Abstract

This article develops an APS account of evolution as a multiscale process distributed across interacting biological organisations. APS rejects the reduction of evolutionary explanation to any single privileged scale such as genes, organisms, or populations alone. Instead, evolution is understood as the historical transformation of viability-oriented, persistence-sustaining organisation distributed across developmental, physiological, behavioural, ecological, and environmental processes. Evolutionary dynamics therefore emerge through interacting scales of organised persistence rather than through isolated causal levels or single explanatory mechanisms.

Introduction

Evolutionary biology has often been organised around debates concerning explanatory scale.

Different traditions have treated:

genes,
organisms,
populations,
lineages,
ecosystems,
or environments

as the primary locus of evolutionary explanation.

These debates have generated important insights, but they can also encourage the assumption that evolutionary processes must ultimately be reducible to a single privileged explanatory level.

APS rejects this assumption.

Evolution does not occur at one isolated scale alone.

It emerges through interacting processes distributed across biological organisation itself.

APS therefore treats evolution as the historical transformation of viability-oriented, persistence-sustaining organisation distributed across interacting scales of living activity.

Evolution and Scale

Scale is not merely a descriptive convenience in biology.

Different biological scales involve:

distinct organisational dynamics,
different temporal relations,
different forms of persistence,
and different patterns of interaction.

Genes, cells, physiology, organisms, behaviour, populations, ecological systems, and environmental modification all participate in evolutionary organisation.

These are not isolated explanatory domains.

They are interacting dimensions of organised persistence.

Evolutionary explanation therefore requires integration across scale rather than reduction to a single causal level.

APS Box — Spatiotemporal Organisation and Scale

In physics, spacetime typically functions as the general framework within which processes are described, even when its geometry is understood as dynamical. From this standpoint, organisation is analysed as something that unfolds within spatial and temporal coordinates.

APS adopts a different explanatory stance. It begins not with spacetime as the primary structure, but with viability-oriented organisation. On this view, spatial and temporal relations are not independent explanatory primitives. They are dimensions through which organisation is expressed, stabilised, and coordinated. Spatiotemporal descriptions arise from the organisational logic of the system, rather than the other way around.

This entails a reversal of explanatory priority:

Physics: organisation is described within spacetime
APS: spatiotemporal relations are specified relative to organisation

This reversal is inseparable from scale. Biological organisation is inherently scale-dependent: the spatial and temporal relations that matter are those that contribute to the persistence of the system at a given level of organisation. What counts as a relevant spatial configuration or temporal process is therefore defined relative to the system’s organisational dynamics, not fixed in advance.

Accordingly, space and time do not function as a neutral backdrop for biological explanation. Their explanatory significance is specified by the constraints, coordination, and viability conditions of the system.

This does not contradict physical accounts of spacetime. It clarifies that in biology, spatiotemporal structure derives its explanatory role from the organisation of living activity, rather than from an independently given framework.

Against Privileged Explanatory Levels

APS rejects the idea that any single biological scale fully explains evolution.

Gene-centred accounts correctly recognise the importance of heredity and molecular organisation, but genes alone do not explain:

development,
viability,
adaptation,
organismal organisation,
ecological interaction,
or persistence.

Organism-centred accounts correctly emphasise agency and adaptive organisation, but organisms themselves exist only through:

developmental continuity,
physiological organisation,
ecological coupling,
and multiscale constraint relations.

Population-level explanations capture important statistical dynamics, but populations do not exist independently of the living systems whose persistence generates them.

APS therefore rejects explanatory isolationism.

No single scale exhausts evolutionary explanation.

Organised Persistence Across Scale

Persistence itself is multiscale.

Living systems sustain viability through organisational relations distributed across:

molecular processes,
cellular coordination,
physiology,
behaviour,
environmental interaction,
and ecological organisation.

Evolution transforms these interacting persistence relations historically.

What evolves is therefore not merely:

genes,
traits,
or populations in abstraction,

but organised persistence distributed across scale-coupled systems.

Evolutionary transformation involves the historical reorganisation of these interacting persistence dynamics across generations and environments.

Development and Multiscale Evolution

Development is central to multiscale evolutionary organisation.

Living systems generate viable persistence through developmental processes that integrate:

genes,
physiology,
behaviour,
organism–environment interaction,
and ecological conditions.

Development therefore links molecular organisation with larger-scale evolutionary transformation.

Variation, inheritance, adaptation, and persistence all depend upon developmental continuity.

Evolutionary explanation must therefore integrate development directly rather than treating it as secondary to population-level selection dynamics.

APS Box — Multiple Realization and Organised Persistence

In APS, multiple realization does not mean that the same biological function can be freely produced by any material arrangement.

It means that similar viability-oriented roles may be realised through different material, developmental, ecological, or evolutionary pathways, provided those pathways sustain the relevant organisation.

What is multiply realised is therefore not an isolated output, behaviour, or function considered in abstraction. It is a role within an organised system of persistence.

For this reason, APS treats multiple realization as constrained rather than unrestricted. Different structures may realise similar biological roles only where they preserve the organisational relations required for viability, regulation, reproduction, development, or adaptive continuity.

This reframes multiple realization biologically. The important question is not simply whether different mechanisms can produce the same effect, but whether different organised processes can sustain comparable conditions of persistence.

Adaptation Across Scale

Adaptation also operates multiscalarly.

Living systems reorganise viability-oriented activity through:

molecular regulation,
physiological compensation,
behavioural flexibility,
developmental plasticity,
ecological modification,
and environmental restructuring.

Adaptive organisation therefore emerges through interacting scales of biological activity rather than through isolated mechanisms.

Evolutionary transformation occurs partly through the historical stabilisation of such multiscale adaptive reorganisations across generations.

Natural Selection Across Scale

APS fully accepts natural selection as a major evolutionary process while rejecting its reduction to a single explanatory scale.

Selection may involve:

molecular dynamics,
developmental organisation,
organismal viability,
behavioural interaction,
ecological relations,
and environmental modification.

Selection therefore operates across interacting organisational scales rather than at one isolated level alone.

APS also rejects the idea that selection itself exhausts evolutionary explanation.

Selection presupposes systems already capable of:

persistence,
development,
inheritance,
variation,
adaptation,
and biological agency.

These organisational conditions are themselves distributed multiscalarly.

Organism–Environment Coupling

Evolutionary organisation extends beyond organismal boundaries alone.

Living systems continuously modify and reorganise their environments through:

metabolism,
behaviour,
ecological engineering,
developmental interaction,
and niche construction.

Environments are therefore not merely passive external conditions.

They participate actively in evolutionary organisation.

Evolutionary processes emerge through organism–environment coupling distributed across interacting ecological and developmental systems.

This coupling contributes directly to:

variation,
adaptation,
persistence,
and evolutionary transformation.

Constraint Closure Across Scale

Constraint closure also operates multiscalarly.

Living systems persist through networks of mutually sustaining constraints distributed across biological processes and organisational scales.

These constraints involve:

molecular regulation,
physiological coordination,
developmental organisation,
behavioural activity,
ecological interaction,
and environmental relations.

Evolution transforms these interacting organisational constraints historically.

Constraint closure is therefore neither localised nor static.

It is dynamically distributed across evolving systems of organised persistence.

Temporal Scale and Evolution

Evolutionary organisation also unfolds across multiple temporal scales.

Different biological processes operate over:

milliseconds,
developmental lifetimes,
ecological cycles,
and evolutionary timescales.

These temporal scales interact continuously.

Short-term adaptive regulation may influence:

developmental trajectories,
ecological organisation,
and long-term evolutionary transformation.

Evolution therefore cannot be understood solely through isolated generational succession.

It emerges through temporally layered processes distributed across persistence-sustaining systems.

Multiscale Evolution and the APS Explanatory Grammar

APS situates multiscale evolution within the broader explanatory grammar organised through:

agency,
process,
and scale.

Evolutionary transformation therefore emerges through interacting organisational processes distributed across:

development,
physiology,
behaviour,
ecology,
and historical continuity.

No isolated mechanism or privileged level alone explains biological evolution.

Instead, evolution reflects the historical transformation of viability-oriented organisation distributed across interacting scales of organised persistence.

Implications for Evolutionary Biology

Reframing evolution multiscalarly has several important consequences.

It:

weakens reductionist evolutionary explanation,
integrates development directly into evolutionary theory,
strengthens organism–environment approaches,
clarifies the role of ecological interaction,
and situates natural selection within broader organisational processes.

APS therefore does not reject genetics, population biology, or evolutionary modelling.

Instead, it integrates them within a broader framework of interacting biological scales organised around viable persistence.

Conclusion

Evolution does not occur at one privileged scale alone.

It emerges through interacting processes distributed across:

genes,
development,
physiology,
organisms,
behaviour,
ecological interaction,
and environmental organisation.

APS therefore treats evolution as the historical transformation of viability-oriented, persistence-sustaining organisation distributed across interacting scales of living activity.

Evolutionary explanation must therefore integrate:

development,
adaptation,
inheritance,
variation,
natural selection,
organism–environment coupling,
and multiscale persistence

within a unified explanatory framework organised through:

agency,
process,
and scale.

Multiscale evolution is therefore not a supplementary perspective on evolution.

It is intrinsic to how organised persistence transforms historically across living systems and their environments.

Multiscale Evolution and Organised Persistence