s40
S27-2
Physiology of perfusion of the secondary circulatory system in fish.
Steffensen. J. F.
Marine Biological Laboratory, University of Copenhagen, DK-3000
Helsingar, Denmark.
Teleosts were believed to possess a lymphatic system similar to that of
mammalians until Vogel & Claviez (1981) via corrosion casts showed
connections between segmental arteries and the hitherto considered lymphatic
system. Since this system did not have any terminal vessels it was termed the
secondary vascular system, and the connections between this system and the
traditionally (primary) vascular system accordingly inter-arterial anastomoses.
In an in vivo morphological study the secondary system in glass catfish
(Cryptopferus bicirrhus) was described by Steffensen & Lomholt (1986).
Plasma skimming was observed to take place in the anastomoses and the
blood in the secondary system had a significantly lower haematocrit than in
the primary system. This has later been confirmed for a number of other
species. Later Steffensen & Lomholt (1992) suggested that the volume of the
secondary system was approximately 50 % larger than the primary system.
The physiological significance of the secondary system is still not clear, but
will be discussed. Among others it may play an important role in distributing
white blood cells to the skin in case of infections, it may be involved in
transporting lipids via the primary system from the intestine, or it may even
functionally work as a lymphatic system.
References:
Steffensen, J. F., Lomholt, J. P. and Vogel, W. 0. P. (1986). In vivo
observations on a specialized microvasculature, the primary and secondary
vessels in fishes. Acta Zool. 67; 193-200.
Steffensen, J. F. & Lomholt, J. P. (1992). The secondary vascular system.
In “Fish Physiology”. Eds.: Randall, D. J. & Farrell, A. P. Academic Press.
Vogel, W. 0. P. & Claviez, M. (1981). Vascular specialization in fish, but no
evidence for lymphatics. Z. Naturforsch. 36C; 490-492.
S27-3
Physiological role of the secondary circulatory system in fish
Heisler, N.‘, Ishimatsu, A.2 and Iwama, G.K.3
’Animal Physiology, Humboldt Universitit zu Berlin, Germany;
‘Univ. of Nagasaki, Japan; ’ Univ. of BC, Vancouver/Canada
Vessels of the secondary circulatory system (SCS) of fish originate
from primary arteries through capillary-sized anastomoses. They are
mainly located close to the body surfaces, often in juxtaposition to
mitochondria-rich cells. Accordingly, they have been suggested as
sites for ionic and acid-base regulation, a proposition supported by
recent studies on fluid composition and flow rate in SCS-vessels.
In rainbow trout exposed to environmental hypercapnia,
[HCO;] in the erythrocyte-poor fluid of the lateral cutaneous SCS
vessel (LCV) is elevated above the level of dorsal aortic (DA)
plasma by 2.2 mM as compared to control conditions, with the
increase being balanced by a reduction in [Cl-]. However, LCV
flow rate has been estimated to be only about 0.3 % of cardiac
output, so that even substantial elevations in LCV-[HCO; ] probably
cannot contribute significantly to the overall compensation.
The contribution of the branchial SCS is considerable on the
basis of a much larger flow rate. As estimated by indicator-
distribution, flow through the branchial vein (BV) as the collecting
SCS vessel of this area is about 7 % of cardiac output. With the
[HCO;] difference between DA and BV rising during hypercapnia
to about the same value as in the LCV, net transfer of bicarbonate
equivalents from the environment amounts to 3 mmol.kg” body
weight during 8 h of hypercapnia. These data indicate that more
than 90% of bicarbonate-equivalents gained from the environment
are transported through the secondary circulatory system. In contrast
to the LCV, the [HCO;] difference between DA and BV is not
balanced by any other ionic concentration change. These data
suggest an electrogenic hydrogen ion pump as the underlying
mechanism, with electroneutrality maintained by passive diffusion
through the epithelial integument related to the primary circulation.
s27-4
The role of pseudohranch and choroid rete for ocular
oxygen supply
Waser. W.P. & Heisler, N.
Dept. Animal Physiology, Humboldt Universitgt Berlin,
Germany
Mechanisms for ocular oxygen concentration, elevating
PO> in the eye above arterial levels, are found in many teleost
fish species. These mechanisms are considered crucial for
sufficient oxygen supply of non-vascularized retinae providing
unusually large diffusion distances. Liberation of oxygen in the
choroid rete mirabile, the ocular counter-current capillary
exchange system, is considered to be performed similar to the
rete mirabife of the swimbladder. In contrast to the
swimbladder, however, the choroid rete mirabile is supplied
with blood only in series with the pseudobranch, a reduced
mandibular gill arch. This close association led Miiller in 1839
to suggest that the pseudobranch may have a role related to
vision.
This allusion was followed up by separating functions of
pseudobranch and choroid rete in isolated preparations. For this
purpose perfusion rate of the system was determined in vivo by
application of a special pulsed Doppler technique to the
pseudobranchial artery and direct calibration of the velocity
signal to flow in situ (745 c 282 ~1 min“ kg-‘). Applying the
obtained blood flow rates to isolated eye preparations,
intraocular I’01 profiles were determined by use of oxygen
microelectrodes. Changes in ocular PO* were related to
electroretinograms as the expression of varied retinal metabolic
activity, and pseudobranch function was simulated by changes
of the perfusing blood. The obtained data support the notion
that the pseudobranch may play a role in preconditioning the
perfusate for liberation of oxygen in the eye.
s27-5
Acid-base relevant metabolism of the pseudobranch
Berenbrink M
Dept. Animal Physiology, Humboldt UniversitLt zu Berlin,
Germany
An isolated saline-perfused preparation of the pseudobranch from
rainbow trout (Uncorb~nchus mykiss) was utilized in order to
investigate its biochemical characteristics and physiological
functions. The experimental arrangement allowed for
determination of glucose and oxygen consumption, as well as of
lactic acid and carbon dioxide production as a function of
perfusate pH. Changes in perfusate pH, induced by variations in
Pco,, provoked significant changes in the rates of carbon dioxide
and lactic acid production. As a consequence, shifts in pH of the
perfusate entering the pseudobranch were not directly conducted
to the perfusate leaving the pseudobranch. Additional data
obtained on the blood-perfused pseudobranch confirm the notion
that metabolic carbon dioxide production is an important factor in
determining the acid-base status of blood leaving the
pseudobranch. Accordingly, this tissue may have a pH-regulatory
function, effectively uncoupling the post-pseudobranchial
circulation from extensive pH-changes frequently encountered in
the pre-pseudobranchial, arterial circulation. Since blood leaving
the pseudobranch is the exclusive blood supply to the choroid
rete ruirahik of the fish eye, this mechanism may have a
protective role for the sensitive pH-dependent counter-current
oxygen concentrating mechanism associated with the choroid rev
mirrrhilr.