Neuroendocrine control of breathing in fish

Beginning with the discovery more than 35 years ago that oxygen chemoreceptors of the fish gill are enriched with serotonin, numerous studies have examined the importance of this, and other neuroendocrine factors in piscine chemoreceptor function, and in particular on the chemoreceptor-mediated reflex control of breathing. However, despite these studies, there is continued debate as to the role of neuroendocrine factors in the initiation or modulation of breathing during environmental disturbances or physical activity. In this review, we summarize the state-of-knowledge surrounding the neuroendocrine control of oxygen chemoreception in fish and the associated reflex adjustments to ventilation. We focus on neurohumoral substances that either are present in chemosensory cells or those that are localised elsewhere but have also been implicated in the direct control of breathing. These substances include serotonin, catecholamines (adrenaline and noradrenaline), acetylcholine, purines and gaseous neurotransmitters. Despite the growing indirect evidence for an involvement of these neuroendocrine factors in chemoreception and ventilatory control, direct evidence awaits the incorporation of novel methods currently under development.