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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.
- Evidence that 1-methyl-l-tryptophan is a food-related odorant for the Senegalese sole (Solea senegalensis)Publication . Velez, Zélia; Hubbard, Peter; Hardege, Jörg D.; Welham, K.; Barata, E. N.; Canario, Adelino V. M.Nocturnal animals often rely heavily on olfactory cues to locate their food. This is especially true for fish whose prey live buried in the substrate. The aim of the current study was to identify non-polar odorants released by the ragworm (Hediste diversicolor) that are detected by the olfactory system of the Senegalese sole (Solea senegalensis) and therefore may be food-related cues. Ragworm-conditioned water was passed through a C18 solid-phase extraction cartridge (to remove polar compounds) and the eluate fractionated by reverse phase high-performance liquid chromatography (HPLC). The fractions were then tested for olfactory activity in the sole by the electro-olfactogram (EOG). Most olfactory activity was found in the first two fractions to elute from the HPLC column (0–4 min). Liquid chromatography-mass spectrometry (LC/MS) of both fractions revealed a base peak with a molecular mass of 219.4 Da and a fragment of 205.3 Da. These data are consistent with a methylated form of tryptophan; synthetic 1-methyl-l-tryptophan had a similar HPLC retention time (2.8 min) and similar LC/MS spectra. Furthermore, sole showed olfactory sensitivity to 1-methyl-l-tryptophan with the lower (left) epithelium being more sensitive than the upper (right). Cross-adaptation (using EOG recording) suggested that the olfactory receptors responding to 1-methyl-l-tryptophan are different in the lower epithelium from the upper. These results suggest that ragworms release 1-methyl-l-tryptophan, or similar tryptophan derivative, and that sole may use this chemical cue to locate and/or identify one of their main prey species. This may help the formation of artificial feeds that are more attractive to sole.
- 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.
- 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.
- Short- and medium-term exposure to ocean acidification reduces olfactory sensitivity in Gilthead SeabreamPublication . Velez, Zélia; Roggatz, Christina C.; Benoit, David M.; Hardege, Jörg D.; Hubbard, PeterThe effects of ocean acidification on fish are only partially understood. Studies on olfaction are mostly limited to behavioral alterations of coral reef fish; studies on temperate species and/or with economic importance are scarce. The current study evaluated the effects of short- and medium-term exposure to ocean acidification on the olfactory system of gilthead seabream (Spares aurata), and attempted to explain observed differences in sensitivity by changes in the protonation state of amino acid odorants. Short-term exposure to elevated PCO2 decreased olfactory sensitivity to some odorants, such as L-serine, L-leucine, L-arginine, L-glutamate, and conspecific intestinal fluid, but not to others, such as L-glutamine and conspecific bile fluid. Seabream were unable to compensate for high PCO2 levels in the medium term; after 4 weeks exposure to high PCO2 , the olfactory sensitivity remained lower in elevated PCO2 water. The decrease in olfactory sensitivity in high PCO2 water could be partly attributed to changes in the protonation state of the odorants and/or their receptor(s); we illustrate how protonation due to reduced pH causes changes in the charge distribution of odorant molecules, an essential component for ligand-receptor interaction. However, there are other mechanisms involved. At a histological level, the olfactory epithelium contained higher densities of mucus cells in fish kept in high CO2 water, and a shift in pH of the mucus they produced to more neutral. These differences suggest a physiological response of the olfactory epithelium to lower pH and/or high CO2 levels, but an inability to fully counteract the effects of acidification on olfactory sensitivity. Therefore, the current study provides evidence for a direct, medium term, global effect of ocean acidification on olfactory sensitivity in fish, and possibly other marine organisms, and suggests a partial explanatory mechanism.
- 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.