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MICROVASCULAR RESEARCH 39, 246-249 (1990)

Venular Vasomotion in the Bat Wing


I. P. TORRES FILHO

Departamento de Ci@ncias Fisiolbgicas, Instituto de Biologia, Universidade do Estado do Rio de


Janeiro, 20551, Rio de Janeiro, RJ, Brazil

Received June 19, 1989

INTRODUCTION
Spontaneous rhythmic constrictions and relaxations (vasomotion) have been
reported for small arteries and arterioles in several tissues (Funk et al., 1983;
Golantuoni et al., 1984; Slaaf et al., 1987; Meyer et al., 1988). This activity has
a frequency of about I-10 cycles/min and may last for long periods of time
(Funk and Intaglietta, 1983). The behavior of venules has been studied relatively
little in comparison with arteriolar and capillary vessels. Rhythmic diameter
changes of venous vessels are rarely reported in microvascular preparations.
However, vasomotion in the venules of the bat wing has been observed in all
species investigated (Nicoll and Webb, 1946; Mislin, 1978). Quantitative de-
scriptions of venomotion are scarce, incomplete, and mostly confined to vessels
larger than 60 pm (Wiedeman, 1959; D’Agrosa, 1970). In this study, we present
quantitative data on the vasomotion in the venous vasculature in the wing of
the intact, unanesthetized bat.

MATERIALS AND METHODS


The experimental data were obtained using Mexican free-tailed bats (Tadarida
brasiliensis maintained in a chronic colony in the laboratory. The
mexicana)
unanesthetized ,bat was enclosed in an acrylic box with one wing spread to the
outside and secured by cotton swabs. The areas of the wing between the third
and fourth or between fourth and fifth fingers were observed with a Leitz In-
travital microscope coupled with a TV camera (Panasonic WV1500N), monitor
(National WV5400BN), and videocassette recorder (Panasonic NV8200). For
transillumination, a tungsten-halogen light source (100 W) was used. Twenty-
eight venules from different areas were randomly chosen and filmed for at least
2 min at a final magnification of 1900 x . The room temperature was 20”-25”.
Diameter measurements were made by successive replays of the films using
a calibrated electronic caliper. Two electronically generated vertical video lines
were superimposed over the image of the internal margins of the vascular wall.
The analog output of the device was linearly related to the horizontal distance
between the lines. By manually keeping the lines over the vessel walls, vessel
diameter changes were continuously recorded in a polygraph (Hewlett-Packard
246
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BRIEF COMMUNICATIONS 247

7602). The number of active and inactive vessels was noted. For each active
venule, the frequency of the venomotion was determined by counting the number
of cycles over the entire period of recording. Spectrum analysis procedures were
not employed since a single frequency could be easily discerned by visual in-
spection of the tracings. The maximum, D,,,, and minimum, Dmin, diameters
were measured, from which we calculated mean diameter, D,,,, (D,,, + Dmin
/2) and absolute (D,,, - Dmin) and relative ((D,,, - Dmin/Dmean) x 100)
amplitudes.
Data are presented as means + 1 standard deviation (SD) and ranges. The
coefficient of variation (CV) was calculated as a percentage ((SD/mean) x 100)
and used as a measure of the variability for each parameter.
To assess the relations between different parameters, the experimental points
were fitted to linear functions by the least-squares method and the significance
of the correlation coefficient was tested. The minimum level of significance
considered was 5%.

RESULTS AND DISCUSSION


Vasomotion is a very prominent phenomenon in the venular network of the
bat wing. However, venules with diameter ranging from 10 to 25 pm did not
show active changes. This group of vessels composed 43% of the venules studied.
We observed that when two venules joined to form a larger one, their fre-
quencies could be similar but the movements were never synchronized. Different
portions of a single vessel usually showed various degress of vasomotion am-
plitude but we did not select any special segment for diameter measurement.
However, all vessels considered inactive did not show vasomotion in any portion.
Table 1 presents the descriptive statistics of venomotion parameters. Vaso-
motion frequency showed the smallest variability (CV = 11%). The variability
of absolute amplitude (CV = 76%) was about twice the variation for mean
diameter. Significant correlations were found for relative and absolute amplitudes
against mean diameter, as well as for relative against absolute amplitudes (Table
2). Logarithmic transformation of the data did not alter the results.
Some quantitative aspects of venular vasomotion presented here are strikingly
different from those reported for arterioles. Frequency has been reported to be
inversely related to mean diameter in arterioles of the hamster skin (Funk et
al., 1983; Colantuoni et al., 1984) and of the rabbit tenuissimus (Meyer et al.,
1988). We found no correlation for those variables (Table 2). Slaaf et al. (1987)
and Bouskela (1989) also reported no variation of frequency for arterioles with
different diameters.

TABLE 1
INFORMATION ON VASOMOTION PARAMETERS OF VENULES (N = 16)

Parameter Unit Means k SD Range

Mean diameter w 53.3 ‘- 21.1 27.7-92.0


Frequency min-’ 18.1 rt 2.0 14.5-21.4
Absolute amplitude pm 15.8 k 12.0 4.8-44.3
Relative amplitude % 27.3 2 12.7 13X-56.5
248 BRIEF COMMUNICATIONS

TABLE 2
LINEAR REGRESSION ANALYSIS OF VENOMOTION PARAMETERS (N = 16)

Correlation Significance
Parameters Intercept Slope coefficient level

Frequency x mean diameter 0.186 -0.01 I 0.12 0.33


Absolute amplitude x mean diameter -11.0 0.50 0.89 <0.001*
Relative amplitude x mean diameter 11.86 0.29 0.48 <0.05*
Absolute amplitude x frequency 50.96 -1.95 0.32 0.11
Relative amplitude x frequency 66.35 -2.16 0.33 - 0.10
Relative amplitude x absolute amplitude 13.86 -0.85 0.81 <0.001*

Regarding the relative amplitudes, large vessels usually present a less pro-
nounced vasomotion than terminal arterioles (Funk et al., 1983; Colantuoni et
al., 1984; Slaaf et al., 1987; Meyer et al., 1988). Conversely, a direct relation
between amplitude and mean diameter was encountered for the bat wing venules
in the present study. Our data are similar to those reported by Colantuoni et al.
(1984) who observed a direct correlation between vasomotion absolute amplitude
and mean arteriolar diameter.
It is instructive to compare our results with previously reported data on quan-
titative aspects of venomotion. Figure 1 presents data collected from different
bat preparations. Note that most previous data refer to vessels larger than 60
pm. One may conclude that venomotion frequency is independent of vessel
diameter. Vasomotion frequency in bat wing venules is usually faster than in
arterioles of the same tissue (Nicoll and Webb, 1946; Bouskela, 1989). It is
interesting to note that bat wing arterioles subjected to a transmural pressure
close to that found in the venous side do not display venomotion characteristics.
Instead, both arteriolar amplitude and frequency decreased (Bouskela, 1989).
However, venomotion is also influenced by temperature and intravascular pres-
sure (Wiederhielm, 1966) and by changes in chemical environment of vessel wall
(D’Agrosa, 1970).

- 24
- A h A n

l THIS WORK
6 DAii FiOi WIEDEMAN. 1959
A DATA FROM D’AGROSA 1970

I
0 40 160 200

FIG. 1. Vasomotion frequency for 69 bat wing venules with different diameters. Data from brown
bat (Myotis) are presented as open symbols.
BRIEF COMMUNICATIONS 249

In most microvascular preparations, vasomotion is not present on the venular


side (Colantuoni et al., 1984). It is possible that venomotion is a unique phe-
nomenon of the bat wing, which might help to return blood toward the heart
against centrifugal forces generated in flight (Kallen, 1978). However, cyclic
changes in venous tension are found in preparations using larger vessels like the
rat portal vein (Hermsmeyer, 1982). Venomotion may be a good model for
studying vasomotion characteristics, and more quantitative data are needed in
order to clarify the significance of vasomotion in venous vessels.

ACKNOWLEDGMENTS
We thank Dr. Eliete Bouskela for collaboration in the’project. Mr. Nelcir Moraes and Mr. Sergio
Bemardes helped in the care of the animals. This work was supported by CNPq.

REFERENCES
BOUSKELA, E. (1989). Vasomotion frequency and amplitude related to intraluminal pressure and
temperature in the wing of the intact, unanesthetized bat. Microvasc. Res., 37, 339-351.
COLANTUONI, A., BERTUGLIA, S., AND INTAGLIETTA, M. (1984). Quantitation of rhythmic diameter
changes in arterial microcirculation. Amer. J. Physiol. 246, H508-H517.
D’AGROSA, L. S. (1970). Patterns of venous vasomotion in the bat wing. Amer. J. Physiol. 218, 530-
535.
FUNK, W., ENDRICH, B., MESSMER, K., AND INTAGLIETTA, M. (1983). Spontaneous arteriolar vaso-
motion as a determinant of peripheral vascular resistance. Int. J. Microcirc. Clin. Exp. 2, 1 l-25.
FUNK, W., AND INTAGLIETTA, M. (1983). Spontaneously arteriolar vasomotion. Prog. Appl. Micro&c.
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vascular muscle. Physiologist 25, 454-465.
KALLEN, F. C. (1978). Overview of circulation in the wing membrane. Experientia 34, 1398-1400.
MEYER, J.-U., BORGSTROM, P., LINDBOM, L., AND INTAGLIETTA, M. (1988). Vasomotion patterns in
skeletal muscle arterioles during changes in arterial pressure. Microvasc. Res. 35, 193-203.
MISLIN, H. (1978). The active venous pulse in the wing circulation of bats (Chiroptera): A contribution
to comparative angiology. Experentia 34, 1391-1398.
NICOLL, P. A., AND WEBB, R. L. (1946). Blood circulation in the subcutaneous tissue of the living
bat’s wing. Ann. N. Y. Acad. Sci. 46, 697-708.
SLAAF, D. W., TANGELDER, G. J., TEIRLINK, H. R., AND RENEMAN, R. S. (1987). Arteriolar vasomotion
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WIEDEMAN, M. P. (1959). Response of subcutaneous vessels to venous distention. Circ. Res. 7,238-
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WIEDERHIELM, C. A. (1967). Effects of temperature and transmural pressure on contractile activity
of vascular smooth muscle. Bib. Amt. 9. 321-327.

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