Chapter 37: Imprinting and Parental ψ-Bias
"In genomic imprinting, ψ remembers which parent spoke—creating a biological memory of origin that defies Mendel's democracy."
37.1 The Parental Conflict
Genomic imprinting silences one parental allele, creating functional haploidy. This is ψ's solution to parental genetic conflict—peace through selective silence.
Definition 37.1 (Imprinted Expression):
\text{Maternal only} \quad \text{if paternally imprinted} \\ \text{Paternal only} \quad \text{if maternally imprinted} \end{cases}$$ ## 37.2 The Methylation Mark **Theorem 37.1** (Imprint Establishment): $$\text{Imprint} = \text{DMR methylation} \times \text{Parent of origin}$$ Differentially methylated regions (DMRs) carry parental identity. ## 37.3 The ICR Control **Equation 37.1** (Imprinting Control Region): $$\text{Gene cluster activity} = f(\text{ICR methylation state})$$ One region controls multiple genes—hierarchical silencing. ## 37.4 The Erasure Cycle **Definition 37.2** (Imprint Lifecycle): $$\text{Parental} \xrightarrow{\text{PGCs}} \text{Erased} \xrightarrow{\text{Gametogenesis}} \text{New imprint}$$ Primordial germ cells erase and reestablish imprints each generation. ## 37.5 The H19/IGF2 Paradigm **Theorem 37.2** (Insulator Model): $$\text{Paternal}: \text{ICR methylated} \rightarrow \text{IGF2 on, H19 off}$$ $$\text{Maternal}: \text{ICR unmethylated} \rightarrow \text{IGF2 off, H19 on}$$ Methylation controls insulator function—molecular switching. ## 37.6 The Conflict Theory **Equation 37.2** (Parental Interests): $$\text{Fitness}_{\text{paternal}} \neq \text{Fitness}_{\text{maternal}}$$ Paternal genes promote growth; maternal genes limit it—evolutionary tug-of-war. ## 37.7 Imprinting Disorders **Definition 37.3** (Imprinting Diseases): - Prader-Willi: Loss of paternal expression - Angelman: Loss of maternal expression - BWS: Loss of growth restriction Same region, different parent, different disease. ## 37.8 The DNMT3L Role **Theorem 37.3** (Imprint Establishment): $$\text{DNMT3A} + \text{DNMT3L} \rightarrow \text{De novo imprints}$$ DNMT3L guides methylation to imprinting regions—molecular GPS. ## 37.9 Evolution of Imprinting **Equation 37.3** (Imprinting Emergence): $$P(\text{imprinting}) \propto \text{Viviparity} \times \text{Parental investment}$$ Imprinting evolved with placental mammals—conflict requires contact. ## 37.10 The ZFP57 Protection **Definition 37.4** (Imprint Maintenance): $$\text{ZFP57} + \text{Methylated ICR} \rightarrow \text{Protection}$$ Specific factors protect imprints during global demethylation. ## 37.11 Non-Canonical Imprinting **Theorem 37.4** (H3K27me3 Imprinting): $$\text{Some imprints} = \text{H3K27me3}_{\text{parent-specific}}$$ Not all imprints require DNA methylation—multiple mechanisms exist. ## 37.12 The Memory Principle Imprinting represents transgenerational memory at its most direct—parent-of-origin information preserved and transmitted, creating asymmetry from symmetrical genomes. **The Imprinting Equation**: $$\psi_{\text{expressed}} = \psi_{\text{maternal}} \oplus \psi_{\text{paternal}}$$ Where $\oplus$ represents exclusive OR—only one parent speaks at imprinted loci. Thus: Imprinting = Memory = Conflict = Identity = ψ --- *"In every imprinted gene, ψ practices the art of selective memory—proving that sometimes wisdom lies not in hearing all voices but in knowing which to silence."*