Many aspects of people-plant relationships have been explored in past years, especially the effects of plants and flowers on the human psyche. Some researchers have suggested that the presence of plants positively affects mood (Larsen et al., 1998; Shibata & Suzuki, 2002; Haviland-Jones et al., 2005) and attention (Herzog et al., 1997; Kaplan & Kaplan, 1995; Kaplan, 1995; Lohr, Pearson-Mims & Goodwin, 1996; Raanaas et al., 2011; Tennessen & Cimprich, 1995), reduces stress (Cackowski & Nasar, 2003; Grahn & Stigsdotter, 2010) and even decreases recovery time after surgery (Ulrich, 1984).
The perceived beauty of flowers might influence the psychological benefits they provide to humans. It is thus reasonable to ask if there exist any common human flower preferences or whether the perceived beauty of flowers depends solely on individual taste. Although several studies targeted on best-selling flower products provide us with some data (Behe et al., 1999; Yue & Behe, 2010), they have two major limitations. First, they typically focus on only one segment of products (such as geraniums or single stem cut flowers), so it is not possible to generalize their results. Moreover, these studies do not attempt to explain the causes of the observed preferences. Second, the studies combine the effect of morphological traits (color, number or size of the flowers on the plant etc.) with the effect of price, product packaging etc.
General color preference may also influence the beauty of many objects with the same color, including flowers. Green and blue colors could be preferred because they signal a rich and safe habitat (lush vegetation, water, clear sky). Brown or yellow are connected with barren land, drought, dead vegetation or feces and could be avoided (Orians & Heerwagen, 1995, pp. 567--569; Palmer & Schloss, 2010). On the other hand, edible fruits and nuts are often yellow or brown, so the predicted avoidance of these colors is somewhat dubious. Red color may signal edible fruits, sexual arousal or blood (Humphrey, 1980). Red objects should be regarded as stimulating, but whether as beautiful is uncertain.
Some studies targeting the behavior of florist shop customers reported red and pink flowers as the most preferred and blue and yellow flowers as the least preferred (Behe et al., 1999; Yue & Behe, 2010). A study examining the beauty of street flowers found equal preference for diverse flower colors (Todorova, Asakawa & Aikoh, 2004). When people rated their favorite color of a tree canopy, they most preferred red (Kaufman & Lohr, 2004; Heerwagen & Orians, 1993). However, in another study, a red canopy was the least preferred and blue had the highest rating (Müderrisoğlu et al., 2009).
Complexity influences the preference for objects (Jacobsen et al., 2006; Reber, Schwarz & Winkielman, 2004), but not linearly. Studies have reported that objects with very low or very high complexity are preferred less than moderately complex ones (Akalin et al., 2009; Hekkert & Wieringen, 1990). People dislike highly complex objects because they cannot be easily and rapidly recognized and categorized, while objects with very low complexity are just boring. It is questionable whether we would observe an effect of boredom in the case of flowers, because even the simplest ones reach a certain base level of complexity.
Symmetrical objects are also considered beautiful (Jacobsen & Höfel, 2002; Jacobsen et al., 2006; Leder et al., 2004). The processing fluency and the preference for objects increase with the number of their axes of symmetry (Evans, Wenderoth & Cheng, 2000; Tinio & Leder, 2009). This implies that radially symmetrical flowers should be preferred more than bilaterally symmetrical flowers. On the other hand, some researchers claim humans have a very strong preference for bilaterally symmetrical objects, which may be a by-product of the selection of partners (Little & Jones, 2003) and the recognition of partners or enemies (Johnstone, 1994; see also Mithen, 2003). According to the habitat selection approach of Heerwagen & Orians (1993), the type of symmetry could provide information about the nutritive value of flowers. Bilaterally symmetrical flowers usually have more nectar than radially symmetrical ones and indicate richer habitats. For this reason, they should be regarded as more beautiful.
These conditions allowed us to reduce the immense number of flowering plants while maintaining a high morphological diversity. The flowers were not absolutely unknown or notoriously familiar to the respondents, as both of these situations could possibly lead to biased results. The flower size limit guaranteed that the shape of the real flowers could be normally seen with the naked eye. The preparation of the flower stimuli set also included the conversion of photographs to a single size, and it was desirable to keep the converted flower size close to the real one. The last condition eliminated possible problems with compact inflorescences, because it is arguable whether we should distinguish the appearance of single flowers in the inflorescence or treat the whole inflorescence as a single flower. The only exceptions to the last condition were the inflorescences of the aster family (Asteraceae). We included aster family members in the stimuli set because they are very common and the vast majority of people (laypersons) perceive their inflorescences as single flowers.
We found all the Czech flowering plant species in the Key to the Flora of the Czech Republic (Kubát et al., 2002). When the flowers met the inclusion criteria, we included them in the working flower set. In the case of genera with very similar species (e.g., Rubus, Taraxacum), we included the flower of just one species in the working set. The working set comprised flowers of 199 species, which we divided into 26 groups according to their shape. From each group we selected two flowers with different color (e.g., Fig. 1A) and added them to the final flower set (see Table 2).
We used Corel Photo Paint X7 to replace the original flower background by a neutral black color. The black background did not favor any flower (flowers are usually seen on a green, brown, grey or blue background) and provided enough contrast for the clear distinction of the flowers. We then centered the flowers and placed them in the same position, the top petal or tepal pointing directly upwards. Finally, we converted all of the flowers to the same size, optimal for displaying on most computer screens (flower = 150 pixels, flower + background = 200 pixels, the three photographs next to each other = 600 pixels). We also copied the final flower set and converted the photographs in it to a sepia tone (Fig. 1B). This new set was thus devoid of colors and helped us to test the influence of color on the rating of flower beauty. We did not use a conversion to a greyscale because grey photographs on a black background seemed somehow gloomy, which could negatively influence their rating.
In the first questionnaire the respondents rated a set of photographs of flowers by their beauty. The questionnaire also contained several sets of questions concerning basic information about the respondents, their attitude towards plants, color preferences and psychological characteristics.
Because the number of the flower stimuli was quite high (52 flowers in color and sepia tone), we decided to show each respondent only half of them (the first flower of each pair in color and in sepia tone, i.e., subset 1, or the second flower of each pair in color and sepia tone, i.e., subset 2). Although the flower stimuli in each subset remained the same, we randomized their display order. To prevent the respondents from rating the colored flower stimuli under the influence of the sepia tone stimuli and vice versa, we randomized the display order of the colored and sepia tone stimuli and also separated their rating by a set of questions.
As a control, we also created another linear model in which the flower hues were represented as levels of a single factor variable and the flowers with a unique or uncertain dominant color were deleted from the dataset. The final reduced model was very similar to the model in which no flowers were excluded from the data set (adjusted R2 = 0.64, F7,38 = 12.50, p < 0.001), and it contained the same variables with similar significant effects (prototypicality: estimate = 0.32, 95% CI [0.23--0.42], t = 7.02, p < 0.001; dominant blue color: estimate = 0.35, 95% CI [0.09--0.62], t = 2.72, p = 0.010; sharp contours: estimate = 0.30, 95% CI [0.076--0.53], t = 2.70, p = 0.010).
According to empirical research on the perceived beauty of simple colors (Camgöz, Yener, & Güvenç, 2002; Ellis & Ficek, 2001; Hurlbert & Ling, 2007; Schloss, Strauss & Palmer, 2013; Zemach, Chang & Teller, 2007); and tree canopies (Müderrisoğlu et al., 2009), blue is the most and yellow the least attractive color. A preference for blue was also reported for pita birds, which are very similar in shape but differ in coloration (Lišková, Landová & Frynta, 2014). We can assume that the general human color preference (as determined in American and European populations) also applies to flowers.
We must point out, however, that the yellow color (least preferred) in our set of stimuli was saturated. It is evident that clear yellow is more related to the sun or ripe fruits than to dead vegetation or drought. In our opinion, the habitat selection theory cannot fully explain the dislike of saturated yellow. Studies based on the ecological valence theory reported a low preference only for dark shades of yellow, whereas saturated yellow had an average preference. If we follow the assumptions of the ecological valence theory, we could argue that the Czech population tends to particularly dislike salient objects that typically have a saturated yellow color. This in turn could lead to a low general preference for saturated yellow and explain the observed dislike of yellow flowers.
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