Introduction Ollaetion Morphological basis an overview Peripheral olfactory organ Central projections Registration of olfactory activities Electroolfactogram (EOG) ExtraceUular recordings of action potentials Electroencephalogram (EEG) Cross-adaptation Responses to chemical stimuli Amino acids Steroid hormones Bile salts Prostaglandins (PGs) Other compounds Gustation Morphological basis an overview Taste buds Gustatory nerves and central projections Recordings of gustatory responses Responses to chemical stimuli Amino acids Carboxylic acids Nucleotides Bile salts CO2 and pH Toxins and other compounds Future prospects and research needs Summary and conclusions Olfaction Gustation Acknowledgements References

The diversity of chemical stimulation in fish olfaction and gustation

Living in an aquatic environment, often devoid of light but rich in dissolved compounds, fish have highly developed chemosensory and chemical signalling systems. The olfactory and gustatory systems comprise the major chemosensory pathways. Despite considerable variations in structural organization of the peripheral olfactory organ throughout fish species, ultrastructural organization of the olfactory sensory epithelium is extremely consistent. The olfactory receptor cell is a bipolar neurone which is directly exposed to the external environment and sends information to the brain by its own axon (cranial nerve I). Four major classes of chemicals have been identified as specific olfactory stimuli and their stimulatory effectiveness characterized: amino acids, sex steroids, bile acids/salts and prostaglandins. Olfactory signals such as those involved in reproduction and feeding may be processed independently through two distinct subsystems: the lateral and medial olfactory systems. The taste buds constitute the structural basis of the gustatory organ. Taste buds may occur not only in the oropharyngeal cavity, but on the whole body surface. Chemical information detected by specialized epithelial cells, gustatory cells, is transmitted to the central nervous system by cranial nerve VII (facial), IX (glossopharyngeal), and X (vagal). Besides diverse sensitivities and specificities for amino acids, fish gustatory receptors detect various organic acids, nucleotides and bile salts. Putative receptors, molecular mechanisms of transduction and the role played by olfaction and gustation in feeding, reproduction, migration and other fish behaviours are discussed.

Olfaction and gustation in fish: an overview

Neurobiology of fish olfaction: a review.

Sexually mature male goldfish release large quantities of androstenedione into the water where it functions as a pheromone.

Pheromones play vital roles for survival and reproduction in various organisms. In many fishes, prostaglandin F2α acts not only as a female reproductive hormone, facilitating ovulation and spawning, but also as a sex pheromone inducing male reproductive behaviors. Here, we unravel the molecular and neural circuit mechanisms underlying the pheromonal action of prostaglandin F2α in zebrafish. Prostaglandin F2α specifically activates two olfactory receptors with different sensitivities and expression in distinct populations of ciliated olfactory sensory neurons. Pheromone information is then transmitted to two ventromedial glomeruli in the olfactory bulb and further to four regions in higher olfactory centers. Mutant male zebrafish deficient in the high-affinity receptor exhibit loss of attractive response to prostaglandin F2α and impairment of courtship behaviors toward female fish. These findings demonstrate the functional significance and activation of selective neural circuitry for the sex pheromone prostaglandin F2α and its cognate olfactory receptor in fish reproductive behavior.

Olfactory receptor for prostaglandin F2α mediates male fish courtship behavior.

To determine whether wild common carp (Cyprinus carpio) employ as pheromones the same gonadal hormones as goldfish, and whether their olfactory sensitivity is influenced by gonadal condition, the o...

Acute olfactory sensitivity of wild common carp, Cyprinus carpio, to goldfish hormonal sex pheromones is influenced by gonadal maturity

Twenty years ago Kittredge et al. (1971) speculated that aquatic organisms may have evolved to use hormonal metabolites as sex pheromones. Their reasoning was simple: these compounds are pre-adapted for this function because they are naturally released to the environment in synchrony with important reproductive (endocrinological) events and their recognition might only require a simple mutation by which endocrine receptors are expressed on chemosensory tissue. Ten years later, Colombo et al. (1980) found that a fish, the male black goby (Gobius jozo) produces and releases a conjugated reduced androgen to water where it attracts ovulated females. Soon thereafter, Van den Hurk and Lambert (1983) reported that a mixture of two conjugated steroids, testosterone glucuronide and estradiol glucuronide, attracts male zebra danio (Brachydanio rerio). These exciting findings prompted us to test whether goldfish (Carassius auratus) also use hormonal metabolites as pheromones. Using electrophysiological measures of olfactory sensitivity (the electro-olfactogram or EOG) as a bioassay we discovered that goldfish are extremely sensitive to the steroidal maturation hormone 17α,20β-dihydroxy-4-pregnen-3-one (17,20βP) (see Sorensen, 1992). Subsequently we discovered that 17,20βP is released to the water where if functions as a preovulatory “primer” pheromone, increasing male gonadotropin (GtH) and milt (sperm and seminal fluid) production. Knowing that ovulated goldfish also release a pheromone which stimulates male behavior, we continued our pursuit of hormonal pheromones wing EOG recording to screen putative hormonal pheromones. We subsequently discovered that metabolites of prostaglandin F2α (PGF2α ) are likely released by recently ovulated fish to function as a postovulatory ‘releaser’ pheromone. A preliminary account of these findings was presented at Chemical Signals in Vertebrates V (Sorensen and Stacey, 1990), along with predictions that many other fish use hormonal metabolites as pheromones.

Electrophysiological Measures of Olfactory Sensitivity Suggest That Goldfish and Other Fish Use Species-Specific Mixtures of Hormones and Their Metabolites as Pheromones

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Acute olfactory sensitivity of wild common carp, Cyprinus carpio, to goldfish hormonal sex pheromones is influenced by gonadal maturity

Electrophysiological Measures of Olfactory Sensitivity Suggest That Goldfish and Other Fish Use Species-Specific Mixtures of Hormones and Their Metabolites as Pheromones