自私的大脑:对能量资源的竞争
The selfish brain: competition for energy resources.
作者信息
Fehm H L, Kern W, Peters A
机构信息
Medizinische Klinik I, Universität Lübeck, Ratzeburger Allee 160, D-23538 Lübeck, Germany.
出版信息
Prog Brain Res. 2006;153:129-40. doi: 10.1016/S0079-6123(06)53007-9.
Although the brain constitutes only 2% of the body mass, its metabolism accounts for 50% of total body glucose utilization. This delicate situation is aggravated by the fact that the brain depends on glucose as energy substrate. Thus, the contour of a major problem becomes evident: how can the brain maintain constant fluxes of large amounts of glucose to itself in the presence of powerful competitors as fat and muscle tissue. Activity of cortical neurons generates an "energy on demand" signal which eventually mediates the uptake of glucose from brain capillaries. Because energy stores in the circulation (equivalent to ca. 5 g glucose) are also limited, a second signal is required termed "energy on request"; this signal is responsible for the activation of allocation processes. The term "allocation" refers to the activation of the "behavior control column" by an input from the hippocampus-amygdala system. As far as eating behavior is concerned the behavior control column consists of the ventral medial hypothalamus (VMH) and periventricular nucleus (PVN). The PVN represents the central nucleus of the brain's stress systems, the hypothalamus-pituitary-adrenal (HPA) axis and the sympathetic nervous system (SNS). Activation of the sympatico-adrenal system inhibits glucose uptake by peripheral tissues by inhibiting insulin release and inducing insulin resistance and increases hepatic glucose production. With an inadequate "energy on request" signal neuroglucopenia would be the consequence. A decrease in brain glucose can activate glucose-sensitive neurons in the lateral hypothalamus (LH) with the release of orexigenic peptides which stimulate food intake. If the energy supply of the brain depends on activation of the LH rather than on increased allocation to the brain, an increase in body weight is evitable. An increase in fat mass will generate feedback signals as leptin and insulin, which activate the arcuate nucleus. Activation of arcuate nucleus in turn will stimulate the activity of the PVN in a way similar to the activation by the hippocampus-amydala system. The activity of PVN is influenced by the hippocampal outflow which in turn is the consequence of a balance of low-affinity and high-affinity glucocorticoid receptors. This set-point can permanently be displaced by extreme stress situations, by starvation, exercise, hormones, drugs or by endocrine-disrupting chemicals. Disorders in the "energy on request" process will influence the allocation of energy and in so doing alter the body mass of the organism. In this "selfish brain theory" the neocortex and the limbic system play a central role in the pathogenesis of diseases, such as anorexia nervosa, obesity and diabetes mellitus type II. From these considerations it appears that the primary disturbance in obesity is a displacement of the hippocampal set-point of the system. The resulting permanent activation of the feedback system must result in a likewise permanent activation of the sympatico-adrenal system, which induces insulin resistance, hypertension and the other components of the metabolic syndrome. Available therapies for treatment of the metabolic syndrome (blockade of alpha- and beta-adrenergic receptors, insulin and insulin secretagogues) interfere with mechanisms, which must be considered compensatory. This explains why these therapies are disappointing in the long run. New therapeutic strategies based on the "selfish brain theory" will be discussed.
尽管大脑仅占体重的2%,但其新陈代谢却占全身葡萄糖利用总量的50%。大脑依赖葡萄糖作为能量底物这一事实使这种微妙的情况更加恶化。因此,一个主要问题的轮廓变得明显:在脂肪和肌肉组织等强大竞争者存在的情况下,大脑如何能维持自身大量葡萄糖的恒定通量。皮质神经元的活动产生一个“按需供能”信号,该信号最终介导从脑毛细血管摄取葡萄糖。由于循环中的能量储备(相当于约5克葡萄糖)也有限,所以需要第二个信号,即“应需供能”信号;这个信号负责激活分配过程。“分配”一词指的是海马体 - 杏仁核系统的输入对“行为控制柱”的激活。就进食行为而言,行为控制柱由腹内侧下丘脑(VMH)和室旁核(PVN)组成。PVN是大脑应激系统、下丘脑 - 垂体 - 肾上腺(HPA)轴和交感神经系统(SNS)的中枢核。交感 - 肾上腺系统的激活通过抑制胰岛素释放和诱导胰岛素抵抗来抑制外周组织对葡萄糖的摄取,并增加肝脏葡萄糖生成。如果“应需供能”信号不足,后果将是神经低血糖症。脑葡萄糖水平降低可激活下丘脑外侧区(LH)的葡萄糖敏感神经元,释放促食欲肽,从而刺激食物摄入。如果大脑的能量供应依赖于LH的激活而不是增加对大脑的分配,体重增加将不可避免。脂肪量的增加会产生如瘦素和胰岛素等反馈信号,激活弓状核。弓状核的激活反过来会以类似于海马体 - 杏仁核系统激活的方式刺激PVN的活动。PVN的活动受海马体输出的影响,而海马体输出又是低亲和力和高亲和力糖皮质激素受体平衡的结果。这个设定点可能会因极端应激情况、饥饿、运动、激素、药物或内分泌干扰化学物质而永久改变。“应需供能”过程中的紊乱将影响能量分配,进而改变生物体的体重。在这个“自私大脑理论”中,新皮质和边缘系统在诸如神经性厌食症、肥胖症和II型糖尿病等疾病的发病机制中起核心作用。从这些考虑来看,肥胖症的主要干扰似乎是该系统海马体设定点的位移。反馈系统由此产生的持续激活必然导致交感 - 肾上腺系统同样持续激活,进而引发胰岛素抵抗、高血压和代谢综合征的其他成分。现有的治疗代谢综合征的疗法(α和β肾上腺素能受体阻滞剂、胰岛素和胰岛素促分泌剂)干扰的是一些必须被视为代偿性的机制。这就解释了为什么从长远来看这些疗法令人失望。将讨论基于“自私大脑理论”的新治疗策略。