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Shifts in thermoregulatory strategy during ontogeny in harp seals (Pagophilus groenlandicus)
Affiliation:1. Jiangsu University of Science and Technology, Zhenjiang, Jiangsu 212018, China;2. The Sericultural Research Institute, Chinese Academy of Agricultural Sciences, Zhenjiang, Jiangsu 212018, China;1. College of Biological Sciences, University of Minnesota–Twin Cities, Saint Paul, MN 55108, USA;2. Graduate Program in Neuroscience, University of Minnesota–Twin Cities, Minneapolis, MN 55455, USA;3. Department of Ecology, Evolution and Behavior, University of Minnesota–Twin Cities, Saint Paul, MN 55108, USA
Abstract:Heat balance can be difficult for young and/or small animals in polar regions because environmental conditions in combination with small body size or physiological immaturity can increase heat loss. We investigated how thermoregulatory patterns change with ontogeny in 5 age classes of harp seal (Pagophilus groenlandicus) from birth to post-molt to further understand the timing of thermoregulatory development in relation to their potential vulnerability to ongoing fluctuations in the extent and stability of Arctic pack ice. We measured changes in the amount, conductivity, and resistance of the seal pups׳ insulative layers (blubber and fur), the potential for endogenous heat-generation by shivering (muscle enzyme activity), and nonshivering thermogenesis (NST; brown adipose tissue (BAT) uncoupling protein 1 (UCP1) expression and mitochondrial density). There was no significant difference in blubber conductivity among age classes, though the amount of blubber insulation significantly increased from birth to weaning. Pelage conductivity was low (0.12±0.01 W m−1 °C−1) except in 9-day old pups (0.40±0.08 W m−1 °C−1); the significantly higher conductivity may signal the beginning of the molt, and this age group may be the most vulnerable to early water entry. Citrate synthase activity significantly increased (49.68±3.26 to 75.08±3.52 μmol min−1 g wet weight−1) in the muscle; however it is unlikely that increasing a single enzyme greatly impacts heat generation. BAT of younger pups contained UCP1, though expression and mitochondrial density quickly declined, and the ability of pups to produce heat via NST was lost by weaning. While total thermal resistance did not differ, neonatal and early nursing animals gained the majority of their thermal resistance from lanugo (82.5±0.03%); however, lanugo is not insulative when wet, and NST may be important to maintain euthermia and dry the coat if early immersion in water occurs. By late nursing, blubber seems sufficient as insulation (75.87±0.01% of resistance after 4 weeks), but high conductivity of fur may be responsible for retention of UCP1 expression. Weaned animals rely on blubber insulation, and no longer need NST, as wetted fur is no longer a threat to euthermia.
Keywords:Brown adipose tissue  Blubber  Lanugo  Phocid  Thermogenesis  Thermal conductivity  BAT"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0010"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  brown adipose tissue  COX"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0020"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  cytochrome c oxidase  CS"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0030"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  citrate synthase  HOAD"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0040"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  β-hydroxyacyl CoA dehydrogenase  LD"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0050"  },"  $$"  :[{"  #name"  :"  text"  ,"  $$"  :[{"  #name"  :"  italic"  ,"  _"  :"  longissimus dorsi  LCT"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0060"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  lower critical temperature  MR"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0070"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  metabolic rate  MT"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0080"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  mitochondria  NST"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0090"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  nonshivering thermogenesis  PWF"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0100"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  post-weaning fast  RMR"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0110"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  resting metabolic rate  SA:V"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0120"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  surface area to volume ratio  UCP1"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0130"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  uncoupling protein 1  TEM"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0140"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  transmission electron microscope  TNZ"  },{"  #name"  :"  keyword"  ,"  $"  :{"  id"  :"  key0150"  },"  $$"  :[{"  #name"  :"  text"  ,"  _"  :"  thermal neutral zone
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