We have spent much of this series following a movement that initially seemed quite simple.
The gene is inherited.
Inheritance carries organisation into the future.
Development transforms inherited organisation into biological form.
Development constrains and enables variation.
Organisms encounter environments.
Those encounters affect which variations matter.
And evolutionary history can alter the developmental and ecological conditions under which subsequent variation becomes possible.
At each stage, however, one thing has become increasingly difficult to ignore.
Nothing evolves in isolation.
A gene does not evolve by itself.
A developmental system does not evolve by itself.
An organism does not evolve by itself.
Even a population does not evolve in an empty space.
Evolution occurs within a web of relations.
And those relations do something more than provide the background against which evolution happens.
They help determine what can happen.
We have therefore reached the point at which our phrase "the transformation of possibility" needs another refinement.
Possibility is not merely developmental.
It is ecological.
The organism never meets an empty world
Imagine an organism encountering its environment.
It is tempting to picture two separate entities:
organism + environment.
The organism possesses traits.
The environment possesses conditions.
The two then interact.
But this picture already divides the world into things before asking what their relations are.
A better starting point is the relation itself.
A wing is not simply a thing that possesses the property "flight".
Flight emerges from a relation among wing, body, air, gravity, muscular organisation and behaviour.
A root does not possess "access to nutrients" independently of soil, water, microorganisms and surrounding plants.
A predator does not possess "predatory fitness" in isolation from prey.
A behaviour becomes advantageous only within an ecological context.
The significance of a biological structure therefore cannot always be located entirely inside the structure.
It emerges through what the structure can participate in.
This is one reason relational ontology has become increasingly useful to us.
The organism is not an isolated object subsequently placed into an environment.
It is already a participant in a world of relations.
From developmental possibility to ecological possibility
In the previous essay, we distinguished potential, possibility and actuality.
A developmental system possesses potentials.
Under particular conditions, some become possible.
Some possibilities become actual.
But now we need to add another layer.
A trait may be developmentally possible without being ecologically significant.
An organism may be capable of producing a particular structure, yet that structure may have no consequence in the environment it inhabits.
Conversely, an environmental change may make an existing capacity suddenly consequential.
This gives us another transformation:
developmental potential → ecological possibility → evolutionary actuality
The distinction matters.
A phenotype does not become evolutionarily important simply because development can produce it.
It becomes consequential through relations.
The environment can therefore function not merely as a filter that selects among already existing traits, but as part of the system that determines which capacities become evolutionarily meaningful.
The niche is not a box
This is why the ordinary image of an ecological niche can be misleading.
A niche can sound like a place into which an organism fits.
There is the organism.
There is the niche.
Evolution somehow makes the organism better suited to the niche.
But the relation is more complicated.
An organism's activities affect the conditions that constitute its niche.
Its morphology determines what resources it can exploit.
Its behaviour changes its surroundings.
Its competitors alter what resources remain available.
Its predators alter its behaviour.
Its symbionts alter its physiology.
Its descendants inherit some of these altered relationships.
The niche is therefore not simply a box waiting to be filled.
It is partly constituted through the activities and relations of organisms.
This is the significance of niche construction.
The organism does not merely adapt to an environment.
It can participate in changing the environment to which subsequent generations are exposed.
A beaver changes the question
The familiar example of the beaver is useful precisely because it makes the relational point so obvious.
A beaver builds a dam.
The dam changes the flow of water.
The altered water flow changes vegetation, sedimentation, temperature and habitat.
Other organisms respond to the changed conditions.
The beaver's descendants therefore encounter an environment that is not simply the environment their ancestors encountered.
Something has been added to the causal history.
The organism has become part of the construction of its own future ecological circumstances.
No foresight is required.
The beaver need not possess an evolutionary theory.
It does not need to intend the eventual consequences for descendants.
It simply performs activities characteristic of its way of life.
Yet those activities alter the world.
And the altered world changes what is possible for subsequent organisms.
This is precisely the kind of process in which the distinction between actuality and possibility becomes illuminating.
An actual biological activity can transform the conditions of future possibility.
The environment becomes historical
We have already argued that organisms inherit history through their biological organisation.
Now we can see that environments can carry history too.
A forest is not merely a collection of present organisms.
It bears traces of previous organisms.
Soil chemistry reflects past biological activity.
Habitats reflect previous construction.
Species distributions reflect historical interactions.
Predator–prey relationships have histories.
Symbiotic relationships have histories.
Even apparently "natural" environments are often the accumulated consequence of innumerable biological processes.
The environment therefore has a kind of historical memory.
Again, this is not memory as representation.
It is persistence.
The past remains present in the organisation of the world.
And that means that organisms inherit not only biological structures but, indirectly and sometimes directly, historically modified environments.
A second inheritance
This gives us a useful distinction.
Genetic inheritance is not the only form of continuity across generations.
There can also be ecological inheritance.
A descendant may encounter environmental conditions partly produced by its ancestors.
Birds inherit nests or nesting sites.
Plants alter soils in ways that affect subsequent generations.
Animals modify habitats.
Social animals transmit patterns of behaviour that alter the environment in which their descendants develop.
Microorganisms transform chemical environments.
None of these examples makes ecological inheritance identical to genetic inheritance.
It does something more interesting.
It shows that biological history can persist through different channels.
The past can reach the future through genes.
It can also reach the future through developmental organisation, behaviour, ecological structures and inherited relationships.
This does not mean that "everything is inheritance".
It means that inheritance itself is more heterogeneous than a simple gene-centred picture suggests.
The organism inherits a world
This leads to a rather striking thought.
An organism does not begin life with a genome and then encounter an otherwise untouched world.
It begins within a world already structured by history.
The developmental system is inherited.
The cellular environment is inherited.
The ecological circumstances may be inherited.
The behaviour of conspecifics may be inherited through social transmission.
The organism therefore enters a world in which many of the conditions for its development are already organised.
This is not Lamarckism.
We do not need acquired characteristics to be written back into the genome.
The point is simply that evolutionary continuity operates through multiple causal pathways.
The world encountered by an organism is partly the world made by previous organisms.
And that means:
the past is not merely inside the organism. It is also around it.
This is an important extension of our earlier argument.
The ecology of possibility
We can now give a more precise meaning to the title of this essay.
An ecology is not merely a collection of organisms.
It is a system of relations among organisms and their environments.
Within that system, some things are possible and others are not.
A predator makes some prey behaviours consequential.
A prey species makes some predator strategies viable.
A pollinator makes certain plant reproductive strategies possible.
A plant provides resources that make certain insect populations possible.
A symbiont can make physiological capacities available to its host.
A competitor can close an ecological opportunity.
The possibilities available to one organism are therefore partly constituted by the activities of others.
Possibility is not merely something an organism carries around with itself.
It is distributed across relations.
Possibility without a possessor
This gives us a subtle shift in language.
We normally ask:
"What can this organism do?"
But sometimes the more illuminating question is:
"What can this organism do here, with these other organisms, under these conditions?"
The second question is not a replacement for the first.
An organism genuinely possesses capacities.
But those capacities become consequential only within relations.
A bird may possess wings.
Whether those wings afford particular forms of movement depends on the organism's body, its musculature, the atmosphere and its environment.
A bacterium may possess a metabolic pathway.
Whether that pathway matters depends on the chemical environment.
A plant may possess a physiological tolerance.
Whether that tolerance affects fitness depends on ecological circumstances.
The capacity is real.
But its possibility is relational.
The grammar of ecological agency
This also brings us back to the question of agency.
We have been careful throughout the series not to describe genes as agents simply because grammatical constructions make them appear so.
But organisms really can act.
They move.
They forage.
They construct.
They reproduce.
They alter environments.
They respond to circumstances.
The important distinction is therefore not between "agency" and "no agency".
It is between different kinds of participation.
A gene is not an agent merely because we say:
"the gene produces..."
An organism may genuinely be an agent because it performs activities that alter its relations with the world.
Yet even organismal agency should not be isolated from the larger system.
The organism acts within conditions it did not create.
Its capacities were historically produced.
Its environment constrains it.
Its actions alter that environment.
Its descendants inherit the consequences.
Agency is therefore embedded.
It is not the return of the hidden designer.
Selection as relation
This gives us another way to think about natural selection.
The language of selection can encourage the image of an external selector.
Something called "the environment" examines organisms and selects the successful ones.
But there is no such examiner.
Selection emerges from differential survival and reproduction within a set of ecological relations.
A phenotype becomes advantageous because of what it enables an organism to do under particular conditions.
Change the conditions, and the advantage can disappear.
Change the organism, and the relation can change.
Change another species, and the consequences can change again.
Fitness is therefore not best thought of as a substance possessed by an organism.
It is a relational outcome.
The organism has capacities.
The environment has conditions.
Their interaction produces consequences for reproduction.
And those consequences feed back into inheritance.
This makes selection look less like a force acting upon organisms and more like a pattern emerging from relations among organisms and worlds.
The world selects nothing
This is worth stating explicitly.
The environment does not decide.
The predator does not vote.
The ecosystem does not compare candidates.
There is no agent called "natural selection" sitting outside the process.
Selection is our abstraction over a pattern of differential consequences.
This is another place where grammatical and conceptual metaphor can mislead.
We say:
"selection favours..."
and the verb gives the process an apparent agent.
But the biological reality is distributed across interactions.
Some organisms reproduce more successfully under particular conditions.
Their hereditary variants become more represented.
The population changes.
No one had to choose.
The apparent agency is an emergent property of the process we are describing.
Relations can create opportunities
The relational perspective also changes how we think about innovation.
A new possibility may arise not because a new intrinsic property appears, but because a new relation becomes available.
A structure that previously had little significance can become important when another species appears.
A behavioural pattern can become advantageous after an environmental change.
A metabolic pathway can become consequential when a new resource becomes available.
A symbiotic relationship can make a physiological capacity possible that neither partner could sustain independently.
This means that evolutionary novelty can arise through reconfiguration of relations.
The world does not have to acquire an entirely new object.
Sometimes the objects are already there.
What changes is what they can do together.
That is a deeply relational conception of becoming.
Symbiosis and the expansion of possibility
Symbiosis makes this especially striking.
Two organisms enter into a relationship.
The relationship changes what each can do.
The resulting system may possess capacities that neither organism could realise independently.
Again, we should avoid romanticising this.
Symbiosis can involve conflict, exploitation and instability as well as cooperation.
The important point is structural.
A relation can generate new biological capacities.
This means that the unit of evolutionary possibility cannot always be the isolated organism.
Sometimes the relevant unit is a relationship.
Sometimes it is a developmental system.
Sometimes an ecological network.
Sometimes a lineage.
The appropriate level depends on the question.
But the recurring lesson is that biological possibility is often organisationally distributed.
The ecology is not a backdrop
We can now see why the conventional separation between organism and environment becomes increasingly difficult to sustain.
If organisms modify environments,
and environments modify selection,
and selection changes organisms,
then organism and environment participate in a feedback system.
The environment is not simply where evolution happens.
It is part of what evolution is changing.
And because organisms modify environments, the process can be self-transforming without being self-directed.
Again, there is no paradox.
A process can alter its own conditions without intending to do so.
Rain alters a landscape.
The altered landscape changes subsequent water flow.
The new water flow alters erosion.
No agent is required.
Evolutionary systems can behave similarly.
The transformation of ecological possibility
We can now extend our central thesis one step further.
Evolution can change developmental possibility.
But organisms can also change ecological possibility.
A new organism can occupy a previously unused ecological relation.
Its activities can create resources or destroy them.
Its presence can make another organism viable.
Its disappearance can remove an ecological opportunity.
The ecological world therefore changes as organisms change.
So:
evolution changes organisms; organisms change ecological relations; ecological relations change future evolutionary possibilities.
This is a feedback loop.
And it means that the transformation of possibility is not confined to the genotype–phenotype relationship.
It occurs across the organism–environment relation as well.
Possibility becomes collective
This is perhaps the most surprising consequence.
Some possibilities do not belong to individual organisms at all.
They belong to systems of relations.
A coral reef creates possibilities for organisms that could not exist in the same form without the reef.
A forest creates possibilities for organisms whose lives depend upon the forest's structure.
A microbial community can create metabolic possibilities for its members.
A social group can create behavioural possibilities for individuals within it.
The possibility belongs neither entirely to the individual nor entirely to the environment.
It emerges from the relation.
This suggests that evolution can generate not merely new organisms but new ecological possibilities.
And once new ecological possibilities exist, they can become the setting for further evolutionary change.
The process is therefore cumulative in a deeper sense than simple genetic accumulation.
From population to ecology
This changes the scale of our investigation.
We began with the gene because Dawkins's question forced us to ask what persists through generations.
We then discovered that the gene participates in a larger developmental organisation.
Now we discover that the organism participates in a larger ecological organisation.
At each step, the previous level remains real.
Genes matter.
Development matters.
Organisms matter.
Populations matter.
Ecologies matter.
But none is sufficient as the complete description of evolutionary becoming.
The interesting phenomena occur partly between levels.
A gene matters through developmental processes.
A phenotype matters through ecological relations.
An ecological change matters through reproductive consequences.
A population change matters through altered future variation.
Evolution is therefore not simply a hierarchy of objects.
It is a network of processes operating at different scales.
The danger of holism
But here again we need discipline.
There is an easy temptation to say:
"Everything is connected."
That is true enough to be almost useless.
Relational ontology should not become a licence for vague holism.
Relations have to be specified.
What relates to what?
Through which process?
At what scale?
With what consequences?
Over what timescale?
Which relations are causally relevant?
Which are merely correlations?
Our conceptual framework is valuable only if it helps us ask these questions more precisely.
The point is not to dissolve biological entities into an undifferentiated web.
The point is to understand how entities and processes acquire their evolutionary significance through particular relations.
SFL and the ecology of process
This is also where our SFL perspective continues to earn its keep.
Language habitually packages processes as things.
We say:
"the adaptation"
"the niche"
"the selection"
"the environment"
"the fitness landscape".
These nominalisations are extraordinarily useful.
They allow us to stabilise complex processes as objects of discussion.
But they can also conceal the relationships that constitute those processes.
"The adaptation" hides adapting.
"The niche" hides the relations through which a niche is constituted.
"The selection" hides differential reproduction.
"The environment" can hide an enormous network of interactions.
Again, the point is not to reject the nouns.
It is to unpack them when necessary.
A process can become a thing in grammar without becoming a thing in nature.
And once we remember that, some of the apparent entities of evolutionary theory become conceptually more transparent.
The world as an affordance structure
We can now return to the idea of affordance.
An organism encounters a world that is structured by possibilities for action.
Some resources are accessible.
Some habitats are inhabitable.
Some predators can be escaped.
Some mates can be found.
Some behaviours can be performed.
But these affordances are not simply "out there".
They arise from relations between organismal capacities and environmental structures.
A branch affords perching to a bird because of the relation between branch and bird.
The same branch may afford something else to another organism.
The environment therefore does not contain a fixed list of affordances.
Affordances emerge relationally.
And because organisms evolve, the set of affordances available to them can evolve too.
This gives us another formulation:
Evolution can transform the affordance structure of a world.
That is a strong claim.
But it follows naturally from what we have already established.
The world becomes different by becoming inhabited differently
Perhaps this is the deepest ecological insight.
The environment of a lineage is not simply a physical stage that remains constant while organisms change.
It is partly the world that those organisms make available to one another.
A change in one species can transform the possibilities of another.
The emergence of flowering plants transformed the ecological possibilities available to pollinators.
The evolution of predators transforms the possibilities available to prey.
The emergence of oxygen-producing organisms transformed the chemical possibilities available to other forms of life.
The arrival of a new species can reorganise an ecosystem.
Again, none of this requires foresight.
It is enough that organisms interact.
Life changes the world in which life subsequently evolves.
Evolution becomes ecological memory
We can now bring together two forms of historical persistence.
The organism carries its history in its inherited organisation.
The environment carries history in its ecological organisation.
The two interact.
So the future organism encounters:
a biological inheritance
within
an ecological inheritance.
Neither is static.
Each changes the other.
This gives us a richer meaning of evolutionary memory.
The past persists not only in genes.
It persists in developmental architecture.
It persists in modified environments.
It persists in species relationships.
It persists in ecological structures.
It persists in the possibilities that previous evolutionary events have opened or closed.
The world itself becomes historical.
A world that remembers without representation
There is something philosophically striking about this.
Nothing needs to represent the past for the past to matter.
A gene sequence can preserve historical consequences.
A developmental architecture can preserve them.
A nest can preserve them.
A dam can preserve them.
A soil community can preserve them.
A symbiotic relationship can preserve them.
History can therefore become materially present.
This is perhaps a more general form of memory than representation.
And it helps explain why evolution can be cumulative without possessing a cumulative plan.
The system does not start from zero at every generation.
It starts from a world already transformed by previous activity.
The ecology of becoming
We can now state the thesis of this essay.
Evolution does not occur within a fixed ecology of possibilities.
It occurs within ecological relations that are themselves historically transformed.
Organisms alter environments.
Environments alter evolutionary consequences.
Relations among organisms generate new capacities and close others.
Ecological structures persist across generations.
And these persistent structures become conditions for subsequent evolution.
Therefore:
the evolution of life can transform not only biological forms but the ecological possibilities within which future forms can arise.
This gives "the transformation of possibility" a second, broader meaning.
It is not only the transformation of developmental possibility.
It is the transformation of ecological affordances.
And the two are inseparable.
Development makes organisms capable of particular things.
Ecology makes those capacities consequential.
Ecology can then alter the conditions of development and selection.
The process feeds back upon itself.
But what is actually evolving?
We have now accumulated enough pieces to ask a slightly unsettling question.
If genes evolve,
and developmental systems evolve,
and organisms evolve,
and ecological relations evolve,
then what exactly is the thing we call evolution?
Perhaps the question itself is revealing.
Maybe evolution is not best understood as a process performed by a particular object.
Perhaps it is a historical process distributed across levels of organisation.
Genes participate in it.
Organisms participate in it.
Populations participate in it.
Environments participate in it.
None is the evolutionary agent.
The process emerges from their interactions.
This returns us to where we began with Dawkins.
The gene was proposed as the replicator.
But perhaps the deeper lesson is that evolutionary continuity cannot be reduced to the persistence of one kind of entity.
What persists is organised through relations.
And those relations themselves change.
The ecology of the possible
We can now see why our original metaphor of possibility has become increasingly difficult to contain.
Possibility is not a box.
It is not a catalogue.
It is not an abstract space waiting to be searched.
It is a property of organised relations.
Development makes some forms possible.
Ecology makes some capacities consequential.
History makes some pathways available.
Existing organisms create conditions for other organisms.
And every actualisation can alter the conditions of subsequent possibility.
So perhaps we should stop imagining evolution as moving through possibility space.
Perhaps evolution is better understood as participating in the ongoing reorganisation of the conditions under which possibilities arise.
That is a much more modest claim than saying that life creates possibilities from nothing.
But it is also much more interesting.
The furrow is no longer only genetic
Our furrow metaphor has now deepened.
At first, the furrow represented inheritance.
The past left a structure that altered the future.
Then it represented developmental constraint.
The inherited organisation shaped the paths available to subsequent variation.
Now it becomes ecological.
Organisms themselves make furrows in the world.
Their activities alter the terrain.
Their descendants inherit a world partly shaped by those earlier movements.
The plough and the field are no longer easily separable.
The field changes the plough's possible paths.
The plough changes the field.
And the next movement occurs in a world altered by both.
This is perhaps why the metaphor has proved so persistent.
The furrow does not tell the plough where to go.
But neither does the plough encounter an untouched field.
The furrow is history made available as constraint and affordance.
The danger of saying "life creates possibility"
At this point, we should be especially careful with our language.
It would be tempting to conclude:
Life creates possibility.
But that statement is too strong.
Life does not create the physical laws.
It does not create all the conditions under which life exists.
Evolution does not have unrestricted freedom.
Many possibilities remain closed by physics, chemistry, development and history.
What organisms do is more specific.
They participate in reconfiguring the actual relations that determine which biological possibilities are accessible.
That is enough.
We do not need cosmic creativity.
We need historical transformation.
A naturalistic becoming
This is perhaps where Conceptual Naturalism reaches its most useful form in the series.
We are not proposing a mysterious metaphysics of becoming.
We are asking whether a familiar set of biological facts can be construed more adequately when we attend to processes rather than only the entities produced by them.
Genes participate in inheritance.
Development participates in the production of organisms.
Organisms participate in ecological relations.
Ecological relations participate in differential reproduction.
Historical actualities alter subsequent conditions.
The conceptual vocabulary of potential, possibility and actuality allows us to describe the temporal structure of those relations.
But the biology remains in charge.
Where the biological evidence does not support the philosophical claim, the philosophical claim must retreat.
That is the discipline we need to preserve.
What we can now see
At the beginning, evolution seemed to present us with a puzzle about design.
How can intricate biological organisation emerge without an organiser?
Dawkins's answer was powerful:
do not look for an organism-level designer; look at differential replication and selection.
We have retained the force of that insight while questioning the agentive language in which it was often expressed.
Then we discovered inheritance.
Inheritance preserves history.
Development transforms inheritance into form.
Development structures variation.
Evolvability shows that evolutionary history can alter the conditions under which future variation arises.
Ecology shows that organisms alter the conditions under which other organisms evolve.
And now the picture has become substantially richer.
Evolution is not merely:
inheritance + variation + selection.
It is a historical process in which inherited organisation, development, organismal activity and ecological relation continually alter one another.
The question that now becomes unavoidable
But this raises a final difficulty.
If evolutionary systems are so deeply relational, how should we understand the organism itself?
We have repeatedly used the organism as though it were a stable unit:
the organism develops;
the organism acts;
the organism encounters an environment;
the organism reproduces.
But what is an organism?
Is it merely a vehicle constructed by genes?
Clearly not.
Is it an autonomous agent?
Not in any absolute sense.
Is it a bounded object?
Physically, perhaps.
Biologically, the boundaries become much stranger.
The organism depends upon symbionts.
It exchanges matter and energy continuously with its environment.
Its development depends upon environmental signals.
Its behaviour modifies its surroundings.
Its evolutionary history is distributed across relationships.
The organism is therefore beginning to look less like an isolated thing and more like a temporally stabilised participant in a network of processes.
That does not make it unreal.
It makes it relational.
And if that is right, then perhaps we have been asking the wrong question again.
Instead of:
What is the organism?
we may need to ask:
What kind of organisation allows an organism to persist as an organism while participating in relations that continually transform it?
That question takes us toward the next stage of the series.
Because the deeper we go, the harder it becomes to locate evolution in a single entity.
Perhaps evolution is not something that happens to organisms.
Perhaps it is something that happens through organised participation.
And if so, the most interesting evolutionary question may no longer be:
What survives?
but:
What kinds of organisation can participate in becoming?
That is where we should go next. 🍷🙂
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