Adaptation Constraint And Learning Limits to Learning and Memory

Of interest to behavioral ecologists is the degree to which learned behavior reflects adaptation by natural selection versus constraints on selection. Generalization (see above), for example, may seem at first to reflect a constraint on learning, but conceivably represents an adaptive mechanism of imprecision. A pollinator, for example, that responded only to the precise odor blend emitted by the first rewarding flower encountered might never visit another flower, owing to among-flower variation in the blend.

A classic case study of limits on learning and memory in nature that interested Darwin himself concerns the tendency for bees, butterflies, and other pollinators to show greater fidelity to one or a few floral species than expected based on the profitabilities of those species. According to one point of view, this so-called floral constancy is dictated by limits on the acquisition, retention, and/or retrieval of stored information about the floral resource.

That foraging success in insects is limited in terms of acquisition and retention seems unlikely at the level of LTM. LTM in insects, as mentioned above, is extraordinarily durable and the amount of information that can be maintained in LTM, as currently understood, is extremely impressive. Butterflies can learn visual cues in two foraging modes (nectar collection and oviposition) simultaneously, showing meaningful responses in each instance in just a single trial. Bees can be trained to distinguish multiple rewarded stimuli from multiple unrewarded ones and to link features of eight or more different flower species to the time of day at which nectar is available. These features include flower color, odor, pattern, and microtexture. In addition, a bee learns the location, profitability, and visual landscape associated with a rewarding patch of flowers, as well as the route between hive and patch and, in conjunction with the sun compass used to navigate, even the pattern of movement of the sun through the sky.

Retention at the level of LTM is similarly impressive. Bees have been shown to retain LTM without reinforcement for several weeks, a period of time comparable to average worker life expectancy. In Tribolium beetles and Drosophila, there is evidence that memory formed in the larval stage persists through metamorphosis.

If pollinators are limited at all in memory, it may be at the level of STM. As noted above, STM is particularly vulnerable to conflicting information; this fact may make it difficult for a bee once fixed on a flower type to switch to a novel one. Alternatively, the key to floral constancy may lie in the retrieval of stored information, specifically a constraint on the minimum time required to activate information stored in LTM and a limited capacity to activate multiple memories at once (together, limits on what for vertebrates has been referred to as working memory).

Learning and Memory as Products of Adaptation

An alternative, albeit not mutually exclusive, view holds that natural selection generates an adaptive balance between activation and suppression of memory, tuning that balance finely to the specific ecological requirements of a given species. For example, floral constancy might conceivably permit workers in a colony to partition floral resources efficiently, in which case the properties of learning and memory that contribute to constancy would be viewed as adaptive. It has even been proposed that memory dynamics in bees are tightly matched to foraging activity rhythms as well as the spatial patterning of the floral resource.

Abundant propositions as to adaptive specialization in learning have been made, especially from a comparative standpoint: "Insects of a given species should be prepared to learn particularly well those stimuli relevant to that species' needs." "Social insects should learn better than solitary ones (owing to the demands of a complex and unpredictable social environment)." "Generalist insects should learn better than specialists." For none of these propositions is there compelling evidence, nor will there be until better descriptions are made of learning in an ecological context, learning protocols are brought closer in rigor to those employed in comparative psychology, and more insect species are evaluated.

For now, the primary comparison to be made is a comparison between learning in insects and in vertebrates. Here, the pattern is one of shared features. Despite significant phylogenetic distance between insects and vertebrates, and despite substantial differences in their underlying physiology, there is a remarkable congruence in the diversity and form of learning processes in these taxa (see above). The similarities may reflect shared ancestry, evolutionary convergence, or both. A finding of evolutionary convergence would imply that certain universal, yet to be clearly defined functional principles govern the evolution of learning and memory processes.

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Brain • Dance Language • Neuropeptides

Further Reading

Abramson, C. I., Yuan, A. I., and Goff, T. (1990). "Invertebrate Learning: A

Source Book." Am. Psychol. Assoc., Washington, DC. Bitterman, M. E. (1996). Comparative analysis of learning in honeybees.

Anim. Learn. Behav. 24, 123—141. Chittka, L., Thomson, J. D., and Waser, N. M. (1999). Flower constancy, insect psychology, and plant evolution. Naturwissenschaften 86, 361-367.

Dukas, R. (ed.) (1998). "Cognitive Ecology." University of Chicago Press, Chicago.

Menzel, R. (2001). Searching for the memory trace in a mini-brain, the honeybee. Learn. Memory 8, 53-62. Papaj, D. R., and Lewis, A. C. (eds.) (1993). "Insect Learning: An Ecological and Evolutionary Perspective." Chapman & Hall, New York. Shettleworth, S. (1998). "Cognition, Evolution and Behavior." Oxford

University Press, Oxford. Smith, B. H. (1996). The role of attention in learning about odorants. Biol. Bull. 191, 76-83.

Tully, T. (1997). Regulation of gene expression and its role in long-term memory and synaptic plasticity. Proc. Natl. Acad. Sci. USA 94, 4239-4241.

LegS

Peter H. Adler

Clemson University

One of the most generally known and oft-repeated facts about insects is that they possess three pairs of legs, one pair each on the prothorax, mesothorax, and metathorax. Indeed, this condition is in the fundamental ground plan of insects and is amply represented in the fossil record. The condition inspired Latreille's taxon Hexapoda (Greek hexa, six, andpoda, foot). Exceptions to the hexapodous condition are found in the apodous, or legless, insects that have secondarily lost their legs, typically as a result of selection for an obligatory parasitic or sedentary existence.

The six-legged condition is derived from an ancestral arrangement in which legs occurred on the majority of body segments. Over evolutionary time, the serially uniform legs became modified in the insectan lineage into the characteristic mouthparts, thoracic legs, and various abdominal appendages, such as cerci and genitalia, while typically becoming lost on other abdominal segments. Further evolution of the basic six-legged condition in the insectan lineage has resulted in an enormous diversity of structure and function. This structural and functional diversity of legs, along with the acquisition of wings without the loss of legs, which is a condition unique to insects, undoubtedly has been a key factor in the numerical success of insects and their representation in nearly every habitat on the planet. The exquisite diversity in leg structure plays an important role in the taxonomy and classification of insects.

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