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1 Invertebrate Olfaction (60/26) -- Introduction to Neuroscience

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1 Invertebrate Olfaction

1 Invertebrate Olfaction Tom Finger EVOLUTION OF TASTE Thomas E. Finger Dept. Cell & Developmental Biology Rocky Mountain Taste & Smell Center Univ. Colorado Medical School Aurora CO 80045 USA Contact: Thomas Finger Univ. Colorado Health Sciences Center Room L18-11118, RC-1 12801 E. 17th Ave. MS 8108 PO Box 6511 Aurora CO 80045 <EMAIL_ADDRESS>KEYWORDS: Amphid Amphioxus Cell type Chemoreception Drosophila Fish Insect Labellum Leech Mammal Nematode Octopus Solitary Chemoreceptor Cell Taste Bud Taste receptor 2 SYNOPSIS: The sense of taste is well-defined only for vertebrates where there are readily identified taste buds. These endorgans always are innervated by particular cranial nerves and always serve in the regulation of feeding and food intake. Since invertebrates, including amphioxus, have no such endorgans, comparisons across groups must be made on the basis of analogy rather than homology. I adopt the term “taste” only for chemosensory endorgans devoted to feeding-related behaviors without regard to endorgan structure, expression of molecular receptors, chemical nature of the stimulus (e.g. volatility), or vehicle in which a stimulus is presented (e.g. air vs water). In that context, nearly all metazoans have a sense of taste, i.e. perioral or intraoral chemoreceptors that regulate feeding. This chapter explores the diversity of such endorgans in both vertebrates and invertebrates. For the vertebrate lineage, this modality is mediated by taste buds innervated only by the facial, glossopharyngeal or vagus nerves. Taste buds in all vertebrates share many common features but two basic forms of organization can be recognized. In non-mammalian vertebrates, taste buds consist of elongate taste cells and Merkel-like basal cells. In contrast, taste buds in mammals contain only elongate cells, but one of these (Type III taste cells) shares many characteristics with the Merkel-like basal cells of non-mammalian vertebrates. 3 01.1 INTRODUCTION We humans recognize the sense of taste as those sensations arising from the oral cavity and indicating information about the chemical quality of potential foodstuffs. The sense of taste uniquely arises from the specialized sensory endorgans for this system: taste buds. In humans, taste sensations are only those which we describe as salty, sweet, sour, bitter, and “umami” (the taste of glutamate). All other oral sensations, e.g. the coolness of mint, the smoothness of fats or the hotness of pepper, arise from the general cutaneous innervation of the epithelium and should not be considered to be “taste”. So for humans and other vertebrates, “taste” is a system defined by the sensory endorgans mediating the sensibility. For humans, and by extension, other vertebrates, taste can be defined by the use of taste buds in the context of food selection. When examining the evolution of the sense of taste, we are met with several difficulties. First, how can the sense of taste be defined for organisms lacking taste buds and
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