Colonial Readiness: Life at the Boundary of the One and the Many: 2 Ability: The Colony’s Distributed Aperture

Colonial organisms such as Volvox are often described as “simple multicellular algae.” This phrase conceals more than it reveals. For what Volvox demonstrates with uncompromising clarity is that the capacity of a biological system—its ability—is never contained in any one of its parts. Ability is a horizon of possibility structured by relation, not a repertoire stored in code.

This is the first pillar of the readiness triad. In the context of colonial life, it becomes the illumination that reveals how collective behaviour, developmental transformations, and evolutionary transitions are even possible.

To understand colonial organisms, we begin not with their cells but with their aperture: the integrated configuration that makes certain enactments possible and others impossible.

Ability is not local: no cell knows how the colony swims

Conventional accounts attribute function to individual cells:

• somatic cells “drive motility,”

• gonidia “manage reproduction,”

• light-sensitive cells “steer phototaxis.”

But none of this explains how Volvox swims. Or turns. Or maintains stability. Or inverts its body plan during development.

No cell possesses the capacity for coordinated rotation or directional swimming. No cell contains the “program” for these behaviours. Cells produce local forces; the colony-level behaviour is an enacted potential of the entire relational configuration.

In other words: the colony swims; cells beat.

The two are not commensurate.

Ability lives at the scale where relational potentials are integrated.

The colony as a hydrodynamic aperture

A Volvox colony is a spherical shell of cells embedded in a dense extracellular matrix (ECM). What this configuration actualises is extraordinary:

• Every somatic cell’s flagellum contributes a tiny, angled stroke.

• Hydrodynamic coupling through the ECM aligns these strokes.

• The spherical geometry funnels these micro-forces into a coherent global motion.

This is not emergence.

This is not “coordination” as a behavioural miracle.

This is structural readiness: the colony’s ability is built into its relational architecture.

If any of these relational constraints were altered—ECM viscosity, cell spacing, geometric curvature—the ability itself would shift. Not because the genome changed, but because the aperture that actualises possibility was rearranged.

This is why ability cannot be located in cells.

It is jointly constituted by:

• spatial arrangement

• ECM mechanics

• hydrodynamic couplings

• polarity axes

• the geometry of the colony itself

The colony does not add these features together; it is these features construed as a horizon of potential.

Ability as structured potential: not function, not instruction

In representational biology, ability is treated as “function”: what the organism does, usually linked to genetic programs or functional modules.

This is the wrong category entirely.

Ability is not a property possessed by a biological entity.

It is the structured set of enactable possibilities defined by the relational configuration.

A single cell can beat a flagellum.

A colony can swim, rotate, stabilise itself, and respond directionally to light.

The colony’s ability is not richer because it has more cells; it is richer because its relational architecture supports modes of enactment that were not available before.

This is the shift from mechanism to readiness.

Why ECM mechanics matter more than genetic detail

One of the most revealing facts about the volvocine algae is that alterations to the ECM produce dramatic shifts in colony-level behaviour—even when the cells themselves remain physiologically normal.

Experiments show:

• Modify ECM stiffness → swimming becomes unstable.

• Alter ECM connectivity → inversion succeeds or fails.

• Change ECM thickness → phototaxis improves or degrades.

This is exactly what a readiness ontology predicts: ability is sensitive to system-level relational constraints, not to the representational content of individual cells.

The ECM is not a scaffold; it is part of the horizon of ability. Its mechanical properties shape the field of possible enactments.

Ability and polarity: the anterior makes sense only because the colony does

The anterior pole of Volvox is packed with cells biased toward enhanced photoreception and directed flagellar activity. But these local enhancements only matter because the colony provides the integrative architecture that makes directional turning possible.

Strip away the colony, and “anterior identity” becomes meaningless.

Polarity is a relational role, not a property of cells.

In readiness terms: local inclinations can only tilt within an ability-field that gives those tilts something to enact.

Ability comes first.

Inclination only modifies it.

Developmental ability: how inversion becomes possible

One of Volvox’s most iconic features is inversion: the colony turns itself inside-out during development so its outward-facing flagella can beat freely.

Conventional accounts treat inversion as a programmed developmental step. But inversion is not an instruction; it is a system-level recutting of readiness.

During inversion:

• ECM linkages loosen in specific regions;

• cell shapes change;

• mechanical stresses propagate globally;

• the entire architecture reconfigures.

Every local action participates in a collective shift of ability—from a configuration that cannot swim to one that can.

This demonstrates the central insight: development is the reorganisation of ability, not the execution of instructions.

Inversion is a re-opening and re-shaping of the colony’s aperture.

Ability and the continuum from unicellular to colonial life

Colonial transitions in the volvocine lineage are often described as “steps toward multicellularity.” But this teleology disguises the deeper truth: what evolves is not an organism but an aperture.

As volvocine species shift from dispersed to tightly integrated colonies, what changes is:

• how potential is distributed,

• how local actions resonate globally,

• how system-level ability becomes increasingly structured.

It is not “the same organism” getting more complex.

It is a different horizon of enactable possibility being formed.

From a readiness perspective:

• a unicell has one aperture;

• a colonial organism has many local loci contributing to a shared aperture;

• a fully multicellular organism is a highly integrated aperture with steep readiness gradients.

This is not hierarchy.

It is relational individuation of ability.

Why ability clarifies seemingly paradoxical behaviours

The readiness lens explains features of Volvox that otherwise appear paradoxical:

• Robust swimming: because ability is distributed, not centralised.

• Predictable turning biases: inclinations act within the colony’s ability-field.

• Fragile developmental inversion: ability must be restructured before behaviour becomes possible.

• Flexible division of labour: individuation tunes local construals of ability.

Every one of these behaviours reflects the colony actualising structured potential.

Where this leads

This post has established the centrality of ability: the colony’s distributed, relational horizon of possibility.

In the readiness triad:

• Ability provides the aperture.

• Inclination provides the tilt.

• Individuation provides the perspective.

The next post will take up inclination: how gradients, positions, and local biases shape the colony’s readiness without invoking instructions, programs, or functional roles.