relative contribution of the anatomical, diffusion, and distribution dead
space volume to the water shunt is difficult to assess. The size of the
anatomical dead space will be related to the arrangement of the fila
ments to each other and to the ventilation volume. The anatomical
dead space is probably larger at high water flow rates. The position of the
filaments is controlled by muscles at their base (Bitjel, 1949 ) which could
play a role in regulating the size of the anatomical dead space (Pasztor and
Kleerekoper, 1962 ). The author has observed movements of trout gills dur
ing normal breathing movements, and these may be caused by changing
water velocities or by the action of the muscles at the base of the filaments.
Whatever the cause, the effect of altering the relative position of filaments
to each other must be to change the size of the anatomical dead space,
which is probably very variable with time in a single fish, as well as be
tween different species of fish. There appears to be a tendency to enlarge
supporting structures and fuse parts of the gill in fish exposed to high ven
tilation volumes. In tuna, the secondary lamellae of adjacent filaments
are used to form a compact sievelike gill structure (Muir and Kendall,
1968 ). This presumably helps to maintain flow between the secondary
lamellae and reduces the size of the anatomical dead space, but it must
also increase the resistance to flow through the gills. Fusion of gill parts,
however, need not always be associated with the maintenance of flow be
tween secondary lamellae. Hughes (1966b ) has suggested that fusion in
Amia serves to prevent collapse of the gill sieve when the animal is in air.
Kylstra et al. (1967 ) have analyzed gas transfer in water-breathing
dogs and in the gills of fish. They related the size of the diffusion dead
space ( V D' d i ff02 ) in the gills of fishes to the rate of diffusion (D), the
time for diffusion (t), and the distance over which diffusion takes place
(a) and have derived the following equation which permits the evalua
tion of the diffusion dead space for oxygen at the gills (Kylstra et al.,
1967 ) :
1
.
VD' diffO,
1 3D.
t/a2
Va
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