Body size and oxygen stores in the blood and muscle set breath-hold limits in marine mammals, yet... more Body size and oxygen stores in the blood and muscle set breath-hold limits in marine mammals, yet these characteristics are understudied in immature cetaceans. We examined body mass and hematology from birth through adulthood in beluga whales (Delphinapterus leucas). At birth, body mass was 8% and 6% of the maximum mass recorded for adult females and males, respectively. Body mass then increased rapidly, approaching an asymptote around 12 yr for females and 18 yr for males. Interestingly, red blood cell counts, hemoglobin content, and hematocrit levels decreased after birth; this neonatal anemia was reversed as levels increased after 2 mo postpartum. Mature levels were obtained at approximately 8, 9, and 11 mo postpartum, respectively. Neonatal mean corpuscular hemoglobin also increased with ontogeny; mature levels were achieved by approximately 13 mo after birth. In contrast, mean corpuscular volume and mean corpuscular hemoglobin concentration demonstrated a significant but subtle increase throughout ontogeny. Our results indicate that postnatal maturation was required and that maturation occurred far earlier than the age at weaning (i.e., 2-3 yr postpartum). This is atypical of marine mammals, which generally achieve mature hemoglobin levels at weaning. Hematological maturation before maternal independence undoubtedly supports the prolonged breath holds of young belugas transiting under sea ice. This assessment enhances our knowledge of cetacean physiology and provides important inputs for determining age-specific dive capacity, yielding insights into age-specific flexibility to alter underwater behaviors, as will be required for future regime shifts and disturbances.
Physiological and Biochemical Zoology, Mar 1, 2020
Muscle biochemistry of aquatic birds and mammals varies in accordance with swimming and diving pe... more Muscle biochemistry of aquatic birds and mammals varies in accordance with swimming and diving performance, as well as with ontogeny. Similar to other odontocetes, the locomotor muscles (longissimus dorsi) of neonatal melon-headed whales (Peponocephala electra) have low myoglobin content (Mb; 1.06±0.20 g Mb/100 g wet muscle mass; mean ± SE; n=2] and low muscle nonbicarbonate buffering capacity (37.78±3.75 slykes; n=2), representing only 16% of adult Mb (6.64±0.33 g Mb/100 g wet muscle mass; n=5) and 56% of adult muscle nonbicarbonate buffering capacities (66.90±4.80 slykes; n=5). By the juvenile stage, Mb (2.75±0.80 g Mb/100 g wet muscle mass; n=3) is still only 41% of adult levels, but nonbicarbonate buffering capacity (65.61±2.62 slykes; n=3) has matured. Despite the observation that Hawaiian melon-headed whales are not deep divers or long-duration divers, their Mb rivals that found in ziphiids that forage in the bathypelagic zone and monodontids that forage under sea ice. The pelagic lifestyle of melon-headed whales likely requires sustained swimming, such that endurance training could elevate Mb in the locomotor muscle. Indeed, elevated Mb in the locomotor muscles of other pelagic odontocetes has been observed. Unlike deep-diving and Arctic-dwelling odontocetes, melon-headed whales do not achieve mature muscle characteristics before nursing. It is likely that early in life, the hydrodynamic benefits of swimming in echelon position with their mothers minimizes the endurance training of the calves that would otherwise promote rapid elevations in Mb.
Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, whi... more Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, which could affect body condition and have population‐level consequences. Body condition has historically been monitored by measuring blubber thickness over the xiphoid process (sternum). This may be an unreliable condition index because blubber at other sites along the body may be preferentially targeted to balance energetic demands. Animals in aquaria provided an opportunity for controlled study of how blubber topography is altered by caloric intake. Morphology, body mass, blubber thickness (21 sites), and caloric intake of five mature, nonpregnant, nonlactating female walruses were measured monthly (12 month minimum). Body condition (mass × standard length−1) was described by a model that included caloric intake and a seasonal effect, and scaled positively with estimates of total blubber mass. Blubber thicknesses (1.91–10.69 cm) varied topographically and were similar to values reported for free‐ranging female walruses. Body condition was most closely related to blubber thickness measured dorsomedially in the region of the anterior insertion of the pectoral flippers (shoulders); sternum blubber thickness was a relatively poor indicator of condition. This study demonstrates the importance of validating condition metrics before using them to monitor free‐ranging populations.
Physiological and Biochemical Zoology, Jul 1, 2021
Fat-level measurements used to indicate individual body condition and fitness are useful only whe... more Fat-level measurements used to indicate individual body condition and fitness are useful only when taken at a region along the body where fat responds to variations in caloric intake. Investigations to identify appropriate species-specific regions are limited, especially for cetaceans that have a specialized fat (blubber) that serves as an energy reserve and provides insulation. Over 18 mo, body mass of six pilot whales varied (range: 50–172 kg), and although caloric intake increased when water temperatures were lower, generally the best-fitting state-space model for length-adjusted mass was based on a single factor, caloric intake. After correcting for body length (range: 330–447 cm), the slope for blubber thickness and “blubber ring” thickness (average blubber thickness along a girth) in relation to body mass was positive and had a P value of <0.10 at six of 16 blubber measurement sites and one of five girth measurement sites, respectively. The slope for body girth (a reflection of changes in underlying blubber thickness) in relation to body mass was positive and had a lower P value (P<0.10) at three of five girth measurement sites. Results indicate that blubber from the anterior insertion of the pectoral fins to the posterior insertion of the dorsal fin is the most metabolically active region. This region includes the midflank site, a location where blubber thickness measurements have historically been taken to monitor cetacean body condition. Conversely, blubber in the peduncle region was comparatively inert. These findings must be considered when measuring blubber thickness and body width (i.e., photogrammetry) to monitor the condition of free-ranging cetaceans.
Synopsis The demands on the locomotor muscles at birth are different for cetaceans than terrestri... more Synopsis The demands on the locomotor muscles at birth are different for cetaceans than terrestrial mammals. Cetacean muscles do not need to support postural costs as the neonate transitions from the womb because water’s buoyant force supports body weight. Rather, neonatal cetacean muscles must sustain locomotion under hypoxic conditions as the neonate accompanies its mother swimming underwater. Despite disparate demands at birth, cetaceans like terrestrial mammals require postnatal development to attain mature musculature. Neonatal cetaceans have a low proportion of muscle mass, and their locomotor muscles have lower mitochondrial density, myoglobin content (Mb), and buffering capacity than those found in the adult locomotor muscle. For example, the locomotor muscle of the neonatal bottlenose dolphin has only 10 and 65% of the Mb and buffering capacity, respectively, found in the adult locomotor muscle. The maturation period required to achieve mature Mb and buffering capacity in the locomotor muscle varies across cetacean species from 0.75 to 4 and 1.17 to 3.4 years, respectively. The truncated nursing interval of harbor porpoises and sub-ice travel of beluga whales may be drivers for faster muscle maturation in these species. Despite these postnatal changes in the locomotor muscle, ontogenetic changes in locomotor muscle fiber type seem to be rare in cetaceans. Regardless, the underdeveloped aerobic and anaerobic capacities of the locomotor muscle of immature dolphins result in diminished thrusting capability and swim performance. Size-specific stroke amplitudes (23–26% of body length) of 0–3-month-old dolphins are significantly smaller than those of &gt;10-month-olds (29–30% of body length), and 0–1-month-olds only achieve 37 and 52% of the mean and maximum swim speed of adults, respectively. Until swim performance improves with muscle maturation, young cetaceans are precluded from achieving their pod’s swim speeds, which could have demographic consequences when fleeing anthropogenic disturbances.
Body size and oxygen stores in the blood and muscle set breath-hold limits in marine mammals, yet... more Body size and oxygen stores in the blood and muscle set breath-hold limits in marine mammals, yet these characteristics are understudied in immature cetaceans. We examined body mass and hematology from birth through adulthood in beluga whales (Delphinapterus leucas). At birth, body mass was 8% and 6% of the maximum mass recorded for adult females and males, respectively. Body mass then increased rapidly, approaching an asymptote around 12 yr for females and 18 yr for males. Interestingly, red blood cell counts, hemoglobin content, and hematocrit levels decreased after birth; this neonatal anemia was reversed as levels increased after 2 mo postpartum. Mature levels were obtained at approximately 8, 9, and 11 mo postpartum, respectively. Neonatal mean corpuscular hemoglobin also increased with ontogeny; mature levels were achieved by approximately 13 mo after birth. In contrast, mean corpuscular volume and mean corpuscular hemoglobin concentration demonstrated a significant but subtle increase throughout ontogeny. Our results indicate that postnatal maturation was required and that maturation occurred far earlier than the age at weaning (i.e., 2-3 yr postpartum). This is atypical of marine mammals, which generally achieve mature hemoglobin levels at weaning. Hematological maturation before maternal independence undoubtedly supports the prolonged breath holds of young belugas transiting under sea ice. This assessment enhances our knowledge of cetacean physiology and provides important inputs for determining age-specific dive capacity, yielding insights into age-specific flexibility to alter underwater behaviors, as will be required for future regime shifts and disturbances.
Physiological and Biochemical Zoology, Mar 1, 2020
Muscle biochemistry of aquatic birds and mammals varies in accordance with swimming and diving pe... more Muscle biochemistry of aquatic birds and mammals varies in accordance with swimming and diving performance, as well as with ontogeny. Similar to other odontocetes, the locomotor muscles (longissimus dorsi) of neonatal melon-headed whales (Peponocephala electra) have low myoglobin content (Mb; 1.06±0.20 g Mb/100 g wet muscle mass; mean ± SE; n=2] and low muscle nonbicarbonate buffering capacity (37.78±3.75 slykes; n=2), representing only 16% of adult Mb (6.64±0.33 g Mb/100 g wet muscle mass; n=5) and 56% of adult muscle nonbicarbonate buffering capacities (66.90±4.80 slykes; n=5). By the juvenile stage, Mb (2.75±0.80 g Mb/100 g wet muscle mass; n=3) is still only 41% of adult levels, but nonbicarbonate buffering capacity (65.61±2.62 slykes; n=3) has matured. Despite the observation that Hawaiian melon-headed whales are not deep divers or long-duration divers, their Mb rivals that found in ziphiids that forage in the bathypelagic zone and monodontids that forage under sea ice. The pelagic lifestyle of melon-headed whales likely requires sustained swimming, such that endurance training could elevate Mb in the locomotor muscle. Indeed, elevated Mb in the locomotor muscles of other pelagic odontocetes has been observed. Unlike deep-diving and Arctic-dwelling odontocetes, melon-headed whales do not achieve mature muscle characteristics before nursing. It is likely that early in life, the hydrodynamic benefits of swimming in echelon position with their mothers minimizes the endurance training of the calves that would otherwise promote rapid elevations in Mb.
Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, whi... more Pacific walruses may be unable to meet caloric requirements in the changing Arctic ecosystem, which could affect body condition and have population‐level consequences. Body condition has historically been monitored by measuring blubber thickness over the xiphoid process (sternum). This may be an unreliable condition index because blubber at other sites along the body may be preferentially targeted to balance energetic demands. Animals in aquaria provided an opportunity for controlled study of how blubber topography is altered by caloric intake. Morphology, body mass, blubber thickness (21 sites), and caloric intake of five mature, nonpregnant, nonlactating female walruses were measured monthly (12 month minimum). Body condition (mass × standard length−1) was described by a model that included caloric intake and a seasonal effect, and scaled positively with estimates of total blubber mass. Blubber thicknesses (1.91–10.69 cm) varied topographically and were similar to values reported for free‐ranging female walruses. Body condition was most closely related to blubber thickness measured dorsomedially in the region of the anterior insertion of the pectoral flippers (shoulders); sternum blubber thickness was a relatively poor indicator of condition. This study demonstrates the importance of validating condition metrics before using them to monitor free‐ranging populations.
Physiological and Biochemical Zoology, Jul 1, 2021
Fat-level measurements used to indicate individual body condition and fitness are useful only whe... more Fat-level measurements used to indicate individual body condition and fitness are useful only when taken at a region along the body where fat responds to variations in caloric intake. Investigations to identify appropriate species-specific regions are limited, especially for cetaceans that have a specialized fat (blubber) that serves as an energy reserve and provides insulation. Over 18 mo, body mass of six pilot whales varied (range: 50–172 kg), and although caloric intake increased when water temperatures were lower, generally the best-fitting state-space model for length-adjusted mass was based on a single factor, caloric intake. After correcting for body length (range: 330–447 cm), the slope for blubber thickness and “blubber ring” thickness (average blubber thickness along a girth) in relation to body mass was positive and had a P value of <0.10 at six of 16 blubber measurement sites and one of five girth measurement sites, respectively. The slope for body girth (a reflection of changes in underlying blubber thickness) in relation to body mass was positive and had a lower P value (P<0.10) at three of five girth measurement sites. Results indicate that blubber from the anterior insertion of the pectoral fins to the posterior insertion of the dorsal fin is the most metabolically active region. This region includes the midflank site, a location where blubber thickness measurements have historically been taken to monitor cetacean body condition. Conversely, blubber in the peduncle region was comparatively inert. These findings must be considered when measuring blubber thickness and body width (i.e., photogrammetry) to monitor the condition of free-ranging cetaceans.
Synopsis The demands on the locomotor muscles at birth are different for cetaceans than terrestri... more Synopsis The demands on the locomotor muscles at birth are different for cetaceans than terrestrial mammals. Cetacean muscles do not need to support postural costs as the neonate transitions from the womb because water’s buoyant force supports body weight. Rather, neonatal cetacean muscles must sustain locomotion under hypoxic conditions as the neonate accompanies its mother swimming underwater. Despite disparate demands at birth, cetaceans like terrestrial mammals require postnatal development to attain mature musculature. Neonatal cetaceans have a low proportion of muscle mass, and their locomotor muscles have lower mitochondrial density, myoglobin content (Mb), and buffering capacity than those found in the adult locomotor muscle. For example, the locomotor muscle of the neonatal bottlenose dolphin has only 10 and 65% of the Mb and buffering capacity, respectively, found in the adult locomotor muscle. The maturation period required to achieve mature Mb and buffering capacity in the locomotor muscle varies across cetacean species from 0.75 to 4 and 1.17 to 3.4 years, respectively. The truncated nursing interval of harbor porpoises and sub-ice travel of beluga whales may be drivers for faster muscle maturation in these species. Despite these postnatal changes in the locomotor muscle, ontogenetic changes in locomotor muscle fiber type seem to be rare in cetaceans. Regardless, the underdeveloped aerobic and anaerobic capacities of the locomotor muscle of immature dolphins result in diminished thrusting capability and swim performance. Size-specific stroke amplitudes (23–26% of body length) of 0–3-month-old dolphins are significantly smaller than those of &gt;10-month-olds (29–30% of body length), and 0–1-month-olds only achieve 37 and 52% of the mean and maximum swim speed of adults, respectively. Until swim performance improves with muscle maturation, young cetaceans are precluded from achieving their pod’s swim speeds, which could have demographic consequences when fleeing anthropogenic disturbances.
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