Andreas Fahlman - Academia.edu (original) (raw)
Papers by Andreas Fahlman
Journal of Zoo and Wildlife Medicine, 2021
Abstract: Lung function (breath duration, respiratory flow [], and tidal volume [VT]), and end-ex... more Abstract: Lung function (breath duration, respiratory flow [], and tidal volume [VT]), and end-expiratory O2 were measured in 19 adult bottlenose dolphins (Tursiops spp.) while at rest in water or beached for up to 10 min. The results show that inspiratory VT, expiratory VT, or inspiratory did not differ on land or in water. The average expiratory for all dolphins on land decreased by 16%, and the expiratory and total breath durations increased by 5% and 4%, respectively, compared with in water. There were temporal changes observed during beaching, where expired and inspired VT and inspired decreased by 13%, 16%, and 9%, respectively, after 10 min on land. These data suggest that dolphins compensate for the effect of gravity by adjusting respiration to maintain alveolar ventilation and gas exchange, but during extended durations, the increased work of breathing may impede ventilation and gas exchange. Continuous monitoring of lung function and gas exchange may help prevent long-term damage during out-of-water medical procedures, optimize animal transport conditions, and improve survival during stranding events.
Journal of Experimental Biology, 2020
Among the many factors that influence the cardiovascular adjustments of marine mammals is the act... more Among the many factors that influence the cardiovascular adjustments of marine mammals is the act of respiration at the surface, which facilitates rapid gas exchange and tissue re-perfusion between dives. We measured heart rate (fH) in six adult male bottlenose dolphins (Tursiops truncatus) spontaneously breathing at the surface to quantify the relationship between respiration and fH, and compared this with fH during submerged breath-holds. We found that dolphins exhibit a pronounced respiratory sinus arrhythmia (RSA) during surface breathing, resulting in a rapid increase in fH after a breath followed by a gradual decrease over the following 15–20 s to a steady fH that is maintained until the following breath. RSA resulted in a maximum instantaneous fH (ifH) of 87.4±13.6 beats min−1 and a minimum ifH of 56.8±14.8 beats min−1, and the degree of RSA was positively correlated with the inter-breath interval (IBI). The minimum ifH during 2 min submerged breath-holds where dolphins exhib...
: The objective of this study is to update a current gas dynamics model with recently acquired da... more : The objective of this study is to update a current gas dynamics model with recently acquired data for respiratory compliance (P-V), and body compartment size estimates in California sea lions. The model will be calibrated against measured arterial and venous PO2 levels from California sea lions, and estimate the error between predicted and observed values. The model will be used to investigate specific scenarios where marine mammals could be particularly prone to decompression sickness (DCS) due to changes in dive behavior or physiology.
Frontiers in Physiology, 2021
Frontiers in Physiology, 2020
Previous reports suggested the existence of direct somatic motor control over heart rate (fH) res... more Previous reports suggested the existence of direct somatic motor control over heart rate (fH) responses during diving in some marine mammals, as the result of a cognitive and/or learning process rather than being a reflexive response. This would be beneficial for O2 storage management, but would also allow ventilation-perfusion matching for selective gas exchange, where O2 and CO2 can be exchanged with minimal exchange of N2. Such a mechanism explains how air breathing marine vertebrates avoid diving related gas bubble formation during repeated dives, and how stress could interrupt this mechanism and cause excessive N2 exchange. To investigate the conditioned response, we measured the fH-response before and during static breath-holds in three bottlenose dolphins (Tursiops truncatus) when shown a visual symbol to perform either a long (LONG) or short (SHORT) breath-hold, or during a spontaneous breath-hold without a symbol (NS). The average fH (ifHstart), and the rate of change in fH...
Frontiers in Marine Science, 2021
Decompression theory has been mainly based on studies on terrestrial mammals, and may not transla... more Decompression theory has been mainly based on studies on terrestrial mammals, and may not translate well to marine mammals. However, evidence that marine mammals experience gas bubbles during diving is growing, causing concern that these bubbles may cause gas emboli pathology (GEP) under unusual circumstances. Marine mammal management, and usual avoidance, of gas emboli and GEP, or the bends, became a topic of intense scientific interest after sonar-exposed, mass-stranded deep-diving whales were observed with gas bubbles. Theoretical models, based on our current understanding of diving physiology in cetaceans, predict that the tissue and blood N2levels in the bottlenose dolphin (Tursiops truncatus) are at levels that would result in severe DCS symptoms in similar sized terrestrial mammals. However, the dolphins appear to have physiological or behavioral mechanisms to avoid excessive blood N2levels, or may be more resistant to circulating bubbles through immunological/biochemical ada...
Frontiers in Physiology, 2021
Sea turtles, like other air-breathing diving vertebrates, commonly experience significant gas emb... more Sea turtles, like other air-breathing diving vertebrates, commonly experience significant gas embolism (GE) when incidentally caught at depth in fishing gear and brought to the surface. To better understand why sea turtles develop GE, we built a mathematical model to estimate partial pressures of N2 (PN2), O2 (PO2), and CO2 (PCO2) in the major body-compartments of diving loggerheads (Caretta caretta), leatherbacks (Dermochelys coriacea), and green turtles (Chelonia mydas). This model was adapted from a published model for estimating gas dynamics in marine mammals and penguins. To parameterize the sea turtle model, we used values gleaned from previously published literature and 22 necropsies. Next, we applied this model to data collected from free-roaming individuals of the three study species. Finally, we varied body-condition and cardiac output within the model to see how these factors affected the risk of GE. Our model suggests that cardiac output likely plays a significant role i...
Ischemic events, such as ischemic heart disease and ischemic stroke, are the number one cause of ... more Ischemic events, such as ischemic heart disease and ischemic stroke, are the number one cause of death globally. Ischemia prevents blood, carrying essential nutrients and oxygen, from reaching tissues and organ systems, leading to cell and tissue death, and eventual organ failure. While humans are relatively intolerant to these ischemic events, other species, such as marine mammals, have evolved remarkable tolerance to chronic ischemia/reperfusion during diving. Here we capitalized on the unique adaptations of bottlenose dolphins (Tursiops truncatus) as a comparative model of ischemic stress and hypoxia tolerance to identify molecular features associated with breath-holding. Using RNA-Seq we observed time-dependent upregulation of the arachidonate 5-lipoxygenase (ALOX5) gene during breath-holding. Consistent with the RNA-Seq data, we also observed increased ALOX5 enzymatic activity in the serum of dolphins undergoing breath holds. ALOX5 has previously been shown to be activated duri...
The Journal of Experimental Biology, 2020
ABSTRACTIn the current study we used transthoracic echocardiography to measure stroke volume (SV)... more ABSTRACTIn the current study we used transthoracic echocardiography to measure stroke volume (SV), heart rate (fH) and cardiac output (CO) in adult bottlenose dolphins (Tursiops truncatus), a male beluga whale calf [Delphinapterus leucas, body mass (Mb) range: 151–175 kg] and an adult female false killer whale (Pseudorca crassidens, estimated Mb: 500–550 kg) housed in managed care. We also recorded continuous electrocardiogram (ECG) in the beluga whale, bottlenose dolphin, false killer whale, killer whale (Orcinus orca) and pilot whale (Globicephala macrorhynchus) to evaluate cardiorespiratory coupling while breathing spontaneously under voluntary control. The results show that cetaceans have a strong respiratory sinus arrythmia (RSA), during which both fH and SV vary within the interbreath interval, making average values dependent on the breathing frequency (fR). The RSA-corrected fH was lower for all cetaceans compared with that of similarly sized terrestrial mammals breathing con...
The Journal of Experimental Biology, 2020
ABSTRACTWe analysed 3680 dives from 23 satellite-linked tags deployed on Cuvier's beaked whal... more ABSTRACTWe analysed 3680 dives from 23 satellite-linked tags deployed on Cuvier's beaked whales to assess the relationship between long duration dives and inter-deep dive intervals and to estimate aerobic dive limit (ADL). The median duration of presumed foraging dives was 59 min and 5% of dives exceeded 77.7 min. We found no relationship between the longest 5% of dive durations and the following inter-deep dive interval nor any relationship with the ventilation period immediately prior to or following a long dive. We suggest that Cuvier's beaked whales have low metabolic rates, high oxygen storage capacities and a high acid-buffering capacity to deal with the by-products of both aerobic and anaerobic metabolism, which enables them to extend dive durations and exploit their bathypelagic foraging habitats.
Ecology and Evolution, 2020
The Journal of Experimental Biology, 2020
ABSTRACT The air volume in the respiratory system of marine tetrapods provides a store of O2 to f... more ABSTRACT The air volume in the respiratory system of marine tetrapods provides a store of O2 to fuel aerobic metabolism during dives; however, it can also be a liability, as the associated N2 can increase the risk of decompression sickness. In order to more fully understand the physiological limitations of different air-breathing marine vertebrates, it is therefore important to be able to accurately estimate the air volume in the respiratory system during diving. One method that has been used to do so is to calculate the air volume from glide phases – periods of movement during which no thrust is produced by the animal – which many species conduct during ascent periods, when gases are expanding owing to decreasing hydrostatic pressure. This method assumes that there is conservation of mass in the respiratory system, with volume changes only driven by pressure. In this Commentary, we use previously published data to argue that both the respiratory quotient and differences in tissue and blood gas solubility potentially alter the mass balance in the respiratory system throughout a dive. Therefore, near the end of a dive, the measured volume of gas at a given pressure may be 12–50% less than from the start of the dive; the actual difference will depend on the length of the dive, the cardiac output, the pulmonary shunt and the metabolic rate. Novel methods and improved understanding of diving physiology will be required to verify the size of the effects described here and to more accurately estimate the volume of gas inhaled at the start of a dive. Summary: Various factors affect our ability to accurately determine lung capacity during diving in air-breathing marine vertebrates; improving our estimates has implications for our understanding of the physiology and ecophysiology of these animals.
Frontiers Research Topics, 2020
ed accelerometer transmissions against outputs from established processing procedures, and (2) as... more ed accelerometer transmissions against outputs from established processing procedures, and (2) assess the validity of previously developed dive foraging indices, calculated solely from time-depth measurements. We found transmitted estimates of PrCA behaviors were generally comparable to those obtained from archival processing, although a small but consistent over-estimation was noted. Following this, dive foraging segments were identified either from (1) sinuosity in the trajectories of high-resolution depth archives, or (2) vertical speeds between low resolution transmissions of key depth inflection points along a dive profile. In both cases, more than 93% of the estimated PrCA behaviors (from either abstracted transmissions or archival processing) fell into inferred dive foraging segments (i.e., “hunting” segments), suggesting the two methods provide a reliable indicator of foraging effort. The validation of transmitted acceleration data and
The Journal of Experimental Biology, 2020
Frontiers for Young Minds, Dec 18, 2017
Frontiers in Ecology and Evolution, 2020
The Journal of Experimental Biology, 2020
Journal of Zoo and Wildlife Medicine, 2021
Abstract: Lung function (breath duration, respiratory flow [], and tidal volume [VT]), and end-ex... more Abstract: Lung function (breath duration, respiratory flow [], and tidal volume [VT]), and end-expiratory O2 were measured in 19 adult bottlenose dolphins (Tursiops spp.) while at rest in water or beached for up to 10 min. The results show that inspiratory VT, expiratory VT, or inspiratory did not differ on land or in water. The average expiratory for all dolphins on land decreased by 16%, and the expiratory and total breath durations increased by 5% and 4%, respectively, compared with in water. There were temporal changes observed during beaching, where expired and inspired VT and inspired decreased by 13%, 16%, and 9%, respectively, after 10 min on land. These data suggest that dolphins compensate for the effect of gravity by adjusting respiration to maintain alveolar ventilation and gas exchange, but during extended durations, the increased work of breathing may impede ventilation and gas exchange. Continuous monitoring of lung function and gas exchange may help prevent long-term damage during out-of-water medical procedures, optimize animal transport conditions, and improve survival during stranding events.
Journal of Experimental Biology, 2020
Among the many factors that influence the cardiovascular adjustments of marine mammals is the act... more Among the many factors that influence the cardiovascular adjustments of marine mammals is the act of respiration at the surface, which facilitates rapid gas exchange and tissue re-perfusion between dives. We measured heart rate (fH) in six adult male bottlenose dolphins (Tursiops truncatus) spontaneously breathing at the surface to quantify the relationship between respiration and fH, and compared this with fH during submerged breath-holds. We found that dolphins exhibit a pronounced respiratory sinus arrhythmia (RSA) during surface breathing, resulting in a rapid increase in fH after a breath followed by a gradual decrease over the following 15–20 s to a steady fH that is maintained until the following breath. RSA resulted in a maximum instantaneous fH (ifH) of 87.4±13.6 beats min−1 and a minimum ifH of 56.8±14.8 beats min−1, and the degree of RSA was positively correlated with the inter-breath interval (IBI). The minimum ifH during 2 min submerged breath-holds where dolphins exhib...
: The objective of this study is to update a current gas dynamics model with recently acquired da... more : The objective of this study is to update a current gas dynamics model with recently acquired data for respiratory compliance (P-V), and body compartment size estimates in California sea lions. The model will be calibrated against measured arterial and venous PO2 levels from California sea lions, and estimate the error between predicted and observed values. The model will be used to investigate specific scenarios where marine mammals could be particularly prone to decompression sickness (DCS) due to changes in dive behavior or physiology.
Frontiers in Physiology, 2021
Frontiers in Physiology, 2020
Previous reports suggested the existence of direct somatic motor control over heart rate (fH) res... more Previous reports suggested the existence of direct somatic motor control over heart rate (fH) responses during diving in some marine mammals, as the result of a cognitive and/or learning process rather than being a reflexive response. This would be beneficial for O2 storage management, but would also allow ventilation-perfusion matching for selective gas exchange, where O2 and CO2 can be exchanged with minimal exchange of N2. Such a mechanism explains how air breathing marine vertebrates avoid diving related gas bubble formation during repeated dives, and how stress could interrupt this mechanism and cause excessive N2 exchange. To investigate the conditioned response, we measured the fH-response before and during static breath-holds in three bottlenose dolphins (Tursiops truncatus) when shown a visual symbol to perform either a long (LONG) or short (SHORT) breath-hold, or during a spontaneous breath-hold without a symbol (NS). The average fH (ifHstart), and the rate of change in fH...
Frontiers in Marine Science, 2021
Decompression theory has been mainly based on studies on terrestrial mammals, and may not transla... more Decompression theory has been mainly based on studies on terrestrial mammals, and may not translate well to marine mammals. However, evidence that marine mammals experience gas bubbles during diving is growing, causing concern that these bubbles may cause gas emboli pathology (GEP) under unusual circumstances. Marine mammal management, and usual avoidance, of gas emboli and GEP, or the bends, became a topic of intense scientific interest after sonar-exposed, mass-stranded deep-diving whales were observed with gas bubbles. Theoretical models, based on our current understanding of diving physiology in cetaceans, predict that the tissue and blood N2levels in the bottlenose dolphin (Tursiops truncatus) are at levels that would result in severe DCS symptoms in similar sized terrestrial mammals. However, the dolphins appear to have physiological or behavioral mechanisms to avoid excessive blood N2levels, or may be more resistant to circulating bubbles through immunological/biochemical ada...
Frontiers in Physiology, 2021
Sea turtles, like other air-breathing diving vertebrates, commonly experience significant gas emb... more Sea turtles, like other air-breathing diving vertebrates, commonly experience significant gas embolism (GE) when incidentally caught at depth in fishing gear and brought to the surface. To better understand why sea turtles develop GE, we built a mathematical model to estimate partial pressures of N2 (PN2), O2 (PO2), and CO2 (PCO2) in the major body-compartments of diving loggerheads (Caretta caretta), leatherbacks (Dermochelys coriacea), and green turtles (Chelonia mydas). This model was adapted from a published model for estimating gas dynamics in marine mammals and penguins. To parameterize the sea turtle model, we used values gleaned from previously published literature and 22 necropsies. Next, we applied this model to data collected from free-roaming individuals of the three study species. Finally, we varied body-condition and cardiac output within the model to see how these factors affected the risk of GE. Our model suggests that cardiac output likely plays a significant role i...
Ischemic events, such as ischemic heart disease and ischemic stroke, are the number one cause of ... more Ischemic events, such as ischemic heart disease and ischemic stroke, are the number one cause of death globally. Ischemia prevents blood, carrying essential nutrients and oxygen, from reaching tissues and organ systems, leading to cell and tissue death, and eventual organ failure. While humans are relatively intolerant to these ischemic events, other species, such as marine mammals, have evolved remarkable tolerance to chronic ischemia/reperfusion during diving. Here we capitalized on the unique adaptations of bottlenose dolphins (Tursiops truncatus) as a comparative model of ischemic stress and hypoxia tolerance to identify molecular features associated with breath-holding. Using RNA-Seq we observed time-dependent upregulation of the arachidonate 5-lipoxygenase (ALOX5) gene during breath-holding. Consistent with the RNA-Seq data, we also observed increased ALOX5 enzymatic activity in the serum of dolphins undergoing breath holds. ALOX5 has previously been shown to be activated duri...
The Journal of Experimental Biology, 2020
ABSTRACTIn the current study we used transthoracic echocardiography to measure stroke volume (SV)... more ABSTRACTIn the current study we used transthoracic echocardiography to measure stroke volume (SV), heart rate (fH) and cardiac output (CO) in adult bottlenose dolphins (Tursiops truncatus), a male beluga whale calf [Delphinapterus leucas, body mass (Mb) range: 151–175 kg] and an adult female false killer whale (Pseudorca crassidens, estimated Mb: 500–550 kg) housed in managed care. We also recorded continuous electrocardiogram (ECG) in the beluga whale, bottlenose dolphin, false killer whale, killer whale (Orcinus orca) and pilot whale (Globicephala macrorhynchus) to evaluate cardiorespiratory coupling while breathing spontaneously under voluntary control. The results show that cetaceans have a strong respiratory sinus arrythmia (RSA), during which both fH and SV vary within the interbreath interval, making average values dependent on the breathing frequency (fR). The RSA-corrected fH was lower for all cetaceans compared with that of similarly sized terrestrial mammals breathing con...
The Journal of Experimental Biology, 2020
ABSTRACTWe analysed 3680 dives from 23 satellite-linked tags deployed on Cuvier's beaked whal... more ABSTRACTWe analysed 3680 dives from 23 satellite-linked tags deployed on Cuvier's beaked whales to assess the relationship between long duration dives and inter-deep dive intervals and to estimate aerobic dive limit (ADL). The median duration of presumed foraging dives was 59 min and 5% of dives exceeded 77.7 min. We found no relationship between the longest 5% of dive durations and the following inter-deep dive interval nor any relationship with the ventilation period immediately prior to or following a long dive. We suggest that Cuvier's beaked whales have low metabolic rates, high oxygen storage capacities and a high acid-buffering capacity to deal with the by-products of both aerobic and anaerobic metabolism, which enables them to extend dive durations and exploit their bathypelagic foraging habitats.
Ecology and Evolution, 2020
The Journal of Experimental Biology, 2020
ABSTRACT The air volume in the respiratory system of marine tetrapods provides a store of O2 to f... more ABSTRACT The air volume in the respiratory system of marine tetrapods provides a store of O2 to fuel aerobic metabolism during dives; however, it can also be a liability, as the associated N2 can increase the risk of decompression sickness. In order to more fully understand the physiological limitations of different air-breathing marine vertebrates, it is therefore important to be able to accurately estimate the air volume in the respiratory system during diving. One method that has been used to do so is to calculate the air volume from glide phases – periods of movement during which no thrust is produced by the animal – which many species conduct during ascent periods, when gases are expanding owing to decreasing hydrostatic pressure. This method assumes that there is conservation of mass in the respiratory system, with volume changes only driven by pressure. In this Commentary, we use previously published data to argue that both the respiratory quotient and differences in tissue and blood gas solubility potentially alter the mass balance in the respiratory system throughout a dive. Therefore, near the end of a dive, the measured volume of gas at a given pressure may be 12–50% less than from the start of the dive; the actual difference will depend on the length of the dive, the cardiac output, the pulmonary shunt and the metabolic rate. Novel methods and improved understanding of diving physiology will be required to verify the size of the effects described here and to more accurately estimate the volume of gas inhaled at the start of a dive. Summary: Various factors affect our ability to accurately determine lung capacity during diving in air-breathing marine vertebrates; improving our estimates has implications for our understanding of the physiology and ecophysiology of these animals.
Frontiers Research Topics, 2020
ed accelerometer transmissions against outputs from established processing procedures, and (2) as... more ed accelerometer transmissions against outputs from established processing procedures, and (2) assess the validity of previously developed dive foraging indices, calculated solely from time-depth measurements. We found transmitted estimates of PrCA behaviors were generally comparable to those obtained from archival processing, although a small but consistent over-estimation was noted. Following this, dive foraging segments were identified either from (1) sinuosity in the trajectories of high-resolution depth archives, or (2) vertical speeds between low resolution transmissions of key depth inflection points along a dive profile. In both cases, more than 93% of the estimated PrCA behaviors (from either abstracted transmissions or archival processing) fell into inferred dive foraging segments (i.e., “hunting” segments), suggesting the two methods provide a reliable indicator of foraging effort. The validation of transmitted acceleration data and
The Journal of Experimental Biology, 2020
Frontiers for Young Minds, Dec 18, 2017
Frontiers in Ecology and Evolution, 2020
The Journal of Experimental Biology, 2020