Colin Brauner


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Dissertations completed in 2010 or later are listed below. Please note that there is a 6-12 month delay to add the latest dissertations.

Patterns of plasma accessible carbonic anhydrase localisation in derived, basal, and developing fishes (2023)

Teleosts comprise half of all vertebrates and have successfully invaded every aquatic habitat on Earth. They owe this success, at least in part, to their unique oxygenation system. This system is comprised of highly pH sensitive hemoglobins (Hb; large Bohr and Root effects), red blood cell (RBC) intracellular pH (pHi) protection by a beta adrenergically stimulated sodium proton exchanger (β-NHE), and a heterogeneous distribution of plasma accessible carbonic anhydrase (paCA; absence at the gills and presence at the tissues). Together, these components enhance oxygen O₂ unloading to the tissues while protecting oxygen uptake at the gills during a generalised acidosis. While the first two components have received much research interest, the third component, a heterogeneous distribution of paCA, has not and is the focus of my thesis. It has been hypothesised that the absence of paCA in the gills of teleosts is due to the presence of the large Root effect and the associated risks to O₂ loading. In my thesis I aimed to address this principal hypothesis with three research chapters. I firstly confirmed the presence of paCA in most tissues, but absence in all four gill arches of the rainbow trout, which until now had been assumed to be true based on data exclusively from gill arch 2. I then investigated rainbow trout through ontogeny, and found that pre-hatch, during embryonic Hb expression branchial paCA is expressed while β-NHE is not; a pattern that exhibits an abrupt switch with hatching. Finally, I investigated several CA characteristics in three basal actinopterygian species for which nothing was known previously. I demonstrated that paCA in the gill was likely lost earlier than predicted by my principal hypothesis, suggesting instead that the loss of branchial paCA is related to the increasing magnitude of the Bohr coefficient and corresponding decrease in Hb buffer capacity. Collectively, my findings suggest that the presence of a large Root effect is unlikely to be the single selective factor for the loss of branchial paCA in teleosts, with implications for our understanding of a group that represents half of all vertebrates.

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Oxygen, carbon dioxide and ammonia exchange in the gastrointestinal tract of teleost fish (2022)

The gastrointestinal tract (GIT) of teleost fish is a multifunctional organ system involved in digestion, assimilation of nutrients, immune responses, neuroendocrine functions and osmoregulation. Many of these processes have implications for gas exchange, acid-base regulation and ion regulation. However, there is a current lack of knowledge on the basic physiology of the three respiratory gases - oxygen, carbon dioxide and ammonia - in the GIT. My thesis expands our current understanding of the basic profiles of the three respiratory gases in the GIT lumen, their potential exchange with the vascular system, and the changes associated with feeding. In addition, the potential influence of osmoregulatory functions on these parameters was investigated by conducting a comparative study between freshwater and seawater species; the rainbow trout (Oncorhynchus mykiss) was used as a freshwater model species for in depth investigations and the English sole (Parophrys vetulus) as a comparative seawater species. A combination of in vivo, in situ and in vitro techniques was used to assess these areas at the whole animal, tissue and cellular levels. Based on findings from 5 research chapters, I conclude that both freshwater rainbow trout and seawater English sole have nearly anoxic, hypercapnic and high ammonia environments in the GIT lumen in both fasting and fed conditions. Feeding increases carbon dioxide and ammonia levels, but the GIT epithelia regulates the diffusion of the three respiratory gases into the vascular system regardless of feeding status. The different osmoregulatory requirements in seawater versus freshwater did not greatly affect these conclusions. Furthermore, the intestinal tissues have the capacity to handle and transport two nitrogenous products: urea and ammonia. Throughout the GIT, there were expressions of both a potential NH₄⁺ transporter (NKCC) and NH₃ channels (Rhbg, Rhcg2), but ammonia transport was independent of the PNH3 gradient. The thesis also highlights the importance of the stomach in absorbing ammonia despite a reverse PNH3 gradient at physiologically relevant pH levels.

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Towards a comprehensive understanding of gas bubble trauma in fish (2022)

Features such as river dams can generate total dissolved gas (TDG) supersaturation in water downstream, which can lead to harmful bubble growth in the tissues of aquatic animals, known as gas bubble trauma (GBT). Despite numerous previous studies, there remain many unknowns about the factors that affect GBT and the indirect effects of GBT on aquatic animals. I conducted a series of studies to determine key factors that affect the induction of GBT and to describe the outcomes in fish. My systematic review and meta-analysis of the relationship between TDG and GBT in fish generated models that identified depth, dissolved oxygen to nitrogen ratios, temperature, body size, and species as important factors affecting the progression of GBT. Depth has a protective effect against GBT by causing bubbles to dissolve in animal tissues, but the empirical relationship was untested. I conducted an experiment to test the relationship between depth, TDG, and GBT in rainbow trout (Oncorhynchus mykiss). My results indicate that 47 cm of depth compensated for 4.1% (±1.3% SE) TDG supersaturation, as predicted. Based on this result, I proposed an equation for the threshold for GBT at depth. Conversely, locomotion is predicted to promote GBT in fish tissues. I tested the relationship between locomotion and GBT in rainbow trout. My results suggest that there is no significant effect of locomotion on the progression of GBT. I also conducted an experiment to determine whether fish can detect and avoid harmful levels of TDG supersaturation. I found no statistically significant difference in the duration spent by fish in a channel with a lethal level of TDG supersaturation compared with a channel containing air-equilibrated water. I summarized the primary factors affecting the progression of GBT on freshwater fish and the direct and indirect effects of GBT that are supported by my data and the literature. I propose directions for future research and recommendations for TDG monitoring and management.

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On the temperature-dependence of h?moglobin-oxygenation and blood-oxygen transport in regionally heterothermic teleosts and sharks (2021)

Hæmoglobin (Hb)-O2 affinity in most vertebrates typically decreases with increasing blood temperature because the oxygenation enthalpy (ΔH') is usually exothermic. However, in regionally heterothermic fishes, such as tunas and some sharks, ΔH' is commonly low or even endothermic, causing a very reduced or reversed effect of temperature on Hb-O₂ affinity. Regionally heterothermic fishes conserve metabolic heat with vascular heat exchangers that prevent circulatory heat loss and establish internal temperature gradients. My objective was to investigate the functional significance of reduced and reverse temperature-dependent Hb-O₂ affinity in regionally heterothermic fishes. I hypothesized that temperature-independent Hb-O₂ affinity conserves heat energy and matches O₂ supply to O₂ demand despite large internal temperature gradients, so I expected this trait to be shared by all regionally heterothermic fishes. I investigated this by (1) examining the effect of temperature on Hb-O₂ affinity in three regionally heterothermic species whose Hb has not been well studied (swordfish, opah, and common thresher shark), and (2) mathematical modelling of O₂ transport to quantitatively assess the relative contributions of Hb concentration and Hb-O₂ affinity to O₂ and heat transport.I found that opah Hb-O₂ affinity is temperature-independent, the temperature-dependence of swordfish Hb-O₂ affinity is pH dependent, becoming temperature-independent at low pH, and common thresher shark Hb-O₂ affinity is temperature-independent. I also found that Hb from bigeye thresher shark, a suspected regional heterotherm, is temperature-independent below 50% Hb-O₂ saturation. Using a mathematical model of O₂ transport I demonstrated that Hb concentration and Hb-O₂ affinity are relatively more important than other factors of O₂ transport in determining maximum O₂ consumption in yellowfin tuna, a regional heterotherm with aiv“high-energy demand.” I also showed that Hb with a reversed temperature-dependence diminishes temperature induced changes to blood O₂ tension and prevents Hb-heat loss, as much as 13% of metabolic heat production.These results provide insight into the functional significance of reduced and reversed temperature-dependent Hb-O₂ affinity in regionally heterothermic fishes. All known lineages of regionally heterothermic fishes have Hb with a low ΔH', and increases to ΔH' have convergently evolved by different molecular mechanisms with underlying dependence on different allosteric effectors.

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Ions before oxygen: ancestral origins of vertebrate gill function (2020)

Gas exchange and ion regulation at gills play many key roles in vertebrate evolution. Current hypotheses assume gills acquired these important functions from the skin along the vertebrate stem, facilitating the evolution of larger, armoured and more active modes of life. However, this assumption lacks functional support from representatives of early vertebrates and their ancestors.To better understand how and when vertebrate gills became the primary site of gas exchange and ion regulation, I characterized gill and skin function in representatives of ancestral vertebrates (lamprey ammocoete, Entosphenus tridentatus), cephalochordates (amphioxus, Branchiostoma floridae) and hemichordates (acorn worm, Saccoglossus kowalevskii). Intraspecific comparisons within ammocoetes tested the effects of size, dermal thickness and activity on gill function, and interspecific comparisons between taxa tested ancestral origins of gill function.For ammocoetes, I measured multiple gas and ion fluxes in vivo at gills and skin of different sized animals (0.02-2.00 g). Gills accounted for ~20% of gas flux in the smallest ammocoetes in normoxia at 10°C, and contributions increased with size, hypoxia and temperature. Conversely, gills accounted for 100% of ion flux in all sizes and conditions.For acorn worms, I exploited their regenerative ability to partition animals into viable halves with and without gills for respirometry. Gills did not enhance oxygen uptake or ammonia excretion despite hypoxic or thermal challenges. Morphometry by others estimates amphioxus gills also contribute negligibly to gas exchange. However, both acorn worm and amphioxus gills displayed elevated signals for ion regulation (NKA/VHA activity; CA/NHE/AE/Foxi expression).This is the first functional support in ancestral representatives for a vertebrate origin of gas exchange at gills associated with increasing size, dermal thickness and activity. However, ion regulation at gills appears unrelated to this vertebrate transition, and results instead suggest a novel and much earlier deuterostome origin near the inception of pharyngeal arches.

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Characterization of a novel androgen membrane receptor in lampreys that may regulate sexual development: Androgen binding in Lampreys may have implications for steroid ligand and receptor evolution in vertebrates (2019)

Lampreys are basal vertebrates, and their physiology may provide insight into the evolution of physiological systems in vertebrates. To date only progestin, corticoid and estrogen receptors, have been identified in the sea lamprey, Petromyzonmarinus. This is remarkable because 1) more derived vertebrates have evolved six nuclear receptors; 2) if androgen nuclear receptors are absent how might male sexual development be regulated in this ancient group of fishes; 3) androgens have been identified in lampreys so the lack of a nuclear androgen receptor may suggest alternative signaling pathways. I test the hypothesis that lampreys have an active androgen steroid receptor.This hypothesis led to three predictions: first that lamprey synthesize the androgens dehydroepiandrosterone (DHEA) and androstenedione (Ad), second that the master sex hormone gonadotropin releasing hormone regulates androgen synthesis in the testes of male lamprey, and third that androgen signaling in the testes occurs via a novel receptor. Using high performance liquid chromatography, thin layer chromatography and radio-immunoassay I demonstrated the presence of the androgenic steroids DHEA and Ad in the circulation and tissues of sea lamprey and Pacific lamprey. Further, incubation of lamprey testes with lamprey specific GnRH I and III resulted in promoting the conversion of DHEA to Ad. Finally I have demonstrated that androstenedione binds to a membrane fraction isolated from the testes of Pacific lamprey, Entosphenus tridentatus, testes suggesting the presence of a putative androgen membrane receptor (mAR). The binding characteristics indicate a high-affinity (Kd = 7.548 +/- 1.455 nM, R​² = 0.9804,) low capacity (Bmax = 0.0.2366 +/- 0.01345 nM/mg of protein), single binding site androgen receptor. The association rate was determined by non-linear analysis to be 10.2 +/- 3.2 min, with maximum binding achieved at approximately 30 min. The dissociation rate was similar: 9.5 +/- 5.0 min, with maximum binding achieved at approximately 30 min. A partial identification of the receptor was achieved through the use of an affinity column and liquid chromatography. Identification of a novel androgen receptor may point to a novel evolutionary pathway for androgen signaling in vertebrates. It’s significant as it may indicate that this pathway is an ancestral state.

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The functional significance of plasma-accessible carbonic anhydrase for cardiovascular oxygen transport in teleosts (2018)

A novel mechanism has recently been discovered in rainbow trout that allows these fish to enhance the partial pressure of O₂ (PO₂) in their muscles. Teleosts have evolved highly pH-sensitive haemoglobins (Hb), where an arterial-venous pH shift (ΔpHa-v) can severely reduce Hb-O₂ binding affinity. Most teleosts create large ΔpHa-v by actively regulating the intracellular (pHi) of their red blood cells (RBC) through adrenergically stimulated sodium-proton exchangers (β-NHE). This creates H⁺ gradients across the RBC membrane that are short-circuited in the presence of plasma-accessible carbonic anhydrase (paCA) at the tissues, to greatly enhance O₂ unloading from pH-sensitive Hb. Thus, I hypothesised that teleosts increase the O₂ capacitance of their blood (βb) by a mechanism of active RBC pHi regulation that is modulated through a heterogeneous distribution of paCA, which has functional significance for O₂ transport in vivo. Mechanistically, I discovered that in rainbow trout, the time-course of β-NHE short-circuiting in the capillaries and the recovery of RBC pHi during venous transit, are consistent with a system that can enhance O₂ unloading at the tissues with every pass through the circulation. Functionally, I discovered that the inhibition of paCA in Atlantic salmon swimming at a low speed or at rest, required a compensatory increase in cardiac output of ~30%, corroborating a role of paCA in O₂ transport over a broad range of conditions. Further, I discovered paCA in the heart lumen of coho salmon; thus, also cardiac O₂ supply in salmonids may rely on β-NHE short-circuiting. In teleosts, the evolution of β-NHE short-circuiting required the loss of paCA at the gills. However, in Antarctic icefish, I propose that the loss of Hb and RBCs released the functional constraint on the expression of paCA at the gills, and allowed for the enzyme to catalyse CO₂ excretion in the absence of RBC CA. Collectively, my findings indicate that an active mechanism at the level of the RBC enhances βb and is an integral part of the salmonid mode of cardiovascular O₂ transport, and perhaps most teleosts, with important implications for the physiology, the conservation and the evolutionary history, of nearly half of all vertebrates.

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Vertebrate preferential intracellular pH regulation during severe acute hypercarbia (2018)

Environmental CO2 tensions reach >8 kPa (ca. 79,000 μatm; hypercarbia) in some habitats and create severe acid-base challenges for vertebrates. Typically, during a hypercarbic-induced respiratory acidosis, changes in blood pH are compensated for, which returns pH to its normal value, and this is coupled to tissue pH (pHi) regulation. However, during acute environmental CO₂ exposure, this process may be limited to 1 pH unit) and can tolerate PCO₂ >3 kPa. I hypothesized that preferential pHi regulation is used by adult fishes and embryonic amniotes during severe acute acid-base disturbances. This was investigated by examining (1) whether preferential pHi regulation is a general response to various types of acid-base disturbances, (2) surveying fishes for the presence or absence of preferential pHi regulation, and (3) whether preferential pHi regulation is used during development in reptiles.Using white sturgeon, I found that preferential pHi regulation is not a general response to both respiratory and metabolic acidoses. Despite a robust capacity for preferential pHi regulation during respiratory acidoses, not all tissues were protected during metabolic acidoses to the same degree. Preferential pHi regulation was observed to be a common pattern of acid-base regulation amongst fishes in response to severe acute hypercarbia. A total of 20 species, ranging from basal (“primitive”) to derived, were examined and 18 were observed to use preferential pHi regulation. Finally, developing amniotes (snapping turtle and American alligator) used preferential pHi regulation during severe acute respiratory acidosis, but the capacity for pHi regulation was progressively reduced throughout development.This thesis demonstrates that preferential pHi regulation is likely a common strategy of acid-base regulation occurring in response to severe acute hypercarbia in adult fishes and possibly amniotes. I propose that preferential pHi regulation is an embryonic vertebrate strategy, that has been retained or lost in adults depending on the environmental acid-base challenges they face.

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Physiological responses associated with aquatic hypercarbia in the CO2-tolerant white sturgeon, Acipenser transmontanus (2011)

Through investigations conducted at the organismal, tissue and cellular levels, this thesis provides clear evidence that the white sturgeon, Acipenser transmontanus, is among the most CO₂ tolerant of all fishes investigated to date. During moderate increases in water CO₂ tension (PCO₂) (≤ 15 mm Hg PCO₂, hypercarbia), white sturgeon exhibited changes in gill morphology and restored blood pH (pHe) through net HCO₃⁻/Cl⁻, a process observed in most fishes (Chapter 3). At CO₂ tensions lethal to other fishes (≥ 22.5 mm Hg PCO₂), white sturgeon completely protected intracellular pH (pHi) of the heart, liver, brain and white muscle (termed preferential pHi regulation), despite a large reduction in pHe (up to 1 pH unit) (Chapter 3, 4). Tissue pHi regulation was activated in heart within minutes of the onset of hypercarbia (measured via NMR, Chapter 5), and completely protected pHi in this tissue even during exposure to potentially lethal CO₂ levels (i.e., 90 mm Hg PCO₂). In hearts examined in situ, maximum cardiac performance was well defended and associated with partial pHi compensation in ventricles (which exhibited only ~40% of predicted acidosis). Preferential pHi regulation was not associated with large increases in metabolic costs, as during exposure to severe hypercarbia (~45 mm Hg PCO₂), heart [ATP] and [CrP] had recovered to pre-exposure levels within 90 min, and whole animal was decreased (30%) when pHi was completely protected. Preferential pHi regulation of this magnitude and rapidity has not been documented before in any vertebrate in response to hypercarbia and represents a novel pattern of acid-base regulation among fishes. White sturgeon represent the first exclusively water-breathing fish to exhibit preferential pHi regulation during hypercarbia. Furthermore, white sturgeon are the most basal vertebrate to demonstrate complete pHi protection during severe pHe depression. As sturgeon may retain ancestral characteristics, I propose that preferential pHi regulation is the basis for enhanced CO₂ tolerance in other tolerant Osteichthyan fishes, and first arose in association with ionoregulatory and respiratory challenges experienced during freshwater invasion in the vertebrate lineage.

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A novel mechanism for enhancing tissue oxygen delivery in teleost fishes (2010)

No abstract available.

Master's Student Supervision

Theses completed in 2010 or later are listed below. Please note that there is a 6-12 month delay to add the latest theses.

Fecal hormone metabolites as indicators of stress in the southern and northern resident killer whale (Orcinus orca) populations in coastal waters of British Columbia, Canada (2022)

Fecal hormone metabolite analysis is increasingly used as a non-invasive method to evaluate physiological stress in free-ranging animals, especially those of conservation concern. In this study, it is applied to investigate stress in the Northeast Pacific southern and northern resident killer whale (Orcinus orca) populations. The southern resident killer whale population is listed as endangered, as the population is small and declining. Despite sharing a similar diet and environment, the northern resident population shows greater reproductive success and has increased in recent years. It has been hypothesized that the southern residents are more physiologically and nutritionally stressed than the northern residents, due to anthropogenic threats of reduced prey availability, acoustic and physical disturbance, and environmental contaminants. Fecal samples were collected from both populations along the coast of British Columbia, Canada, in July and August of 2018-2019. I first validated four commercially available enzyme-linked immunosorbent assays for quantifying steroid (progestogens, androgens, and glucocorticoid) and thyroid hormone metabolites in killer whale feces, then compared concentrations of fecal glucocorticoid and thyroid metabolites between the southern and northern resident populations as indicators of stress. Mean fecal progestogen metabolites (fPM) were highest in pregnant females, higher in females (non-pregnant) than males, and were successfully used to determine pregnancies. Mean fecal androgen metabolites (fAM) were highest in pregnant females and sprouter (adolescent) males, and were greater in males relative to non-pregnant females. Mean fecal glucocorticoid metabolites (fGCM) were higher in pregnant females than non-pregnant females and males, but were not significantly different between the southern and northern resident populations. There were no significant differences in mean fecal thyroid metabolites (fTHM) among any demographics, but mean fTHM was significantly lower in the southern resident population than the northern resident population (P
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The effects of temperature acclimation and heating rate on the thermal tolerance of juvenile white sturgeon (Acipenser transmontanus) (2022)

Freshwater fish such as white sturgeon (Acipenser transmontanus) are particularly vulnerable to the effects of anthropogenic global warming; however, little is known about how acclimation to higher temperatures or rate of temperature increase affects their thermal tolerance. The Kenney dam on the Nechako River is home to the northern-most population of white sturgeon and is mandated to maintain water temperatures below 20°C for migrating sockeye salmon, but it remains unclear whether 20°C is an appropriate threshold for developing white sturgeon. To address this, 37-51 days post hatch (dph) and 66-80 dph juvenile white sturgeon were acclimated to one of four ecologically relevant temperatures (15°C, 18°C, 21°C, and 24°C) for two weeks, following which thermal tolerance (CTmax), size, condition factor, and survival were assessed. White sturgeon displayed highly plastic CTmax in response to acclimation, illustrated by a positive relationship between acclimation temperature and CTmax and large acclimation response ratios compared to other fish species. Acclimation to temperatures above 18°C was found to negatively affect condition factor, which suggests the presence of a sub-lethal threshold between 18°C and 21°C. Their highly plastic response to temperature was further demonstrated when the effect of heating rate (0.3°C/min, 0.03°C/min, 0.003°C/min) on thermal tolerance, somatic indices, and Hsp mRNA expression was assessed. White sturgeon CTmax was highest in the slowest heating rate, contrary to what has been observed in most other fish species. Hepatosomatic index decreased in all heating rates relative to control fish, indicative of the metabolic costs of thermal stress. Expression of Hsp70 mRNA was increased in all heating rates relative to controls, whereas expression of Hsp90a and Hsp90b mRNA only increased in the two slower trials. Together these data indicate that while white sturgeon have a very plastic thermal response, acclimation to temperatures above 18°C may negatively affect overall health, indicated by lower condition factor. As such, in the best interest of white sturgeon conservation, the operators of the Kenney dam may want to reconsider whether the 20°C threshold is appropriate.

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On the teleost system for enhanced oxygen unloading: time course following catecholamine removal and an investigation into other teleosts (2019)

Recent studies suggest that teleost fishes may be able to greatly enhance the amount of oxygen (O₂) unloaded to the tissues during a generalized acidosis associated with stress. This mechanism relies on pH-sensitive hemoglobin, a red blood cell (RBC) β-adrenergic Na⁺/H⁺ exchanger (β-NHE) activated by catecholamines to protect RBC pHi, and plasma-accessible carbonic anhydrase (paCA) at the tissues, but not at the gills, to short-circuit the β-NHE, acidify the RBC, and unload O₂ from hemoglobin. This system has been shown to increase tissue PO₂ (∆PO₂) by up to 30 Torr and has been proposed to play an important role both during early teleost evolution, as well as in modern teleost physiology. To date, most studies regarding this system have been conducted in the context of high stress and circulating catecholamines; little is known about this system under low catecholamine levels, such as following stress. In addition, despite being often extrapolated to all teleosts, this system has only been studied within the salmonids. Here, I investigate the time course of β-NHE short-circuiting by paCA in rainbow trout blood following in vitro stimulation by isoproterenol, a synthetic catecholamine, as well as natural catecholamines in vivo to determine how long after adrenergic stimulation this system may remain operational. A significant increase in plasma ∆PO₂ due to β-NHE short-circuiting by paCA was found in rainbow trout blood stimulated in vitro up to one hour after removal of isoproterenol; this was reduced to less than 30 min under natural catecholamines. A significant ∆PO₂ of approximately 5 Torr was determined even 6 h after stimulation, as well as in pharmaceutically blocked blood, suggesting that enhanced O₂ unloading may be operational in resting fish. The system was also investigated in two teleost species distantly related to the salmonids, cobia (Rachycentron canadum) and mahi-mahi (Coryphaena hippurus). In cobia and mahi-mahi, evidence of a RBC β-NHE was found, and it is predicted that cobia may exhibit up to a 61% increase in enhanced O₂ unloading with no change in blood flow through short-circuiting of RBC β-NHE, a value consistent with salmonids. This second phylogenetically distant group provides support that this system may be functional in teleosts in general.

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The effect of salinity and photoperiod on thermal tolerance and growth of Atlantic and coho salmon reared from smolt to adult in recirculating aquaculture systems (2019)

Land-based, closed containment salmon aquaculture involves rearing salmon from smolt to adult in recirculating aquaculture systems (RAS). Unlike in open-net pen aquaculture, rearing conditions can be specified in RAS in order to decrease physiological stress. The environmental conditions that yield optimal growth and physiological stress tolerance in salmon are, however, unknown. To address this knowledge gap, we reared Atlantic (Salmo salar) and coho (Oncorhynchus kisutch) salmon in 7 separate RAS for 400 days post-smoltification under 2 photoperiods (12 or 24 hours of light) and 4 salinities (2.5, 5, 10 or 30 ppt) and assessed the effects of these conditions on growth and thermal tolerance. We found that salinity and photoperiod had significant effects on growth of Atlantic and coho salmon, but optimal conditions were not determined. Secondly, we found Atlantic salmon generally grew best under 24 hours of light until day 400 when the trend was lost and that coho salmon grew best under 12 hours of light in the freshwater (2.5 ppt) treatment. Finally, we found higher levels of maturation in Atlantic salmon reared under 24 hours of light, whereas the 12:12 photoperiod triggered greater levels of early maturation in coho. In order to evaluate the effects of photoperiod and salinity on the physiological stress tolerance we used critical thermal maxima or CTmax tests as a performance proxy. We found that over the first 120 days post-smoltification, rearing coho under a 24 hour photoperiod resulted in a ~2°C lower CTmax than in coho reared under a 12:12 photoperiod. This photoperiod effect did not persist at 200 and 400 days, which was coincident with an overall decrease in CTmax in coho salmon relative to 120 days. Finally, Atlantic salmon had a higher CTmax (~28°C) compared to coho (~26°C) at 400 days post-smoltification. Overall, these findings are important for the future implications of RAS and for the aquaculture industry to help identify physiologically sensitive time stages.

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The effects of embryonic incubation temperature on subsequent development, growth, and thermal tolerance through early ontogeny of white sturgeon (2019)

An organism’s phenotypic characteristics can be altered by environmental variations experienced during embryonic development. These changes can persist into adulthood. Increasing global temperatures are a current concern that may be particularly acute for species already threatened or endangered, such as the white sturgeon, Acipenser transmontanus. Given scant information on the effects of early embryonic temperatures on subsequent physiological parameters such as thermal tolerance, development, and growth in this species, the effects of global climate change on the future of white sturgeon populations is uncertain. To address this, white sturgeon embryos were incubated at 12, 15, and 18 °C until hatch, after which fish were reared at a common 15 °C for 80 days post-hatch. I conducted acute thermal tolerance tests through early ontogeny to determine how relative thermal tolerance (CTmax) changed and differed during this period while also sampling and evaluating various morphometric characteristics to determine the effects of embryonic temperatures on growth and development. Embryonic incubation at 12 °C with subsequent rearing at 15 °C increased larval development rate, growth rate, and the development of thermal tolerance; however, once developed CTmax averaged 29.9 for all embryonic rearing temperatures. For white sturgeon, embryonic temperatures appear to have discernable effects on growth and development up to 80 days post-hatch but not on thermal tolerance.

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Carbonic anhydrase in the gills and blood of chondrichthyan fishes (2018)

Carbon dioxide (CO₂) is continuously produced as a result of aerobic respiration and must be excreted to maintain internal acid-base balance. Most CO₂ is carried in the blood as HCO₃- and must be converted back to molecular CO₂ at the respiratory surface to diffuse out into the environment. The uncatalyzed rate of HCO₃- dehydration is too slow to excrete CO₂ at a physiologically relevant rate and therefore it must be catalyzed by the enzyme carbonic anhydrase. The distribution of carbonic anhydrase in the blood and gills of fish therefore provides important information about general patterns of gas exchange and acid-base balance. Teleost fishes have a fast CA in the red blood cell (RBC), no extracellular CA activity, an endogenous plasma CA inhibitor and a relatively low plasma buffer value so HCO₃- dehydration is largely restricted to the RBC. Pacific spiny dogfish (Squalus suckleyi), however, have a slow RBC CA, extracellular CA activity, no endogenous plasma CA inhibitor, plasma accessible CAIV at the gills and a relatively high plasma buffer value, implying that both the RBC and plasma compartments may contribute to HCO₃- dehydration. This thesis uses biochemical assays, subcellular localization and immunohistochemistry on blood and gill samples from 13 chondrichthyan species to examine whether the characteristics of the dogfish model of CO₂ excretion apply to chondrichthyan fishes in general. Overall, the results of this study were consistent with the proposed chondrichthyan model of CO₂ excretion because most chondrichthyans had lower RBC CA activity than teleosts, some extracellular CA activity, no endogenous plasma CA inhibitor, higher plasma buffer values and type IV-like CA at the gills. Pacific spiny dogfish had 3x more microsomal CA activity (183 ± 13.2 µmol CO₂ min-1 mg protein-1) in the gills than the other three species examined for this trait, indicating that dogfish may not be a representative species to compare with other vertebrate groups. Overall, the results of this thesis suggest that all chondrichthyans have the capacity to use both the plasma and RBC compartments for CO₂ excretion and these data provide important information about general patterns of gas exchange and acid-base balance in fishes.

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Effects of salinity and photoperiod on growth, aerobic scope, and hypoxia tolerance of Atlantic and coho salmon in recirculating aquaculture systems (2018)

Recirculating aquaculture systems (RAS) are an emerging technique in aquaculture to rear salmon in land-based facilities, but the systems are also associated with high costs, so rearing fish under optimal conditions for maximum growth is required for profitability. However, few systematic studies have been conducted to determine optimal conditions for growth of salmon from smolt through to market size in RAS and the related effects on physiological performance. To address this knowledge gap, the first part of my thesis investigated the effect of salinity on growth, metabolism and hypoxia tolerance of Atlantic and coho salmon. Smolts were reared at salinities of 0, 5, 10, 20 and 30 ppt under 24 h of light in RAS for up to 460 days. Between Days 200 and 400, respirometry was conducted to measure routine, maximum metabolic rate and aerobic scope, while time to loss of equilibrium at 10% air saturation was measured to determine hypoxia tolerance. The initial effect of salinity on growth was found in Atlantic and coho salmon at Day 295 and 59, respectively, after which growth was generally enhanced at intermediate salinities of 5 and 10 ppt. No clear relationship was found between salinity and metabolic measurements in either species. Hypoxia tolerance of Atlantic salmon was enhanced at 5 and 10 ppt, but salinity had no effect on hypoxia tolerance of coho salmon. The second part of this thesis aimed to (1) determine metabolism and hypoxia tolerance of coho salmon during their early growth stages where the effect of salinity on growth was the most profound and (2) explore the interactive effect of photoperiod. A new cohort of coho salmon smolts were reared in RAS at 2.5, 5, 10 and 30 ppt under 12:12 and 24:0 (light:dark) photoperiods, while respirometry and hypoxia trials (15% air saturation) were conducted at Day 60 and 120. No effect of salinity and photoperiod was found on metabolic measurements and hypoxia tolerance in the younger coho salmon during these periods. Overall, my data suggest that there is some potential to enhance growth of salmon by manipulating environmental conditions in RAS without compromising other physiological performance.

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The effects of salinity and photoperiod on growth and swimming performance in Atlantic and coho salmon raised in recirculating aquaculture systems from smolt to adult (2018)

Salmon are among the most popular seafood products and their culture continues to expand with improved aquaculture technology. Typical aquaculture production rears salmon from smolt to market size in net-pens in oceans, but this practice has been criticized due to potential environmental concerns such as eutrophication and interactions between escaped cultured fish and wild populations. Rearing fish in recirculating aquaculture systems (RAS) is a new technology to address some of the concerns over net-pen aquaculture, as well as enhance production of salmonids and other fish species. Salmon can inhabit a wide range of salinities with different osmoregulatory costs, but presumably these costs can be reduced if fish reside in water isosmotic to their blood plasma. Photoperiod has been shown to affect growth rates in salmon at different life stages but can also affect early maturation in salmon. To examine the effects of salinity and photoperiod on the growth of salmon, seven replicate RAS with salinities of 2.5, 5, 10 and 30ppt under 12:12 and 24:0 light:dark photoperiod were used to rear Atlantic and coho salmon from smolts onwards for 120 days. Salinity and photoperiod had an effect on Atlantic salmon growth, with those reared at 10ppt in 24:0 light showing the highest growth rates. However, neither photoperiod nor salinity affected coho salmon growth. To understand the effects of salinity and photoperiod on swimming performance and hematology, coho salmon from two separate studies underwent repeat maximum swimming speed (Umax) tests. In the first study, Umax was assessed in coho salmon that were reared for 350 days in 0 and 10ppt. In the second study, Umax was assessed in coho salmon that were reared for 60 and 150 days in 2.5, 10 and 30ppt under 12:12 and 24:0 light:dark photoperiod. In the first study, salinity had significant effects on resting plasma osmolality and chloride concentration. In the second year, salinity affected first Umax, but neither salinity nor photoperiod affected repeated Umax and recovery ratio. There were also significant effects of salinity on the hematocrit, hemoglobin concentration, MCHC and plasma osmolality and chloride concentration of exhausted salmon after repeated swimming tests.

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The relationship between thermal tolerance and hypoxia tolerance in Amazonian fishes (2018)

The Amazon contains 20% of the world’s freshwater fish species that are predicted to experience an increase in temperature by up to 2.2 to 7˚C within the next century. An increase in temperature will likely be associated with an increase in the frequency, duration, and magnitude of hypoxic bouts, creating an even greater challenge. Thermal tolerance may be limited by the ability to supply and deliver enough oxygen to tissues at critical temperatures, as is the case for hypoxia tolerance, thus both may be associated with similar mechanisms in fish. A direct relationship between thermal and hypoxia tolerance however, has not yet been investigated in a wide range of fish species. To address this, I conducted acute thermal tolerance (CTMax) and hypoxia tolerance (% air saturation at loss of equilibrium) assays in 20 species that spanned a broad phylogenetic range. In fish acclimated to the temperatures within the current temperature range of the Amazon River (28 or 31˚C), I found a positive relationship between CTMax and hypoxia tolerance. In fish acutely transferred to higher temperatures of 33 or 35˚C, there was a reduction in hypoxia tolerance relative to that at 28 or 31˚C. Acclimation (10 days or 4 weeks) to 33 or 35˚C did not increase hypoxia tolerance, and in some species there was a further reduction in hypoxia tolerance. Acclimation to 33 or 35˚C (10 days or 4 weeks) and exposure to hyperoxia (>200% air saturation) increased CTMax, although in most species only moderately. One of the most significant findings of my thesis was that most species failed to acclimate to the higher temperatures: of the 13 species investigated, 2 species did not survive 10 days or 4 weeks (chronic lethal maximum) at 31˚C, 9 species did not survive at 33˚C, and only 2 species survived 35˚C. Overall, acclimation to higher temperatures that are predicted to occur within the next century had little or no effect on thermal tolerance and reduced hypoxia tolerance indicating that the high fish biodiversity of the Amazon may be at risk given the predicted changes in temperature and hypoxia associated with climate change.

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The effects of water ionic composition on acid-base regulation in rainbow trout, Oncoryhnchus mykiss, during hypercarbia at rest and during sustained exercise (2015)

Rising atmospheric carbon dioxide (CO₂) and by association pCO₂ in aquatic habitats (hypercarbia) has put increased focus on understanding the underlying mechanisms of acid-base regulation in fish. Aquatic hypercarbia results in a blood acidosis in fish, which is compensated for by the exchange of Na+ and Clˉ for its acid/base counterpart (H+, HCO₃ˉ) across the gill epithelium. Surprisingly, there are no studies on how a single species, capable of inhabiting both fresh and saltwater, responds to hypercarbia, and no existing studies examining how sustained exercise may affect hypercarbia recovery. The goal of this thesis was to examine how changes in ambient water ionic composition (soft-, hard-, and saltwater) affect the rate and degree of acid-base compensation in a rainbow trout, Oncorhynchus mykiss, during hypercarbia, at rest and during sustained exercise. Additionally, I sought to determine the effect of sustained exercise on the rate and degree of acid-base compensation during hypercarbia. Trout were acclimated to soft-, hard-, or saltwater and acid-base relevant blood parameters were measured during a 1% CO₂ hypercarbia exposure, both at rest and during sustained exercise (~60% Ucrit). After 48h of hypercarbia, resting hard-, and saltwater acclimated trout had fully restored blood pH, however soft water acclimated trout were only 60.6±10.5% recovered, and in all fish recovery was associated with an increase in plasma [HCO₃ˉ] and a decrease in plasma [Clˉ] of similar magnitude. Trout exposed to hypercarbia during sustained exercise had a similar response, and following 8h the saltwater acclimated fish had fully restored blood pH, while soft-, and hard water fish were 42±18.1 and 64±6.8% recovered, respectively. Furthermore, following 8h of hypercarbia there was a significant effect of exercise on the degree of recovery compared with resting fish, suggesting that sustained exercise results in a more rapid recovery from hypercarbia in trout, relative to rest. These results provide intra-specific support to previous studies that demonstrate marine fish compensate for hypercarbia faster than freshwater fish. This thesis not only demonstrates an important link between ambient water ion levels and the ability to recover from acid-base disturbances, and presents novel data suggesting sustained exercise enhances acid-base regulation.

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The Effect of Climate Change-Related Environmental Acidification on the Growth, Development and Energetics of the Early Life Stages of Pink Salmon (Oncorhynchus gorbuscha) y (2014)

As a consequence of increasing atmospheric CO₂, the partial pressure of CO₂ (pCO₂) in the oceans is rising, causing a decrease in pH and carbonate ions known as ocean acidification (OA). Since freshwater systems have the same potential for atmospheric equilibrium with CO₂, similar scenarios of acidification will occur in freshwater, yet little is known about the potential impacts of climate change-related acidification on freshwater species and ecosystems. Moreover, virtually no research has investigated the effect of oscillating pCO₂ tensions on fish, which are more likely to reflect natural coastal conditions. The goal of this thesis is to address some of these knowledge gaps by studying the potential effects of climate change-related acidification on pink salmon (Oncorhynchus gorbuscha) at a sensitive and critical life-stage during development in both freshwater and seawater under future elevated and fluctuating pCO₂ tensions (freshwater: 400 μatm, 1000 μatm, 2000 μatm, 400-2000 μatm (over 24hr); seawater: 400 μatm, 1600 μatm, 400-1600 μatm). Growth, production efficiencies and aerobic scope were measured starting 2 weeks pre-hatch in freshwater up until 2 weeks post-seawater transfer. Growth was reduced during freshwater rearing and following seawater transfer. Specifically, size, production efficiencies and absolute growth rates were reduced at freshwater tensions of 1000 and 2000 μatm and seawater tensions of 1600 μatm. However, no significant effects on growth were seen in response to oscillating pCO₂ tensions. Similarly, aerobic scope was reduced at high pCO₂ following seawater transfer through a reduction in maximal oxygen consumption rate (ṀO₂max) (but not routine (ṀO₂routine)), indicating that exercise in pink salmon fry may be particularly affected by climate change-related acidification. Given that control fish exhibit a dramatic increase in ṀO₂max 7 days post-seawater transfer, which is likely associated with a change in life history from a sedentary to a migratory stage, elevations in seawater pCO₂ may have implications for their seaward migration success. Overall, this thesis demonstrates that pink salmon, under predicted future increases in pCO₂, may be faced with sublethal impacts of acidification on various aspects of their physiology at a very critical time in their life history.

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The ontogeny of sodium balance of rainbow trout (Oncorhynchus mykiss) and pink salmon (Oncorhynchus gorbuscha) during post-embryonic development in freshwater (2014)

This thesis contributes to our general knowledge of ionoregulatory function in developing fish by characterizing the functional ontogeny of sodium (Na⁺) balance in salmonids reared in freshwater during early development. Chapter 2 investigated the plasticity of Na⁺ balance and transport capacity during a critical developmental transition from cutaneous-dominated to gill-dominated ionoregulation in the model teleost fish, the rainbow trout. Fish experienced very high resting unidirectional Na⁺ uptake rates early in development which were reduced to values typical of adults following yolk absorption. Maximal uptake rate (Jmax) for Na⁺ was high during early development and decreased following yolk absorption while uptake affinity decreased (Km increased) following hatch and increased following yolk absorption. It appeared that early in development, high Na⁺ uptake rates across cutaneous ionocytes were driven by high maximal uptake rate, while the gill ionocytes that dominate ionoregulation post-yolk absorption had an increased affinity for Na⁺. Following hatch, when ionoregulation occurs predominantly across cutaneous ionocytes, larval fish exhibited little ionoregulatory plasticity in their Na⁺ uptake rates and Na⁺ uptake kinetics. As ionoregulation shifts to the gill, developing fish exhibited increased uptake affinity for Na⁺ in low-[Na⁺] environments as observed in adult fish; however, maximal uptake rate for Na⁺ did not increase in low-[Na⁺] environments as seen in adult fish, suggesting that the capacity to overcome Na⁺-poor environments may be limited and still developing at this stage. Chapter 3 contributed to our understanding of Na⁺ transport during early salmonid development and explored Na+ transport characteristics employed by the early-migrating anadromous salmonid, the pink salmon. It was clear that heightened and increasing whole-body [Na⁺] during yolk absorption in freshwater was not a unique characteristic of developing pink salmon associated with preparation for early ocean entry. This trait was shared by the non-anadromous rainbow trout. Interestingly, the mechanism by which pink salmon and rainbow trout achieved high whole-body [Na⁺] during early development did not appear to be the same. Rainbow trout experienced increasing Na⁺ uptake rates during development while pink salmon did not alter Na⁺ uptake rates during development, suggesting that pink salmon regulated whole-body [Na⁺] via modulation of Na⁺ efflux rates

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The Development of Salinity Tolerance in Juvenile Pink Salmon (Oncorhynchus Gorbuscha) (2012)

Following yolk-sac absorption and gravel emergence pink salmon (Oncorhynchus gorbuscha) migrate into seawater (SW) at as small as 0.2 g. This life-history strategy is in contrast with most anadromous salmonid species that generally spend 1-2 years growing in fresh water (FW) and physiologically preparing for life in SW before they migrate to SW as smolts. This study characterized for the first time the ontogeny of SW tolerance in pink salmon around the period of yolk-sac absorption. Post-hatch juvenile pink salmon were either held in FW for 26 weeks or transferred to SW every two weeks for 20 weeks to follow % survival, whole body (WB) Na+ and water content, as well as changes in wet and dry mass, gill Na+K+ATPase (NKA) activity and α1a and α1b mRNA isoform expression. An increase in gill NKA activity and the ratio of the α1b/α1a isoform expression, a plateau in WB water and Na+ levels, and the switch from catabolic to anabolic growth were all observed at the time of yolk-sac absorption in fish retained in freshwater. At this time, morbidity following subsequent SW transfer fell to 0% from a high of 100% for newly hatched alevins, but then rose to 25% in older fry, suggesting that a window of increased salinity tolerance exists for pink salmon at the time of yolk-sac absorption. This proposed window of SW tolerance is similar to the smolt window that has been identified for other salmonids; but in pink salmon appears to be endogenously mediated, as fish were reared under constant (12L:12D) photoperiod and at 5˚C throughout the study. Moreover, smoltification is incomplete since transfer to SW further elevated gill NKA activity and increased gill NKA α1b/α1α isoform expression ratio 8-fold at yolk-sac absorption. Thus, even the most SW-tolerant fish were not fully prepared for SW before entry, but responded directly to SW by further increasing hypo-osmoregulatory ability. This study filled the previously existing void of knowledge regarding the acquisition of salinity tolerance in juvenile pink salmon.

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Regulation of intracellular pH during exposure to hypercapnia in a rainbow trout hepatoma cell line, RTH 149, and in white sturgeon, Acipenser transmontanus, primary liver cells (2011)

No abstract available.

Of saline and sea lice: hydromineral challenges and osmoregulatory strategies associated with early ocean entry of juvenile pink salmon (Oncorhynchus gorbuscha) (2010)

Pink salmon (Oncorynchus gorbuscha) enter seawater (SW) following gravel emergence at a body mass of 0.2 g. Two hydromineral challenges associated with this remarkable early ocean entry were investigated: (1) initial exposure to a hyper-osmotic environment and (2) sea louse (Lepeophtheirus salmonis) parasitism. To survive SW, pink salmon were hypothesized to develop hypo-osmoregulatory abilities as larval alevins prior to natural SW entry as post-larval fry. To test this, alevins and fry were transferred from freshwater (FW) in darkness to SW under a simulated natural photoperiod (SNP). Ionoregulatory status was assessed at 0, 1 and 5 days post-transfer. Alevins showed no evidence of hypo-osmoregulation, marked by a loss of water balance, a 35% increase in body [Cl-], and no change in gill Na⁺/K⁺-ATPase (NKA) activity. Conversely, fry maintained water balance and increased gill NKA activity by 50%. Fry gill NKA activity also increased by 50% following exposure to SNP in FW, providing the first evidence of photoperiod-triggered smoltification for pink salmon. A 15% increase in fry body [Na⁺] was observed as well, perhaps representing a novel mechanism for maintaining water balance during ocean entry. Physical damage to the host epidermis is a primary proximal effect of louse infection. Such damage may exacerbate existing hydromineral flux in SW. To test this, ionoregulatory status was measured in pink salmon of varying size with and without attached-stage lice. In laboratory-infected fish (~1 wk SW; 0.2-0.4 g), body [Na⁺] increased by 12% when infected with 1 chalimus IV louse, and by 23% with 2-3 chalimus III lice. Mortality was 6%. In wild-infected fish (~4-12 wks SW; 0.5-1.5 g), body [Na⁺] did not differ from controls. Combining data sets revealed a “no effect” fish size threshold of 0.5 g for 1 chalimus IV louse. This threshold is partly due to increasing hypo-osmoregulatory ability. Pink salmon thus appear to possess a novel hypo-osmoregulatory strategy where ion balance is sacrificed to maintain water balance prior to maximum ion excretion capacity. Out-migrating fish are particularly vulnerable to sea louse parasitism at this time, and as such, BC fish farms have relocated to minimize interactions during this critical period.

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