Intercohort cannibalism and parturition-associated behaviour of captive-bred swordtail, Xiphophorus helleri (Pisces: Poeciliidae)
- Jones, Clifford Louis Wilshire
- Authors: Jones, Clifford Louis Wilshire
- Date: 2003
- Subjects: Poeciliidae -- Behavior Xiphophorus helleri Fish culture
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5351 , http://hdl.handle.net/10962/d1007812
- Description: Adult fish that belong to the family Poeciliidae cannibalise juveniles, both in the wild and under captive conditions, but this behaviour has only been partly investigated in the Poeciliidae in some of the commercially valuable species. The objective of the research is to develop an understanding of intercohort cannibalism and parturition-associated behaviour in captive-bred swordtail (Xiphophorus helleri), with applications to industry and future research of other poeciliids. Experiments investigating the effect of adult stocking density and sex ratio on the production of juveniles were used to determine if cannibalism occurs under culture conditions. The average rate of intercohort cannibalism ranged from (5.5 to 53.9%), and was positively density dependent and independent of sex ratio, indicating that males and females were probably equally cannibalistic. The highest number of juveniles (1725.7±141.4) produced per tank over 70 days was obtained from two males and eight females. To develop a better understanding of adult and juvenile behaviour during parturition, fish were observed under controlled laboratory conditions using video and behaviours such as attack (burst of speed by an adult in the direction of a juvenile), escape (avoidance of cannibalism after attack) and cannibalism (predation of a live juvenile by an adult), for example, were identified. Under laboratory conditions most young escaped in downward direction after attack (49%) and most utilised the refuge made that was made available. Furthermore, most attacks (62-65%) and cannibalism (57-84%) occurred at the bottom. Since the presence of refuge significantly increased the rate of juvenile survival under culture conditions, it was hypothesised that the rate of cannibalism could be reduced under farming conditions if juveniles were protected when they escaped downwards. This hypothesis was accepted as it was found that refuge at the bottom of the water column or the inclusion of a false-bottom reduced the rate of cannibalism by 49% and 72%, respectively. Similarly, the hypothesise that the rate of cannibalism could be reduced if juveniles where given protection when escaping sideways (32% of juveniles escaped sideways in the laboratory) was also accepted when tested under farm-scale conditions because a false-side reduced the rate of cannibalism by an average of 45%. Since males and females were equally responsible for cannibalising juveniles in the laboratory, it was hypothesised that the rate of cannibalism would decrease proportionately with the removal of males (Le. 20% of the cannibals) from the population; the removal of males under farming conditions resulted in a 19.5% reduction in the rate of cannibalism. Since older juveniles were better able to escape cannibalism than neonates and since adults habituate to stimuli that previously resulted in attack behaviour, it was hypothesised that the rate of cannibalism would remain unaffected by the length of time that juveniles were exposed to adults in the breeding tanks. This hypothesis was also accepted when tested under farm conditions. However, some hypotheses based on laboratory observations were not accepted. For example, a constant low light intensity did not appear to decrease the rate of cannibalism under farm conditions; also, the occurrence of dead and deformed juveniles went unnoticed in the laboratory, and under farm conditions, where adults did not have access to the bottom of the tank, 10% of the harvest consisted of dead and deformed juveniles. It is concluded that technologies, such as bottom-refuge or a false-side, that increase the size of the liveharvest and allow for the removal of potentially less viable offspring are recommended for the commercial production of poeciliids. The overall similarity of X. helleri behaviour between the laboratory experiments and the farm-scale trials suggests that the post-partum behaviour of X. helleri remains consistent under these different conditions; thus, behaviour under one set of conditions may be used to predict behaviour under other conditions. The application and significance of extrapolations to industry and future research of X. helleri and possibly other poeciliids were discussed and the most applicable laboratory observations with the highest extrapolation capacity were proposed. Furthermore, techniques were developed to aid industry and future researchers in making predictions relating to behaviour of X. helleri under different conditions based on laboratory observations. The results were used to develop a model indicating that selection pressures against cannibalism are not likely to exist at the rate of cannibalism observed here since the potential genetic gain through kin survival and inclusive fitness was shown to be greater than any potential genetiC loss experienced by a victim of cannibalism. The model was successfully tested under a range of social conditions. Other possible explanations for cannibalism in poeciliids, such as parental manipulation, nutritional advantages, opportunistic predation and the recovery of energy are discussed. It is suggested that the most likely proximate cause of cannibalism under captive conditions is opportunistic predation. The theory that cannibalism ensures that only viable genes of the victim are expressed, through inclusive fitness, is a possible ultimate cause of cannibalism, which may have been inherited from feral ancestors of captive-bred X. helleri.
- Full Text:
- Authors: Jones, Clifford Louis Wilshire
- Date: 2003
- Subjects: Poeciliidae -- Behavior Xiphophorus helleri Fish culture
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5351 , http://hdl.handle.net/10962/d1007812
- Description: Adult fish that belong to the family Poeciliidae cannibalise juveniles, both in the wild and under captive conditions, but this behaviour has only been partly investigated in the Poeciliidae in some of the commercially valuable species. The objective of the research is to develop an understanding of intercohort cannibalism and parturition-associated behaviour in captive-bred swordtail (Xiphophorus helleri), with applications to industry and future research of other poeciliids. Experiments investigating the effect of adult stocking density and sex ratio on the production of juveniles were used to determine if cannibalism occurs under culture conditions. The average rate of intercohort cannibalism ranged from (5.5 to 53.9%), and was positively density dependent and independent of sex ratio, indicating that males and females were probably equally cannibalistic. The highest number of juveniles (1725.7±141.4) produced per tank over 70 days was obtained from two males and eight females. To develop a better understanding of adult and juvenile behaviour during parturition, fish were observed under controlled laboratory conditions using video and behaviours such as attack (burst of speed by an adult in the direction of a juvenile), escape (avoidance of cannibalism after attack) and cannibalism (predation of a live juvenile by an adult), for example, were identified. Under laboratory conditions most young escaped in downward direction after attack (49%) and most utilised the refuge made that was made available. Furthermore, most attacks (62-65%) and cannibalism (57-84%) occurred at the bottom. Since the presence of refuge significantly increased the rate of juvenile survival under culture conditions, it was hypothesised that the rate of cannibalism could be reduced under farming conditions if juveniles were protected when they escaped downwards. This hypothesis was accepted as it was found that refuge at the bottom of the water column or the inclusion of a false-bottom reduced the rate of cannibalism by 49% and 72%, respectively. Similarly, the hypothesise that the rate of cannibalism could be reduced if juveniles where given protection when escaping sideways (32% of juveniles escaped sideways in the laboratory) was also accepted when tested under farm-scale conditions because a false-side reduced the rate of cannibalism by an average of 45%. Since males and females were equally responsible for cannibalising juveniles in the laboratory, it was hypothesised that the rate of cannibalism would decrease proportionately with the removal of males (Le. 20% of the cannibals) from the population; the removal of males under farming conditions resulted in a 19.5% reduction in the rate of cannibalism. Since older juveniles were better able to escape cannibalism than neonates and since adults habituate to stimuli that previously resulted in attack behaviour, it was hypothesised that the rate of cannibalism would remain unaffected by the length of time that juveniles were exposed to adults in the breeding tanks. This hypothesis was also accepted when tested under farm conditions. However, some hypotheses based on laboratory observations were not accepted. For example, a constant low light intensity did not appear to decrease the rate of cannibalism under farm conditions; also, the occurrence of dead and deformed juveniles went unnoticed in the laboratory, and under farm conditions, where adults did not have access to the bottom of the tank, 10% of the harvest consisted of dead and deformed juveniles. It is concluded that technologies, such as bottom-refuge or a false-side, that increase the size of the liveharvest and allow for the removal of potentially less viable offspring are recommended for the commercial production of poeciliids. The overall similarity of X. helleri behaviour between the laboratory experiments and the farm-scale trials suggests that the post-partum behaviour of X. helleri remains consistent under these different conditions; thus, behaviour under one set of conditions may be used to predict behaviour under other conditions. The application and significance of extrapolations to industry and future research of X. helleri and possibly other poeciliids were discussed and the most applicable laboratory observations with the highest extrapolation capacity were proposed. Furthermore, techniques were developed to aid industry and future researchers in making predictions relating to behaviour of X. helleri under different conditions based on laboratory observations. The results were used to develop a model indicating that selection pressures against cannibalism are not likely to exist at the rate of cannibalism observed here since the potential genetic gain through kin survival and inclusive fitness was shown to be greater than any potential genetiC loss experienced by a victim of cannibalism. The model was successfully tested under a range of social conditions. Other possible explanations for cannibalism in poeciliids, such as parental manipulation, nutritional advantages, opportunistic predation and the recovery of energy are discussed. It is suggested that the most likely proximate cause of cannibalism under captive conditions is opportunistic predation. The theory that cannibalism ensures that only viable genes of the victim are expressed, through inclusive fitness, is a possible ultimate cause of cannibalism, which may have been inherited from feral ancestors of captive-bred X. helleri.
- Full Text:
On the use of metabolic rate measurements to assess the stress response in juvenile spotted grunter, Pomadasys commersonnii (Haemulidae, Pisces)
- Authors: Radull, John
- Date: 2003
- Subjects: Fishes -- Metabolism Fishes -- Physiology Pomadasys -- Physiology Grunts (Fishes) -- Physiology Stress (Psychology) Stress (Physiology)
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5350 , http://hdl.handle.net/10962/d1007564
- Description: Quantitication of stress requires the use of a stress indicator that is easy to measure, and which can be readily interpreted in terms of the potential long-term effects to an organism. This study evaluates the suitability of metabolic rate as an indicator of the stress response in fish. By comparing the metabolic with the cortisol stress response, the most commonly used indicator of stress in fish, it was possible to assess the suitability of metabolic rate as a stress indicator. Changes in metabolic rate were used to predict the long-term effects of transport-related stressors. This study also detennined the baseline metabolic rates of the tish. The standard and the active metabolic rates of juvenile P. cummersonnii were 0.16 ± 0.02 (mean ± S.D, n = 6) mg O₂g⁻¹h⁻¹, and 0.56 ± 0.04 mg O₂g⁻¹h⁻¹, respectively, whereas the routine metabolic rate for the fish was 0.25 ± 0.03 mg O₂g⁻¹h¹. The relationship between metabolic rate and body weight was described by the equation ϺO₂ = 0.64 W⁻°·³⁸. 24-h oxygen consumption measurements showed that juvenile P. commersonnii exhibited diel rhythmicity in oxygen consumption rate, the higher rates occurring at night and the lower rates during the daytime. The higher nocturnal metabolic activity may have been due to increased activity induced by an endogenous rhythm related to feeding. Diel rhythmicity has direct implications for the measurement of baseline metabolic rates since it could result in overestimation or underestimation of these rates. 24-h continuous oxygen consumption measurements enabled the detection of the rhythmicity in oxygen consumption rate, and thereby ensured a greater degree of accuracy in the estimation of these parameters. The metabolic stress response in juvenile P. commersonnii was best described by the equation, y = -0.0013 x² + 0.0364 x ÷ 0.3052, where x = time after application of stressor, and y = oxygen consumption rate. Using the derivative of this equation, the metabolic stress response was estimated to peak approximately 14 min after application of a simulated capture and handling stressor. Oxygen consumption increased by about 300 % as a result of the stress. Approximately 15 min after application of a similar stressor, plasma cortisol levels in stressed fish was 200 % higher than baseline levels. However, cortisol levels in fish sampled 30 min after the disturbance was similar to the baseline cortisol levels, indicating that full recovery had occurred. Although the patterns in the metabolic and cortisol stress responses were similar, metabolic rate could be measured continuously, thereby ensuring accurate interpretation of the data. Furthermore, increases in metabolic rate during the stress response are a culmination of physiological events from the primary to the tertiary levels of biological organization and are, therefore, easier to interpret in terms of long-term effects on the fish. Different transportation procedures elicited variable degrees of stress in juvenile P. commersonnii. The cost of metabolism attributed to the effects of capture and handling was twice as much as that attributed to acute temperature elevation. Acute temperature decrease resulted in a signiticant reduction in the oxygen consumption rate (ANOVA, P < 0.05). Oxygen consumption by the fish was not affected by fish density (ANOVA: F = 2.002, P = 0.5), or by oxygen depletion at dissolved oxygen concentrations above the critical level. Below this level, however, oxygen consumption decreased linearly with further decrease in dissolved oxygen concentration. These results showed that the highest energetic cost to juvenile P. commersonnii was incurred as a result of capture and handling. The results also showed that by subjecting fish to different stressors, it was possible to categorize them according to their relative metabolic costs to the fish. At 25º C, the effective concentration of 2-phenoxyethanol to fully anaesthetize (Stage IV, McFarland 1960) juvenile P. commersonnii was 0.4 ml l⁻¹ and the most appropriate concentration for deep sedation (Stage II, McFarland 1960) of the fish for at least 24 h was 0.2 ml l⁻¹. A maximum of 3 minutes was required by the fish to recover from the effects of the anaesthetic. There was no correlation between fish weight and the rate of induction of anaesthesia (r² = 0.001, p = 0.3). At the peak of the metabolic stress response, oxygen consumption was twice as high in the un-anaesthetized fish compared to the fish anaesthetized after the application of the simulated capture and handling stressor, suggesting that anaesthetization with 2-phenoxyethanol may have reduced the effect of the disturbance on the fish. Similar oxygen consumption rates for the fish anaesthetized prior to capture and the non-stressed fish suggested that the increases in metabolic rate could be linked to the struggling associated with attempts by fish to escape from the perceived stressor. Anaesthetization of juvenile P. commersonnii with 0.3 ml l⁻¹ 2-phenoxyethanol resulted in a more than 200 % increase in plasma cortisol concentration. The elevated levels of plasma cortisol in the anaesthetized fish suggested a manifestation of 2-phenoxyethanol as a stressor. At the time of capture, cortisol levels in fish that were anaesthetized prior to capture were the same as those measured in the disturbed fish at the peak of the stress response (ANOVA, p = 0.95), suggesting that the anaesthetized fish were already experiencing considerable stress at the time they were captured. Undisturbed juvenile P. commersonnii that were anaesthetized for 1 h also had cortisol levels that were five times higher than those measured in undisturbed-unanaesthetized fish, indicating that the duration of exposure to the anaesthetic had a significant effect on plasma cortisol levels. The results presented in this study demonstrate the usefulness of metabolic rate as an indicator of acute stress in fish. This was achieved by comparing the metabolic and the cortisol stress responses. The ease and accuracy with which oxygen consumption of fish could be measured made it possible to measure the stress response more accurately than by plasma cortisol concentration. It was also possible to monitor metabolic rate continuously over a long duration using polarographic oxygen sensors, thus enabling a better evaluation of the stress response. These results, thus, suggest that metabolic rate measurements could be a more practical way to quantify the effects of acute stressors on juvenile fishes. By detailing the profile of the metabolic stress response in P. commersonnii, this study makes a contribution towards understanding the physiological effects of stress in fishes. The study also contributes towards the quantification of baseline metabolic rates of this species under captivity. This study also contributes towards understanding the effects of 2-phenoxyethanol on the stress physiology of fish. By anaesthetizing fish under different conditions of stress, it was possible to evaluate the effect of 2-phenoxyethanol on the metabolic stress response. The ability of 2-phenoxyethanol to reduce physical activity of the fish, and thereby reduce the impact of acute stress on the metabolic stress response, makes it a good agent for the mitigation of stress during the capture and handling of fish. However, the increase in plasma cortisol concentration during prolonged anaesthetization using this drug suggests that the anaesthetic might be a stressor to fish and may, therefore, not be suitable for long-term sedation.
- Full Text:
- Authors: Radull, John
- Date: 2003
- Subjects: Fishes -- Metabolism Fishes -- Physiology Pomadasys -- Physiology Grunts (Fishes) -- Physiology Stress (Psychology) Stress (Physiology)
- Language: English
- Type: Thesis , Doctoral , PhD
- Identifier: vital:5350 , http://hdl.handle.net/10962/d1007564
- Description: Quantitication of stress requires the use of a stress indicator that is easy to measure, and which can be readily interpreted in terms of the potential long-term effects to an organism. This study evaluates the suitability of metabolic rate as an indicator of the stress response in fish. By comparing the metabolic with the cortisol stress response, the most commonly used indicator of stress in fish, it was possible to assess the suitability of metabolic rate as a stress indicator. Changes in metabolic rate were used to predict the long-term effects of transport-related stressors. This study also detennined the baseline metabolic rates of the tish. The standard and the active metabolic rates of juvenile P. cummersonnii were 0.16 ± 0.02 (mean ± S.D, n = 6) mg O₂g⁻¹h⁻¹, and 0.56 ± 0.04 mg O₂g⁻¹h⁻¹, respectively, whereas the routine metabolic rate for the fish was 0.25 ± 0.03 mg O₂g⁻¹h¹. The relationship between metabolic rate and body weight was described by the equation ϺO₂ = 0.64 W⁻°·³⁸. 24-h oxygen consumption measurements showed that juvenile P. commersonnii exhibited diel rhythmicity in oxygen consumption rate, the higher rates occurring at night and the lower rates during the daytime. The higher nocturnal metabolic activity may have been due to increased activity induced by an endogenous rhythm related to feeding. Diel rhythmicity has direct implications for the measurement of baseline metabolic rates since it could result in overestimation or underestimation of these rates. 24-h continuous oxygen consumption measurements enabled the detection of the rhythmicity in oxygen consumption rate, and thereby ensured a greater degree of accuracy in the estimation of these parameters. The metabolic stress response in juvenile P. commersonnii was best described by the equation, y = -0.0013 x² + 0.0364 x ÷ 0.3052, where x = time after application of stressor, and y = oxygen consumption rate. Using the derivative of this equation, the metabolic stress response was estimated to peak approximately 14 min after application of a simulated capture and handling stressor. Oxygen consumption increased by about 300 % as a result of the stress. Approximately 15 min after application of a similar stressor, plasma cortisol levels in stressed fish was 200 % higher than baseline levels. However, cortisol levels in fish sampled 30 min after the disturbance was similar to the baseline cortisol levels, indicating that full recovery had occurred. Although the patterns in the metabolic and cortisol stress responses were similar, metabolic rate could be measured continuously, thereby ensuring accurate interpretation of the data. Furthermore, increases in metabolic rate during the stress response are a culmination of physiological events from the primary to the tertiary levels of biological organization and are, therefore, easier to interpret in terms of long-term effects on the fish. Different transportation procedures elicited variable degrees of stress in juvenile P. commersonnii. The cost of metabolism attributed to the effects of capture and handling was twice as much as that attributed to acute temperature elevation. Acute temperature decrease resulted in a signiticant reduction in the oxygen consumption rate (ANOVA, P < 0.05). Oxygen consumption by the fish was not affected by fish density (ANOVA: F = 2.002, P = 0.5), or by oxygen depletion at dissolved oxygen concentrations above the critical level. Below this level, however, oxygen consumption decreased linearly with further decrease in dissolved oxygen concentration. These results showed that the highest energetic cost to juvenile P. commersonnii was incurred as a result of capture and handling. The results also showed that by subjecting fish to different stressors, it was possible to categorize them according to their relative metabolic costs to the fish. At 25º C, the effective concentration of 2-phenoxyethanol to fully anaesthetize (Stage IV, McFarland 1960) juvenile P. commersonnii was 0.4 ml l⁻¹ and the most appropriate concentration for deep sedation (Stage II, McFarland 1960) of the fish for at least 24 h was 0.2 ml l⁻¹. A maximum of 3 minutes was required by the fish to recover from the effects of the anaesthetic. There was no correlation between fish weight and the rate of induction of anaesthesia (r² = 0.001, p = 0.3). At the peak of the metabolic stress response, oxygen consumption was twice as high in the un-anaesthetized fish compared to the fish anaesthetized after the application of the simulated capture and handling stressor, suggesting that anaesthetization with 2-phenoxyethanol may have reduced the effect of the disturbance on the fish. Similar oxygen consumption rates for the fish anaesthetized prior to capture and the non-stressed fish suggested that the increases in metabolic rate could be linked to the struggling associated with attempts by fish to escape from the perceived stressor. Anaesthetization of juvenile P. commersonnii with 0.3 ml l⁻¹ 2-phenoxyethanol resulted in a more than 200 % increase in plasma cortisol concentration. The elevated levels of plasma cortisol in the anaesthetized fish suggested a manifestation of 2-phenoxyethanol as a stressor. At the time of capture, cortisol levels in fish that were anaesthetized prior to capture were the same as those measured in the disturbed fish at the peak of the stress response (ANOVA, p = 0.95), suggesting that the anaesthetized fish were already experiencing considerable stress at the time they were captured. Undisturbed juvenile P. commersonnii that were anaesthetized for 1 h also had cortisol levels that were five times higher than those measured in undisturbed-unanaesthetized fish, indicating that the duration of exposure to the anaesthetic had a significant effect on plasma cortisol levels. The results presented in this study demonstrate the usefulness of metabolic rate as an indicator of acute stress in fish. This was achieved by comparing the metabolic and the cortisol stress responses. The ease and accuracy with which oxygen consumption of fish could be measured made it possible to measure the stress response more accurately than by plasma cortisol concentration. It was also possible to monitor metabolic rate continuously over a long duration using polarographic oxygen sensors, thus enabling a better evaluation of the stress response. These results, thus, suggest that metabolic rate measurements could be a more practical way to quantify the effects of acute stressors on juvenile fishes. By detailing the profile of the metabolic stress response in P. commersonnii, this study makes a contribution towards understanding the physiological effects of stress in fishes. The study also contributes towards the quantification of baseline metabolic rates of this species under captivity. This study also contributes towards understanding the effects of 2-phenoxyethanol on the stress physiology of fish. By anaesthetizing fish under different conditions of stress, it was possible to evaluate the effect of 2-phenoxyethanol on the metabolic stress response. The ability of 2-phenoxyethanol to reduce physical activity of the fish, and thereby reduce the impact of acute stress on the metabolic stress response, makes it a good agent for the mitigation of stress during the capture and handling of fish. However, the increase in plasma cortisol concentration during prolonged anaesthetization using this drug suggests that the anaesthetic might be a stressor to fish and may, therefore, not be suitable for long-term sedation.
- Full Text:
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