheffield for several valuable dis-
cussions on the anatomical and physiological aspects of renal function. We are
grateful to Drs. G. Oster, R. A. Riemke and R. B. Kelman for a number of
helpful comments on the manuscript.
 FLUID THROUGH A PERMEABLE TUBE 473

APPENDIX
DERIVATION OF THE PRESSURE AND VELOCITY FIELD SOLUTIONS
The derivation of the pressure and velocity field solutions (11)i, 2. a is effected in the fol-
lowing manner. The elimination of vr and v z from (2)1 and (2)2 using (3) gives
~2p 1 ~p a2p
V2p _- ~-W + 7 }-7 + T ~ = o. (A.1)

A variables separable solution to (A.1) of the form

p(r, z) -- PT = R(r)Z(z), (A.2)
is sought. The physical concept t h a t the fluid flows through a tube in the direction of
decreasing pressure suggests that if the fluid flow is in the positive z direction then the
pressure should be non-periodic and decreasing as z increases. A solution satisfying this
requirement which is bounded as r -+ 0 is

p(r, z)--pT = (A cosh--Z+a B s i n h -~) J0(-~), (A.3)

in which A and B are integration constants, ~ is a separation parameter, and Jo is a Bessel

function of zero order.
The suhstitution for p from (A.3) into (2)2 gives
a2v~ l ~vz ~%z ~ { ~ Az'~j f;tr~
&-'--~+ r -~r + ~ - A sinh + B cosh a f ~ f" (A.4)

I t is required that vz is bounded at r = 0 and satisfies the condition

v~(a, z) = 0. (4)
Consideration of (A.4) shows that the only admissible variables separable form for vz
is
vz(r,z)= S(r){Asinh--aZ+a B c o s h - ~ } 9 (A.5)

By substituting (A.5) into (A.4) and (4) it is found t h a t S(r), which is required to be bounded
at r = 0, is a solution of the ordinary differential equation
d2S 1 dS ~2 h . I~tr\,

which satisfies the boundary condition

S(a) = O. (A.7)
A particular solution, SH, of the homogeneous equation
L{~} = 0,
is
SH(r) = Jo a " (A.8)

The general solution of (A.6) is then obtained b y substituting

8(r)= H(~-~)S~(r), (A.9)

where H is a function of r to be determined, into (A.6).

474 E..4. MARSHALL AND E. A. TROWBRIDGE
The first-order ordinary differential equation for H ' which results from this operation
is

dH" ~2J"o I_.~H,= a (A.10)

d(~r/a-"-"~+ ~,Jo + (,~r/a)] ~'
where the prime denotes differentiation with respect to the argument ,~r/a. By using the
results

f r dw 89 + J~(w)] + constant,
(A. 11)
Jl(to) = - s J0,

the solution
r r o( )1 dw
H = 4~ + ~ J tO[jo--~j doJ + D 1 (oJ~)(oJ-'--"~
D2, (A.12)
where D1, D2 are integration constants, is obtained.
I t is noted that
~,/a rJ'o(O,)] a ~ Ar Jx(Ar/a) AUr2
f
and that Sx'ta [dw]wd](w)] becomes unbounded as r --> 0. The use of these results to-
gether with (A.9) and (A.12) gives
a (~ (~r)a [Jl(~)l,l~r\~
S(r) = r, _ ij_. j.,oLz]p
where J1 is a Bessel function of first order. The axial velocity component vz is then found
by combining (A.5) and (A.14), and is

a r ~r Jl(~) Ar . Xz eosh ~ . (A.15)

By substituting v~ into (3) (Section 2) and using the results

f co2Jl(oJ) dw = 2wJl(oJ) -- to2do(tO) + constant,

(A.16)

f OJJo(w) dw = eoJl(r + constant,

it is found that

a At./At\ eosh-~ B sinh ~-), (A.1 7)

bhe integration constants from (A.16) having been set equal to zero since it is required that
v, is bounded at r = 0.
The constants A and B are evaluatmd by using (A.3) and (A.15) together with (5) and (6)
of Section 2. From (A.3) and (6) it is found that

A = ~(Po - p~) (AAS)

2JI(A)
 FLUID THROUGH A PERMEABLE TUBE 475

whilst from (A.15) and (5) we obtain

(2h2Q9)
,flu a
B -- (A.19)
)tJo(,~) Jl(h)l'7"7~,~, -

F o r brevity we denote

VO = Qo (12)
,fin--"~ ,

(13)
C -- 2/~Vo ~,gl(h) + Jo(h)
then, b y combining (A.3), (A.15), (A.17), (A.18), (A.19), (12), (13), equations (11)1.2,a of
Section 2 are recovered.

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RECEIVED 10.10-73