George Hughes and the history of fish ventilation: From Du Verney to the present

This paper traces the research history of fish ventilation from its origins in the early 1700s to the present with emphasis on the work of George M. Hughes, who is considered by many to be the founder of the modern era of fish respiratory science. A particularly important year in the timeline for fish respiratory studies was 1960, when Hughes presented the currently accepted biomechanical model driving fish ventilation. He showed that both bony and cartilaginous fishes breathe through the use of a dual-pumping mechanism: a buccal or orobranchial pressure pump to force water over the gills and an opercular or parabranchial suction pump to pull water through the branchial chambers. Hughes divided this mechanism into four stages and demonstrated that during each the pressure of the buccal cavity usually exceeded that of the opercular chamber, thus indicating the continuous, or nearly continuous, nature of the ventilatory stream. Studies by Hughes and later researchers focused on variation in the four stages and related these to interspecific differences in fish habitat and activity level. Differences noted in the respiration of pelagic and benthic species largely led to the description and quantification of ram ventilation. Hughes further made significant contributions to the correlation of gill structure and function and was one of the first to examine gill morphometrics in relation to the ventilatory stream and the diffusivity of oxygen from the water into the blood. Such pioneering measurements paved the way toward the modern analyses of gill hydromechanics and the modeling of respiratory gas exchange in fishes.