186 Specialia ]~XPERIENTIA32/2

crushing, n o t o n a b s o l u t e g r o u n d s (per g tissue), b u t o n a

r e l a t i v e basis (expressed as t h e r a t i o of free g l u t a m a t e /
0.25 -- ?
t o t a l free a m i n o acid c o n c e n t r a t i o n s ) . P e r h a p s , h o w e v e r ,
a s i g n i f i c a n t free g l u t a m a t e c o n c e n t r a t i o n decrease could
b e d e t e c t e d a t t i m e s earlier t h a n 25 d a y s if a v e r y large
s a m p l e size was used w i t h d a t a e x p r e s s e d p e r g tissue

I
0.20--
only. A l t h o u g h t h i s m a y s u g g e s t t h a t t h e r e is a decrease
7' in free g l u t a m a t e in t h e dorsal s e n s o r y n e u r o n following
? iiljufy, some a d d i t i o n a l difficulties m u s t be c o n s i d e r e d

o.i~-o,io_ ? _L,
_L
0,05 --
X • ~0( ~OC( )O0(
0
DR1 DR,?. I}R3 DRI DP,2 DR3 DR1 DR2 DR3 DR1 DR2 DP~
2 DAY 4 DAY 10 ~AY
Fig. 3. The free glutamate/total free amino acid concentration ratio
is shown on the Y-axis. Each tissue is shown at 2, 4, 10 and 25-day
periods following crushing (between DR1 and DR2 as in Figure 1)
on the X-axis. The arrow points to the control value glutamate/free
amino acid concentration ratio obtained in each particular time peri-
od. The bar for each tissue on the X-axis represents the glutamate/free
amino acid concentration ratio, and the line above this is the standard
deviation. The significance of the difference between control and
crush tissue was tested by the Student's t-test. The number of X's
at the base of each bar represents the level of significance of this
difference; X = 5%, XX = 2.5% and XXX = 0.5% level of
significance.
I
I before t h i s conclusion is reliable. D i s t a l t o t h e i n j u r y t h e r e
i s c o m p l e t e d e g e n e r a t i o n of t h e s e p a r a t e d a x o n s 1~ w i t h
p h a g o c y t i c satellite cells b r e a k i n g d o w n a x o n a l d e b r i s is.
T h e p r o t e o l y t i c a c t i v i t y of t h e satellite cells m a y cause
free g l u t a m a t e increases w h i c h cancel a n y d e c r e a s e d
a x o n a l g l u t a m a t e w h e n m e a s u r e d o n a tissue basis.
P r o x i m a l t o t h e i n j u r y , t h e i n t a c t a x o n s r e g r o w in a n
a t t e m p t t o r e i n n e r v a t e t h e e n d o r g a n lL I t is n o t y e t
k n o w n w h e t h e r t h e t o t a l free a m i n o acid increase ob-
s e r v e d p r o x i m a l l y here is n e u r o n a l or e x t r a n e u r o n a l in
n a t u r e , a l t h o u g h free g l u t a m a t e is d e c r e a s i n g r e l a t i v e t o
t o t a l free a m i n o acids a t a g r e a t d i s t a n c e p r o x i m a l to t h e
crush. Therefore, t h e results of t h i s study~, a n d t h e f a c t
t h a t g l u t a m a t e is s u c h a n a c t i v e s u b s t a n c e of w i d e s p r e a d
25 DAY i m p o r t a n c e 6, i n d i c a t e t h a t one m u s t a s k t h e following
t h r e e q u e s t i o n s c o n c e r n i n g s u c h d a t a . 1. H o w i m p o r t a n t
are s u c h o b s e r v a t i o n s in Showing w h e t h e r or n o t g l u t a m a t e
is a s y n a p t i c t r a n s m i t t e r ? 2. H o w specific are s u c h c r u s h
effects o n free g l u t a m a t e levels w h e r e one is r e a s o n a b l y
sure it is a s y n a p t i c t r a n s m i t t e r as o p p o s e d t o w h e r e i t is
n o t a t r a n s m i t t e r ? 3. W h a t a m i n o acid c h a n g e s following
a x o n a l c r u s h or section are associated w i t h t h e a x o n s a n d
w h i c h are associated w i t h satellite cells ?
IT j. Z. YouNG, Adv. Surg. 1, 165 (1949).
18 M. SINGER and M. C. STEINBERG,Am. J. Anat. 133, 51 (1972).

Long-Chain (Z)-9-Alkenes are 'Psychedelics' to Houseflies with Regard to Visually Stimulated

Sex Attraction and Aggregation

I. lP~ICHTER, H. KRAIN a n d H. K. MANGOLD

Bundesanstalt /fir FeH/orschung, Piusallee 68-76, D-44 Miimter (German Federal Republic, BRD), 28 July 1975.

Summary. T h r e e d i f f e r e n t t e s t s on houseflies (Musca domestics L.) r e v e a l e d t h a t b o t h p h e r o m o n e - f r e e pseudoflies a n d

m a l e p a r t n e r flies e x h i b i t in t h e p r e s e n c e of m i x t u r e s of l o n g - c h a i n (Z)-9-alkenes or p u r e (Z)-9-tricosene e n h a n c e d
releasing effects for t w o o p t i c a l cues, w h i c h s t i m u l a t e m a l e h o u s e f l i e s t o m a t i n g s t r i k e s a n d houseflies of b o t h sexes
to aggregation.

B e h a v i o r a l responses of l o n g - c h M n (Z)-9-alkenes in t h e dishes2,a, 1~ in w h i c h one p s e u d o f l y is e x p o s e d t o 2 m a l e

m a l e housefly, Musca domestics L. a r e b e l i e v e d t o b e
sex a t t r a c t i o n l-a, sex s t i m u l a t i o n 8, e x c i t e m e n t , m a t i n g
a n d o r i e n t a t i o n ~. F u r t h e r m o r e , a g g r e g a t i o n of b o t h sexes
of houseflies is a t t r i b u t e d t o (Z)-9-tricoseneb,% Several
h o m o l o g o u s c o m p o u n d s show p h e r o m o n e a c t i v i t y 3,*,
especially a 7 : 3 m i x t u r e of (Z)-9-tricosene a n d (Z)-9-
h e n e i c o s e n e 4. A l t h o u g h it is well k n o w n t h a t v i s u a l
s t i m u l i elicit s t r o n g sex a t t r a c t i o n ~ a n d a g g r e g a t i o n s,
the mutual interaction between visual and olfactory
s t i m u l i in t h e h o u s e f l y h a s n o t b e e n e s t a b l i s h e d 9.
I n t h e p r e s e n t s t u d y , c o m b i n a t i o n s of several (Z)-9-
alkenes were t e s t e d t o e l u c i d a t e t h e i r effect o n t h e be-
h a v i o r of houseflies in c o n j u n c t i o n w i t h v i s u a l stimuli.
3 t y p e s of t e s t s were carried out, e a c h ill t h e a b s e n c e of
p h e r o m o n e s ( ' s t a n d a r d s ' ) a n d in tile p r e s e n c e of p h e r o -
m o n e s : 1. T h e well e s t a b l i s h e d p s e u d o f l y t e s t in P e t r i
houseflies; 2. a m o d i f i e d p s e u d o f l y t e s t in P e t r i dishes,
in w h i c h two pseudoflies are exposed t o 1 m a l e h o u s e f l y ;
a n d 3. a large-scale t e s t u s i n g p s e u d o f l y - f l y p a p e r strips.
I n 2. as well as in 3. pseudoflies c o n t a i n i n g n o p h e r o -
m o n e s were e m p l o y e d as ' c o n t r o l s ' in t h e p r e s e n c e of
pseudoflies i m p r e g n a t e d w i t h p h e r o m o n e s .
Material and methods. T h e t e s t m a t e r i a l s (I-VI), as
s h o w n in T a b l e I, c o n t a i n e d v a r i o u s levels of biologically
a c t i v e h y d r o c a r b o n s , s u c h as (Z)-9-heneicosene a,4, (Z)-
9-docosene 3, ~, (Z)-9-tricosene l-e, (Z) - 9 - t e t r a c o s e n e 8, 4, a n d
(Z)-9-pentacosene ~.
1. T h e s h o r t r a n g e a c t i o n of l o n g - c h a i n (Z)-9-alkenes
o n t h e sex a t t r a c t i o n of m a l e houseflies was s t u d i e d
u s i n g t h e p s e u d o f l y t e s t in P e t r i dishes 2,3,1~ Covered
P e t r i dishes, 9 c m in d i a m e t e r , e a c h w i t h 2 m a l e h o u s e -
flies, 5 t o 10 d old, f r o m l a b o r a t o r y c u l t u r e a n d 1 ' p s e u d o -
15.2. 1976 Specialia 187

fly' ( k n o t m a d e f r o m b l a c k shoelace), p l a c e d in t h e center, flies was i n v e s t i g a t e d u s i n g ' p s e u d o f l y - f l y p a p e r ' s t r i p s

were a r r a n g e d o n a t a b l e in 3 groups. Tile t e s t s were car- as t r a p s . I n t h i s t e s t s y s t e m t h e pseudoflies w e r e p r i n t e d
ried o u t a t 25~ b e t w e e n 14.00 a n d 16.00 h u n d e r b r i g h t o n Aeroxon| f l y p a p e r s t r i p s (4 • 80 era} in t h e f o r m of
d a y l i g h t a n d e a c h fly was u s e d for a single t e s t only. A 300 b l a c k spots of 5 m m d i a m e t e r , p e r strip. F o r assays,
g r o u p of 3 o b s e r v e r s c o u n t e d , e a c h a t 6 dishes in a n y b a t c h e s of 5,000 t o 12,500 houseflies f r o m l a b o r a t o r y
single 30 m i n session, t h e n u m b e r of m a t i n g s t r i k e s f r o m c u l t u r e s were k e p t o n a s u g a r plus w a t e r d i e t in a 142 m a
one fly t o t h e a c c o m p a n y i n g fly as well as t o t h e p s e u d o - t e s t r o o m w i t h 4 windows. W h e n t h e flies were 5 t o 7 d a y s
fly, w h i c h was p r e t r e a t e d w i t h t h e t e s t m a t e r i a l . old, 10 p s e u d o f l y - f l y p a p e r s t r i p s were h u n g f r o m t h e
2. T h e e x t e n t of s h o r t r a n g e o r i e n t a t i o n of m a l e h o u s e - ceiling (3 m high), s e p a r a t e d b y 1 m f r o m e a c h other. I n
flies t o o l f a c t o r y s t i m u l i b y l o n g - c h a i n (Z)-9-alkenes e a c h t e s t series, 10 t r a p s were exposed once t o t h e flies
was a s s a y e d ill a m o d i f i e d p s e u d o f l y t e s t in P e t r i dishes. b e t w e e n 10.00 a n d 16.00 h a t a b o u t 23~ E a c h g r o u p of
Male houseflies were' t e s t e d i n d i v i d u a l l y ill P e t r i dishes, flies was t a k e n for 2 c o n s e c u t i v e t e s t s only, I n t h e a b -
9 c m in d i a m e t e r , c o n t a i n i n g 2 k n o t s m a d e f r o m b l a c k sence o f (Z)-9-alkenes, 10 s t a n d a r d series were r u n in
shoelace, w h i c h were s e p a r a t e d b y 5 c m f r o m e a c h o t h e r . o r d e r to d e t e r m i n e t h e b a s i c a v e r a g e p e r c e n t a g e s of
I n h a l f of t h e dishes, b o t h pseudoflies were t r e a t e d w i t h c a p t u r e s a t e a c h of t h e t e n t r a p p o s i t i o n s . T h e s e q u e n c e
10 ~1 of h e x a n e a n d t h e s o l v e n t was allowed t o e v a p o r a t e . of p s e u d o f l y - f l y p a p e r strips, i m p r e g n a t e d w i t h t h e dif-
Ill e a c h of t h e r e m a i n i n g dishes, one k n o t was t r e a t e d f e r e n t m i x t u r e s of (Z)-9-alkenes a n d of c o n t r o l s t r i p s
w i t h 10 ~1 p u r e h e x a n e a n d t h e o t h e r w i t h 100 vg of i n v e s t i g a t e d s i m u l t a n e o u s l y , was c h a n g e d in all four t e s t
(Z)-9-tricosene ~2 d i s s o l v e d in 10 ~1 h e x a n e . A f t e r eva- series.
p o r a t i o n of t h e s o l v e n t , t h e t e s t s were c a r r i e d o u t a t Results. 1. T h e d a t a o b t a i n e d in t h e t e s t s w i t h p s e u d o -
28~ u n d e r t h e s a m e c o n d i t i o n s as above. T h e n u m b e r f l y - P e t r i dishes, as g i v e n in T a b l e I I , show t h a t in s t a n -
of m a t i n g s t r i k e s t o 'left' a n d ' r i g h t ' pseudoflies was d a r d s t h e a v e r a g e n u m b e r of h o m o s e x u a I m a t i n g s t r i k e s
c o u n t e d s e p a r a t e l y b y a g r o u p of 4 observers, e a c h a t was 2 t i m e s h i g h e r t h a n c o p u l a t i v e j u m p s o n t h e p s e u d o -
6 dishes in a n y single 30 mill session. fly. I n t h e p r e s e n c e of l o n g - c h a i n {Z)-9-alkenes, h o w e v e r ,
3. T h e influence of l o n g - c h a i n (Z)-9-alkenes o n t h e t h e n u m b e r of m a t i n g strikes o n t o m a l e flies was in-
a g g r e g a t i o n b e h a v i o u r of m a l e a n d female (1:1) h o u s e - creased 2 t o 3.6 t i m e s a n d o n t o pseudoflies 4.4 t o 6.6
times. T h u s , t h e t o t a l n u m b e r of m a t i n g s t r i k e s b e t w e e n
fly-fly a n d f l y - p s e u d o f l y r e a c h e d t h e s a m e a b s o l u t e level.
2. I n t h e m o d i f i e d v e r s i o n of t h e t e s t s ill p s e u d o f l y -
Table I. Composition of mixtures of the (Z)-9-alkenes tested P e t r i dishes, 192 m a l e s were a s s a y e d i n d i v i d u a l l y for
s t a n d a r d s , b o t h pseudoflies ill e a c h dish b e i n g free of
Test Percentagesof(Z)-9-alkenes (Z)-9-alkenes. T h e t o t a l n u m b e r of m a t i n g s t r i k e s t o
material. 'left' a n d ' r i g h t ' pseudoflies was 600 a n d 535, respectively.
C16 el? C18 Cl~ ego C21 C22 C2a C24 C25 T h e d o u b l e d u m m y t e s t s o n 192 m a l e flies, e a c h ill P e t r i
dishes c o n t a i n i n g one pseudofly, i m p r e g n a t e d w i t h
100 ~g of (Z)-9-tricosene 12, a n d t h e second p s e u d o f l y for
I 2 1.5 2.5 88 6
control, were r u n s i m u l t a n e o u s l y . T h e t o t a l n u m b e r of
II 0.5 0.5 54 38 7
III 1.5 97 1.5 c o p u l a t i v e strikes was 2,043 o n k n o t s t r e a t e d w i t h (Z)-
IV 4 22 11 59 4 9-tricosene, a n d 1,236 o n k n o t s for control. T h u s , t h e
V 0.5 1 60 33 5.5 n u m b e r of m a t i n g strikes c o m p a r e d t o s t a n d a r d s , r u n in
VI Trace 3 96 1 t h e a b s e n c e of (Z)-9-tricosene, w a s i n c r e a s e d 3.6 a n d
2.2 times, respectively. I n one f o u r t h of all t e s t s (48
males) carried o u t w i t h (Z)-9-tricosene, t h e n u m b e r o f
"Prepared from the seed oils of Brassicc~ napus (I and IV), Sim-
mondsia chinensis (II and V) and Tropaeo[um mains (III and VI) 1I. copulative strikes on the pseudofly impregnated with
(Z)-9-tricosene (606 j u m p s ) was a l m o s t e q u a l t o t h a t o n
t h e c o n t r o l d u m m y (575 j u m p s ) ,
3. All p s e u d o f l y - f l y p a p e r s t r i p s a g g r e g a t e d b o t h sexes
Table II. Effect of long-chain (Z)-9-alkenes on male houseflies in of houseflies, m a l e s p r e d o m i n a n t l y a t t h e u p p e r p a r t a n d
tests in pseudofly-Petri dishes" females p r e d o m i n a n t l y a t t h e lower p a r t . T a b l e I I I shows
t h a t in e a c h case t h e n u m b e r of m a l e s t r a p p e d w a s 2.7

Test materiaD Number of mating strikes o on

Male fly Pseudofly 1 D. A. CARLSOIq, ~[. S. MAYER, D. L. SILHACEK, J. D. JAMES, M.

BEROZAand B. A. BIERL, Science 774, 76 (1971).
Standard 268 107 2 W. M. ROGOFF, G. H. GEETZ, M. JACOBSO~r and M. BEROZA,
I 542 (2.0) 526 (5.0) Ann. entom. Soc. Am. 66, 739 (1973).
IV 657 (2.5) 701 {6.6) 3 D. A. CARLSON', R. E. DOOLITTLE, ~t~. BEROZA, W. IV[. ROGOFF
and G. H. GRETZ, J. agric. Food Chem. 22, 194 (1974).
Standard 363 149 4 A. MA~SINGH, R. W . STEELE, B. N. SMALLMAN, 0. MERESZ and
i~ 816 (2.2) sos (5.4) C. MOZOGAI,Can. Entom. 104, 1963 (1972).
V 774 (2.1) 652 (4.4) 5 D. A. CARLSON"and M. BEROZA, Envir. Entom. 2, 555 (1973),
Standard 189 130 6 p. B. ~/[ORGAN, I. I-I. GILBERT and R. L. ]~YE, Fla. Entom. 57,
Iii 684 (3.6) 693 (5.3) 136 (1974).
VI 597 (3.1) 501 (3.9) 7 G. VOGEL, Z. Tierpsychol. 14, 309 (1957).
8 R. WIESMANX, lVfitt. Schweiz. entom. Ges. 33, 121 (1960).
9 I. RICHTER, Naturwissellschaften 61, 365 (1974).
"Tests according to=,a, 1~ bl0 [xl hexane (for standards) or 100 p.g lO W. M. ROGOFF, A. D. BELTZ, J. O. JOllNSE~ and F. W. PLACe,
(Z)-9-alkenes dissolved ill 10 ~1 hexane were applied to each pseudo- J. Insect. Physiol. 10, 239 (1964).
fly and the solvent was allowed to evaporate, eCounted for 30 min 11 H. K. MAgOOLD, H. BECKER and I. RICHTER, in preparation.
using 168 male flies; in parentheses, times increased compared to 1= I. RlCllTER and H. K. I~AN'GOLD, Chem. Phys. Lipids 11, 210
standards tested simultaneously. (1973).
188 Specialia EXPERIENTIA32/2

Table III. Effect of long-chain (Z)-9-alkenes on both sexes (1 : 1) of orientation of male houseflies to the source of odor in
houseflies in ~ests with pseudofly-flypaper strips t h e modified pseudofly tests using P e t r i dishes, a l t h o u g h
i t is well known t h a t houseflies p a y a t t e n t i o n to n e a r l y
Test- No. of TotM number Of Ratio Times increase a n y odor at a short distance s. H o w e v e r , in t h e presence
m/~te~alb traps flies captured (male/female) ofcaptures of long-chain (Z)-9-alkenes the optical stimuli of sex
(%) a t t r a c t i o n b y u n t r e a t e d pseudoflies (controls) exerted
an enhanced releasing effect, c o m p a r e d to those in
Standards 100 4248 2.7 s t a n d a r d tests run in the absence of pheromones.
The c o n v e n t i o n a l tests w i t h pseudofly-Petri dishes
Controls 16 1312 3.0 128
I" 4 467 5.7 2.4 also showed unequivocally, t h a t in t h e presence of long-
II 4 679 3.3 2,5 chain (Z)-9-alkenes the optical stimuli of sex a t t r a c t i o n
HI 4 442 2.7 2.4 b y male p a r t n e r flies were enhanced, c o m p a r e d to those
IV 4 398 3.5 2.5 in t h e s t a n d a r d tests which were run in the absence of
V. 4 462 3.4 1.7 pheromones. Similar observations h a v e been m a d e using
VI 4 657 3.9 2.7 benzene e x t r a c t s of female houseflies 1~
Similarly, in the tests using pseudofly-flypaper strips,
~Tests on 55000:houseflies (total) at 22 to 24~ in 14 six-hour test tile visual stimuli for aggregation p r o v i d e d b y the un-
series using 10 flypaper strips, each, wit-h 300 black spots (5 mm fg). t r e a t e d strips (controls) were p o t e n t i a t e d in the presence
b2.5 ml hexane, for standards (10 test series) or 125 mg (Z)-9-alkenes of high dosages of long-chain (Z)-9-alkenes c o m p a r e d to
dissolved in 2.5 ml hexane, for tests (4 test series) and 2;5 ml hexane, standards run in the absence of pheromones.
for control strips ruI1 Simultaneously, were applied to each trap and On the basis of all these findings, it is concluded t h a t
the solvent was allowed to evaporate. the threshold for optical cues s t i m u l a t i n g houseflies to
sex a t t r a c t i o n or aggregation is decreased in the presence
of long-chain (Z)-9-alkenes in the test systems. Thus,
to 5.7 times higher t h a n t h a t of femMes, t h o u g h in t h e long-chain (Z)-9-alkenes c a n be considered 'psychedelics'
tGSt r o o m t h e s~arting ratios of t h e sexes was I : I. I n t h e to houseflies wi• regard ~o visually stimnlated sex at-
presence of 10ng-chain (Z)-9-alkenes, b o t h traps impreg- t r a c t i o n and aggregation. The state of e x p a n d e d sensoric
n a t e d w i t h t h e s e : c o m p o u n d s and those not i m p r e g n a t e d susceptibility is a characteristic effect of psychedelic
c a u g h t m o r e flies c o m p a r e d t o standards run in the drugs.
absence of the u n s a t u r a t e d hydrocarbons, The efficiency Sex a t t r a c t i o n of male houseflies b y m o v i n g pseudo-
of the pseudofly-flypaper strips loaded w i t h test m a t e r i a l flies is n o t changed in the presence of long-chain (Z)-9-
V, which contained the smallest proportion of biologi- alkenes 9, because t h e releasing effect of such fast m o v i n g
cally active (Z):~9-alkenes 3,4 w a s increased b y a factor d u m m i e s is greatly increased c o m p a r e d to static female
of 1.7, whereas t h a t of the controls, run simultaneously, fly modelsL I n tests using plexiglas olfactometers 1,8,4
w a s increased b y a factor o f 1.8. The percentages of or o t h e r types of traps 5, ~ for the assay of pheromones on
c a p t u r e s of all o t h e r traps i m p r e g n a t e d w i t h (Z)-9- houseflies, visual stimuli m a y h a v e beert i n v o l v e d to
alkenes w e r e increased 2.4 to 2.7 times. v a r y i n g extents.
The results obtained in tests using pseudofly-flypapers
Discusslo:n. The results o b t a i n e d w i t h houseflies show show poor captures c o m p a r e d to t h e t o t a l n u m b e r of
t h a t a w i d e s p e c t r u m of long-chain (Z)-9-alkenes, in flies present, a l t h o u g h b o t h olfactory and visual stimuli
v a r y i n g c o m b i n a t i o n s , exhibit, b o t h q u a l i t a t i v e l y and were provided. Therefore, the use of long-chain (Z)-9-
q u a n t i t a t i v e l y ; the same b e h a v i o r a l responses, depending alkenes or other olfactory stimuli s seems to be of minor
u p o n t h e different visual signals in the tests using Value for control of the housefly.
p:seUdofly-Petri dishes and pseudofly-flypaper strips.
R e c e n t l y , (Z)'9'tri60sene has been characterized as a
short r a n g e sex p h e r o m o n e 1~. I n t h e present s t u d y (Z)- 13 p. A. LANGLEY, R. W. PIMLEY and D. A. CARLSON,Nature,
9-tricosene did not give clear evidence for short range Lond. 254, 51 (1975).

Sodium Pumps and Galactose Transfer in the Short-Circuited Small Intestine

J~ F. UI)E and R. J. C. BARRX~
Department o/Physiology, University o/Ibadan, Ibadan (Nigeria), and Department o/Physiology, University o/Sheffield,
Western Bank, She/field $10 2 T N (England), 10 June ~975.
Summary. I n a short-circuited p r e p a r a t i o n of r a t j e j u n u m , there are two sodium pumps, one etectrogenic and t h e other
neutral, W h e n energy sources are limited, t h e t o t a l sodium transfer is limited. I n t h e presence of a n o n - m e t a b o l i z e d
a c t i v e l y transferred hexos e, the electrogenic p u m p is preferentially used. The neutral sodium p u m p is only able to
function when additional energy is availab!e.

The absorption of Na and certain hexoses b y the small
intestine are closely linked 1-8. The situation in the r a t
j e j u n u m in v i t r o i s complicated b y there being two dis-
t i n c t N a pumps, one which is electrogenic and t h e other
which is non-electr0genie ~, 9. N a transferred b y the elec-
trogenic m e c h a n i s m g i v e s rise ~o a t r a n s m u r a l p o t e n t i a l
difference: and is linked iI1 p a r t to active hexose transfer,
while t h a t t_ransferred b y t h e non-electrogenic m e c h a n i s m
is linked to hexose m e t a b o l i s m and causes all increase in
fluid absorption. The c a p a c i t y of t h e j e j u n u m to transfer
N a and the relationship between these two t y p e s of Na
p u m p has been investigated using a short-circuiting
technique.
Materials and methods. W h i t e male rats (220-250 g)
were anaesthetised w i t h i.p. p e n t o b a r b i t o n e sodium
(Nembutal). The ~mal! intestine w~ts r e m o v e d and