Heaven and earth have their great beauties but do not speak of them; the four seasons have their clear-marked regularity but do not discuss it; the ten thousand things have their principles of growth but do not expound them.
天地有大美而不言，四时有明法而不议，万物有成理而不说。
— Chuang Tzu, Knowledge Wandered North
In classical Daoist thought, beauty is neither a human invention nor a symbolic construct imposed upon the world. It is not crafted, declared, or even necessarily reflected upon. Rather, it arises spontaneously from the patterned regularities of nature, which governing the rhythms of seasons, the organization of living forms, and the unfolding of change itself. Although articulated in a metaphysical idiom, Chuang Tzu’s insight poses a question that resonates strongly with contemporary anthropological and evolutionary inquiry: Are aesthetic capacities best understood as a product of cultural construction, or as an expression of evolved perceptual and cognitive adaptations to ecological environments?
For a long time, particularly under the influence of the European Enlightenment, aesthetics has been viewed as a uniquely human achievement—a high-level hallmark of culture involving abstract thought and “disinterested” appreciation. However, when we look back at the deep past and across the vast natural world, we see prehistoric rock art, tribal rituals, ornamental traits in animals, and the elaborate courtship dances of birds. It becomes clear that the “love of beauty” is woven into the very fabric of life’s long evolutionary history. It is not a cultural luxury, but a survival script written into our biology by ancestors who navigated the rigors of nature, mate choice, and social cooperation.
To understand human aesthetic minds, we must first break free from the shackles of “art objects.” Aesthetic experience is not limited to gazing at masterpieces in museums; it is a multisensory system shaped by the coevolution of signals and evaluative mechanisms that engages vision, touch, and sound, and that encompasses complex experience, from the beautiful to the ugly, the comic to the tragic . Art, in this sense, is not defined by particular objects, but as a universal behavioral tendency termed “making special” or “artification” . This behavior involves the deliberate use of operations like formalization, repetition, and exaggeration to transform ordinary actions and materials into something extraordinary, thereby attracting attention and sustaining interest, particularly within ritualistic contexts.
Exploring human aesthetic minds, therefore, is not simply about how we appreciate a painting or a piece of music; it is fundamentally about understanding how the human mind has been shaped over millions of years of hunter-gatherer life. If aesthetic capacity is the product of a coevolution between genes and culture, then to find its origin, we must return to the very source of human creation.
The dawn of art in the early paleolithic
Archaeological evidence from the Lower Paleolithic suggests that the cognitive foundations of human aesthetics and art began to emerge long before the appearance of fully symbolic behavior. In the absence of explicit symbolic artefacts, such early developments are inferred indirectly from patterns in stone tool production.
The earliest stone tools, associated with the Oldowan tradition and dating to ~2.5 million years ago, consist primarily of flakes and cores produced by direct percussion of stone nodules using hammerstones . These artefacts, made from materials such as basalt, flint, and limestone, were used in a variety of practical contexts, including the processing of animal carcasses, nut cracking, plant cutting, bark stripping, digging, honey harvesting, and possibly throwing at prey. However, Oldowan tools lack any imposed overall shape: their final form largely reflects the original geometry of the raw material and the number of flakes removed, rather than an internally represented design.
A major shift occurs around 1.4 Ma with the emergence of Acheulean (named after stone tools found near St Acheul in France, but produced a million year earlier in Africa already) bifacial tools, commonly referred to as handaxes . Unlike Oldowan artefacts, early Acheulean handaxes exhibit deliberately imposed overall form and bilateral symmetry. These forms become increasingly pronounced in the later Acheulean, particularly between ~500–300 ka, when handaxes display high degrees of congruency and consistency in symmetry that exceed what is strictly required for functional tasks such as cutting or butchery. In addition to their carefully controlled form, many handaxes were produced from visually distinctive raw materials, such as stones with smooth, lustrous surfaces or fossil inclusions . These Acheulean handaxes may therefore constitute some of the earliest material evidence for the emergence of human aesthetic preference, suggesting that early hominins were attentive to how artefacts appeared, not merely to what they did.
The extraordinary temporal and geographic stability of Acheulean handaxes over more than 1.5 Ma is difficult to reconcile with cultural transmission models, which typically predict copying error, innovation, and regional divergence . Such long-term consistency suggests that key aspects of handaxe production may have been under partial genetic constraint, comparable to genetically nest building by birds. At the same time, the aesthetic elaboration of handaxes depended on advances in cognitive capacities . Producing consistent bilateral symmetry and controlled overall form required coordination across multiple viewpoints, mental rotation, basic spatial reasoning, and stable representations of tools as structured objects. These shared cognitive prerequisites enabled early hominins to reliably reproduce similar forms across individuals and generations.
Similar to aesthetic preferences observed in other animals, such forms can be understood as arising from sexual selection. Finely made and highly symmetrical handaxes may have functioned both as costly signals in the sense of the handicap principle and as aesthetic displays that exploited evolved perceptual sensitivities to symmetry. Through this dual mechanisms, handaxes acted as honest signals of individual quality, conveying multiple adaptive dimensions, including planning ability, motor control, access to resources, and social competence . Therefore, Acheulean handaxes are best understood not primarily as culturally symbolic artefacts, but as material expressions of evolutionarily stabilized aesthetic predispositions, grounded in both inherited biases and emerging cognitive abilities.
Acheulean handaxes (Mithen, 2003)
Evidence from the Lower Paleolithic is not limited to the aesthetic elaboration of utilitarian tools. At the Homo erectus site of Trinil in Java, a freshwater mussel shell dated to ~500 ka was found bearing a deliberately engraved zigzag-like geometric pattern . These incisions were intentionally produced by hominins, rather than resulting from natural abrasion or taphonomic processes. Although the function or meaning of the engraving cannot currently be determined, the presence of such deliberate geometric incisions, often considered markers of modern cognition and behavior, provides tentative evidence that Homo erectus may already have possessed an emerging capacity for symbolic meaning and rudimentary aesthetic capacity of symbolic art. In this early stage, aesthetic preferences also appear to have been expressed as an interest in visually unusual objects. Archaeological evidence indicates that early hominins, including Homo erectus and Neanderthals, collected and transported items such as crystals, fossils, or stones with distinctive shapes .
The engraving on shells by the Homo erectus at the Trinil site on Java (Joordens et al., 2015)
Taken together, early hominins had already developed sensitivities to symmetric pattern and visual form. While isolated instances of abstract mark-making may have occurred, aesthetic preference at this stage appear to have remained grounded in biologically predispositions, shaped by genetic inheritance rather than cumulative cultural transmission.
The emergence of symbols in the middle paleolithic
The Middle Paleolithic (~300–50 ka) witnessed the emergence of modern humans (Homo sapiens) and major advances in symbolic art. At the same time, other hominin lineages, such as Neanderthals, also exhibited indications of symbolic behavior. During this period, archaeological evidence begins to demonstrate innovations in decoration, self-expression, and ritual practices among humans.
One of the clearest archaeological indicators of this shift is the widespread use of mineral pigments, particularly red ochre (iron oxide rich minerals). Across multiple sites in Africa and the Near East, ochre pieces bearing traces of grinding, scraping, and heat treatment document deliberate pigment production rather than incidental use . At 92,000-year old Qafzeh Cave, red ochre appears to have been deliberately selected and deposited within mortuary contexts, interpreted as culturally structured color symbolism . Evidence from the Levant indicates that during the mid–Middle Paleolithic (~130–80 ka), different Homo groups converged on a shared set of behaviors, including similar lithic technologies, increased reliance on large-game hunting, and socially elaborated practices such as intentional burial and the use of ochre in mortuary contexts . In this context, mineral pigments functioned not merely as aesthetic substances, but as socially meaningful media embedded in practices of decoration, identity signaling, and ritual action, marking an important transitional stage between biologically grounded aesthetic sensitivities and culturally evolved symbolic expression.
Different types of ochre from Tinshemet Cave and their association with human and animal bones (Zaidner et al., 2025)
Personal ornamentation provides the complementary evidence for the emergence of symbolic aesthetics during the Middle Paleolithic. Unlike pigment use, which could be applied in multiple contexts, ornaments constitute durable, portable objects designed to be displayed on the body, making them particularly well suited for social signaling. The earliest known personal ornaments date to this period: at Bizmoune Cave in Morocco, perforated marine shell beads dated to ~150–142 ka exhibit systematic perforation and use-wear, indicating that they were strung and worn as body adornments . At Blombos Cave in South Africa, perforated marine shell beads recovered dated to ~75–77 ka constitute the evidence for the habitual use of personal ornaments . These beads represent standardized and repeatable visual elements, marking a shift from isolated collections of visually unusual objects toward the deliberate production of patterned symbolic forms. Such ornaments are widely interpreted as media of non-verbal communication, conveying information about identity, group affiliation, or social relationships . Their appearance therefore signals not only advances in abstract representation, but also changes in social organization, including expanded social networks and increased interaction beyond immediate kin groups.
Perforated _N. kraussianus_ beads from the Middle Stone Age of Blombos Cave (Henshilwood et al., 2004)
Early Homo sapiens also produced deliberately engraved abstract markings alongside the use of personal ornaments. Evidence from Blombos Cave and Diepkloof Rock Shelter in southern Africa includes engraved ochre pieces with regular geometric. A particularly well-known ochre fragment from Blombos Cave, dated to ~75 ka, features intersecting linear motifs and is widely considered the earliest secure evidence of intentional geometric marking .
Engraved ochres from Blombos Cave (Henshilwood et al., 2016)
Taken together, during the Middle Paleolithic, humans developed a broader range of aesthetic engagement, including the use of color painting, the production of personal ornaments to signal identity, the engraving of abstract marks, which may have expressed emotions and beliefs through ritual and mortuary practices. A generalized tendency to adorn the body and the environment with ordered visual elements and patterns appears to be characteristic of Homo sapiens, setting them apart from earlier hominins and other animals. These behaviors reflect advances in cognitive capacities, such as abstract thinking and symbolic cognition, as well as increasing complexity in social organization, including larger group sizes and more deeply established cultural traditions.
The art explosion in upper paleolithic
The Upper Paleolithic (~50–10 ka) is widely regarded as a period of intensified artistic production. During this time, modern humans dispersed across Africa and Eurasia, leaving a rich archaeological record that includes cave paintings, rock engravings, portable sculptures, personal ornaments, and evidence of music and dance. The diversity and formal complexity of these aesthetic engagement represent a peak in prehistoric artistic activity.
Cave art represents one of the most prominent forms of Upper Paleolithic aesthetic engagement. More than 200 caves in Europe, particularly in France and Spain, preserve paintings and engravings dated primarily between ~40 and 10 ka . Iconic examples include the animal paintings of Chauvet Cave (~37 ka), the bison imagery of Altamira (~35 ka), and the large-scale polychrome compositions of Lascaux (~17 ka). These works are characterized by highly naturalistic and dynamic depictions of animals, accompanied by abstract signs and occasional human figures, and widely interpreted as reflecting ritual, mythological, or cosmological dimensions of Upper Paleolithic societies. Upper Paleolithic cave art extends beyond Europe, with early figurative examples also documented in Indonesia, such as the Sulawesi warty pig painting from Leang Bulu’ Sipong 4 dated to at least 43.9 ka . Therefore, the origins of complex art are not confined to any single region but may have emerged simultaneously across the globe or through multiple independent developments.
Dated painting of a pig at Leang Balangajia 1 (Brumm et al., 2021)
In addition to cave paintings, the Upper Paleolithic is characterized by a rich tradition of portable art, including small carvings and sculptures made from stone, bone, ivory, and clay . Among the most prominent examples are the so-called Venus figurines, typically depicting highly stylized female forms with exaggerated breasts, abdomens, and hips, and minimal facial detail. These figurines are widely distributed across Europe and date primarily to between ~35 and 25 ka, with examples including the ivory Venus from Hohle Fels in Germany (~35 ka) and the ceramic figurine from Dolní Věstonice (~27 ka) , to date the oldest ceramic documented. While their precise meanings remain debated, these objects are also often interpreted as symbolic representations related to fertility, embodiment, mythological thought, or social identity rather than purely decorative artifacts.
(a) Fragmentary Dolní Věstonice Venus figurine; (b) Bear figurine; (c) Lioness figurine head from Dolní Věstonice (Vandiver, 1989)
Aesthetic engagement during this period also extended beyond the visual domain to include music and performative practices. Archaeological evidence for early musical traditions comes from bone and ivory flutes dated to ~35–40 ka, discovered at sites in southwestern Germany such as Hohle Fels and Geissenklösterle . These instruments, crafted from bird bones and mammoth ivory and featuring carefully spaced finger holes, indicate the production of structured melodies and reflect sophisticated technical and acoustic knowledge (. Music is widely interpreted as having played an important role in Upper Paleolithic social life, potentially supporting ritual activities, group coordination, and social bonding . Although dance has no direct material evidence in the archaeological record, indirect evidence, combined with ethnographic analogy from recent hunter-gatherer societies, suggests that coordinated movement and communal performance likely formed part of collective ceremonies, contributing to emotional synchronization, narrative transmission, and the reinforcement of group identity .
Bone flute from Hohle Fels archaeological horizon Vb (Conard et al., 2009)
Taken together, aesthetic engagement in the Upper Paleolithic has remarkable diversity and formal richness, representing a peak in prehistoric cultural production. These developments reflect both advances in human cognitive and imaginative capacities and increasing social complexity and cultural transmission. From spiritually charged imagery in deep cave contexts, to finely crafted portable sculptures, and to the production of music through bone instruments, Upper Paleolithic aesthetic engagement reveal creative impulses and symbolic concerns that are continuous with those of modern humans.
Why aesthetics emerges and evolves
The anthropological evidence reviewed above suggests that the emergence of aesthetics is not accidental, but rather the product of both biological evolution and cultural evolution. This process can be analyzed along three complementary dimensions: its biological foundations, its cultural foundations, and the interactive dynamics between the two.
Biological foundations: The driving force of survival and reproduction
Sexual selection and the “taste for the beautiful”. In The Descent of Man, Darwin proposed that sexual selection (primarily female choice) is the driving force shaping animal ornamental traits (such as the peacock’s tail), a preference he termed the “taste for the beautiful” . Prum further developed this theory, positing that aesthetic evolution is a coevolutionary process between a signal and an evaluator . In this process, certain traits are retained because they are sensorily attractive and capable of stimulating pleasure (a phenomenon also called “sensory exploitation”, even if they do not necessarily indicate a survival advantage (i.e., “arbitrary beauty”). Human preferences for universal, specific body proportions largely stem from this ancient mechanism of sexual selection. Just as peacocks display their ornamental traits during courtship, early hominins may have attracted mates by producing and displaying “overly” refined handaxes signaling genetic quality and health condition . Female mate preferences could, in turn, have reinforced such aesthetic displays, contributing to the evolution of preferences for particular formal features, such as symmetry and of course mate choice in humans is famously bidirectional.
Natural selection and sensory preferences. Aesthetic preferences also derive from perceptual biases that originally evolved for survival. For instance, primates’ sensitivity to red hues likely evolved in the context of foraging for ripe fruits , and such pre-existing biases could later be co-opted for aesthetic preferences, such as the preferential selection of red ochre . Artistic forms often operate by creating extraordinary stimuli , thereby strongly activating ancient sensory–reward systems. The markedly exaggerated female features of the Hohle Fels Venus figurine provide a clear example of such sensory amplification . Natural selection has also shaped preferences for specific environmental features. For example, human preferences for certain landscapes (such as open prospects, water sources, and refuge) likely stem from ancestral choices regarding survival environments (the Savanna Hypothesis) , and aesthetic responses to animals (such as large felines or young) due to Kindchenschema . Whether the sense of the sublime arising from fear or the affinity elicited by cuteness possess inevitable evolutionary roots.
Metabolic efficiency and perceptual economy. Aesthetic preferences are further shaped by metabolic efficiency, reflecting evolutionary pressures to minimize energetic costs in perception and cognition. The expensive tissue hypothesis proposes that the high metabolic demands of an enlarged brain were offset by reductions in other costly organs, particularly the gut . Under such energetic constraints, selection would have favored not only energetically efficient diets, but also perceptual and cognitive systems optimized for low-cost information processing. Preferences for regularity, symmetry, and predictable structure can therefore be understood as part of a broader evolutionary strategy of energy minimization . Archaeological patterns, such as the persistent emphasis on symmetrical form in Acheulean handaxes, are consistent with this efficiency-oriented framework, linking metabolic constraints, cognition, and material aesthetics in early human evolution . A recent work combining neuroimaging and computational modeling provides convergent evidence that contemporary human aesthetic preferences are negatively correlated with the metabolic expense of visual processing, supporting the idea that aesthetic preferences functions as an energy-saving affective heuristic .
Neural co-option. Another biological explanation is that aesthetic perception likely relies on or co-opts other evolutionarily derived systems. Research in neuropsychology and affective neuroscience indicates that appreciating art can activate emotional and reward circuits in the brain, similar to other adaptive behaviors such as social bonding or parent-child attachment . For instance, listening to music or synchronous dancing releases oxytocin and endorphins, generating feelings of pleasure and social connection, essentially co-opting the neurochemical mechanisms of intimacy and trust . This suggests that early forms of artistic activities (rhythmic drumming, rhythmic movement, melodic singing) may have originated from the biological exploitation of mother-infant bonding systems or group cohesion systems. In fact, Dissanayake’s “artification” hypothesis posits that art “piggybacks” on ancient attachment and emotional circuits: for example, the way mothers naturally sing, speak with exaggerated intonation (motherese), and play with infants possesses aesthetic qualities (rhythm, repetition, variation), which likely facilitated emotional bonding and thus held reproductive value. Over the long course of evolution, these proto-aesthetic behaviors may have developed into music and dance .
Cross-species continuities in aesthetic preferences. The biological foundations of aesthetic preferences are not unique to humans but show continuity across species. Male bowerbirds construct and decorate elaborate bowers to attract mates, selectively arranging objects by color and form, thereby demonstrating aesthetic agency in non-human species . Similarly, black kites (Milvus migrans) have been shown to decorate their nests with conspicuous white materials, such as plastic, functioning as visual signals of territory quality and competitive ability . Even chimpanzees display intrinsic interest in visual mark-making under captive conditions, producing paintings in the absence of explicit external rewards, despite lacking comparable material affordances in the wild . Together, these findings suggest that the biological foundations of aesthetic preferences are evolutionarily ancient and continuous across species and the hominin lineage, rather than a uniquely human innovation.
Cultural evolution: the social brain and group survival
Although biology provides the foundation for human aesthetic capacities, the remarkable diversity and complexity of artistic and aesthetic traditions can only be fully explained through cultural evolution. Over the long history of human evolution, relaxed selection reduced constraints on gene functions, allowing symbolic systems to emerge through the recombination and synergistic integration of preexisting cognitive capacities . On the other hand, changes in diet and energy availability provided the physiological foundation for increased brain size and complexity for social brain . The social brain hypothesis proposes that primate brains, including those of humans, contain specialized neural mechanisms dedicated to perceiving, interpreting, and predicting the behavior and intentions of others . Once humans acquired the cognitive capacities necessary for symbolic understanding and complex social life, artistic activities likely assumed critical social functions and became subject to cultural selection.
Social brain and group cohesion. As human brain size increased, social groups became progressively more complex. Aesthetic activities, such as singing, dancing, and bodily or material decoration in ritual contexts, function to “make special” ordinary actions, transforming them into structured and emotionally salient events . Such ritualized practices exploit biologically grounded sources of pleasure, including rhythm and synchrony, to induce neurophysiological coordination among participants and generate experiences of communitas, thereby strengthening group cohesion and alleviating existential and survival-related anxieties. In non-literate societies, art and ritual thus function not merely as expressions of emotion, but as key media for the transmission of social norms, shared values, and collective knowledge . This function is also evident in religious contexts. The evolution of art and aesthetics is closely intertwined with the evolution of human religious behavior . Participation in religious artistic practices may also have enhanced group survival by fostering social cohesion and facilitating coordinated collective action. During activities such as hunting or territorial defense, ritual performances could help align group members’ shared goals, strengthen commitment to the collective, and reduce free-riding behavior .
Identity signaling and group distinction. At the group level, aesthetic preferences also function as tools for cooperation and competition. The evolutionary theories suggest that during the Late Pleistocene, as human group sizes expanded, new mechanisms transcending kinship ties were required to sustain cooperation, and artistic rituals, such as drumming, chanting, masquerades, and cave paintings in public spaces, likely served as identity markers and coalitionary signals . Adorning oneself with identical body paint or ornaments, chanting shared clan songs, and participating in the same initiation rites all signaled who belonged to “us.” Studies demonstrate that the emergence of standardized ornaments in the African Middle Stone Age, such as those at Bizmoune Cave, correlates with climatic shifts that forced populations into closer proximity, and helped individuals “express their ethnicity or identity to indicate that they belonged to a specific area” during periods of resource scarcity, thereby mitigating conflict through visible symbols of affiliation, thereby mitigating conflict through visible symbols of affiliation . In contexts of ecological uncertainty or resource scarcity, reinforcing social networks through material symbols may be critical for group survival. Demographic modeling further supports that high population density was a prerequisite for the sustainability of such symbolic complexity . Analyzing the distribution of these beads, Vanhaeren & d’Errico (2006) conclude that they functioned as ethno-linguistic markers, allowing groups to visually demarcate boundaries and assert territorial identity . The standardized morphology and repetitive use of specific shell types suggest that these ornaments were not merely expressions of personal aesthetic preference, but signals of adherence to shared social conventions. Thus, the use of beads reflects an identification with a collective symbolic system, wherein conformity to standardized forms served as a marker of group belonging and social commitment.
External symbolic storage. Long before the advent of writing, art likely functioned as a “memory system” and knowledge repository across various cultures. Anthropologists observe that indigenous oral societies utilize mediums such as song, rock art, carving, and dance to encode and transmit information . Myths and epic tales, essentially forms of oral art, encapsulate practical knowledge regarding flora, fauna, and moral codes within emotionally engaging narratives that facilitate retention. Visual art also serves as a mnemonic device. For instance, Australian Aboriginal Songlines and rock art simultaneously encode cartographic data of waterholes alongside spiritual knowledge . Upper Paleolithic cave paintings may have served, in part, to record seasonal animal migrations, hunting techniques, or cosmologies for collective “decoding” and recall . Thus, art emerged as a system of external symbolic communication (external symbolic storage or collective memory), enabling increasingly complex societies to share abstract concepts and maintain cultural continuity.
Co-evolution: the dual inheritance of genes and culture
Human aesthetic experience also emerges from the co-evolution between biological evolution and cultural evolution . Biologically evolved sensory biases shape which artistic forms attract attention and are culturally transmitted, while culturally stabilized aesthetic practices feedback to structure learning environments, social rewards, and mate preferences. Through mechanisms such as sensory exploitation and prestige-biased transmission, cultural selection amplifies certain aesthetic forms far more rapidly than genetic evolution, while biological constraints canalize and stabilize this diversity. Human aesthetic experience thus reflects a dynamic feedback loop in which genes shape culture, culture reshapes selection, and both are jointly inherited across generations.
Conclusion
When modern humans look up at the night sky and feel moved by its beauty, similar emotions may well have been experienced by our ancestors hundreds of thousands of years ago under the same stars. Such shared aesthetic experiences across evolutionary time connect individual cognition to collective human history, revealing how the human mind has long been attuned to finding meaning and emotion in the world. Human aesthetics and art should therefore not be understood as luxuries of late civilization, but as products of the coordinated interaction among natural regularities, biological evolution, and cultural evolution. From the perception of order in the natural world to the emergence of complex symbolic rituals, aesthetic capacities have played a role throughout the evolutionary trajectory by which humans diverged from other animals. These capacities are grounded in genetic inheritance, and are subsequently elaborated and transformed through cultural transmission. From this perspective, the study of aesthetics offers a powerful window onto the cognitive and cultural mechanisms underlying human evolution.
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