Chemical defense and evolutionary trends in biosynthetic capacity among dorid nudibranchs (Mollusca: Gastropoda: Opisthobranchia) |
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Authors: | Guido Cimino Michael T Ghiselin |
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Institution: | (1) Istituto per la Chimica di Molecole di Interesse Biologico, CNR, Via Toiano 6, I-80072 Arco Felice (Napoli), Italy, e-mail: cim@cim.icmib.na.cnr.it, IT;(2) Department of Invertebrate Zoology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, USA, e-mail: mghiselin@alacademy.org, US |
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Abstract: | Summary. An evolutionary scenario incorporating recent advances in phylogenetic research begins with an opisthobranch-pulmonate common
ancestor that was herbivorous and had some diet-derived chemical defense. The Nudibranchia and their closest relatives, the
Notaspidea, form a lineage the ancestors of which had switched to feeding upon sponges and deriving protection from metabolites
contained in them. Subsequently there have been repeated shifts in food and defensive metabolites, and trends are evident
in the ability to detoxify, sequester and utilize metabolites from food, as well as to synthesize defensive compounds de novo. The Notaspidea display a minor adaptive radiation that foreshadows a more extensive one in the various lineages of nudibranchs.
This review emphasizes changes that have occurred within the Holohepatica, or dorid nudibranchs (order Doridacea). Their sister-group,
the Cladohepatica, consists of three other orders, Dendronotacea, Arminacea, and Aeolidiacea, in which there has been a shift
from sponges to Cnidaria as food. The Dendronotacea often feed upon Octocorallia, which combine spicules, chemical defense,
and stinging capsules and thereby suggest a transition from feeding on sponges. A previous diet of Octocorallia is suggested
by the defensive use of prostaglandins in the dendronotacean Tethys fimbria, which eats crustaceans. A shift to bryozoans in some Arminacea is accompanied by use of different metabolites. Dorid nudibranchs
evidently began as sponge-feeders, but some lineages have shifted to a variety of other food organisms, and others have specialized
in the kind of sponges they feed on and how they do it. There have been shifts to bryozoans (Ectoprocta) and ascidians (Chordata:
Urochordata) that track metabolites rather than the taxonomy of the food. There is a crude correlation between the genealogy
and the defensive metabolites of the sponge-feeding dorids. De novo synthesis is well documented in this order and the metabolites are appropriately positioned so as to have an adaptive effect.
The hypothesis that the capacity for de novo synthesis was acquired by gene transfer across lineages is rejected, partly on the basis of different chirality of metabolites
in the nudibranchs and their food organisms. Instead it is proposed that there has been a preadaptive phase followed by evolution
in a retrosynthetic mode, with selection favoring enzymes that enhance the yield of end products that are already present
in the food.
Received 5 February 1999; accepted 26 July 1999 |
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Keywords: | : chemical defense biosynthesis Gastropoda Opisthobranchia Notaspidea Nudibranchia Doridacea Porifera evolutionary trends |
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