Open Access Article
Christos Vatistas
*,
Dafni Despoina Avgoustaki and
Thomas Bartzanas
Department of Natural Resources and Agricultural Engineering, Agricultural University of Athens, Iera Odos 75, Athens 118 55, Greece. E-mail: vatistasx@aua.gr
First published on 24th April 2026
Controlled Environment Agriculture (CEA) and vertical farming (VF) enable urban food production through artificial lighting and soilless cultivation, reducing transport distances and supporting year-round supply. However, sustainability is strongly affected by the energy demand of LED lighting, making it critical to improve yield per unit of supplied light and energy. Microgreens are well suited to VF systems because they are harvested at an early developmental stage, typically 7 to 21 days after sowing, allowing short production cycles. In this study, nine LED spectra (UV, blue, green, orange, red, far-red, blue-red, blue-red-far-red, and cool-white) were tested under 4, 8, and 16 h photoperiods at constant light intensity, resulting in different daily light integral (DLI) levels, to evaluate their effects on seedling emergence, early development, biomass accumulation, and final microgreen yield in cabbage and arugula. Higher emergence values for cabbage were observed under 16 h blue light, while arugula showed the highest emergence under 8 h green light. In both species, 16 h blue-red-far-red treatment was associated with more compact seedlings and higher dry to fresh biomass ratios. In a second phase, emergence enhancing spectra were combined with the biomass enhancing blue-red-far-red treatment to evaluate microgreen production on soil. This two-stage lighting strategy yielded the highest fresh biomass in both species, 2.189 g for cabbage and 12.56 g for arugula. These findings indicate that tailoring the early light environment through species-specific emergence-enhancing spectra can improve microgreen uniformity and yield in vertical farming systems, while simultaneously enhancing energy-use efficiency and light-use efficiency.
Sustainability spotlightEnergy demand from artificial lighting remains a key challenge for the sustainability of urban vertical farming. This study demonstrates that appropriate lighting during the early cultivation stage can increase the number of successfully established seedlings, while targeted lighting during subsequent growth can accelerate development and increase microgreens biomass production in cabbage and arugula. By combining an emergence-enhancing light treatment with a biomass-enhancing spectrum in a stage-specific strategy, higher yields were achieved compared with conventional production practices. These productivity gains were associated with improved light-use and energy-use efficiency, highlighting the potential of targeted lighting management to support more sustainable and resource-efficient urban food production. |
The ease of mass production has led to inefficient management of production volumes, resulting in the overexploitation of natural resources, increased energy consumption, and the accumulation of waste. These effects contribute to ecosystem degradation, greenhouse gas emissions, and water and soil pollution.3 According to the Food and Agriculture Organization (FAO), 14% of food produced by agriculture is lost between harvest and before reaching consumers, mainly due to inefficiencies in harvesting, transportation, storage and processing.4 A promising approach to addressing these challenges and supporting urban food production is vertical farming (VF).5 This method employs hydroponic systems and vertically stacked cultivation units, allowing efficient use of space. The use of artificial lighting enables precise control of the Daily Light Integral (DLI) which can enhance biomass production and shorten growth cycles compared with open-field or greenhouse systems.6 In addition, automation and environmental control systems allow stable cultivation conditions and improved resource use efficiency.7
Microgreens constitute a category of crops with high commercial and research interest in VF systems. They are tender seedlings grown mainly from vegetable seeds and harvested once the cotyledons are fully developed, typically before the first true leaves appear. Their rising popularity is due to several factors that make them attractive for production.8 These include their suitability for cultivation under artificial lighting, their adaptability to urban environments, their year-round production potential, and their use for both personal consumption and educational purposes.9 Additional advantages include their high nutritional value, intense flavor, and wide diversity in color, taste, and morphology, which contribute to their increasing demand and added value.10 Microgreen cultivation requires relative low inputs, such as water, substrate, and space, while minimizing waste by producing mainly edible biomass. Furthermore, local production close to consumers, reduces post-harvest losses and transportation-related emissions, supporting more sustainable food systems.8,11
In microgreens production, early-stage growth is particularly critical due to the very short cultivation cycle. Uniform and rapid seedling emergence is essential for achieving homogeneous canopies and synchronized harvest, while hypocotyl elongation directly affects product quality and marketability. In addition, maximizing biomass production within a limited time frame is a key objective, closely linked to light use efficiency and overall energy consumption in controlled-environment systems. Therefore, optimizing light conditions during the early developmental stages represents an important challenge for improving both productivity and resource efficiency in microgreens cultivation.9,11
In this context, lighting management becomes a critical factor in microgreens production systems, particularly in vertical farming environments. One of the main challenges for the sustainability, profitability and viability of VFs is the intense energy demand required for lighting operation. Although the use of LED lamps significantly reduces energy consumption, lighting costs remain high and can reach up to 80% of the total production cost in a VF.12 Research on the optimal light intensity and spectrum for each crop, the optimal photoperiod, and improving LEDs efficiency are key areas for reducing costs, increasing production, and enhancing the sustainability of these cultivation systems.13
Plants utilize light in the 400–700 nm range, known as Photosynthetically Active Radiation (PAR), to support growth and development.14 Light perception is mediated by photoreceptors, which regulate molecular and physiological processes throughout the plant life cycle.15 In plants, major photoreceptor groups include phytochromes (detect light at red/far-red parts of spectrum), cryptochromes (detect light at blue/UV-A parts of spectrum), phototropins (blue light receptors associated with stomatal opening), UV-B receptors (UVR8) (detect light in the UV-B range) and ZTL-like proteins, which detect light in the blue range of the spectrum and are associated with the regulation of the circadian clock.16 In seeds, phytochromes play a key role in regulating germination by controlling hormonal balance, promoting gibberellin (GA) synthesis and reducing abscisic acid (ABA) levels under favorable light conditions.15,17 Photoreceptor activity influences signaling pathways involved in photomorphogenesis and seed germination, although responses vary among species and wavelengths.11,14,18–20 While some species exhibit light-sensitive germination, many cultivated vegetables germinate equally well in darkness, with light primarily affecting seedling emergence and early post-germinative development. Seed size has also been linked to light sensitivity, as smaller seeds, with limited nutrient reserves, tend to respond more strongly to light cues that facilitate emergence and the transition to photosynthetic growth.21,22
Although previous studies have investigated the effects of light spectrum and photoperiod on plant growth, limited information is available on how early-stage light conditions affect seedling emergence and subsequent microgreen yield, particularly in controlled-environment systems. In addition, the potential benefits of applying different lighting conditions during successive developmental stages remain insufficiently explored. Understanding seedling responses not only to blue and red wavelengths, which are strongly associated with photomorphogenic regulation and photosynthetic activity, but also to other spectral regions such as green, orange, UV, and far-red radiation, is of particular interest. While green and orange wavelengths fall within the PAR range, UV and far-red lie outside it and are known to influence plant development through distinct photoreceptor-mediated pathways.15–17
The aim of this study was to evaluate the effects of different light spectra and photoperiods, applied under constant light intensity (different DLI), on seedling emergence, early seedling development, and subsequent microgreen production of cabbage and arugula under controlled-environment conditions. Faster and more uniform emergence can shorten the overall production cycle and improve cultivation process efficiency, while early-stage light conditions strongly influence seedling morphology and subsequent biomass accumulation. In this context, the second phase of the study focused on identifying the light spectrum that maximizes biomass production and on evaluating whether a two-stage lighting strategy, in which the emergence-enhancing spectrum is applied during the first days after sowing and the biomass-enhancing spectrum is used during subsequent growth, can enhance final microgreen yield while enabling a more efficient use of supplied light and energy. Based on current knowledge of plant responses to light, it was expected that both spectral quality and daily light exposure would influence seedling emergence and early development, and that combining different lighting conditions across developmental stages could provide advantages over the use of a single lighting regime.
Artificial lighting was provided using LED light sources emitting relatively narrow-band radiation centered on specific wavelengths. Spectral measurements were conducted using a spectroradiometer (UPRTEK PG200N, measurement range 350–800 nm), which was used to determine the peak wavelength and approximate emission range of each LED spectrum. Although LED sources were selected to target specific spectral regions, measured emission profiles indicated partial overlap between adjacent wavelength bands; however, each treatment was characterized by a dominant peak within the intended spectral region. The spectra studied were: (1) UV-A (measured emission ∼350–400 nm, peak 364 nm), (2) blue (∼410–500 nm, peak 448 nm), (3) green (∼470–570 nm, peak 516 nm), (4) orange (∼550–640 nm, peak 594 nm), (5) red (∼580–670 nm, peak 630 nm), (6) far-red (∼650–770 nm, peak 729 nm), (7) a combination of blue and red (1
:
1), (8) a combination of blue-red-far-red (1
:
1
:
1), and (9) cool-white LED (∼410–750 nm, peak 448 nm) (Table 1). Mixed spectra were generated using the corresponding monochromatic LED sources described above. The photosynthetic photon flux density (PPFD) was set to 30 µmol m−2 s−1 for all PAR spectra (400–700 nm). For the UV and far-red treatments, which fall outside the PAR range, intensity was measured as photon flux density (PFD). The UV spectrum had a PFD value of 0.29 µmol m−2 s−1, while the far-red treatment was applied at a photon flux level comparable to the PPFD used for PAR spectra. Each treatment lasted for 7 days (168 h).
| Photoperiod | Spectra | DLIa (mol per m2 per day) | ||||||||
|---|---|---|---|---|---|---|---|---|---|---|
| UV | Blue | Green | Orange | Red | Far-red | Blue-red | Blue-red-farred | Cool-white | ||
a DLI values refer to the daily light integral corresponding to a PPFD of 30 µmol m−2 s−1 across the three photoperiods applied. In the case of UV lighting, DLI values were significantly lower due to the very low PFD, measuring 0.0042, 0.0084, and 0.0167 mol per m2 per day for the 4-, 8-, and 16 h photoperiods, respectively. All PAR spectra were applied at a constant PPFD of 30 µmol m−2 s−1, while UV and far-red treatments were expressed as PFD. Mixed spectra were applied at equal photon ratios (blue : red = 1 : 1 and blue : red : far-red = 1 : 1 : 1). |
||||||||||
| 4 hours | 4hUV | 4hB | 4hG | 4hOr | 4hR | 4hFR | 4hBR | 4hBRFR | 4hCW | 0.432 |
| 8 hours | 8hUV | 8hB | 8hG | 8hOr | 8hR | 8hFR | 8hBR | 8hBRFR | 8hCW | 0.864 |
| 16 hours | 16hUV | 16hB | 16hG | 16hOr | 16hR | 16hFR | 16hBR | 16hBRFR | 16hCW | 1.728 |
For the first method (M1), seeds were kept in darkness for 3 days and then transferred to artificial lighting with a blue-red-far-red (BRFR) spectrum under a 16 h photoperiod, a regime commonly used for microgreens cultivation.25,26 In the second method (M2), seeds were placed under the same BRFR spectrum with a 16 h photoperiod from the first day of cultivation. This spectrum was selected because, in the first phase of the experiment, it produced seedlings with the highest dry-to-fresh biomass ratio in both species. In the third method (M3), cabbage seeds were exposed to blue light under a 16 h photoperiod for the first five days. Arugula seeds received green light under an 8 h photoperiod for the first five days. After this initial stage, both species were transferred to BRFR lighting under a 16 h photoperiod for the remainder of the cultivation period. These initial lighting conditions were selected because they produced the highest seedling emergence and the fastest cotyledon-stage development during the first experimental phase. In the fourth method (M4), seeds were exposed to the same initial spectrum used in M3, but only for the first day. On the following day, seedlings were transferred to BRFR lighting under a 16 h photoperiod for the remainder of the cultivation period. This treatment was included to examine whether a brief, 1 day exposure to the emergence-enhancing spectrum could provide an early light cue that supports the initiation of germination and subsequent seedling emergence. The light intensity applied during the initial lighting stage (prior to exposure to the growth spectrum) was 30 µmol m−2 s−1 (M3 and M4). The growth-spectrum intensity was set to 120 µmol m−2 s−1, and the total cultivation period lasted 12 days.
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At the end of each lighting treatment, the developed seedlings were placed on graph millimeter paper to measure shoot length. Measurements were taken in millimeters (mm) and included hypocotyl and cotyledon length, as well as cotyledon area, which was quantified in square millimeters (mm2). Morphological measurements were obtained from the seedlings that emerged within each container. Seedlings within the same container were treated as subsamples rather than independent biological replicates, and sample size varied among treatments according to emergence percentage.
Fresh and dry biomass of the developed shoots and cotyledons were measured for each lighting treatment. Biomass values presented in the results correspond to the seedlings that successfully emerged within each container. Seedlings were weighed using the aeAdam LTB 3602i precision scale, which has a measurement error of ±30 mg. Because some of the samples had masses close to the instrument's error margin, each sample was weighed five times to reduce random measurement variability. The mean of these repeated measurements was used as the final value for each sample. The drying of the fresh biomass was carried out in a drying oven at 80 °C for 24 h. Then the value of the dry to fresh biomass was divided to calculate the DW
:
FW biomass ratio. The same procedure was followed during the second experimental phase involving microgreens production. The mean value was calculated from the biomass measurements of the four containers per cultivation method. This mean value represents the fresh and dry biomass of the above-ground parts (shoots and cotyledons).
These calculations were conducted solely to facilitate a direct comparison among the four cultivation methods assessed in this study. In practical large-scale microgreens production, overall efficiency depends on multiple factors – including light intensity, lamp efficiency, light mounting height, cultivation area, and seeding density – which may lead to substantially different EUE and LUE values. Therefore, the values reported here should be interpreted as method-to-method comparative indicators rather than absolute measures of production efficiency.
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| Fig. 1 Seedling emergence (%) of cabbage under different lighting conditions: (a) continuous darkness (0 h photoperiod), (b) 4 h photoperiod, (c) 8 h photoperiod and (d) 16 h photoperiod. | ||
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| Fig. 2 Seedling emergence (%) of arugula under different lighting conditions: (a) continuous darkness (0 h photoperiod), (b) 4 h photoperiod, (c) 8 h photoperiod and (d) 16 h photoperiod. | ||
Under continuous darkness, only 5% of the seeds developed both a root and a shoot by the end of the 7 days period, representing the lowest emergence percentage among all treatments. A similarly low seedling emergence percentage (5%) was also observed under the 4 hR treatment. During the 4 h lighting application, seedling emergence ranged between 5% and 35%, with the 4hOr and 4hBRFR treatments showing the highest emergence (35%). Under the 8 h photoperiod, emergence varied from 10% (8hOr) to 45%, with the 8hUV treatment exhibiting the highest value. Under the 16 h photoperiod, the UV, far-red and blue-red-far-red spectrum produced the lowest emergence (10%), whereas blue light resulted in the highest percentage (55%). The remaining spectra within this photoperiod produced emergence levels ranging from 20% to 40% (Fig. 1).
Under the 4 h and 8 h photoperiods, the first seedlings emerged at 120 h, whereas under the 16 h photoperiod – specifically in the 16 hB, 16hFR, 16hBR and 16hCW treatments – the first seedlings emerged earlier, at 96 h.
In the case of arugula under dark conditions, the seedling emergence rate was 50%. The lowest seedling emergence rate among the lighting treatments was observed under the 16hCW spectrum (25%). All other light treatments resulted in emergence rates above 50%, with the highest value (85%) recorded under the 8 hG treatment. Specifically, the 4 h photoperiod produced emergence rates ranging from 65% to 75%, the 8 h photoperiod from 50% to 85%, and the 16 h photoperiod from 25% to 80% (16hFR) (Fig. 2).
Under the 4 h and 16 h photoperiods, seedling emergence was observed as early as 48 h after sowing, whereas under the 8 h photoperiod the first seedlings appeared at 72 h, similar to the dark treatment.
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| Fig. 3 Morphological characteristics of the developed (a) cabbage and (b) arugula seedlings, under the different lighting treatments. | ||
| Photoperiod | N | Mean SL ± STDEV (mm) | Mean LA ± STDEV (mm2) |
|---|---|---|---|
| a LA corresponds to cotyledon area. | |||
| Cabbage | |||
| 4 hours | 38 | 25.8 ± 9.1 | 27.4 ± 12.1 |
| 8 hours | 40 | 14.1 ± 5.0 | 21.3 ± 9.6 |
| 16 hours | 45 | 18.4 ± 3.9 | 45.2 ± 14.5 |
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| Arugula | |||
| 4 hours | 123 | 25.4 ± 7.6 | 32.4 ± 10.9 |
| 8 hours | 116 | 24.0 ± 9.1 | 41.3 ± 12.5 |
| 16 hours | 104 | 21.3 ± 7.8 | 42.2 ± 11.7 |
Differences among the applied light spectra were observed in the morphology of cabbage and arugula seedlings (Table 3). For cabbage, under the 4 h photoperiod, the greatest shoot length was recorded under the red spectrum (47 mm – only one seedling developed under this treatment), while the largest mean cotyledon area occurred under the cool-white spectrum (47.5 mm2). Under the 8 h photoperiod, the longest mean shoot length was observed under UV light (17.7 mm), although cotyledon development was minimal under this spectrum (10.7 mm2). The BR spectrum produced the largest cotyledon area within this photoperiod (34 mm2). For the 16 h photoperiod, the green spectrum resulted in the longest mean shoot length (22 mm), whereas the BRFR spectrum produced the largest cotyledon area (58 mm2), representing the highest value recorded for cabbage across all lighting treatments.
| Light treatments | Cabbage | Arugula | ||||
|---|---|---|---|---|---|---|
| N | Mean SL ± STDEV (mm) | Mean LA ± STDEV (mm2) | N | Mean SL ± STDEV (mm) | Mean LA ± STDEV (mm2) | |
| a One-way ANOVA indicated significant differences in shoot length and leaf area among light spectra within each photoperiod (P < 0.05). However, in some treatments, emergence rates were too low to yield enough seedlings for valid post hoc comparisons. LA corresponds to cotyledon area. | ||||||
| 4hUV | 6 | 31.2 ± 7.9 | 17.7 ± 6.2 | 13 | 24.6 ± 6.9 | 15.4 ± 2.5 |
| 4 hB | 2 | 27.5 ± 0.7 | 39.0 ± 1.4 | 14 | 30.3 ± 6.1 | 39.7 ± 7.9 |
| 4 hG | 5 | 13.6 ± 2.1 | 15.8 ± 5.5 | 15 | 30.9 ± 6.3 | 24.9 ± 8.3 |
| 4hOr | 7 | 20.7 ± 5.5 | 15.1 ± 4.7 | 14 | 24.7 ± 6.6 | 34.0 ± 4.3 |
| 4 hR | 1 | 47.0 ± 0.0 | 42.0 ± 0.0 | 13 | 23.4 ± 6.9 | 21.3 ± 5.9 |
| 4hFR | 5 | 31.0 ± 2.9 | 36.0 ± 3.4 | 14 | 26.6 ± 5.9 | 38.0 ± 6.8 |
| 4hBR | 3 | 37.0 ± 2.6 | 38.0 ± 4.0 | 13 | 25.2 ± 8.3 | 39.5 ± 8.6 |
| 4hBRFR | 7 | 21.1 ± 5.3 | 34.7 ± 6.2 | 14 | 21.9 ± 5.8 | 43.4 ± 5.5 |
| 4hCW | 2 | 32.0 ± 4.2 | 47.5 ± 3.5 | 13 | 19.9 ± 6.4 | 35.0 ± 5.3 |
| 8hUV | 9 | 17.7 ± 6.2 | 10.7 ± 3.9 | 12 | 31.5 ± 4.7 | 27.0 ± 2.0 |
| 8hB | 5 | 16.4 ± 3.6 | 26.4 ± 7.9 | 12 | 15.9 ± 3.4 | 49.3 ± 7.0 |
| 8hG | 3 | 7.3 ± 0.6 | 12.7 ± 5.0 | 17 | 31.4 ± 8.7 | 46.9 ± 9.1 |
| 8hOr | 2 | 17.4 ± 2.8 | 26.0 ± 5.7 | 10 | 27.7 ± 6.6 | 19.5 ± 4.4 |
| 8hR | 3 | 8.3 ± 5.1 | 27.3 ± 2.5 | 14 | 27.5 ± 7.1 | 45.1 ± 4.3 |
| 8hFR | 3 | 8.6 ± 1.5 | 25.3 ± 7.6 | 10 | 25.1 ± 7.2 | 37.1 ± 6.7 |
| 8hBR | 4 | 16.3 ± 2.9 | 34.0 ± 9.1 | 14 | 22.5 ± 7.9 | 44.7 ± 8.6 |
| 8hBRFR | 4 | 14.3 ± 1.0 | 27.0 ± 7.4 | 15 | 12.4 ± 5.0 | 53.3 ± 9.5 |
| 8hCW | 7 | 13.0 ± 1.4 | 18.6 ± 6.5 | 12 | 22.3 ± 5.9 | 42.3 ± 4.4 |
| 16hUV | 2 | 16.0 ± 2.8 | 6.0 ± 1.4 | 12 | 23.9 ± 3.4 | 37.4 ± 3.4 |
| 16hB | 11 | 15.7 ± 2.8 | 47.5 ± 9.2 | 10 | 17.7 ± 4.9 | 50.3 ± 8.9 |
| 16hG | 4 | 22.0 ± 2.4 | 21.3 ± 5.4 | 14 | 25.8 ± 5.4 | 29.5 ± 7.7 |
| 16hOr | 8 | 21.4 ± 3.4 | 27.0 ± 8.5 | 13 | 28.1 ± 2.5 | 32.0 ± 8.6 |
| 16hR | 6 | 20.3 ± 3.7 | 29.8 ± 4.5 | 11 | 21.6 ± 4.5 | 40.3 ± 6.7 |
| 16hFR | 2 | 16.5 ± 3.5 | 16.0 ± 5.7 | 16 | 23.1 ± 3.7 | 47.6 ± 5.3 |
| 16hBR | 5 | 16.8 ± 2.2 | 44.0 ± 8.9 | 11 | 12.9 ± 3.4 | 38.4 ± 3.9 |
| 16hBRFR | 2 | 14.5 ± 0.7 | 58.0 ± 2.8 | 12 | 19.8 ± 4.6 | 61.2 ± 6.7 |
| 16hCW | 5 | 19.2 ± 5.1 | 40.6 ± 7.6 | 5 | 14.4 ± 2.1 | 53.4 ± 4.7 |
For arugula, under the 4 h photoperiod, the monochromatic blue and green spectra produced the longest seedlings, with mean shoot lengths of 30.3 mm and 30.9 mm, respectively. Under the 8 h photoperiod, the UV and green spectra were associated with the greatest shoot elongation, with mean lengths of 31.5 mm and 31.4 mm, respectively. Under the 16 h photoperiod, the orange spectrum produced the longest seedlings, with a mean shoot length of 28.1 mm. Across all photoperiods, the BRFR spectrum generated seedlings with the largest cotyledon area, measuring 43.4 mm2 under the 4 h photoperiod, 53.3 mm2 under the 8 h photoperiod, and 61.2 mm2 under the 16 h photoperiod.
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Fig. 5 Ratio of dry weight to fresh weight (DW : FW) biomass of seedlings which was produced from each lighting treatment for (a) cabbage and (b) arugula seeds. | ||
The differences in biomass values are attributed not only to variations in successful germination but also to distinct biochemical activities occurring in the seedlings, modulated by the lighting conditions. This is also reflected in the differences observed in the DW
:
FW biomass ratios across treatments.
DW
:
FW ratios ranged from 0.04 under the 4 hG treatment to nearly 0.18 under the 16hBRFR treatment. The BRFR spectrum also produced the highest ratio under the 8 h photoperiod (0.17). Under the 4 h photoperiod, the highest ratio was achieved with the FR spectrum (0.09). Ratios above 0.10 were recorded for all spectra under the 16 h photoperiod except UV and Orange, whereas under the 4 h photoperiod, ratios for all spectra remained below 0.10.
Regarding the DW
:
FW biomass ratio, the lowest value was observed under the 4hUV treatment (0.06), while the highest was recorded in the 16hBRFR treatment (0.15). Across all spectra, the DW
:
FW ratio tended to be higher under the 16 h photoperiod.
:
FW ratio for the third method was the lowest among the four (0.079), whereas the ratios for the other three methods ranged narrowly between 0.092 and 0.096.
For arugula, the fourth method resulted in the highest FW and DW biomass production. A one-day exposure to the spectrum associated with the highest emergence rate yielded 12.56 g of fresh biomass and 1.54 g of dry biomass. The third method also produced higher biomass than the first two methods; however, its FW biomass was 12.98% lower than that of the fourth method, and its DW biomass was 5.2% lower. Fresh biomass in the fourth method was 22.4% higher than in the first method and 22.5% higher than in the second, while DW biomass was 22.1% and 18.2% higher, respectively. The DW
:
FW ratios of the first, second, and fourth methods were similar (0.123–0.129), whereas the third method showed a slightly higher ratio of 0.134.
For cabbage, all light treatments resulted in higher seedling emergence percentages (except 4hR) compared to those observed under dark conditions (5%). For each light spectrum tested, differences were observed among the three photoperiods: in some cases, longer exposure increased the number of emerged seedlings, whereas in others, shorter photoperiods were more favorable. Overall, when the emergence values were summed across all spectra, the 16 h photoperiod was associated with the highest total number of seedlings, exceeding both the 4 h and 8 h treatments. Among all treatments, 16 h of blue light showed the highest emergence percentage, reaching 55%. The increased number of emerged seedlings is directly associated with the fact that this treatment also yielded the highest number of germinated seeds (55%), all of which successfully developed shoots by the end of the treatment. This response is consistent with previous studies reporting that exposure to blue light can enhance germination in various species, including cabbage (Brassica oleracea) seeds and stevia (Stevia rebaudiana Bertoni) seeds.27,28
For arugula, only the 16hCW treatment (25%) produced an emergence percentage lower than that recorded under darkness (50%), while all other treatments yielded equal or higher values. The highest overall seedling output was observed under the 4 h photoperiod, where all spectra produced emergence percentages between 65% and 75%. All treatments exhibited high germination percentages (above 65% and higher than in darkness), which contributed to the elevated emergence values. However, not all germinated seeds completed shoot development within the 7 days period. Although several spectra resulted in high emergence percentages, the highest value (85%) was observed under an 8 h application of green light. Green light is not commonly used, nor widely investigated, as a germination-promoting wavelength; nevertheless, similarly positive responses have been reported by Vogel & Makedo in their study on orchid (Cyrtopodium glutiniferum Raddi) seeds.29 Opposing findings have also been documented. In their work on Vigna unguiculata (L.) Walp seeds, Lal and Sachan found that red light supported the highest germination percentage, whereas green light did not promote germination.18 Conversely, Hernandez-Adasme et al. reported that blue light enhanced germination in two lettuce cultivars (‘Levistro’ and ‘Carmoli’), while red light reduced it.19 They additionally observed that a mixed spectrum containing 52% blue, 27% green, and 21% red accelerated germination and increased the proportion of germinated seeds, resulting in a greater number of developed seedlings.
In the experiments with mixed light spectra, arugula seedlings had shorter stem lengths compared to monochromatic spectra (except 8 hB and 16 hB), across all photoperiods. For cabbage, this was observed only under the 16 h lighting application. In most experiments, mixed light spectra were associated with higher DW
:
FW biomass ratio compared to single spectrum treatments, indicating lower water content and increased solid compound concentration, which are desirable characteristics for microgreens. Similar characteristics were also observed in seedlings grown under blue light. The blue spectrum has been reported to reduce water retention, enhance the accumulation of phenolic compounds, and consequently increase dry biomass production.19,34 In the case of cabbage, far-red light produced seedlings with morphological characteristics similar to those grown under the blue spectrum, while the stem and leaves developed a reddish coloration. This reddish pigmentation is likely due to the activation and production of anthocyanins.35 The DW
:
FW ratio under far-red light was higher than blue light for the 4 and 8 h photoperiods.
Monochromatic light treatments, in both species examined, generally produced seedlings with elongated stems and reduced cotyledon expansion. Because microgreens have a very short growth cycle, light exposure strongly influences photomorphogenesis, which in turn regulates key developmental traits such as stem elongation and leaf enlargement – both directly linked to photosynthetic performance.10,36 When post-germination light conditions fail to adequately activate the relevant photoreceptors, seedlings exhibit characteristics resembling skotomorphogenesis: they prioritize stem elongation and produce small, pale leaves, a morphological strategy that enables them – under natural conditions – to reach light above the soil surface.37,38 This phenomenon was clearly reflected in the contrasting morphological patterns observed between the 4 h and 16 h photoperiods. Under the 4 h photoperiod, both cabbage and arugula developed longer shoots yet smaller cotyledons, whereas the 16 h photoperiod produced seedlings with more expanded cotyledons and shorter stems. These opposite trends demonstrate that targeted manipulation of early-stage light quality and duration can be used strategically to direct seedling morphology – either promoting shoot elongation or enhancing cotyledon growth, depending on the desired production traits. In short-cycle crops such as microgreens, improvements in seedling emergence and early morphological development directly determine the number and uniformity of plants entering the growth phase, and therefore strongly influence final yield and production efficiency.
Fig. 8 illustrates the developmental progression of cabbage (top row) and arugula (bottom row) seedlings under three representative treatments: continuous darkness, the light spectrum that produced the highest number of emerged seedlings (16 h blue light for cabbage and 8 h green light for arugula), and the 16 h blue-red-far-red spectrum, which yielded the highest DW
:
FW biomass ratio. Although growth in darkness does not represent a typical developmental pathway, it is included here because it reflects a common practice in microgreens production, where seeds are kept without light for the first days after sowing until radicle emergence and early shoot formation occur. Presenting these treatments side by side provides a visual comparison that complements the quantitative results and highlights how light quality and photoperiod shape early seedling development.
In both species, seedlings that developed under complete darkness exhibited pale, undersized cotyledons and elongated hypocotyls-morphological traits characteristic of skotomorphogenesis and the absence of photosynthetic light. When seeds are intentionally kept in darkness until the radicle and hypocotyl emerge, as is common practice in microgreens production, it is essential to monitor the timing of shoot emergence carefully so that seedlings can be transferred promptly to growth lighting, preventing excessive elongation and weakened early development.
Cabbage seedlings exposed to blue light exhibited faster shoot development, shorter and more compact hypocotyls, and larger, intensely green cotyledons. These traits reflect the strong influence of blue wavelengths on photomorphogenesis and photosynthetic activation during early seedling growth. In arugula exposed to green light, an intermediate pattern was observed: seedlings displayed moderate hypocotyl elongation – resembling a partial skotomorphogenic response – while cotyledons exhibited a more distinct green coloration and greater surface area compared to those developed in darkness. This suggests that, although green light is generally considered less effective than blue or red wavelengths in driving seedling development, it still contributes to early-stage growth to some extent. Under the mixed BRFR spectrum, both species produced seedlings with compact morphology, well-expanded cotyledons, and strong green pigmentation. These traits correspond with the higher DW
:
FW ratios measured under this treatment, indicating a more balanced allocation of biomass and improved structural development during the early seedling stage.
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FW biomass ratio, microgreens production increased in both species. This sequential lighting approach (emergence-enhancing spectrum during the first days followed by a biomass-enhancing growth spectrum) was more effective than the conventional practice of keeping seeds in darkness for several days before exposure to growth lighting. Specifically, for cabbage, the third method (M3), which included a five-day initial exposure to blue light under a 16 h photoperiod, produced the highest yield (2.189 g). The fourth method (M4), involving only one day of initial exposure to the emergence-enhancing spectrum, resulted in a yield of 1.692 g. In arugula, the green-light treatment applied for 5 days in M3 (8 h photoperiod) produced 10.93 g of microgreens, whereas M4 resulted in the highest yield (12.56 g). The conventional approach used in microgreens production (M1) – in which seeds are kept in darkness for approximately three days before exposure to growth lighting, produced lower yields in both species.25,26 For cabbage, M1 resulted in a yield approximately 34% lower than the recommended M3 and around 14% lower than M4. For arugula, the yield obtained with M1 was about 11% lower than M3 and roughly 22% lower than M4. The second method (M2), where seeds were placed under the growth spectrum from the start, also improved microgreens production compared to M1 for cabbage. For arugula, however, M1 and M2 produced similar yields.Early light exposure also helps explain why methods M3 and M4, which applied an emergence-enhancing spectrum before switching to BRFR growth lighting, resulted in higher microgreens yields compared with method M2, where seeds were placed directly under 16hBRFR. The initial wavelength–photoperiod combination used in M3 and M4 likely provided a stimulus that triggered earlier germination and supported faster early development, increasing the number of seedlings entering the growth stage and ultimately enhancing total microgreens yield.
Additionally, the large difference observed between cabbage and arugula microgreens yield is partly explained by the sowing strategy: in all treatments, seeds were distributed based on weight (0.85 g per tray). Because arugula seeds have lower individual seed mass – and also showed higher emergence capacity – each tray contained a substantially greater number of viable arugula seeds, resulting in a higher number of seedlings and, consequently, greater total microgreens production.
The reason why method M4 (1 day initial lighting) produced the highest LUE in arugula, whereas cabbage reached its highest LUE under method M3 (5 days initial lighting), is linked to species-specific differences in the speed of early development. Arugula seeds germinated within the first 24 h and initiated shoot development as early as 48–72 h, meaning that even a single day of exposure to the emergence-enhancing spectrum was sufficient for seedlings to receive the necessary stimulus before transitioning to the growth spectrum (16hBRFR). As a result, arugula entered the growth-lighting phase earlier, enabling faster biomass accumulation under method M4. In contrast, cabbage exhibited slower radicle and shoot development; therefore, a longer initial exposure period, as applied in M3, was required for the spectrum to exert its beneficial effects before seedlings were transferred to the growth phase. Consequently, cabbage achieved higher LUE under M3 rather than M4.
Additionally, because PPFD was constant across treatments, these observed differences reflect the combined effect of lighting duration and daily light integral (DLI), rather than photoperiod alone. This is particularly relevant for the UV treatment, where the substantially lower applied light intensity resulted in correspondingly lower DLI values compared to the PAR spectra, limiting direct comparability with the other lighting conditions. Therefore, the morphological characteristics observed in seedlings developed under UV radiation should be interpreted within the context of the applied irradiation level, and not as an indication of reduced biological relevance for seedling development.
Future studies would benefit from the use of a greater number of seeds per treatment and increased replication, as well as the evaluation of a broader range of environmental conditions (e.g., temperature and relative humidity), in order to more comprehensively assess the effects of light spectrum on seedling emergence and development. Further research across a wider range of light intensities and spectral combinations would contribute to refining species-specific lighting strategies and improving early seedling performance in CEAs.
These findings suggest that identifying species-specific lighting requirements is particularly important for improving production efficiency in microgreens cultivation, which is characterized by short production cycle. Optimizing early-stage lighting may contribute to shorter cultivation time and increased yield uniformity. Moreover, targeted manipulation of light quality may provide a practical means to guide seedling morphology toward desirable traits without increasing overall energy input. Together, these findings highlight the potential of stage-specific lighting strategies to support more sustainable microgreens production in urban controlled-environment systems.
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