Preface: Why Bees?
If we want to understand how complex forms of collective life emerge and endure, it makes sense to begin not with language or culture, but with systems in which coordination happens without them. One such system is the eusocial insect colony. Here, across thousands or millions of individuals, we find a remarkable form of distributed organisation: structured division of labour, elaborate reproductive and defensive systems, and real-time adaptation to changing environmental conditions.
Among these insects, bees offer a particularly compelling case. A forager bee departs the hive not as a representative of a collective, but as an individual organism — one guided by its own history, physiology, and sensory-motor capacities. And yet, what it does in the world, and how it returns, contributes directly to the stability and ongoing function of the hive. Perception, navigation, memory, and behaviour are all involved — and they are all grounded in the ongoing regulation of what matters.
In this series, we’ll trace a path from the value-guided behaviour of individual foragers to the large-scale coordination of activity within the colony. Along the way, we’ll distinguish between types of behaviour — those that are continually updated in response to local conditions, and those that are routinised over time. We’ll look at how these patterns emerge, stabilise, and participate in larger dynamics of social organisation.
Our concern here is with systems of regulation, differentiation, and coordination. Bees, in this sense, are not just fascinating creatures — they are living systems in motion, through which we can begin to explore how the local acts of individuals become integrated into a broader, self-regulating whole.
1 The Forager as Value System in Motion
We begin with a single bee in flight. She has left the hive not with instructions or orders, but with a readiness to act — a system of sensitivities and tendencies shaped by her physiology, her developmental history, and the broader conditions of the colony. She is not searching for 'information' in the abstract, but for what matters: nectar, pollen, water, resin, or a suitable site. Her movements are guided by value — by the differentiation of what is salient, what is to be approached, avoided, or ignored.
This is what Gerald Edelman called perceptual categorisation: a continual coordination of sensation and action, governed by the organism’s internal structure and its history of interactions with the environment. In this view, perception is not the passive reception of sensory data, but an active, selective process — a coupling of what is sensed with what can be done. A flower is not simply seen; it is foraged.
At every moment, the forager is engaged in a dynamic regulation of her own viability. Her sensory systems are tuned not to general truths but to immediate relevance: the colour contrast of a petal, the humidity of a potential water source, the direction and strength of the wind. These are not interpreted, but acted upon — within a tightly coupled system where perception and behaviour are entangled.
And yet, her activity is not isolated. Though she flies alone, her behaviour contributes to a larger system — the needs of the hive, the rhythms of the season, the distribution of resources in the environment. In tracing her path, we begin to see how the value-based activity of a single organism can serve as an anchor point for more complex patterns of coordination.
Before we reach the hive, we stay with the forager: a living system in motion, engaging the world not as a blank slate, but as a differentiated field of affordances — a world made actionable through the ongoing regulation of what matters.
2 From Perception to Behavioural Routine
A single foraging flight is a contingent act, guided by the conditions of the moment. But over time, repetition carves a path — not just in the environment, but in the organism. What was once novel becomes familiar; what was uncertain becomes anticipated. Patterns of value-guided action, once fluid, may settle into more stable forms.
This is not memory in the human sense, nor habit as mere repetition. It is a process by which experience reshapes responsiveness. A bee that has successfully located a nectar source may return not by calculation, but by the activation of a coordinated behavioural routine — a sequence of actions no longer dependent on moment-to-moment recalibration.
Behaviour, in this sense, becomes routinised when the conditions that support it are stable enough that continual updating is no longer necessary. The transition from fluid categorisation to stable response is not a shift from intelligence to automation, but a shift in the kind of constraint at play: from flexible regulation to efficient entrainment.
Such routinisation plays a crucial role in colonial life. It enables the colony to stabilise its internal operations — food collection, brood care, ventilation — without requiring every individual to evaluate each situation anew. Not all behaviour can be routinised, but where it can, efficiency and coordination improve.
We might think of this as a gradient: at one end, fully contingent behaviour shaped by immediate value distinctions; at the other, routinised sequences carried out with minimal need for updating. Between these poles lie a range of adaptive strategies — and it is this range that allows a colony to operate as a layered system of distributed responsiveness.
In tracing how perception can stabilise into patterned action, we begin to see how value-guided activity does not remain at the level of the individual. It becomes, through routine, part of a larger structure of coordination — one in which the past reshapes the present without the need for representation.
3 The Hive as a Distributed Value System
The individual forager, though autonomous in action, is never acting in isolation. Her value-guided behaviour takes place within a larger field of relations — the colony — in which the activities of thousands of individuals are likewise constrained, differentiated, and coordinated. What emerges is not a single mind, but a distributed system of regulation: one that maintains its coherence without recourse to central control.
Pheromones play a central role here. They do not convey meaning in the way language does, but they modulate the behavioural tendencies of others. A queen mandibular pheromone, for example, inhibits ovarian development in workers; alarm pheromones trigger defensive mobilisation; trail pheromones support the alignment of foraging activity. Each acts not by informing, but by shifting what matters — adjusting the value gradients that shape perception and action.
Within this system, division of labour arises not through fixed assignments, but through the modulation of responsiveness. Age, experience, nutritional status, and colony needs all interact to shape who does what, when. These dynamics are flexible, but not arbitrary. They are stabilised through feedback: when certain behaviours succeed, their likelihood increases; when needs are met, others recede.
The hive thus functions as a self-organising system of differentiated roles and responsivities. Its internal structure is not imposed from above, but emerges from the coupling of individual value systems through environmental and chemical mediation. It is this coupling — constantly shifting, but highly constrained — that enables the colony to operate as a coordinated whole.
What we see, then, is a distributed form of homeostasis: not the maintenance of a fixed state, but the ongoing regulation of viability across multiple scales. This regulation does not rely on deliberation or symbolic mediation. It is enacted through the structuring of sensitivities — a choreography of value systems tuned to one another through shared participation in a dynamic environment.
4 Routinised vs. Updated Behaviours
Not all behaviours in the hive are alike. Some must remain flexible, constantly tuned to momentary variation — while others can be repeated with minimal adjustment, stabilised across time and individuals. Understanding this distinction is key to grasping how the colony operates as a layered, adaptive system.
Updated behaviours are those shaped in real time by perceptual input and internal state. A forager adjusting her flight in crosswinds, or shifting her search pattern in response to floral density, engages in behaviour that must remain sensitive to ongoing conditions. These acts are situated, contingent, and dynamically regulated.
Routinised behaviours, by contrast, unfold within more stable parameters. Brood care, food exchange, and many aspects of nest maintenance follow well-established sequences. These are not fixed in the sense of mechanical reflex, but they are constrained in form and deployment. They do not require moment-to-moment recalibration.
Crucially, both kinds of behaviour are value-guided — but the structure of that guidance differs. In updated behaviour, value distinctions are re-evaluated continuously. In routinised behaviour, the relevant distinctions have been settled in advance, entrained through past success or genetic predisposition.
This division of behavioural labour allows the colony to manage complexity without overburdening any one part. Continuous responsiveness is preserved where necessary, but routinisation conserves energy and ensures reliability where variability is low or risk is high. The two forms are not opposed, but complementary.
Indeed, the boundaries between them can shift. What begins as updated behaviour — a novel foraging technique, a shift in thermoregulation — may, through reinforcement and stability, become a routinised pattern. Conversely, routinised behaviours may dissolve under pressure, requiring renewed responsiveness.
This interplay is not merely an optimisation strategy; it is a foundation for adaptability. The colony survives not by rigid control, but by maintaining the right mix of behavioural stability and flexibility — ensuring that what matters is acted on appropriately, whether through fresh adjustment or well-tuned repetition.
Conclusion: Social Life Before Meaning
In tracing the life of the bee colony, we have seen how value-guided behaviour in individual organisms becomes integrated into a larger system of coordination. From the forager’s continual adjustment to local conditions, to the stabilisation of behaviour into routinised routines, to the distributed regulation mediated by pheromones, the colony operates through layers of differentiated responsiveness.
This system is not controlled by a central authority or mediated by symbolic communication. Instead, it emerges through the dynamic coupling of individual value systems, each constrained and shaped by environmental and social feedback. Through this ongoing choreography, the hive maintains its viability and adaptability.
By focusing on these foundational processes—perception as action guided by what matters, behavioural routines as efficient entrainments, and social organisation as distributed regulation—we have taken a crucial step toward understanding how complex collective life is maintained.
This understanding sets the stage for further exploration. Future inquiry will consider how, from these layers of value and coordination, more structured systems of interaction might arise. But for now, we rest with the remarkable reality of social life before the emergence of meaning in any semiotic sense.
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