The exercise physiology of snapper (Pagrus auratus): implications for the better commercial harvesting of an iconic New Zealand finfish
Degree GrantorUniversity of Canterbury
Degree NameDoctor of Philosophy
Worldwide, an increasing demand for fish and fisheries products, together with socioeconomic pressure for industry expansion, is placing considerable pressure on wild fish stocks – more than 80% of which are considered by the Food and Agriculture Organisation of the United Nations (FAO) to be either maximally- or over-exploited. Adding value to the existing catch and/or improving the sustainability of current wild capture methods may offer a means of providing industry growth while negating the need for increased landings. In particular, the peri-mortem condition of a fish plays an integral role in the condition of the tissues post-mortem and hence in product quality, with harvesting techniques that result in stress or fatigue yielding a lower quality product. An understanding of the physiology of the target species and its response to harvest is therefore essential to implementing targeted improvements in harvesting technologies. For species harvested using trawl-based technologies, this includes knowledge of their exercise physiology, in particular their swimming capacity, since this is a key determinant of the interaction between fish and trawl gears, and hence of the nature and severity of stress experienced and of the condition of fish at landing.
This thesis describes a series of discrete studies relating to the exercise physiology of juvenile snapper, Pagrus auratus, an iconic New Zealand finfish that comprises important recreational and commercial fisheries. In particular, we sought to characterise the capacity of snapper for sustained swimming activity, including how performance may differ between fish of different size or with environmental temperature; to determine the consequences of exhaustive exercise for both subsequent swimming activity, an important determinant of survival in escaping or discarded catch, and for tissue biochemistry, which ultimately determines product quality in harvested fish; to validate the use of laboratory-based simulations for the study of capture-related stress by comparing the response of laboratory-exercised snapper with commercially caught fish; and to determine the tolerance of snapper to environmental hypoxia, and further, the possible consequences of hypoxia for swimming capacity and for recovery in fish retained for subsequent rested-type harvest.
The capacity of snapper for sustained swimming activity was characterised through the use of incremental exercise tests to determine critical swimming speeds, Ucrit. Juvenile snapper (94-107 mm length, 16-157 g mass) demonstrated a strong swimming capacity, with individual fish attaining critical swimming speeds of up to 7.1 body lengths per second (bl s⁻¹). Swimming performance demonstrated an allometric association, with absolute critical speeds increasing with fish size, whilst relative performance favoured smaller fish. The relation was described by the function Ucrit (m s⁻¹) = 0.003412 [length (mm)] + 0.2669. Critical swimming performance also exhibited variation in response to environmental variables. Thermal performance curves were evident in snapper acclimated to 12, 18 and 24 °C, with the suggestion of optimal performance at acclimation temperatures between 18 and 24 °C. Critical swimming performance was also significantly reduced during exposure to ambient oxygen tensions below 80 mmHg; at 40 mmHg, snapper attained only 21% of the critical swimming speeds observed under normoxic (150 mmHg) conditions.
In juvenile snapper (~75 g), exhaustive exercise resulted in severe metabolic, acid-base, haematological and hormonal perturbations, the nature of which were similar to those classically demonstrated in other strong-swimming fish species, especially salmonids. These included the depletion of glycogen from within the white muscle (WM) and the concomitant production of lactate, with a resultant lactacidosis of the plasma; recruitment of erythrocytes from the spleen; and the release of cortisol to the plasma. The recovery of these disturbances required 6 hours under laboratory conditions. As the stresses experienced by fish during commercial capture are often considered to be greater than those which can be induced during laboratory-based simulations, it was necessary to investigate whether the magnitude of the perturbations observed in laboratory-exercised snapper were an appropriate model of those of trawl-caught fish. In trawl-caught snapper (1100 g, 38 cm) obtained under commercially-relevant conditions (tow speed ~3.0 knots; duration 2.25-2.75 hours), the magnitude of the perturbations were greater than for laboratory-exercised fish. While the recovery of some metabolites was evident within the first 18 hours post-capture, their recovery was prolonged relative to laboratory-exercised fish; other metabolites, namely muscle glycogen and plasma cortisol, exhibited no signs of recovery. These observations suggest that the response of snapper to exhaustive exercise within the laboratory may underestimate the severity of the response induced by commercial harvest. This is further suggested by post-capture mortality rates of 14%, whereas no mortality was observed following fatigue at Ucrit.
Exhaustive exercise also resulted in the impairment of subsequent critical swimming performance. Immediately following fatigue, snapper (85-160 g) were capable of sustained swimming activity at speeds of up to 60-70% Ucrit; however, critical swimming performance was reduced 30%, presumably due to limitations in WM function. There was no suggestion of the recovery of WM function within the first 30 minutes post-fatigue; thereafter, Ucrit was progressively restored, such that snapper were able to repeat their initial swimming performance in a second Ucrit test performed 2 hours after the conclusion of the first.
Snapper were moderately tolerant of hypoxia, oxygen-regulating at reduced oxygen tensions (<100 mmHg) by virtue of increased ventilatory rate and stroke volume, with a distinct bradycardia developing at PO₂ below 60 mmHg. Larger snapper appeared to possess a greater hypoxia tolerance than did smaller fish, with Pcrit resolved to 77 in 20 g fish, and 50 mmHg in 150 and 230 g fish. Exposure to moderate hypoxia (60-80 mmHg) during recovery from an exhaustive exercise event constrained MO₂ max to 78% of that of normoxic fish, however did not appear to impede the return of MO₂ to routine levels.
The present study is the first to examine in detail the swimming performance of snapper, and the consequences of exhaustive exercise for physiological condition. By understanding the swimming capacities of snapper, it may be possible to refine harvesting practices (i.e. tow speeds) or utilise technologies (i.e. net design) such that the water velocities through the trawl net are within the range at which the fish can swim sustainably, minimising the extent of stress and fatigue experienced by fish, and hence their effects on both quality and survival. The study also demonstrates that whilst snapper experience significant physiological disturbance during commercial harvesting, including significant mortality, some fish demonstrate the potential for metabolic recovery, which may permit their retention in an on-board tank facility for subsequent rested-type harvest. Finally, the present work highlights a number gaps in our understanding of the link between harvesting conditions and fish condition, and makes a number of suggestions for future studies or directions.