We have arrived at an unusual point in our journey.
We began with inheritance.
Dawkins asked how biological information persists.
The answer appeared to be the replicator: something capable of being copied, with variation and differential persistence.
But the further we have followed the question, the less satisfactory it has become to imagine inheritance as the transmission of a thing.
Genes are inherited, certainly.
But genes do not develop into organisms by themselves.
Development transforms inherited organisation.
Organisms participate in environments.
Environments carry histories.
Organisms modify those environments.
Ecological relations alter subsequent possibilities.
And those altered possibilities affect what can be inherited, selected and developed in future generations.
So perhaps we need to ask the question differently.
Not:
What is inherited?
but:
What does inheritance make possible?
And then one step further:
Can what is inherited be understood as a way of participating in the world?
This is where the distinction between inheritance and becoming begins to dissolve.
Inheritance is not a photocopier
The metaphor of copying has been extraordinarily influential in evolutionary thought.
A replicator is copied.
Copies resemble their predecessors.
Occasionally they differ.
Selection acts upon the differences.
The process accumulates changes.
There is an obvious truth here.
Biological reproduction does preserve organisation across generations.
But "copying" can suggest something more passive and exact than biological inheritance actually is.
A biological lineage does not simply reproduce an object.
It reproduces under conditions.
The inherited material enters a developmental system.
The developmental system interacts with an environment.
The resulting organism behaves.
That behaviour changes circumstances.
The circumstances affect reproduction.
The next generation therefore begins not merely with a copy of the previous generation, but with a new configuration of inherited organisation and altered conditions.
Inheritance is therefore not the preservation of an object against change.
It is a mechanism through which continuity becomes available for transformation.
What persists?
This gives us a deceptively difficult question.
What exactly persists across generations?
A nucleotide sequence may persist.
But so may developmental organisation.
So may a bodily architecture.
So may a behavioural tendency.
So may an ecological structure.
So may a relationship between organisms.
Some of these are genetically transmitted.
Some are environmentally transmitted.
Some are reconstructed afresh in each generation.
Some depend upon social learning.
Some depend upon developmental conditions.
The mechanisms differ enormously.
We should therefore resist the temptation to call all of them "replication".
The important common feature is not that they are copies.
It is that some organisation of the past contributes causally to the organisation of the future.
That is a broader and more useful conception of inheritance.
Inheritance as continuity of organisation
Perhaps we can therefore formulate a provisional definition:
Inheritance is the persistence of organisation across generations in a form that can contribute to subsequent development.
This deliberately says nothing about genes being the sole vehicle.
It also says nothing about exact copying.
The organisation may be altered.
It may be recombined.
It may interact with new circumstances.
It may generate something unprecedented.
But enough continuity remains for history to matter.
Without continuity, there could be no cumulative evolution.
Without transformation, there would be no evolution.
Inheritance is therefore poised between the two.
It is continuity that permits novelty.
The paradox of inheritance
This gives inheritance a curious structure.
If inheritance preserved everything exactly, evolution would stop.
If inheritance preserved nothing, evolution could not accumulate.
Evolution therefore depends upon a balance:
enough persistence for history to matter; enough variation for history to change.
This is not a compromise between two independent processes.
The two are coupled.
Variation matters because there is inheritance.
Inheritance matters because variation occurs within a historical continuity.
The future can differ from the past precisely because it does not begin from scratch.
That may be one of the deepest facts about evolution.
Inheriting constraints
We have already seen that inherited organisation constrains development.
But perhaps "constraint" sounds more negative than it should.
A constraint closes some possibilities.
But by doing so, it can also stabilise others.
A developmental system cannot produce every conceivable form.
That is not simply a limitation.
It is what makes organised form possible.
A language cannot contain every possible sequence of sounds if it is to remain recognisable as a language.
A musical form cannot permit every possible transition and still retain its identity.
A developmental system cannot generate every imaginable organism.
The constraints are part of what make structured novelty possible.
Inheritance therefore passes forward not only capacities but constraints on capacities.
And those constraints can themselves evolve.
Inheriting affordances
The same point applies to ecological possibility.
An organism inherits a developmental organisation.
That organisation gives it certain capacities.
Those capacities make certain interactions with the environment possible.
But the environment is itself historically structured.
So the organism effectively inherits an affordance structure.
Not because affordances are encoded in genes.
Rather, because inherited organisation and inherited ecological circumstances combine to make some actions possible and others difficult.
A bird inherits wings.
Its lineage also inherits an ecological history in which wings can be useful.
A plant inherits a developmental system.
Its descendants may encounter soils partly altered by previous organisms.
A social animal inherits behavioural capacities into a social environment shaped by previous members of its species.
Inheritance therefore becomes layered.
The organism inherits a body.
It may also inherit a world already partly structured by other organisms.
The inheritance of a practice
This becomes particularly clear when we consider behaviour.
A behaviour can be repeated across generations without being genetically encoded as a detailed instruction.
An animal learns from others.
The learned behaviour alters the environment.
Subsequent individuals encounter that altered environment.
They learn the behaviour again.
The pattern persists.
Here there is continuity without genetic replication.
We should not therefore call every persistent behaviour a "meme".
That would simply reproduce the problem in another vocabulary.
But we can recognise something important:
a pattern can persist historically because organisms reconstruct it through participation.
The continuity lies not in the copying of a static object but in the repeated enactment of a pattern.
This is inheritance as reconstruction.
Reconstruction rather than transmission
The word "transmission" can also mislead.
It suggests something travelling from one location to another.
A packet is transmitted.
A message is transmitted.
A genetic sequence can be transmitted.
But a practice is often not transmitted in that sense.
It is learned.
Reproduced.
Modified.
Adapted.
Re-enacted.
Its continuity depends upon participants reconstructing it.
Language provides a familiar example.
A child does not receive a language as a finished object.
The child enters a semiotic system and develops the capacity to participate in it.
The language persists because speakers continually instantiate its potential in new acts of meaning.
This is not exact copying.
It is continuity through participation.
Biological inheritance is stranger than cultural inheritance
This analogy also helps us see something about biology.
We often imagine cultural inheritance as complicated because it involves learning and interpretation.
Biological inheritance seems simpler.
But biological development is itself an extraordinarily active process.
The inherited genome does not contain a miniature organism.
It participates in a developmental system.
Cells communicate.
Gradients form.
Structures differentiate.
Feedback occurs.
Environmental conditions matter.
The organism emerges through a process.
In that sense, biological inheritance also involves reconstruction.
The inherited organisation does not simply reappear.
It is instantiated anew in each generation.
That SFL distinction is important here.
A potential is instantiated as an instance.
An inherited organisation provides conditions for developmental processes.
The resulting organism is not a copy in the everyday sense.
It is a new instance of an inherited organisation under particular conditions.
Inheritance and instantiation
This allows us to make a careful connection with our earlier use of SFL.
We should not say that genes "realise" organisms.
In SFL, realise has a specific meaning concerning relationships between levels of language and context—for example, context is realised through language, which is realised through phonology and related systems.
Here we are talking about something different.
A biological potential can be instantiated in a particular developmental event or organism.
That distinction matters.
The gene is not a higher-level semiotic system that is realised by the organism.
Nor is the organism simply a phonological realisation of a genetic meaning.
The analogy would be misleading.
What we can say is more modest:
an inherited potential can be instantiated differently under different conditions.
That is the conceptual relation we need.
The organism as a new instance
Every organism is therefore both continuous with its lineage and irreducibly particular.
It inherits.
It develops.
It encounters circumstances.
It acts.
It changes.
It reproduces.
The descendant is recognisably of the same lineage.
But it is not numerically identical with its ancestor.
This is why inheritance and novelty are not opposites.
The descendant can be new because it inherits.
The inherited organisation provides the continuity within which difference can matter.
Without inheritance, difference would have no accumulated history.
Without difference, inheritance would have nothing to transform.
Inheriting a way of becoming
We can now return to the title.
What does an organism inherit?
Not simply a body.
Not simply a sequence.
Not simply a collection of traits.
It inherits a developmental way of becoming an organism.
The inherited organisation constrains how development can unfold.
It makes some forms accessible.
It makes others inaccessible.
It interacts with environmental conditions.
It permits some variations to be produced.
It makes some responses possible.
In this sense, inheritance passes forward not merely an outcome but a structured potential for becoming.
That is a very different image from the gene as blueprint.
A blueprint specifies a finished object.
A developmental organisation provides conditions under which an organism can emerge.
The grammar of "programming"
This is another place where metaphor can mislead.
We say:
"Genes program development."
The expression is convenient.
But what does "program" suggest?
It suggests an instruction sequence whose execution produces a predetermined result.
That is not a good description of biological development.
The genome participates in development.
It does not contain the organism as a completed specification.
The developmental process is interactive, recursive and context-sensitive.
The inherited organisation provides constraints and capacities.
The organism emerges through the interaction of those conditions.
Again, the problem is not the noun program itself.
It is what the metaphor encourages us to imagine.
Conceptual Naturalism asks a simple question
This is precisely where our method of Conceptual Naturalism becomes useful.
When a metaphor is attractive, we should ask:
What does it allow us to see?
and then:
What does it conceal?
"Program" makes inherited specificity visible.
It reminds us that heredity matters.
But it conceals development.
"Blueprint" makes structural correspondence visible.
But it conceals interaction.
"Copy" makes continuity visible.
But it conceals reconstruction.
"Replicator" makes differential persistence visible.
But it can conceal the processes in which the gene participates.
No metaphor is simply true or false.
The question is what kind of construal it affords.
The inheritance of constraints
Perhaps the most important inherited property is therefore not a particular trait but a constraint structure.
A lineage inherits developmental possibilities.
It inherits limitations.
It inherits dependencies.
It inherits sensitivities.
It inherits ways of regulating internal conditions.
It inherits relationships with other organisms.
Over time, evolutionary change can modify those constraints.
A constraint that once closed a possibility can be altered.
A developmental pathway can be opened.
A dependency can become a resource.
A relationship can become a new source of variation.
Evolution therefore changes the structure of the possible.
This is the point at which our central thesis becomes more concrete.
The future is inherited as possibility
We can now formulate the relationship between inheritance and possibility more precisely.
The past does not determine the future by containing it.
It constrains the future by shaping the conditions under which possibilities can arise.
An inherited developmental organisation gives the future a topology.
Some paths are near.
Some are remote.
Some are effectively closed.
Some become accessible under particular environmental conditions.
Evolution modifies that topology.
A new structure can open possibilities.
A lost structure can close them.
A new ecological relation can make an existing capacity consequential.
A changed developmental system can make a previously inaccessible variation producible.
The future is therefore neither predetermined nor unconstrained.
It is historically structured.
The ecology of inheritance
At this point, genetic and ecological inheritance begin to converge conceptually.
A gene persists across generations.
So can an altered environment.
A developmental organisation persists.
So can a social practice.
A symbiotic relation persists.
So can an ecological niche.
These mechanisms are not identical.
We must not collapse them into one universal process.
But they share a structural property:
the present carries forward conditions produced by the past.
This is perhaps the most general meaning of inheritance we have found.
Inheritance is not necessarily the transmission of information.
It can be the persistence of conditions.
And conditions can themselves be transformed.
Inheriting the world made by others
Consider again the beaver.
A descendant inherits genes.
It also enters a landscape shaped by previous beavers.
The dam is not genetically encoded.
Yet it can affect the conditions under which the descendant lives.
The descendant therefore inherits a world.
Not in the sense that the world is handed to it as property.
In the sense that historical activity has altered the conditions it encounters.
This is ecological inheritance.
And it reveals something important about the phrase "environmental selection".
The environment encountered by an organism can itself be an historical product of organisms.
The organism is therefore inheriting into a world that is partly biologically authored—although we should immediately add that "authored" is metaphorical and potentially misleading.
No one designed the world.
It was accumulated through participation.
Inheriting relationships
There is an even subtler possibility.
Sometimes what persists is not a structure but a relation.
A host and symbiont can persist together.
A pollinator and plant can become ecologically interdependent.
A predator and prey can become locked into reciprocal evolutionary dynamics.
A social animal can inherit a behavioural environment constituted by conspecifics.
In such cases, what persists is partly a pattern of participation.
This is why our title says "a way of becoming" rather than "a thing".
The continuity of a lineage may depend upon relationships that are repeatedly reconstructed.
Evolution can therefore preserve and transform patterns of participation.
The inheritance of participation is not a meme
At this point, our original subject—memetics—returns quietly.
It would be tempting to say:
"So these inherited practices are really memes."
But that would miss the point.
The meme was proposed as a theoretical unit of cultural evolution.
We have instead been asking a broader conceptual question.
What kinds of things can persist across generations?
How does continuity work?
What transforms continuity into novelty?
The answer need not be a universal replicator.
Indeed, our investigation suggests that there may be no single mechanism of inheritance across all evolutionary systems.
There may be several.
The interesting commonality lies not in the existence of one universal unit but in the relationship between persistence and transformation.
From replicators to reproductive organisation
This suggests that the replicator may be better understood as one special case of a broader phenomenon.
A replicator is a structure capable of contributing to its own persistence through reproduction.
But biological evolution involves more than replication.
It involves development.
Ecology.
Behaviour.
Interaction.
Construction.
Reproduction.
Selection.
Inheritance.
Perhaps the more fundamental concept is therefore not the replicator but reproductive organisation.
A system persists because some organisation is reproduced across generations.
That organisation can be genetically encoded in part.
But its actualisation depends upon a larger system.
This is not an argument against genes.
It is an argument for locating genes within the organisation that makes inheritance biologically effective.
The lineage remembers through reconstruction
A lineage therefore does something remarkable.
It preserves a pattern without reproducing it exactly.
Each generation reconstructs the organisation.
Each reconstruction occurs under new conditions.
Each can differ.
But enough continuity remains for history to accumulate.
This is a form of memory.
Not memory as a representation stored somewhere.
Memory as structured persistence.
The lineage remembers because its past constrains its future.
And it can change because that constraint is never absolute.
This is perhaps the most naturalistic form of memory we have encountered.
Biological evolution as historical learning?
We should be careful here.
It would be tempting to say that evolution "learns".
But the metaphor is dangerous.
Evolution does not possess a mind.
There is no central learner.
Yet there is a structural analogy worth retaining.
A population can acquire historical organisation through differential reproduction.
That organisation can make subsequent survival and reproduction more likely under recurring conditions.
In that limited sense, evolutionary history can behave as though it has accumulated information about the conditions under which the lineage persists.
But we should not mistake the metaphor for a mechanism.
The information is not necessarily represented.
It is embodied in organisation.
The past as a constraint on the future
This gives us a formulation that avoids both teleology and determinism:
Evolution does not know the future; it carries the consequences of the past into conditions in which the future becomes possible.
That may be the best way to describe historical inheritance.
The lineage has no plan.
But it has history.
The history does not prescribe the future.
But it structures the possible futures.
And every actualised future becomes part of the history that structures what comes next.
This is the recursive architecture of evolution.
Actuality becomes inheritance
We can now complete the cycle.
A possibility becomes actual.
The actual organism participates in the world.
Its activity alters conditions.
Some consequences persist.
Those persistent consequences become part of the conditions inherited by the future.
So:
possibility → actuality → altered conditions → inherited possibility
This is a profoundly different picture from a one-way transmission model.
Evolution is not simply:
inheritance → variation → selection.
It is a recursive process in which actuality can become part of the inherited conditions of future possibility.
The future is therefore shaped not only by what was inherited but by what previous organisms made actual.
The transformation of possibility becomes cumulative
This is why evolutionary history can generate increasing complexity without requiring a direction towards complexity.
A new actualisation can alter the conditions of subsequent possibility.
Some new possibilities become available.
Others disappear.
Further variation occurs within the transformed conditions.
The process can therefore accumulate historical structure.
No foresight is needed.
No goal is required.
All that is required is:
persistence,
interaction,
variation,
differential consequence,
and the capacity of actual conditions to affect subsequent possibilities.
This is enough for history to become cumulative.
The organism inherits a possibility space—but not a map
Perhaps we can now rehabilitate the phrase "possibility space", but only with caution.
An organism does not inherit a map of every possible future.
It inherits a developmental and ecological organisation that makes some futures more accessible than others.
The space is therefore not fixed.
It changes as evolution changes the organisation.
And it is not merely abstract.
It is embodied in actual developmental and ecological relations.
So the organism inherits something like a structured field of possibility.
It does not inherit the future.
It inherits conditions under which futures can emerge.
The furrow again
The furrow now becomes almost unavoidable.
The past leaves a furrow.
But the furrow is not a railway track.
It constrains without determining.
It channels without commanding.
It makes some paths easier than others.
Yet the plough can deepen it, leave it, branch from it or create another.
And the field itself changes as the plough moves.
Inheritance is like the furrow.
It is the persistence of historical organisation.
But becoming is the movement through and transformation of that organisation.
The future therefore inherits the furrow without being condemned to repeat it.
Perhaps that is the most concise formulation of our entire argument:
Evolution inherits the furrow and transforms the field.
What, then, is inherited?
We can now answer our opening question.
What is inherited?
Sometimes genes.
Sometimes developmental organisation.
Sometimes ecological structures.
Sometimes behaviours.
Sometimes relationships.
Sometimes environmental conditions.
But the deepest commonality is not a particular substance or unit.
It is historically persistent organisation capable of contributing to subsequent becoming.
That organisation is never simply reproduced.
It is instantiated anew.
It enters new relations.
It encounters new conditions.
It can be modified.
It can open new possibilities.
And those new possibilities can become the conditions for further inheritance.
Inheriting a way of becoming
We can finally give the title its full meaning.
A lineage does not inherit a finished identity.
It inherits a way of producing and maintaining identity.
It inherits a developmental organisation.
It inherits constraints and capacities.
It inherits ecological relations.
It inherits possibilities shaped by history.
It inherits, in short, a way of becoming.
That way of becoming is not a destiny.
It is not a programme.
It is not a blueprint.
It is a structured potential for producing organisms under conditions.
And because conditions change, the inherited way of becoming can become something new.
From inheritance to evolution
We can therefore see why inheritance alone cannot explain evolution.
Inheritance explains continuity.
Evolution requires continuity plus transformation.
But transformation is not simply random deviation from inheritance.
It occurs within the organisation inherited from the past.
Development constrains variation.
Ecology makes variation consequential.
Organisms modify ecological conditions.
Those conditions alter subsequent evolutionary trajectories.
Inheritance carries the consequences forward.
The process repeats.
So evolution is not the opposite of inheritance.
It is what happens when inheritance participates in becoming.
The question beneath the question
This takes us back to the question with which Dawkins began.
How can complex biological organisation arise and persist without a designer?
We can now answer in a rather different vocabulary.
Because biological systems inherit organisation.
Because inherited organisation constrains development without determining it.
Because development produces organisms capable of participating in environments.
Because those participations alter ecological conditions.
Because differential reproduction preserves some consequences.
Because what is preserved becomes part of the conditions for future variation.
No designer is required.
But neither is a simple copying machine sufficient.
What we need is a historically organised system capable of carrying possibility forward while transforming it.
That is evolution.
The strange continuity of life
There is something almost paradoxical about this.
Life persists by not remaining the same.
A lineage survives because its organisation can be reconstructed.
But reconstruction always occurs under new circumstances.
The continuity is therefore dynamic.
The identity is historical.
The inheritance is transformative.
The future is constrained by the past without being contained in it.
This is perhaps why biological evolution is such a profound challenge to simple metaphysical categories.
It is neither pure persistence nor pure novelty.
It is novelty through persistence.
And perhaps this is what "becoming" means
We began this series wondering whether "the transformation of possibility" was merely a beautiful phrase borrowed from our existing conceptual vocabulary.
It now seems to have acquired a more disciplined meaning.
Possibility is structured by inherited organisation.
Actuality occurs through development and participation.
Actuality changes ecological and developmental conditions.
Those changed conditions become part of what future organisms inherit.
And so the possible is continually reconfigured by what becomes actual.
This is not a metaphysical principle imposed upon biology.
It is a way of describing the recursive historical structure we have uncovered.
The possibility of the future is transformed by the actuality of the past.
One final caution
We should nevertheless resist the temptation to turn this into a grand theory of everything.
Not every biological process is best described in terms of possibility.
Not every inherited feature is a "way of becoming".
Not every ecological relation transforms evolutionary possibility in a significant way.
And not every useful metaphor should become a theoretical entity.
Our conceptual tools remain tools.
SFL helps us notice how grammatical choices construe processes.
Conceptual Naturalism helps us ask what metaphors reveal and conceal.
Relational ontology helps us avoid treating participants as isolated substances.
The distinction between potential, possibility and actuality helps us describe different moments in the emergence of biological form.
None of these replaces evolutionary biology.
They help us see its conceptual architecture.
That distinction remains essential.
What we can now see
At the beginning, inheritance looked like the preservation of a thing.
Now it looks more like the preservation of conditions for becoming.
At the beginning, variation looked like departure from a copied form.
Now it looks like transformation within inherited organisation.
At the beginning, the organism looked like the product of inheritance.
Now it looks like a participant that can alter the conditions of subsequent inheritance.
At the beginning, the environment looked like the background against which selection occurred.
Now it looks like a historical structure partly produced by the organisms that inhabit it.
And at the beginning, possibility seemed like a space into which evolution moved.
Now possibility looks more like something that is continually reconfigured by the interaction of inheritance, development and participation.
That is a considerable change in perspective.
But one question remains.
If inheritance is not simply the transmission of a thing, and if what persists can be a developmental organisation, an ecological condition, a behaviour or a relationship, then perhaps the deepest evolutionary unit is not an entity at all.
Perhaps it is a pattern that can be re-enacted.
And that takes us remarkably close to where our journey began—with memes, symbols and cultural inheritance.
But now the direction of inquiry has reversed.
We no longer need biology to explain culture.
We can ask whether our investigation of culture has taught us something about biology:
Perhaps evolution is not fundamentally about copies that survive.
Perhaps it is about patterns of organisation that can be inherited, re-enacted, transformed—and thereby make new forms of life possible.
And once we put it that way, one final question becomes unavoidable:
What happens when evolution itself changes the kinds of organisation capable of inheriting, participating and becoming?
That is where the next essay must take us. 🍷🙂
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