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Chapter 40: Homomeric vs Heteromeric ψ-Coding

"In oligomerization, ψ explores symmetry and asymmetry—proteins assembling with identical copies or different partners, each choice encoding different functional possibilities."

40.1 The Oligomerization Dichotomy

Protein oligomerization represents ψ's exploration of combinatorial assembly—homomeric complexes achieving symmetry through self-association, heteromeric complexes creating diversity through different subunit combinations.

Definition 40.1 (Oligomer Types): Homomeric=An\text{Homomeric} = \text{A}_n Heteromeric=AαBβ...Zζ\text{Heteromeric} = \text{A}_\alpha\text{B}_\beta...\text{Z}_\zeta

Identity versus diversity in assembly.

40.2 Symmetry Advantages

Theorem 40.1 (Homomeric Benefits):

  • Genetic economy (one gene → multiple subunits)
  • Error correction through averaging
  • Cooperativity through symmetry

Efficiency through repetition.

40.3 The Dihedral Symmetry

Equation 40.1 (D_n Symmetry): Dn=Cn+nC2\text{D}_n = \text{C}_n + n\text{C}_2

N-fold rotation plus perpendicular 2-folds.

40.4 Isologous vs Heterologous

Definition 40.2 (Interface Types): Isologous:Same surface on both subunits\text{Isologous}: \text{Same surface on both subunits} Heterologous:Different surfaces\text{Heterologous}: \text{Different surfaces}

Symmetry of interaction surfaces.

40.5 Domain Swapping

Theorem 40.2 (3D Domain Swapping): MonomerclosedDimerswapped\text{Monomer}_{\text{closed}} \rightleftharpoons \text{Dimer}_{\text{swapped}}

Exchange of identical structural elements.

40.6 Heteromeric Complexity

Equation 40.2 (Subunit Diversity): Functions=f(iSubunitiαi)\text{Functions} = f(\prod_i \text{Subunit}_i^{\alpha_i})

Emergent properties from subunit combination.

40.7 The Hemoglobin Model

Definition 40.3 (α₂β₂ Tetramer): Hb=α2β2\text{Hb} = \alpha_2\beta_2 Interfaces:α1β1,α1β2,α1α2\text{Interfaces}: \alpha_1\beta_1, \alpha_1\beta_2, \alpha_1\alpha_2

Multiple interface types in one complex.

40.8 Evolution of Quaternary Structure

Theorem 40.3 (Gene Duplication): HomomericDuplication + DivergenceHeteromeric\text{Homomeric} \xrightarrow{\text{Duplication + Divergence}} \text{Heteromeric}

Asymmetry evolving from symmetry.

40.9 Allosteric Regulation

Equation 40.3 (Symmetry and Cooperativity): Hill coefficientnsubunits\text{Hill coefficient} \leq n_{\text{subunits}}

Maximum cooperativity limited by oligomeric state.

40.10 Assembly Specificity

Definition 40.4 (Partner Selection): Kdcorrect<<KdincorrectK_d^{\text{correct}} << K_d^{\text{incorrect}}

Discrimination ensuring proper assembly.

40.11 Stoichiometry Determination

Theorem 40.4 (Balanced Expression): Stoichiometry=f(Expression levels,Kd values)\text{Stoichiometry} = f(\text{Expression levels}, K_d\text{ values})

Cellular mechanisms ensuring correct ratios.

40.12 The Coding Principle

Homo- and heteromeric assemblies embody ψ's different strategies for creating functional complexity—symmetry providing robustness and cooperativity, asymmetry enabling specialization and regulation.

The Oligomerization Equation:

\text{Symmetry}(\psi^n) \quad \text{Homomeric} \\ \sum_i f_i(\psi_i) \quad \text{Heteromeric} \end{cases}$$ Different assembly logics for different needs. Thus: Oligomerization = Choice = Symmetry/Asymmetry = Function = ψ --- *"In choosing between homomeric and heteromeric assembly, ψ reveals that both unity and diversity have their place—that repetition creates strength while variation enables sophistication. Each oligomer encodes its functional logic in its assembly pattern."*