高压对象系统的压力测试(二):线粒体极端突变率下核心架构的不可塑性 对MACSM 数学总纲 2.0接口匹配瓶颈的独立系统验证

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Abstract (English)

The mitochondrial genome provides an independent high-pressure system, complementary to the Y chromosome, for testing the extrapolation of microevolutionary mechanisms to macroevolutionary timescales. Mitochondria exhibit a point mutation rate roughly an order of magnitude higher than the nuclear genome, while maintaining a very small effective population size. Under the strong version of the expectation that linearly extrapolates the effects of high mutation rate, small effective population size, and absence of recombination to the core architectural level, this combination should constitute an ideal setup for accelerated evolution—weaker purifying selection and easier accumulation of deleterious mutations. However, existing cross-species comparative data show that the core functional architecture of mitochondria (the catalytic core of the oxidative phosphorylation complexes and the mitochondrial genetic coding system) has remained highly conserved over the long evolutionary history of eukaryotes, creating a significant tension between the observed conservation of core sites and the extrapolated expectations based on mutation rate and effective population size. This “high mutation rate—high conservation” tension constitutes a second independent stress test, distinct from the Y chromosome, for the extrapolation of classical microevolution.

Using the parameterized framework of the MACSM Mathematical Framework 2.0, this paper explicitly quantifies the strong-version expectations of the classical extrapolation and confronts them with cross-species mitochondrial comparative genomic data, deep mutational scanning results, and clinical genetic evidence from human mitochondrial diseases. The results indicate that the elevated mutation rate only improves path accessibility in the single dimension of “candidate input,” whereas the joint constraints of the interface compatibility factor κ, the intermediate-state load function S(σ), and the synergistic complexity k are no more relaxed in the core mitochondrial system than in the nuclear genome. The multiplicative superimposition of multiple constraints means that the core architecture shows no empirical pattern of being continuously reshaped by zero-bridge mutational paths over deep timescales. Mitochondria thus provide a second set of independent empirical support for the core thesis of MACSM: increased mutational supply does not automatically translate into architectural innovation capacity; in core systems with highly coupled interfaces and highly synergistic functions, the strength of architectural constraints is determined by the internal organizational logic of the system, not by the rate of variational input.