Neuroscientific research on aesthetic processes in the visual system has offered evidence for a correspondence between certain properties of artworks and organizational principles in the brain (Zeki, 1999; Kawabata and Zeki, 2004; Chatterjee, 2011; Chatterjee and Vartanian, 2014; Bao et al., 2017a). Aesthetic experience partly relies on a visual analysis of an artwork based on distinct and specific visual attributes in color, line, texture, and form (Cinzia and Vittorio, 2009; Palmer et al., 2013). Indeed, some researchers (e.g., Latto, 1995; Bao et al., 2017b) have argued that artists implicitly tap into propensities of the human nervous system when generating aesthetic appeal. Piet Mondrian’s paintings are exemplary of visual art that excites orientation-selective neurons in the primary visual area. These neurons respond selectively to dots and straight lines, especially horizontal and vertical ones. Zeki (1999) principal idea is that different kinds of artworks excite different groups of neurons in the brain, leading to differential bottom-up processes.

Drawing upon complementarity as a basic principle of biological functions (Bao et al., 2017a), aesthetic experience relies both on higher-level cognitive processes and lower-level visual analyses to enable object identification, which is necessarily both bottom-up and top-down (Zhou et al., 2016). Some of the most studied modulators of the top-down system are individual expectations, prior knowledge, social context, and cultural background (Silveira et al., 2012, 2015b,c; Graupmann et al., 2013; Leder, 2013), which also applies to other cultural domains, like poetry (e.g., Zhao et al., 2018) or even religious beliefs (Silveira et al., 2015a). It has recently been suggested that aesthetic experience is partially top–down and varies between individuals according to their cultural experience (Redies, 2015). A match of internal and external information as represented in feedforward and feedback signals of the two complementary systems plays an important role in creating perceptual stability and even pleasure (von Holst and Mittelstaedt, 1950). Human cognitive architecture is presumably built to predict representations of the world and efficiently minimize prediction errors (Clark, 2013). Conceptually overlapping with this notion, art theory describes an aesthetic experience as an interplay between internal and external perspectives (Bao et al., 2017a), i.e., by dissolving the subject/object dichotomy, the observer experiences a sense of immersion (Heidegger, 1986).

In an attempt to resolve this bi-directionality or complementarity (Bao et al., 2017a), one approach is to define an artwork as a medium capturing both an artist’s understanding of a viewer’s experiences and a viewer’s understanding of an artist’s intentions (Jucker and Barrett, 2011; Tinio, 2013). This perspective considers aesthetic appreciation to be an interaction between an artwork’s objective properties and the viewer’s processing characteristics of those properties. This idea is consistent with suggestions that beauty is not “put in” the artwork by the artist as a distinct entity (Zaidel, 2015), but rather “an emergent property in the brain of the beholder,” where the beholder can include the artist, as well as the viewer (Redies, 2015). From a neuroimaging perspective, this phenomenon might be related to brain activation, which corresponds to the so-called mirror-neuron system and embodied simulations of emotions and actions (Freedberg and Gallese, 2007). It has repeatedly been found that positive aesthetic appreciation goes along with neural activation in parietal and sensory-motor brain regions (e.g., Lutz et al., 2013), which has been interpreted as either an empathic resonance with the painting’s content or imagery and mimicry of artistic gestures. Referring back to early psychological theories, Helmholtz (1868/1995) combined the ideas that artists test and explore the visual system with a theory of vision. Artists, he claimed, are particularly good at observing their sensuous impressions (data of sensation) and at figuring out which patterns trigger which interpretations. Although many visual scientists have given up Helmholtz’s theory of vision, the idea that artists test and explore the visual system has not really been abandoned (Pöppel et al., 2013). On the contrary, it allows for a more detailed and discriminating version of the same idea.

At the neural level, a considerable number of brain imaging studies on aesthetic experience demonstrate the involvement of brain regions, like the ventral striatum and medial prefrontal cortex, which are activated by reward and positive emotion (Cinzia and Vittorio, 2009; Chatterjee, 2011; Nadal and Pearce, 2011; Pearce et al., 2016). The default-mode network (DMN) is also involved when processing paintings of high aesthetic preference (Vessel et al., 2012) or of higher predictability in inferring meaningful content (Silveira et al., 2012) and has been proposed as a delayed aesthetic network (Cela-Conde et al., 2013). The DMN displays the highest activation during the resting state and task-dependent, lower activation levels (Raichle and Snyder, 2007). Overlapping with the cortical midline structures, parts of the DMN may be neural correlates for self-relevant processes. Particularly posterior parts, i.e., the posterior cingulate cortex (PCC) and adjacent precuneus, supposedly correspond to mental processes while integrating external information into a self-referential context (Northoff and Bermpohl, 2004). These processes seem to be culturally sensitive (Han and Northoff, 2008). According to recent enactive accounts of aesthetic experience (Xenakis and Arnellos, 2014, 2015), aesthetic experience arises as a result of the interaction between the viewer and the object. Aesthetic experience is also an embodied phenomenon directly linked to adaptation and aesthetic perception and helps cope with the environment. Aesthetic experience arises when the viewer interacts with both uncertain physical and cultural contexts. A more inclusive understanding of aesthetic experiences in diverse cultures has to be developed. Cultural aesthetics requires an empirical inquiry into the kinds and varieties of experiences associated with artistic activities as they are understood in different cultures.

Regarding cultural aspects, Western and Eastern artists tend to use different perspectives to represent the visual world. Western landscape paintings have been rather precise reproductions of visual surroundings since the Renaissance (Kubovy, 1986), while Eastern landscape paintings, such as Chinese paintings, have a dynamic quality that integrates successive time windows (Bao et al., 2015) and are expected to convey the experience of “being in nature” rather than “seeing nature.” This has led to an arrangement of spatial information in a vertical manner with multi-layers on top of each other in a scroll form (Sullivan, 1984; Cameron, 1993; Law, 2011; Tyler and Chen, 2011). Considering cultural and historical contexts in which painting styles evolved, Western artists applied more logic and mathematics (Kline, 1964), like geometric perspective. Prevailing Buddhist and Taoist influences among Chinese artists could adequately explain the emergence of landscape painting in the Southern Song period and its persistence throughout the Ming dynasty (Golas, 2014). Besides the different artistic approaches employed by Western and Eastern painters, viewers from these two cultural groups also have distinctly different aesthetic reactions to the same artistic visual representations. Both Western and Chinese participants gave higher aesthetic ratings to paintings from their own compared to the other culture (Bao et al., 2016). One explanation might be that repeated exposure to artworks (Zajonc, 2001) promotes the development of concepts of beauty, but also of culturally transmitted values and beliefs. A given cultural value system is internalized by members of the culture, and those who internalize that system display habitual ways of thinking and behaving. Certain artworks can trigger a culturally specific feeling of identity (Pöppel, 2010). We suggest that the cultural environment in which the individual is socialized can account for the production and appreciation of art.

The current study aimed at further investigating aesthetic processes in the context of culture. We used cross-cultural neuroimaging to measure neural activity in native Chinese and Europeans living in Germany while they viewed Western and traditional Chinese landscape paintings. Based on behavioral findings of cross-cultural differences, we predicted that distinct brain-activation patterns can be observed which correspond to aesthetic preference of one’s own vs. the other culture.

We investigated the neural correlates of viewing Western and Eastern landscape paintings in Western and Eastern participants with fMRI. We demonstrated a cultural difference in neural responses underlying aesthetic appreciation; participants showed stronger brain activation to artistic expressions from their own cultural systems. On the behavioral level, participants from the different cultural groups expressed distinct aesthetic judgments from the two different cultures. The behavioral results, however, did not support our expectation that participants would show an overall more positive aesthetic appreciation for landscape paintings from their own culture. European participants rated Western landscape paintings when compared to Eastern landscape paintings higher on valence, preference, beauty, relaxation, empathy, object-related absorption, and lower on arousal. Chinese participants did not express a general preference for Eastern landscape paintings; they gave compared to Western paintings only higher ratings in the domain of relaxation. Despite the absence of a behavioral evidence of reversed aesthetic preference patterns (which does not prove the evidence of absence), we observed greater brain activation in participants for landscape paintings from their own culture. The absence of an effect when comparing the upside-down versions of the paintings highlights the content dependency of the viewed paintings. One can conclude that the observed differences are not based on bottom-up processing of the physical properties of the paintings, but that pictorial properties of the paintings extracted on a level of higher cognitive processing are responsible.

When the European participants viewed original Western as compared to Eastern landscape paintings, the analysis revealed higher BOLD activation levels in the primary visual cortex, the SMA, the PCC, and the hippocampus. The SMA, which has motor and sensory functions (Jeannerod, 2001), counts as one of the commonly activated brain structures when viewing artworks (Vartanian and Skov, 2014; Boccia et al., 2016). Our results correspond to previous evidence relating exposure to aesthetic vs. non-aesthetic visual stimuli to sensory-motor processes (Freedberg and Gallese, 2007; Lutz et al., 2013) and may well indicate empathic resonance with paintings from one’s own cultural background. An enactive understanding of perception (Xenakis and Arnellos, 2014, 2015) suggests aesthetic behavior as emerging from the structural coupling of the viewer with his/her environment. The higher neural activation when viewing Western paintings is related to the viewer’s embodiment, as posited by enactivists, and to the whole spectrum of abilities available in human socio-cultural practices.

The most likely explanation for why Western paintings elicited higher activation in the fusiform gyrus in the European group is that this structure is known to mediate object recognition. When observers view a work of art, they do not see, for instance, a canvas or a statue on an abstract level; rather, they react to what it represents (e.g., a seascape, a tower). Compared to traditional Chinese landscape paintings, Western landscape paintings have clear-cut shapes similar to hard-edge paintings. In Chinese paintings, an object’s edge is ill-defined and disappears or fades into another object or into the background (Cahill, 1960; Zong, 2007). According to Wölfflin (1950), soft-edge paintings involve blurred boundaries and ill-defined forms, whereas hard-edge paintings involve well-defined forms and rich narrative details. Since the left fusiform gyrus probably plays a role in shape processing (Starrfelt and Gerlach, 2007), viewing Western landscape paintings invoked more fine-grained shape analysis than Chinese paintings, thus causing greater activation in the fusiform gyrus. This explanation is consistent with previous observations that activation in the fusiform gyrus can be attributed to object recognition in realistic paintings, including faces (Grill-Spector and Malach, 2004) and pictorial scenes (Siebörger et al., 2007).

The right cuneus, the bilateral calcarine cortex, and the left lingual gyrus were activated in the comparison of aesthetic appreciation of Eastern paintings vs. Western paintings by Chinese participants. Activation of the lingual gyrus has been linked to the encoding of complex images. Chatterjee et al. (2009) argued that activation of the lingual gyrus represents its sensitivity to “beauty,” which corresponds with the present findings. Mizokami et al. (2014) showed that the left lingual gyrus and the bilateral cuneus may be associated with aesthetic judgments of representational paintings, which also supports our findings. Previous studies have related the cuneus to the aesthetic appreciation of beauty (Cupchik et al., 2009; Mizokami et al., 2014). The stronger activation in these brain structures may be explained by higher aesthetic sensitivity to stimuli with high visualization demands. Of particular interest is activation specific to the Chinese paintings localized within the calcarine sulcus corresponding to the primary visual cortex. More specifically, activation was localized to the mid- to anterior parts of the calcarine sulcus, corresponding to the mid-level and peripheral visual field representations (Sereno et al., 1995).

The higher activation levels in the primary visual cortex in both cultural groups not only indicate stronger bottom-up visual processing of paintings from one’s own culture. The simultaneous involvement of the PCC in the European group, which has cortical connections to visual areas in feedforward, as well as feedback projections (Keil et al., 2009), also suggests stronger top-down processing, i.e., higher-level cortical areas modulate neural activity on the sensory-input levels. For the Chinese group, stronger activation was localized in the superior parietal lobule, a neural region known to support the manipulation of information in working memory (Koenigs et al., 2009). Top-down processes originating in superior parietal areas may contribute to the aesthetic perception of paintings and their maintenance in the “mind’s eye” (Mechelli et al., 2004).

Previous studies suggested that aesthetic experience is a function of the interaction between top-down and bottom-up processing (Leder et al., 2004; Cupchik et al., 2009; Redies, 2015). However, it is difficult to delineate the exact projecting pathway of feed-forward and feedback signals with the current experimental settings. Support for a stronger involvement of top–down information processing for paintings from one’s own cultural background is the additional engagement of the hippocampus, a region that has consistently been related to episodic memory retrieval (Eichenbaum et al., 1996, 2007; Brown and Aggleton, 2001). Paintings from one’s own culture might be integrated into a frame of prior experiences or expectations to a higher degree due to their fit with culturally imprinted cognitive schemata, thereby linking aesthetic appreciation to one’s own previous experiences. This could determine the relationship between cognition and action, thereby generating a value-oriented experience of high complexity (e.g., aesthetic appreciation), which motivates human social behavior to be related to certain types of socio-cultural expressions (e.g., appreciating or even creating/producing artworks).

We observed activation in the PCC in European participants when comparing Western to Eastern landscape paintings. This region has emerged as a key component of the DMN (Mason et al., 2007; Christoff et al., 2009; Andrews-Hanna et al., 2010), a highly interconnected network of brain regions that has been associated with self-referential mental processing (Northoff et al., 2006). More specifically, the PCC is active during tasks that involve integrating external stimuli in a self-referential manner (Northoff and Bermpohl, 2004), as well as jn autobiographical memory (Buckner and Carroll, 2007). The PCC responds to self-relevant information even when there is no explicit requirement to evaluate self-relevance (Moran et al., 2009; Reniers et al., 2012). In aesthetic experiences, individual taste in art can be considered as highly subjective, connected with personal pleasure, and can be considered a part of the personal self (Zaytseva et al., 2014). Previous research also provides evidence of higher activation levels within the DMN associated with intense aesthetic experience and interpreted as personal relevance of these stimuli (Vessel et al., 2012). When analyzing temporal dynamics of aesthetic appreciation, it has been suggested that the DMN involvement is a core part of general aesthetic experience, as represented in a delayed aesthetic network (Cela-Conde et al., 2013). The current study demonstrated that an inclusion of a cultural context was a modulator of these processes. Our results support previous findings of culturally sensitive information processing within the DMN (Han and Northoff, 2008). In Chinese group, we found evidence for increased activation in the right postcentral gyrus. This is noteworthy given that previous studies revealed the right postcentral gyrus to be activated in response to self-vs. other-perspective taking in social cognition (Ruby and Decety, 2004; Adams et al., 2010). Because the postcentral gyrus mediates somatosensory experience, the results might be explained by motor resonance created in the viewer when viewing paintings from his/her own culture. This activation, however, was not in a common area found in the previous tests of aesthetic appreciation and requires further empirical corroboration before firm conclusions can be drawn.

Why didn’t Chinese participants show any activation related to self-referential information processing? There are two possible explanations. Our experimental paradigm was comparatively subtle. Pronounced cultural differences in the neural processing may emerge under more demanding conditions, verifying the expected bias of the Eastern cultural group to show preferential processing of artworks in self-related brain regions similar to the Western groups. An alternate explanation is that Asian societies are changing rapidly, and that the young Chinese participants have internalized Western values up to the point that they no longer display behavioral patterns which are characteristic for Asian cultures. Even if this is the case, the neuroimaging results clearly demonstrate different patterns for Eastern and Western participants, with a bias toward higher brain activation for one’s own cultural artworks, a finding supporting the cultural/aesthetic framework proposed by Bao et al. (2016).

Our findings provide neuroimaging evidence for cultural biases in the processing of visual artworks. Visual aesthetic studies make us aware that “seeing” has a history; sensory experiences are socially, culturally, and historically embedded. It is not possible to speak of pure perception as sensation untouched by past experiences, education, and cultural background. Since our self-identity is influenced by the cultural background we inhabit, the “belonging” moments should occur with greater frequency when artists of the presented paintings and viewers who appreciate the paintings share the same cultural background. Members of different cultural groups are repeatedly exposed to various examples of visual images from their respective cultures, and they may implicitly gain knowledge (Pöppel and Bao, 2011) about the dominant aesthetic representation of the world. As mental representations of the world influence the perception of visual information, a cultural framing effect is probably implemented on the neural level and determines the implicit information processing of visual stimuli in particular and sensory stimuli in general. In this way, different cultural groups produce a “sensory model” of the meanings and values associated with their sensory perceptions. We propose that culturally distinct behaviors and thoughts can be construed as differences in aesthetic appreciation that affect neural functions. The authenticity of aesthetic experience, through its directness and immediacy, provides a powerful means of reappraising cultural experience by digging beneath the layers of accrued meanings and cognitive habits.

In conclusion, our results highlight culture as a modulating factor in the visual perception of aesthetic stimuli. Aesthetic experiences are aligned with the human cognitive architecture which is based on the principles of efficiency and prediction. This indicates a cultural framing effect, which corresponds to the mind, as well as to brain states, and determines implicit information processing. Our findings support the notion that visual experiences are biased by cultural factors. We engage aesthetically with artwork in its different expressions (e.g., visual art, poetry, music) and also with different physical environments (e.g., untouched nature, architecture, or different kinds of urban surroundings). Environments define specific perceptual features as characteristic for a particular human culture at a given time resulting in typical aesthetic appreciation. Beauty really is in the “eye” of the beholder.

This study was approved by the ethical committee of Ludwig-Maximilian University Munich, in agreement with the Declaration of Helsinki.

YB and EP conceived and designed the study. TY, SS, and AF acquired the data. TY and SS analyzed and discussed the data. TY and SS drafted the manuscript. MP, TS, YB, and EP critically reviewed the manuscript.

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.