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- Identification, release and olfactory detection of bile salts in the intestinal fluid of the Senegalese sole (Solea senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Welham, K.; Hardege, Jörg D.; Barata, E. N.; Canario, Adelino V. M.Olfactory sensitivity to bile salts is wide-spread in teleosts; however, which bile salts are released in suYcient quantities to be detected is unclear. The current study identiWed bile salts in the intestinal and bile Xuids of Solea senegalensis by mass spectrometry–liquid chromatography and assessed their olfactory potency by the electro-olfactogram.
- Adaptation to reduced salinity affects the olfactory sensitivity of Senegalese sole (Solea senegalensis Kaup 1858) to Ca2+ and Na+ but not amino acidsPublication . Velez, Zélia; Hubbard, Peter; Barata, E. N.; Canario, Adelino V. M.The Senegalese sole is a marine flatfish, which often penetrates into estuarine waters to feed. It cannot, however, survive in full freshwater. The current study investigated the effect of adaptation to low salinity (10‰) on olfactory responses to changes in environmental [Ca2+] and [Na+] and amino acids by the electro-encephalogram (EEG) recorded from the olfactory bulb. The sole showed olfactory responses to increases in environmental [Na+] and decreases in environmental [Ca2+]; sensitivity to Na+ was greater at 10‰ whereas sensitivity to Ca2+ was greater at 35‰. Decreased environmental [Na+] increased sensitivity to changes in [Ca2+] whereas increased environmental [Ca2+] decreased bulbar responses to changes in [Na+]. Sensitivity to amino acids was unaffected by external salinity. However, the absence of external Na+ strongly decreased bulbar responses to amino acids in fish adapted to 35‰ seawater but not in those at 10‰. The absence of external Ca2+ had no such effect at either salinity. This suggests that odorant-receptor binding and/or olfactory transduction is reliant on external Na+ (but not Ca2+) at higher salinities but the olfactory system is able to adapt to lower environmental [Na+]. Taken together, these results suggest that reductions of external salinity modulate olfactory sensitivity to environmental Ca2+ and Na+ but not amino acids. However, at low salinities, olfactory sensitivity to amino acids is maintained by decreasing reliance on external Na+.
- Functional asymmetry in the olfactory system of a flatflish (Solea Senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Barata, E. N.; Canario, Adelino V. M.The two olfactory epithelia of the family Soleidae are exposed to differentwater sources; the upper (right) epitheliumsamples thewater column whereas the lower (left) samples interstitial water. Given that one of the major prey species of the Senegalese sole (Solea senegalensis) is the ragworm Hediste diversicolor which lives in the substrate, we hypothesized that the lower epitheliummay have evolved sensitivity to odorants released by the ragworm whereas the upper epithelium may be more involved in chemical communication and therefore is more sensitive to odorants released by conspecifics.
- Prey odour enhances swimming activity and feed intake in the Senegalese solePublication . Barata, E. N.; Hubert, F.; Conceição, L. E. C.; Velez, Zélia; Rema, P.; Hubbard, Peter; Canario, Adelino V. M.Olfaction is important in many aspects of the life-history of fishes including feeding, and more so in nocturnal benthic feeders. In the current study we assessed the importance of olfaction in food-search behaviour of the Senegalese sole, an economically important marine species both as farmed and wild-caught. Whole-body homogenates of the polychaete Diopatra neapolitana were fractionated by solid-phase extraction (SPE) using C18 cartridges and the olfactory potency of the resultant fractions (hydrophobic eluate and hydrophilic filtrate) was assessed by the electro-olfactogram in juvenile sole. In addition, the effect of both the homogenate and SPE fractions on sole locomotion was assessed in a flow-through tank (fluviarium). Finally, whole-body homogenate was added to commercial feed pellets and tested whether it could enhance food consumption by sole. The SPE hydrophilic filtrate contained the majority of the olfactory activity found in the whole-body homogenate. Both the homogenate and filtrate, but not the eluate, increased number of movements, time moving, linear velocity, distance travelled and time swimming upstream of sole in the fluviarium; ablation of the olfactory epithelia disrupted these behavioural responses to the homogenate. Intact sole consumed more pellets flavoured with worm homogenate than those without. These results show that olfaction plays an important role in food-search behaviour of the Senegalese sole and that the hydrophilic fraction of D. neapolitana whole-body homogenate contains key substances affecting sole search behaviour; moreover, ingestion by sole was enhanced by addition of worm homogenate to the dry feed pellets.
- The contribution of amino acids to the odour of a prey species in the Senegalese sole (Solea senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Hardege, Jörg D.; Barata, E. N.; Canario, Adelino V. M.For many fish, olfaction is important in food search and consumption. Amino acids are known to elicit feeding behaviour in several species. The aim of the current study was to evaluate the contribution of amino acids to the odour of a natural prey organism of the Senegalese sole (Solea senegalensis). Both whole-body macerates and substances released to the water by living ragworms (Hediste diversicolor) were fractionated by molecular weight filtration followed by solid-phase extraction (SPE), and the olfactory activity of the resultant fractions was assessed by the electro-olfactogram (EOG) in the sole. The amino acid concentrations of the macerate and water were determined by gas chromatography and mass spectrometry (GC–MS). In the macerate, the majority of odorants were small molecular weight compounds (b500 Da) which were not retained by C-18 SPE cartridges. An artificial mixture of amino acids at the same concentrations as found in the macerate had similar olfactory potency. The odorants released to the water by living ragworms were also small molecular weight compounds (b500 Da) but the majority of olfactory activity could be extracted by C-18 SPE cartridges. The concentrations of amino acids in these samples were too low to contribute greatly to its olfactory potency. These results suggest that, whilst olfactory sensitivity to amino acids may explain most of the potency of the macerate, living ragworms are releasing additional odorants other than amino acids which may be equally important in chemosensory food location in the sole.
- Differential detection of conspecific-derived odorants by the two olfactory epithelia of the Senegalese sole (Solea senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Barata, E. N.; Canario, Adelino V. M.The two olfactory epithelia of members of the family Soleidae sample two distinct water sources; the upper (right) side is in contact with the open water column whilst the lower (left) side is in contact with interstitial water. To evaluate whether there are differences in the sensitivities, and therefore functional roles, of the two epithelia the olfactory activity of conspecific-derived odorants was assessed in both using the electro-olfactogram (EOG). The upper nostril was significantly more sensitive to conspecific bile fluid, intestinal fluid and mucus than the lower nostril. Crude fractionation of these samples (solid-phase extraction with C18 and C2/ENV+ cartridges) revealed that olfactory activity in each body fluid was likely due to a mixture of compounds. In each case, the upper olfactory epithelium was significantly more sensitive than the lower. Similarly, olfactory sensitivity to a range of C24 and C27 bile acids was greater in the upper epithelium. These results suggest that intra-specific chemical communication is mediated mainly, if not entirely, by the upper olfactory epithelium. The odorants involved, and their functional roles, remain to be established.
- Evidence for functional asymmetry in the olfactory system of the senegalese sole ( solea senegalensis )Publication . Velez, Zélia; Hubbard, Peter; Barata, E. N.; Canario, Adelino V. M.The two olfactory epithelia of flatfish of the family Soleidae are essentially in contact with two distinct environments; the upper (right) side samples open water while the lower (left) side samples interstitial water. This study assessed whether there are differences in the responsiveness of the two epithelia by use of the electro‐olfactogram in the Senegalese sole (Solea senegalensis). The upper epithelium was significantly more responsive to the basic amino acids (l‐lysine and l‐arginine), glycine, and l‐threonine than the lower epithelium. The lower epithelium was significantly more responsive to aromatic amino acids (l‐tryptophan, l‐tyrosine, l‐DOPA, and l‐phenylalanine), l‐leucine, and l‐asparagine than the upper. Both epithelia had similar responsiveness to the sulphur‐containing amino acids (l‐cysteine and l‐methionine), l‐alanine, l‐serine, and l‐glutamine. Neither side was responsive to the acidic amino acids (l‐aspartate and l‐glutamate) or the D‐isomers of any amino acid tested. The upper olfactory organ was much more responsive to conspecific‐derived stimuli (bile and intestinal fluid) than the lower organ. We suggest that these differences in responsiveness may be related to different functional roles of the upper and lower epithelia in feeding and chemical communication.
- Electro-olfactograms in response to chromatographic fractions of food-related odorants in the Senegalese sole, Solea senegalensisPublication . Velez, Zélia; Hubbard, Peter; Barata, E. N.; Hardege, Jörg D.; Bubblitz, R.; Canario, Adelino V. M.The Bohr effect, which can be most generally defined as the influence of pH on the oxygen binding affinity of proteins, is a common feature of respiratory pigments, ranging from the haemocyanins of molluscs and crustaceans to the haemoglobins of vertebrates. Its physiological role is generally seen in the facilitation of oxygen release from respiratory pigments during tissue acidosis. The magnitude of the effect can be influenced by a multitude of factors such as temperature, carbon dioxide, chloride ions, organic phosphates and the investigated pH range. Here we present data on the maximal alkaline Bohr effect in haemoglobins from a large number of species covering all vertebrate classes, obtained at physiological temperatures in the presence of 100 mM chloride ions and the absence of carbon dioxide and organic phosphates.
- Functional asymmetry in the olfactory system of a flatfish, the Senegalese sole (Solea senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Welham, K.; Barata, E. N.; Hardege, Jörg D.; Canario, Adelino V. M.The two olfactory epithelia of the family Soleidae are exposed to different water sources; the upper (right) epithelium samples the water column whereas the lower (left) samples interstitial water. Given that one of the major prey species of the Senegalese sole (Solea senegalensis) is the ragworm Hediste diversicolor which lives in the substrate, we hypothesized that the lower epithelium may have evolved sensitivity to odorants released by the ragworm whereas the upper epithelium may be more involved in chemical communication and therefore is more sensitive to odorants released by conspecifics.