Development in colonial organisms is often described as a sequence of morphological steps. For Volvox, this narrative centres on one of the most striking events in biology: inversion. Traditional accounts frame inversion as a programmatic manoeuvre—a preordained movement of cells that flips the colony inside-out so that flagella face outward. But in the lens of readiness, inversion is not a prewritten instruction; it is a relational reconfiguration of the colony’s potential.
Inversion illustrates development as recutting of readiness fields. The colony does not “execute” inversion. It becomes inverted when the distributed field of potentials—comprising cellular inclinations, ECM mechanics, and perspectival loci—reorganises to open a new horizon of enactable possibilities. Development is not an assembly from parts. It is a transformation of the aperture itself.
ECM as the medium of potential
The extracellular matrix (ECM) is more than structural glue. It is the medium through which the colony’s potential is distributed. In inversion:
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local ECM tensions shift,
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linkages loosen or tighten,
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mechanical stresses propagate through the sphere,
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and the colony’s relational geometry re-partitions its readiness.
A single cell cannot invert the colony; it cannot even “know” what inversion is. Instead, its local construal—its perspectival enactment of what the colony can do—is constrained and directed by ECM-mediated couplings. Inversion is the system-level unfolding of readiness, orchestrated through relational feedback rather than representational programming.
Recutting as distributed negotiation
Each cell’s local perspective participates in a negotiation with its neighbours. Somatic cells at the anterior bend, posterior gonidia shift, and flagella realign. No cell dictates the outcome; no external agent imposes the flip. Instead, the colony actualises inversion through the alignment of perspectival loci across the ability and inclination fields.
The “rules” are not rules at all. They are constraints embedded in the relational configuration. The colony does not follow a script; it resolves into a configuration that makes inversion possible—an event emergent from the interplay of local inclinations, distributed abilities, and partial individuation.
Development as dynamic restructuring of potential
Inversion exemplifies a broader principle: development is not a linear execution of pre-specified steps. It is the dynamic restructuring of the colony’s readiness, where the field of potential is continuously cut, re-cut, and re-partitioned:
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Flagellar orientation is reshaped by local inclinations.
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Cell shape changes redistribute mechanical forces.
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ECM elasticity modulates which potential paths can be actualised.
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Perspectival loci (cells) adjust their enactments in response to these shifts.
Every developmental transition is therefore a relational reconfiguration: a colony-level reorganisation that makes new behaviour possible without invoking instruction, plan, or central control.
Why inversion is a test case for readiness
Inversion’s elegance lies in its clarity: it is visually dramatic, mechanically intricate, and relationally dense. If colonial life were reducible to mechanism or blueprint, inversion would be inscrutable without mapping every genetic and molecular interaction. But readiness reframes it:
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The colony can only invert because the ECM couples local forces across the sphere.
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The colony can only invert because cells enact local construals aligned with the distributed readiness.
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Inversion is not caused by genes; it is allowed by the relational configuration they partially enable.
The event is a direct manifestation of relational potential, not an execution of a prewritten developmental program.
Developmental flexibility and evolutionary insight
Because development is enacted readiness, it is plastic and evolvable:
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Modifying ECM stiffness or connectivity changes the colony’s developmental repertoire.
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Shifting cell inclinations can alter inversion timing or trajectory.
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Perturbations reveal which local constraints are essential to the collective potential.
This perspective reframes evolutionary questions: transitions to multicellularity or colonial complexity are not merely about novel genes or “steps toward the organism.” They are about reshaping relational fields of ability, inclination, and individuation—creating new apertures of potential actualisable by a colony.
Where this leads
Inversion demonstrates that colonial development is a process of relational reconfiguration, where new possibilities emerge not through instruction but through the coordinated adjustment of perspectival loci across structured potential. The colony develops not by executing a program but by recutting its own readiness.
The next post will explore the evolution of colonial life: how shifts in ability, inclination, and individuation shape transitions from unicellular ancestors to integrated colonies, and how readiness landscapes frame evolutionary trajectories without recourse to teleology.
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