Division of Bone and Mineral Diseases, Department of Internal Medicine, Washington University, Saint Louis, MO 63110, USA.
Department of Molecular & Integrative Physiology, University of Michigan, Ann Arbor, MI 48105, USA.
Bone. 2019 Jan;118:32-41. doi: 10.1016/j.bone.2018.01.016. Epub 2018 Jan 31.
Bone marrow adipose tissue (BMAT) is preserved or increased in states of caloric restriction. Similarly, we found that BMAT in the tail vertebrae, but not the red marrow in the tibia, resists loss of neutral lipid with acute, 48-hour fasting in rats. The mechanisms underlying this phenomenon and its seemingly distinct regulation from peripheral white adipose tissue (WAT) remain unknown. To test the role of β-adrenergic stimulation, a major regulator of adipose tissue lipolysis, we examined the responses of BMAT to β-adrenergic agonists. Relative to inguinal WAT, BMAT had reduced phosphorylation of hormone sensitive lipase (HSL) after treatment with pan-β-adrenergic agonist isoproterenol. Phosphorylation of HSL in response to β3-adrenergic agonist CL316,243 was decreased by an additional ~90% (distal tibia BMAT) or could not be detected (tail vertebrae). Ex vivo, adrenergic stimulation of lipolysis in purified BMAT adipocytes was also substantially less than iWAT adipocytes and had site-specific properties. Specifically, regulated bone marrow adipocytes (rBMAs) from proximal tibia and femur underwent lipolysis in response to both CL316,243 and forskolin, while constitutive BMAs from the tail responded only to forskolin. This occurred independently of changes in gene expression of β-adrenergic receptors, which were similar between adipocytes from iWAT and BMAT, and could not be explained by defective coupling of β-adrenergic receptors to lipolytic machinery through caveolin 1. Specifically, we found that whereas caveolin 1 was necessary to mediate maximal stimulation of lipolysis in iWAT, overexpression of caveolin 1 was insufficient to rescue impaired BMAT signaling. Lastly, we tested the ability of BMAT to respond to 72-hour treatment with CL316,243 in vivo. This was sufficient to cause beiging of iWAT adipocytes and a decrease in iWAT adipocyte cell size. By contrast, adipocyte size in the tail BMAT and distal tibia remained unchanged. However, within the distal femur, we identified a subpopulation of BMAT adipocytes that underwent lipid droplet remodeling. This response was more pronounced in females than in males and resembled lipolysis-induced lipid partitioning rather than traditional beiging. In summary, BMAT has the capacity to respond to β-adrenergic stimuli, however, its responses are muted and BMAT generally resists lipid hydrolysis and remodeling relative to iWAT. This resistance is more pronounced in distal regions of the skeleton where the BMAT adipocytes are larger with little intervening hematopoiesis, suggesting that there may be a role for both cell-autonomous and microenvironmental determinants. Resistance to β-adrenergic stimuli further separates BMAT from known regulators of energy partitioning and contributes to our understanding of why BMAT is preserved in states of fasting and caloric restriction.
骨髓脂肪组织(BMAT)在热量限制状态下得以保留或增加。同样,我们发现,在大鼠急性 48 小时禁食时,尾椎骨中的 BMAT 抵抗中性脂质的丢失,而胫骨中的红骨髓则不抵抗。这种现象的发生机制及其与外周白色脂肪组织(WAT)的明显不同调节机制仍不清楚。为了测试β-肾上腺素能刺激的作用,这是脂肪组织脂解的主要调节剂,我们研究了 BMAT 对β-肾上腺素能激动剂的反应。与腹股沟 WAT 相比,BMAT 在用全β-肾上腺素能激动剂异丙肾上腺素处理后,激素敏感脂肪酶(HSL)的磷酸化减少。β3-肾上腺素能激动剂 CL316,243 对 HSL 的磷酸化反应降低了另外约 90%(胫骨远端)或无法检测到(尾椎骨)。体外,纯化的 BMAT 脂肪细胞中的肾上腺素能刺激脂解作用也明显小于 iWAT 脂肪细胞,并且具有特定的部位特异性。具体而言,来自胫骨近端和股骨的调节性骨髓脂肪细胞(rBMAs)对 CL316,243 和 forskolin 均发生脂解反应,而来自尾骨的组成性 BMAs 仅对 forskolin 发生反应。这与 iWAT 和 BMAT 脂肪细胞之间相似的β-肾上腺素能受体基因表达的变化无关,并且不能通过 caveolin 1 解释为β-肾上腺素能受体与脂肪分解机制的偶联缺陷。具体而言,我们发现,虽然 caveolin 1 对于介导 iWAT 中最大的脂解刺激是必需的,但 caveolin 1 的过表达不足以挽救受损的 BMAT 信号传导。最后,我们测试了 BMAT 在体内用 CL316,243 治疗 72 小时的能力。这足以导致 iWAT 脂肪细胞的米色化和 iWAT 脂肪细胞细胞大小的减小。相比之下,尾骨 BMAT 和胫骨远端的脂肪细胞大小保持不变。然而,在远端股骨中,我们鉴定了一群经历脂质滴重塑的 BMAT 脂肪细胞。这种反应在女性中比男性中更为明显,类似于脂解诱导的脂质分配,而不是传统的米色化。总之,BMAT 有能力对β-肾上腺素能刺激做出反应,但是其反应被抑制,并且 BMAT 通常抵抗脂质水解和重塑,相对于 iWAT。这种抵抗在骨骼的远端区域更为明显,其中 BMAT 脂肪细胞较大,并且几乎没有中间的造血作用,这表明可能存在细胞自主和微环境决定因素的作用。对β-肾上腺素能刺激的抵抗进一步将 BMAT 与已知的能量分配调节剂区分开来,并有助于我们理解为什么 BMAT 在禁食和热量限制状态下得以保留。