If we remove the imaginary agency from evolutionary explanation, the gene becomes a rather different kind of thing.
It is no longer the little strategist of the organism.
It does not want to replicate.
It does not compete in the ordinary sense.
It does not formulate plans.
It does not know what the organism is doing.
And yet genes plainly matter.
They are involved in inheritance. They participate in the production of organisms. Differences in genetic sequences can have consequences for development, phenotype and reproductive success. Their transmission across generations is central to evolutionary biology.
So removing agency does not make the gene disappear.
It leaves us with a more interesting question:
What kind of participant is a gene?
This question takes us beyond the problem of anthropomorphic language and toward something more fundamental.
Perhaps the difficulty was never that we attributed too much agency to genes.
Perhaps it was that we were tempted to attribute too much independence to them.
The extraordinary success of the gene
There is a reason the gene became such a powerful concept in twentieth-century evolutionary biology.
Genes provide a way of connecting heredity with evolutionary change.
A population contains heritable variation.
Some variants are transmitted more successfully than others.
Over generations, those differences can accumulate.
The gene therefore offers a powerful handle on the continuity between generations.
It is tempting, however, to turn an explanatory handle into an explanatory foundation.
Once the gene becomes the central object of evolutionary explanation, we can begin to imagine that the gene is the thing that fundamentally evolves, while organisms and environments become secondary contexts in which genes happen to operate.
The metaphor of the selfish gene reinforces this impression.
If the gene is the "selfish" entity, then the organism can begin to look like a temporary vehicle.
If the gene is the replicator, the organism becomes the machinery through which replication occurs.
If the gene is the fundamental unit of selection, other biological levels can appear derivative.
There is something illuminating in this perspective.
But there is also a danger.
The gene is being detached conceptually from the very relations that make its biological significance intelligible.
A gene does not exist by itself
Imagine, for a moment, a gene completely isolated from everything else.
No cell.
No genome.
No developmental machinery.
No organism.
No population.
No environment.
No reproduction.
No historical lineage.
Would it still be an evolutionary gene?
The question is revealing because the answer is not simply "yes".
We can certainly identify a sequence of nucleotides in isolation.
But its significance as a gene is not exhausted by its molecular sequence.
A gene participates in a biological system.
Its effects depend upon cellular machinery.
Its expression depends upon regulatory conditions.
Its consequences depend upon developmental processes.
Its phenotypic effects depend upon interactions with other genes and with the organism's environment.
Its evolutionary significance depends upon inheritance and differential reproduction in populations.
The gene is therefore not an autonomous little agent whose properties explain everything else.
It is a participant in a network of processes.
This does not diminish the gene.
It tells us why the gene is biologically interesting in the first place.
The gene and the genome
Even before we leave molecular biology, the solitary gene begins to dissolve.
A gene is situated within a genome.
Its effects can depend upon other sequences.
Regulatory regions matter.
Gene expression is coordinated.
Products interact.
Networks of regulation can alter what a particular sequence does in a particular context.
So even at the molecular level, the question:
"What does this gene do?"
can be incomplete.
We may need to ask:
What does this gene do in this system?
That final phrase changes the character of the question.
A gene does not possess a single context-free repertoire of effects waiting to be expressed.
Its consequences emerge through interactions.
The same sequence can have different consequences in different genetic, developmental or environmental contexts.
This is one reason why a purely atomistic picture of biological inheritance can become misleading.
The gene is real.
But its causal significance is relational.
From genotype to organism
The next relation is even more important.
Genes do not simply become organisms by themselves.
Between genotype and phenotype lies development.
There are processes of transcription, translation, regulation, signalling, differentiation, growth and interaction.
The organism emerges through an organised history.
This matters because it interrupts a tempting picture:
gene → trait
as though the gene were a little instruction and the trait were its inevitable output.
Sometimes that shorthand is useful.
But it conceals the developmental machinery through which genetic differences acquire phenotypic consequences.
The gene participates in development.
It does not stand outside development and issue commands to it.
And once we see this, the language of the "gene for" something begins to require care.
A gene associated with a trait is not necessarily a miniature blueprint for that trait.
Its consequences arise within an organised developmental system.
The problem with the blueprint
The blueprint metaphor is extraordinarily seductive.
A blueprint contains a representation of a structure.
The builder follows the blueprint.
The finished building corresponds to the plan.
If the genome is treated as a blueprint, the organism appears to be the construction project and development the execution process.
But biological development is not construction in this sense.
The genome does not contain a complete representation of the adult organism waiting to be read off.
Development is dynamic.
It is interactive.
It involves feedback.
Cells respond to signals.
Structures influence subsequent development.
The organism's environment matters.
And the developing organism changes the conditions under which its own subsequent development occurs.
The blueprint metaphor therefore gets something right—it draws attention to the importance of inherited organisation—but it can obscure something more important:
inheritance does not transmit a finished organism; it transmits conditions within which an organism can develop.
That distinction will become central to our next essay.
For now, it already tells us something about the gene.
The gene does not contain the future organism in miniature.
It participates in a process through which a future organism can arise.
Actor, Medium, participant
Our SFL distinction becomes particularly useful here.
Consider:
The gene produces the protein.
The grammar makes the gene an Actor.
But we can ask what kind of process this representation is construing.
The gene is not an intentional producer in the ordinary sense.
Nor is it necessarily useful to imagine the gene as the sole causal source of the protein.
The molecular process involves transcriptional and translational machinery, cellular conditions and regulatory interactions.
The gene participates.
It is one component of an organised process.
This is where the ergative perspective can be illuminating.
The same event can be construed not simply by asking "who did it?" but by asking how the process unfolds and what participant functions as Medium through which it occurs.
That shift is conceptually important.
Instead of asking:
Which thing caused the outcome?
we can sometimes ask:
Through what organisation of relations did this outcome occur?
That does not eliminate causes.
It changes the explanatory picture from isolated agents to participating processes.
The gene in the organism
There is an even larger relation.
A gene is not simply situated inside an organism as a passenger.
Its effects contribute to the organisation of the organism.
But the relationship runs in both conceptual directions.
The organism provides the developmental and physiological context within which genes have effects.
The organism also reproduces.
The organism interacts with its environment.
The organism behaves.
And those organism-level processes affect the reproductive consequences associated with genetic variants.
So we get something like:
gene ↔ organism
rather than simply:
gene → organism.
This arrow is not meant to suggest that causation is literally symmetrical in every case.
It is a warning against treating one direction of explanation as the whole story.
Genes contribute to organisms.
Organisms provide the conditions under which genetic differences have evolutionary consequences.
The relation is therefore not one of simple command.
The organism in the population
Now widen the frame again.
A gene's evolutionary significance depends upon what happens to its variants across generations.
That means population structure matters.
A variant can have one consequence in one population and another in a different genetic or ecological context.
Its frequency can change.
Its effects can depend upon other variants.
Its persistence can depend upon reproductive patterns.
The evolutionary gene is therefore inseparable from population history.
The phrase "the gene succeeds" can be useful shorthand.
But what actually happens?
A lineage containing a particular hereditary variant leaves descendants at a different rate from relevant alternatives under particular conditions.
The frequency of that variant changes.
That is a population process.
The gene is involved.
But it is not alone.
And then there is the environment
The relational picture becomes impossible to ignore once we introduce ecology.
Suppose a genetic variant affects a trait.
Whether that trait matters depends upon the environment.
A thick coat is not intrinsically advantageous.
It depends upon conditions.
A particular colouration is not intrinsically protective.
It depends upon what surrounds the organism.
A metabolic capacity does not have a fixed reproductive value independent of circumstances.
The environment changes the consequences of phenotypic differences.
So:
gene → trait → fitness
is already too simple.
We need something more like:
gene ↔ development ↔ organism ↔ environment ↔ reproduction ↔ population
Again, this is not a claim that all these relations are identical or interchangeable.
It is a reminder that evolutionary significance emerges across them.
Relational does not mean everything causes everything
At this point, we need a safeguard.
It would be easy to respond to reductionism by embracing an equally unhelpful holism.
If genes are relational, perhaps everything affects everything.
If organisms depend on environments, perhaps genes no longer matter.
If development matters, perhaps selection is irrelevant.
If ecology matters, perhaps molecular biology is merely one perspective among many.
That would be a mistake.
Relational thinking does not mean that all causes are equal.
It means that the causal significance of a participant depends upon the system of relations in which it participates.
A gene can have a highly specific molecular effect.
A developmental pathway can impose a strong constraint.
An environmental change can alter selection pressures.
A population bottleneck can alter genetic frequencies.
Different relations matter at different scales.
The point is not to flatten those differences.
It is to stop treating one level as though it were the whole biological story.
The gene is not a fiction
There is another danger.
Once we emphasise relations, someone might object:
"So the gene isn't really real?"
But that is not where the argument leads.
Relational ontology does not require us to deny entities.
It asks us to understand entities through their relations.
A gene is real.
An organism is real.
A population is real.
An environment is real.
The question is what kind of reality each possesses and how each acquires explanatory significance.
A gene is not less real because it participates in a genome.
A human being is not less real because they participate in a society.
A word is not less real because its meaning depends upon a language.
Indeed, in many cases, relations are precisely what make entities intelligible.
The error would be to imagine that being relational means being unreal.
The gene and the metaphor of independence
This brings us back to language.
There is a subtle conceptual pattern in expressions such as:
the gene acts;
the gene replicates;
the gene competes;
the gene survives;
the gene is selected.
Each formulation makes the gene grammatically available as a participant in an action.
Again, none is necessarily false.
But taken together, they can construct a remarkably independent image of the gene.
The gene becomes the thing that persists while everything else becomes context.
That is the conceptual move we need to examine.
The gene is not simply a sequence that happens to be located somewhere.
It is a hereditary participant whose significance arises through a biological organisation.
The very idea of a "gene" presupposes a system capable of inheritance, expression and reproduction.
The gene therefore cannot be understood as the smallest possible independent piece of evolutionary reality merely because it is a useful unit of analysis.
A unit of explanation is not necessarily an ontological atom.
What survives?
This becomes especially interesting when we ask what it means to say that a gene "survives".
The phrase sounds straightforward.
But a gene does not survive in the way an organism survives.
An organism survives by continuing to function as an organised living system.
A genetic sequence persists when copies—or descendants carrying corresponding hereditary information—continue through successive generations.
The forms of persistence are different.
And that difference matters.
When Dawkins called genes "replicators", he was trying to capture a genuine feature of heredity: some structures have a special capacity for being copied with sufficient fidelity for cumulative selection to occur.
But once again, a useful explanatory distinction can become an ontology.
The replicator begins to look like the thing that evolution is fundamentally "about".
We should resist that conclusion.
The gene's capacity for hereditary persistence is real.
But the conditions under which that persistence occurs are distributed across a much larger biological organisation.
From replicator to participant
This gives us a subtle reformulation.
Perhaps we should not ask:
Is the gene the agent of evolution?
It isn't.
Nor:
Is the gene the only unit that matters?
It isn't.
Nor even:
Is the gene the fundamental object of evolution?
That question may be too blunt.
A better question is:
What does the gene participate in that makes hereditary difference evolutionarily consequential?
Now the answer begins to unfold.
It participates in:
- inheritance;
- development;
- organismal organisation;
- reproduction;
- population dynamics;
- ecological relations;
- historical lineages.
And the significance of any particular genetic difference depends upon how these processes are organised.
The gene therefore remains central without becoming sovereign.
A different picture of evolutionary causation
We can now replace one image with another.
The first image is hierarchical:
gene → organism → environment
The gene appears at the bottom of a causal command chain.
The second is relational:
gene ↔ genome ↔ development ↔ organism ↔ environment ↔ population
This second diagram should not be read as a claim that all arrows are equally strong, reversible or simultaneous.
It is a conceptual map.
It reminds us that evolutionary causation is distributed across interacting processes.
The gene contributes to development.
Development determines how genetic differences become phenotypic differences.
Phenotypes participate in organism–environment relations.
Those relations affect reproduction.
Reproduction alters the representation of genetic variants in populations.
Population history changes the context in which future genetic variants will have effects.
And now we can see something that was difficult to see when the gene was imagined as an autonomous replicator.
The consequences of a genetic difference can reach forward into the future while also changing the conditions under which future differences matter.
That is where inheritance begins to acquire a richer temporal structure.
The past does not simply copy itself
We began this essay by asking what kind of participant a gene is.
We can now offer a provisional answer.
A gene is not an agent.
It is not an independent evolutionary strategist.
It is not a miniature blueprint containing the organism's future.
It is a hereditary component of a biological system whose consequences arise through relations among molecular, developmental, organismal, ecological and population processes.
And that immediately changes how we should think about inheritance.
If a gene is inherited, what exactly is being carried from the past into the future?
A sequence?
Yes.
But the sequence does not arrive in an empty universe.
It arrives in a cell.
Within a genome.
Within a developmental system.
Within an organism.
Within an environment.
Within a lineage.
Inheritance therefore cannot be adequately understood as simple transmission of an isolated object.
What is transmitted becomes part of the conditions from which subsequent biological organisation can arise.
That is a much more interesting conception of heredity.
The past does not merely reproduce itself.
It enters into the conditions of the future.
And possibility begins to appear
This is our first genuine encounter with the central theme of the series.
A genetic change can produce a new actuality.
But its evolutionary significance may extend beyond that actuality.
It may alter developmental trajectories.
It may alter ecological interactions.
It may alter reproductive consequences.
It may alter what variations are subsequently useful.
It may alter what kinds of structures can be built upon it.
In other words:
what becomes actual can alter what subsequently becomes possible.
We should not yet elevate this into a grand principle.
But we can see why it matters.
Evolutionary history is not merely a record of things that happened.
It is a history in which what happened changes the conditions under which something else can happen.
And that is beginning to look rather different from simple replication.
The next question
We began with the gene as apparent agent.
We have ended with the gene as participant.
That is already a substantial conceptual shift.
But it leaves us with an even more difficult problem.
If inheritance is not simply the copying of an isolated entity, then what exactly does inheritance carry forward?
What is the relationship between preservation and transformation?
For the organism of today inherits structures from the organism of yesterday—but it does not simply become yesterday's organism again.
Something has been conserved.
Something has changed.
And the conserved organisation does not merely preserve the past.
It helps determine the range of forms through which the future can unfold.
The gene therefore leads us somewhere unexpected.
Not toward a more sophisticated gene-centred theory.
But toward a question about time.
How does the biological past remain active in the present?
And how does what is inherited become part of the conditions from which an evolutionary future can emerge?
That is where we go next. 🍷🙂
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