1. Introduction
Known as “the queen of fruits”, the strawberry is a photophilous, and shade-tolerant herbaceous plant [
1]. It is widely favored by consumers due to its adorable colors, sweet, sour tastes, and high nutritional value. With increasing demands in both the quantity and quality of strawberries in the market, strawberry cultivations are shifting from natural surroundings to artificial greenhouses, due to the merits of environmental controllability and economic efficiency for the latter [
2,
3]. Being an essential environmental factor, light is indispensable to the pho-to-morphogenesis and photosynthesis of plants in both natural and artificial conditions [
4,
5,
6,
7,
8]. More than providing the energy for photosynthesis, light also dictates specific signals which regulate plant development, shaping, and metabolism, driven by light colors [
6,
9,
10]. However, adverse weather like overcast, snow, rain, and fog in winter and spring always leads to inadequate light exposure, resulting in declined productivity of economic plants. Furthermore, because of the aging effect of plastic material and accumulated layers of soil in cultivation greenhouses, transmitted light levels are normally below the natural need for light for plants [
11]. Hence, the introduction of artificial supplemental lights to supply adequate illumination for the plants in facility agriculture is clearly needed.
In recent years, various factors of supplemental light are investigated to explore their influences on the quality of strawberries. For example, studies show that luminous intensity affects the growth speed and nutrient contents of strawberries [
12,
13,
14,
15]. Photoperiod could influence the photosynthesis, flowering stage, and yield of strawberries [
2,
16,
17]. Up to date, more and more attention has been paid to investigations of plant growth and photomorphogenesis of strawberries affected by the color of light [
18,
19,
20,
21]. For example, Takeda found that a red-light treatment could reduce flower falling and a high ratio of far-red light to visible light reaching the crown plays a role in floral bud induction [
22]. Xu found that the treatment using blue light is helpful to maintain the flavors and nutrition of harvested strawberries [
10]. Nhut found that a ratio of 7:3 between the red light and the blue light is optimal for the growth of plantlets [
18]. Li reported that the strawberry plants greatly benefited from a color ratio of 3:2:1 between the orange, red, and blue lights [
23]. Furthermore, in terms of the fruit quality, different proportions of colors are also tried to find the optimal light condition. For instance, Wu found that a ratio of 1:1:1 between the red LEDs, blue LEDs, and white LEDs gave rise to an addition of average weight for single strawberries and an improvement in fruit flavor [
24].
Although there are certain studies on the improvement of fruit quality and quantity of strawberries using supplemental light, selections of colored lights are more empirical or qualitative, from an aspect of photometry or colorimetry. Terms such as "red," "green," and "blue" commonly used in these studies cannot be precisely defined in the field of color or optics sciences. The lack of precision might cause poor repeatability and stability for the fruit quality and production of strawberries in greenhouses during implementations of supplemental lights. Here, we have employed a digitally col-or-coding method (DCCM) to precisely control chromatic parameters of supplemental light in terms of chromaticity coordinates or correlated color temperatures (CCTs) [
25,
26]. We have systematically investigated their influences of these chromatic parameters on the fruit quality of strawberries. The DCCM we have utilized, along with the results regarding the chromatic effects on strawberry fruit quality, can be widely applied to global greenhouses for artificial cultivation and mass production of high-quality strawberries and other fruits in markets.
4. Discussion
Different light qualities and intensities have a wide range of regular effects on plant growth and development. Light quality affects hormone balance in plants through the action of related pigments, which in turn impacts growth, development, and production [
27]. Light intensity directly influences plant reproduction and fruit ripening [
28,
29].
The height of strawberry plants under supplemental lights is higher than those in the reference group. Specifically, the height of strawberry plants with supplemental lights at illuminances of 1000 lx and 600 lx is about 22% and 10% higher than the reference group, respectively (
Figure 3), while chromatic effects on plant height are not so significant. Other studies have found that additional supplemental light on strawberry plant growth is markedly better than natural light [
24]. The maximum plantlet height was observed at illuminances of 45 µmol·m
-2·s
-1 and 75 µmol·m
-2·s
-1 [
18]. Various experiments have been conducted to study the effects of different light qualities on plant height. For example, the tallest plant height was observed when plantlets were grown under 100% red light, while other combinations of red and blue light did not result in significant differences in height [
18]. The plant height can be influenced by the rated power of the light source, with a larger rated power contributing to greater plant height when the same color mixing ratio was used [
23]. When the rated power remains constant, adding orange-color light to the red and blue color mix has been shown to impact the plant height of strawberries [
23]. Additionally, the tallest strawberry plants were obtained when grown under blue light [
5]. A higher plant height was observed when the ratio of red LED to blue LED was 10:1, and no significant difference was observed compared to a 19:1 red to blue light ratio [
30]. These findings indicate that the specific light quality and color mixing ratios can play a role in influencing plant height, although the overall impact of these factors may vary depending on the plant species and environmental conditions.
When comparing the single weight of strawberries with additional supplemental lights to normal natural lights, it nearly stays the same level. A higher illuminance (1000 lx) of supplemental light results in a smaller fluctuation in response to different colors. Furthermore, chromatic effects on the single weight of strawberries are not significant, as shown in
Figure 4. These results agree with previous findings that white LEDs do not increase the single weight of strawberries compared to the reference group [
5].
For the hardness of strawberries, significant variation can be observed under different CCTs and illuminance treatments. Hardness could be significantly reduced by using CCTs ranging from 2400 K to 4000 K. At the meantime, a higher illuminance could enhance such chromatic dependences, as illustrated in
Figure 5. These results agree with previous study [
14]. There are some studies recommend a mixed ratio of 1:1:1 among red, blue and white color supplemental lights to reduce the hardness of strawberries, but their quantitative chromatic parameters are not provided [
24].
The soluble solid content of strawberries shows significant improvements using the treatment of supplemental light. In experimental groups, the soluble solid content is generally higher than in reference groups, as depicted in
Figure 6. These findings are consistent with previous studies, which concluded that the quality of strawberry fruits could be significantly improved by supplemental light treatment, including increased soluble solid content and soluble sugar content [
24]. However, other research suggests that variance of illuminance levels doesn’t cause significant changes in soluble solid content [
14]. Regarding the chromatic effect, our results have a fair agreement with other studies [
31], that is no significant differences in soluble solid content are found between different combinations of monochromatic LED lights (red, green, and blue) and polychromatic (W-R: G: B, 1:1:1) lights [
31].
The titratable acid content of strawberries can be significantly influenced by LED supplemental lights. Maximum titratable acid content can be achieved if the CCT is within the range from 2250 K to 2600 K, and from 4000 K to 4500 K. At the meantime, a higher illuminance level could enhance such chromatic dependences, as shown in
Figure 7. Similar study found that the titratable acid content could be enhanced by following a treatment of supplemental light but in an insignificant manner.
In general, the light intensity should not be a great concern for supplemental light. This is because the dependences of five quality parameters on the light intensity are much weaker than on the CCTs, even the intensity could slightly strengthen, weaken, or shift the corresponding chromatic dependences. Another reason is that the intensity effect can be compensated by other lighting factors, for example, lighting period, to some extent.
For future work, we plan to conduct more specific experiments from the following aspects:
a) Change the CCT and illuminance of supplemental luminaires in smaller steps, so that more continuous chromatic and intensity dependences could be observed.
b) Measure other important quality parameters like vitamins, soluble sugar, anthocyanidin, and so on.
c) Record the total weight of the strawberries from different experimental group.
d) Investigate the impact of supplemental light with varying parameters such as photoperiod and timing on quality parameters of the strawberries.
e) Extend the related techniques to other kinds of fruits.