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The photograph at left is of a sediment core retrieved
by divers from Explorers Cove, Antarctica. Extending from the sediment
surface are tree-like foraminifera called Notodendrodes antarctikos.
This species, like certain other larger agglutinated foraminifera from
Antarctica, may represent a "missing link" in the evolution
of shelled protists. Molecular phylogenetic studies indicate that the
ancestral foraminifera evolved during the Proterozoic (ca. 850 million
years ago), before cells began building skeletonized structures. The modern
analog to these ancestral forms are the allogromiids, i.e., foraminifera
with an organic wall covering the cell body rather than a hard shell.
Notodendrodes has such an allogromiid body that is free
to move within the cavities of its larger agglutinated shell (see diagram,
left). In essence, it is possible that Notodendrodes represents
a primitive allogromiid that has "learned" to build a hard shell
made of cemented sand grains. The emergence of agglutinated foraminifera
is recorded in the fossil record from Early- to Mid-Cambrian strata. The
Explorers Cove foraminifera are an important living resource for understanding
the physiological processes used by their ancient ancestors to establish
hard skeletons. In learning this, we will better define the cellular basis
for the evolution of skeletonization - the key event in the emergence
of "higher" organisms, including man.
Notodendrodes antarctikos
is also interesting to ecologists because it beautifully illustrates the
principle of "trophic plasticity" that makes foraminifera such
a successful group of protists. We believe that Notodendrodes
evolved from ancestors that had a more conventional shape; the other foram
in this genus, Notodendrodes hyalinosphaira (left),
is found both in a "tree" morphology and as a simple sphere.
Genetic studies have also identified other, non-tree-forming relatives,
such as Rhabdammina cornuta (see phylogenetic tree to left.)
The complex shape of the "tree" helps
Notodendrodes to exploit food sources ranging from dissolved
nutrients in the sediment (taken up by pseudopodia in the root-like structures)
to the algae raining down from the underside of the ice cover (captured
by pseudopodia suspended from the branch-like tree structures). The interplay
between pseudopodia - the organelles that do the work of food collection
- and the shell - the nonliving platforms upon which the pseudopodia are
deployed - is a neglected aspect of foraminiferal biology. We believe
that understanding this interplay is crucial for explaining the evolutionary
success of these important marine organisms, and current work in our lab
is designed to help fill this gap in knowledge.
Notodendrodes
bibliography:
Bowser, S.S., Gooday, A.J., Alexander, S.P., Bernhard,
J.M. (1995) Larger agglutinated foraminifera of McMurdo Sound, Antarctica:
Are Astrammina rara and Notodendrodes antarctikos
allogromiids incognito? Mar. Micropal. 26:75-88.
DeLaca, T.E., Lipps, J.H., Hessler, R.R. (1980)
The morphology and ecology of a new large agglutinated antarctic foraminifer
(Textulariina: Notodendrodidae nov). Zool. J. Lin. Soc. 69:205-224.
DeLaca, T.E. (1982) The use of dissolved amino acids
by the foraminifer Notodendrodes antarctikos. Am. Zool.
22:683-690. |