Wetlands Ecology and Management 9: 211–218, 2001.
© 2001 Kluwer Academic Publishers. Printed in the Netherlands.
211
Gas space and oxygen exchange in roots of Avicennia marina (Forssk.)
Vierh. var. australasica (Walp.) Moldenke ex N. C. Duke, the Grey
Mangrove
William G. Allaway, Mark Curran, Lauren M. Hollington, Malcolm C. Ricketts & Nich-
olas J. Skelton
School of Biological Sciences A12, The University of Sydney, NSW 2006, Australia, Fax: +61 2 9351 4771; E-mail:
allaway@bio.usyd.edu.au
Key words: aerenchyma, diffusion, lysigenous, pressure, schizogenous, subrisule
Abstract
Aerenchyma pervades the cortex of the whole root system in Avicennia marina var. australasica plants grown for 3
to 5 years in artificial-tidal tanks. The gas spaces appear to be schizogenous in all but the finest roots, where some
apparently lysigenous gas spaces develop. Gas spaces in the pith are small. Pneumatophores examined at the time
the tips are growing possess subrisules on the tips as well as lenticels on mature regions. At other times subrisules
are not seen. When the tide floods the pneumatophores, gas pressure and oxygen concentration go down – pressure
by 1.7 kPa or less, but oxygen by as much as 3 mol m
-3
, or more in some cases. On draining, pressure recovers
immediately to atmospheric, but oxygen slowly rises to a plateau below the concentration in the atmosphere. The
changes in oxygen concentration are consistent with oxygen supply by diffusion. The contribution of oxygen made
by the small influx of air on pressure recovery is only a minor fraction of the respiratory oxygen requirement. Wet
weather at low tide restricts gas exchange, affecting both pressure and oxygen. The pressure and oxygen changes
are repeated over many tidal cycles. The results are compared with those for other species and situations in the
literature.
Introduction
Avicennia marina has an adventitious root system
(Baylis, 1950): roots emerge from the hypocotyl, and
after briefly growing downwards they become hori-
zontal. These develop into the cable roots which can
spread out for many metres from the stem. Vertical,
upward-growing pneumatophores arise from the cable
roots and emerge from the mud. They are exposed
to the air at low tide but covered by water when
the tide is high. Cable roots and bases of pneuma-
tophores produce fine roots, with several orders of
branching, beneath the mud surface. Pneumatophores
are aerenchymatous with lenticels (Haberlandt, 1884;
Baker, 1915), and abundant gas spaces occur through-
out the root system (Baylis, 1950). Root architecture
is broadly similar – although with differences of de-
tail – in other species of Avicennia that have been
examined (see Chapman, 1976). The oxygen supply
function of this aerenchymatous root system was sug-
gested in the 19th century and demonstrated in the
20th (see Haberlandt, 1884; Chapman, 1976). In a
landmark field study Scholander et al. (1955) demon-
strated the decline of oxygen concentration inside the
root during high tide and its recovery during low tide,
together with a gradual reduction in gas pressure in-
side pneumatophores while covered by the water and
a very quick recovery on emergence. The low pressure
could induce bulk flow of air into the pneumatophore
(Scholander et al., 1955). Oxygen concentrations in-
side roots were elegantly demonstrated in situ by
Andersen and Kristensen (1988).
We have grown A. marina from propagules in
artificial-tidal tanks. These plants produced root sys-
tems as described above: in Sydney, pneumatophores
appeared after about one year (Curran, 1985; Curran et