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ISSN : 1225-5009(Print)
ISSN : 2287-772X(Online)
Flower Research Journal Vol.30 No.3 pp.82-91

Potassium Sulfate Supply under Elevated CO2 Improves Flower Development and Photosynthesis of Phalaenopsis

Ah Ram Cho, Sun Woo Chung, Yoon Jin Kim*
Department of Horticulture, Biotechnology and Landscape Architecture, Seoul Women's University, Nowon-gu, Seoul, Korea

* Corresponding author: Yoon Jin Kim Tel: +82-2-970-5614 E-mail:
05/07/2022 07/09/2022 13/09/2022


Although the effect of elevated carbon dioxide (CO2) on Phalaenopsis plant flowering, biomass, and photosynthesis has received intensive study, whether elevated CO2 affects plant requirements and sensitivity to potassium sulfate (SOP) during the reproductive growth stage remains unclear. To evaluate the combined effect of CO2 and SOP provision on crassulacean acid metabolism orchids, we cultivated Phalaenopsis Queen Beer ‘Mantefon’ under ambient and elevated CO2 treatments (≈ 400 or ≈ 720 μmol×mol-1, respectively) and four levels of SOP supply for 20 weeks after treatments (WAT): potassium and sulfate levels by 10.41 and 1.96 mmol·L-1 (SOP1), 5.98 and 0.90 mmol·L-1 (SOP2), 12.80 and 1.96 mmol·L-1 (SOP3), and 14.83 and 3.16 mmol·L-1 (SOP4), respectively. The number of floral buds and flowers decreased in the plants grown under elevated CO2 than in those grown under ambient CO2, regardless of the SOP level; however, the reduced production of floral buds and flowers did not affect the dry mass of shoot, root, and spike at 20 WAT. There were significant interactive effects of CO2 and SOP on root biomass accumulation and net CO2 uptake. The stimulation of biomass partitioning on the root, as a sink source, observed due to the uptake of elevated CO2 was improved under increased SOP supply. Under ambient CO2, the leaf critical SOP level was SOP1 for root and spike biomass accumulation. Plants grown under elevated CO2 were more sensitive to SOP treatments, with higher essential leaf levels of SOP.



    Potassium (K) is one of the major mineral nutrients impacting orchids’ growth, flower initiation, and flower development of Phalaenopsis (Wang 2007), Cymbidium (An et al. 2012), and Dendrobium (Bichsel et al. 2008). Phalaenopsis ‘Taisuco Kochdian’ fertigated with 300 mg·L-1 K had longer leaf length and spike and more flowers than those with 50 or 100 mg·L-1 K (Wang 2007). Translocation from source to sink in Cymbidium was promoted more in the plants with 100 and 200 mg·L-1 K than in the K-deficient plants at the flower initiation stage (An et al. 2012). Symptoms of the K deficiency in Phalaenopsis when fertigating 50 or 100 mg·L-1 K included rusty or bronze patches, leaf yellowing, and necrosis on leaf tips (Wang 2007). K-deficient leaves of cotton plants decreased photosynthesis to partitioning assimilates in plant tissue, resulting from low chlorophyll content and poor chloroplast ultrastructure (Zhao et al. 2001). Plants show K deficiency symptoms much earlier under limited nitrogen (N) levels (Gartner 1969). Salt stress causes nutrient deficiencies due to the competition of Na+ and Cl- (Abdullakasim et al. 2018). Still, the application of potassium sulfate (SOP) counteracted salt-induced reduction in the growth and yield of rice plants (Din et al. 2001). Foliar application of SOP significantly improved the growth, achene yield, photosynthetic and transpiration rates, stomatal conductance, and water-use efficiency of the salt-stressed sunflower plants (Akram et al. 2009).

    Elevated atmospheric CO2 enhances leaf and canopy photosynthesis and productivity in C3 crops because the present atmospheric CO2 concentration is insufficient to saturate Rubisco and elevated CO2 inhibits the competing process of photorespiration (Bowes 1991). CAM (crassulacean acid metabolism) induced Rubisco can be activated more in CAM Mesembryanthemum crystallinum than in C3M. crystallinum (Davies and Griffiths 2012). Numerous studies have demonstrated that CAM plants’ growth and yield are favored by elevated CO2 (Croonenborghs et al. 2009;Nobel and Israel 1994;Weiss et al. 2010). The cladodes of Opuntia ficus-indica were developed more when the CO2 concentration was doubled (Nobel and Israel 1994). Aechmea ‘Maya’, ornamental bromeliads, showed at both CO2 concentrations an equal biomass enhancement throughout the experimental period. Still, the total leaf area and thickness of Guzmania ‘Hilda’ increased in the plants grown under elevated CO2 than those under ambient CO2 (Croonenborghs et al. 2009). Fruit fresh mass of Selenicereus megalanthus increased by 63% in response to elevated CO2 than those exposed to ambient CO2 (Weiss et al. 2010).

    Like other CAM ornamental plants, Phalaenopsis growth, flowering, and photosynthesis respond well to elevated atmospheric CO2 concentration (Cho et al. 2019;Song et al. 2019;Yun et al. 2018). Yun et al. (2018) found that Phalaenopsis grown under 1600 μmol·mol-1 CO2 had a higher number of leaves, net CO2 uptake, transpiration rate, and stomatal conductance than plants grown under 450 μmol·mol-1 CO2 conditions. However, rapid growth and increased productivity of plants grown under elevated CO2 require more K supply. The leaf critical K level of cotton was 12 g·kg-1 under ambient CO2 for photosynthesis, main-stem elongation, and biomass accumulation, whereas 18-19 g·kg-1 under elevated CO2 of 720 μL·L-1 (Reddy and Zhao 2005). The K uptake was significantly reduced in potato tubers under elevated CO2 of 680 μL·L-1 (-6.7%) compared with ambient CO2 (3.2%) (Fangmeier et al. 2002). The photosynthesis rate under elevated CO2 of 600 ± 10 μmol·mol-1 increased by 15.6% in the plants under 200 mg·L-1 K fertilization than those under K deficiency in wheat (Asif et al. 2017). The potential yield, the sum of the number of visible floral buds and flowers, number of lateral branches, and water-use efficiency of Phalaenopsis Queen Beer ‘Mantefon’ were higher in the plants exposed to nutrient electrical conductivity (EC) of 2.0 dS·m-1 (N 98 mg·L-1, P 66 mg·L-1, K 202 mg·L-1) than those to EC 1.0 dS·m-1 (N 220 mg·L-1, P 149 mg·L-1, K 454 mg·L-1) under elevated CO2 (Cho et al. 2020).

    We hypothesize that elevated atmospheric CO2 concentration may increase the growth and development of Phalaenopsis plants to the CO2. A sufficient K supply using SOP fertilizer is needed for the synergistic effect of elevated atmospheric CO2 concentration. The objectives were to determine the interactive effects of elevated CO2 concentration and SOP supply on Phalaenopsis Queen Beer ‘Mantefon’ flowering, biomass, and photosynthesis characteristics under controlled environmental conditions.

    Materials and methods

    Plant materials and growth conditions

    Clones of 18 months after deflasking, Phalaenopsis ‘Queen Beer ‘Mantefon’ plants were purchased in June 2018 and transplanted into 120 mm pots (0.5L volume) with a medium containing a 100% dried Chilean sphagnum moss (Lonquen Ltd., Puerto Montt, Chile). The plants were grown in a Phalaenopsis nursery in Guangzhou, China, during the vegetative stage (from January 2017 to June 2018). During this period, average temperatures, relative humidity, and photosynthetic photon flux in the greenhouse were 30 ± 2°C, 70 ± 10%, and 250 ± 50 μmol·m-2·s-1, respectively.

    The experiment was conducted at the plastic greenhouse of Bulmuri Orchid Nursery in Paju, Korea (37°N, 126°E) from 8 Jun to 7 Dec 2018. Plants were cultivated at an average temperature of 23 ± 2/18 ± 1°C (day/night) and the relative humidity of 50/80%, monitored by a data logger (AM-21A, WISE Sensing Inc., Yongin, Korea). A 50% shading screen surrounded the plants in the greenhouse with an eave height of 2.5 m and a ridge height of 3.0 m. PPF of full sunlight at Bulmuri Orchid Nursery was 1,400 μmol·m-2·s-1 in June and 1,000 μmol·m-2·s-1 in December 2018.

    CO2 and potassium sulfate treatments

    Eight treatments included two levels of CO2: ≈ 400 μmol·mol-1 (Ambient CO2) and ≈ 720 μmol·mol-1 (Elevated CO2) during 22:00 - 04:00 HR (Fig. 1), which were controlled using a CO2 analyzer (SH-VT250, Soha-Tech Co. Ltd., Seoul, Republic of Korea), and four levels of potassium sulfate (SOP) supply. The CO2 treatments were imposed from spike emergence through final harvest 20 weeks after treatments (WAT). The stock solutions used for all four different SOP treatments were prepared with commercial agricultural fertilizers. The four SOP treatments were used by mixing 13.7N-0P-46K (Krista K, Yara, Oslo, Norway) 267 mg·L-1, 12N-61P-0K (Krista MAP, Yara) 733 mg·L-1, 15.5N-0P-0K (Calcinit, Yara) 466 mg·L-1, and four different concentrations of 0N-52P-43K (SOP, Yara) at four additional (SOP) concentrations: (1) SOP1 level (Control, 100 mg·L-1 K) throughout the experiment; (2) SOP2 level (300 mg·L-1 K); (3) SOP3 level (500 mg·L-1 K); and (4) SOP4 level (700 mg·L-1 K), until final harvest (20 WAT). The EC, pH, and molecular components of each SOP level are indicated in Table 1. The magnesium and phosphorus concentrations were controlled at 0.00 and 2.74 mmol·L-1. Fertigation was applied 200 mL per pot by hand-drip irrigation once a week.


    The first and second flowers spike length, the number of lateral branches, and floral buds were recorded every two weeks for 20 WAT on 11 plants in each treatment. Flower spikes, lateral branches, and floral buds with lengths of ≥ 5 mm were counted for each plant. Spike length was measured from the node at the base to the topmost. The number of flower spikes and flowers was recorded only at 20 WAT. Then, plants were divided into three parts leaves, spikes, and roots, which were dried at 80°C for seven days and weighed for biomass.

    Gas exchange was measured from 22:00 to 01:00 HR in the uppermost mature leaves at 20 WAT on three plants in each treatment. Net CO2 uptake (PN), stomatal conductance (g s), transpiration rate (t r), and water-use efficiency (WUE) were determined using a portable infrared gas analyzer (LI-6400, Li-Cor, Lincoln, NE, USA) equipped with a LED lighting system (6400-02B, Li-Cor). The gas exchange parameters were set up as 0.6 ± 0.4 kPa vapor pressure deficit and 500 mL·min-1 air flow rate. In the leaf chamber, the relative humidity ranged from 55 to 70%, and the temperature was kept at 18°C. The CO2 concentration inside the leaf chamber was maintained at approximately 400 and 720 μmol·mol-1, which equals the level in the greenhouse for ambient and elevated CO2 treatments, respectively. The value of WUE was obtained by dividing the value of PN by E (Xu et al. 2014).

    The petal color of the surface of three spots of two different petals from each plant was measured using a color meter (CR-10 Plus, Minolta Co., Osaka, Japan) and recorded using the CIE (L*, a*, and b*) from uniform color space at 20 WAT. The L* scale ranged from no reflection (L* = 0; black) to perfect diffuse reflection (L* = 100; white), the a* scale ranged from negative values for green to positive values for red, and the b* scale ranged from negative values for blue to positive values for yellow.

    Experimental design and data analysis

    The experiment was a completely randomized block design with 11 plants. Eight treatments were randomly arranged in the greenhouse during the experiment. All flowering and physiological measurements were made on 3 or 11 plants for each treatment.

    Data were statistically analyzed by the ANOVA procedures in the SAS system for Windows version 9.4 (SAS Inst. Inc., Cary, NC, USA) to determine the interactive effects of the two factors of CO2 and SOP nutrition on plant flowering, dry matter accumulation, gas exchange, and petal color. The means were compared by Duncan’s multiple range test at p < 0.05 and showed using Sigma Plot version 10.0 (Systat, Software, Inc., San Jose, CA, USA).


    Flower spike length and number of branches, floral buds, flower spikes, and flowers

    The first flower spike length was 1.08 – 1.34 and 1.08 – 1.19 times increased more in the plants grown under ambient and elevated CO2 with SOP2 than those with SOP1, respectively, from 8 to 20 WAT (Fig. 2A). The first and second flowers spike length did not differ among CO2 treatments from 8 to 20 WAT (Figs. 2A and B). The first and second flowers spike length were increased by 8.3% and 13.9% in the plants grown under elevated CO2 with SOP3 than those with SOP1 at 20 WAT, respectively. The number of lateral branches and floral buds had significant (p < 0.05) CO2 × SOP interactive effects (Figs. 2C and D). The number of lateral branches increased more in the plants grown under elevated CO2 with SOP1-3, ranging from 1.0 to 1.9, than those grown under ambient CO2, ranging from 0.5 to 0.8 at 12 WAT (Fig. 2C). The number of lateral branches was not significantly different between CO2 treatments from 16 to 20 WAT, regardless of SOP levels. The plants grown under ambient CO2 with SOP3 had more floral buds than those under ambient CO2 with SOP1 at 12 WAT. The number of floral buds increased more in the plants grown with SOP3 than those grown with SOP1 at ambient CO2 conditions, but it decreased when plants were grown under elevated CO2 conditions with SOP3 at 20 WAT (Fig. 2D).

    There were no significant differences between the number of flower spikes according to CO2 and SOP treatments (Table 2). However, the number of flowers decreased in the plants grown under elevated CO2 with SOP2-4 than those grown under ambient CO2.

    Biomass partitioning

    Elevated CO2 and SOP treatments not affected not only shoot biomass but also spike biomass, but the interactive effects of CO2 and SOP treatments on root biomass were significant (p < 0.01) (Fig. 3). Plants receiving SOP1 under ambient CO2 had a higher root and spike biomass than those grown with SOP2-4 under ambient CO2 (Figs. 3B and 3C). Elevated CO2 significantly increased biomass accumulation in roots, and the root biomass had the greatest increase in the plants grown with SOP2, while shoot and spike biomass altered least by SOP level (Fig. 3).


    Both CO2 and SOP treatments significantly impacted PN of Phalaenopsis plants (p < 0.001 and p < 0.05, respectively), and their interactive effects on PN were also detected (p < 0.05) (Fig. 4A). The stimulation of PN due to elevated CO2 increased as SOP supplied more from SOP1 to SOP3. Plants grown under elevated CO2 with SOP1-4 had a 33, 174, 82, and 36% higher PN, respectively. g s and t r did not differ between plants grown under ambient and elevated CO2 (Figs. 4B and 4C). A significant difference (p < 0.05 and p < 0.001) was detected in WUE among CO2 and SOP treatments, respectively (Fig. 4D). The plants receiving SOP3 under elevated CO2 had a higher WUE than those under ambient CO2.

    Petal color

    No significant differences existed between L* according to CO2 treatments (Table 2). However, a* decreased in the plants grown under elevated CO2 with SOP1, SOP2, and SOP4 than those under ambient CO2, indicating that the petal color was less red. High values for a* were observed in plants grown under elevated CO2 with SOP4 treatments.


    CO2 is the main component of biomass in plants, and an elevated CO2 level increases the total biomass in plants (Walia et al. 2022). The root biomass was 2 - 16% increased more in Phalaenopsis ‘Fuller’s Pink Swallow’ under elevated CO2 of 800, 1600, and 2400 μmol·mol-1 than those grown under ambient CO2 of 450 μmol·mol-1 (Kim et al. 2017). Using elevated CO2 and K conditions in cotton, biomass accumulation and portioning are promoted by higher K levels in leaves, which act synergistically with elevated CO2 (Reddy and Zhao 2005). The WUE improved at a higher K supply of 10 mM K than those at 0 mM K in Hibiscus (Egilla et al. 2005). The PN and WUE were improved more in Phalaenopsis Queen Beer ‘Mantefon’ grown under elevated CO2 of 800 μmol·mol-1 than those under ambient CO2 of 450 μmol·mol-1 at the vegetative growth stage (Yun et al. 2018). As WUE of plants affected by CO2 and SOP levels at p < 0.05 and < 0.001, respectively (Fig. 4), the elevated CO2 with SOP2 and SOP3 used here increased root biomass in Phalaenopsis Queen Beer ‘Mantefon’ at the reproductive growth stage.

    Elevated CO2 increases PN in CAM plants because higher CO2 increases both Rubisco-mediated daytime CO2 uptake and phosphoenolpyruvate carboxylase (PEPCase)-mediated nighttime CO2 uptake, nocturnal malate accumulation, and carbon assimilated into carbohydrates for plant growth and development (Drennan and Nobel 2000). Elevated CO2 of 700 – 800 μmol·mol-1 induces floral transition with enhanced sugar through increased photosynthesis in ornamental flowering plants: 6 days in Gailardia pulchella, 7 days in Lotus corniculate and Phytolacca americana, and 20 days in Phalaenopsis at a light level of 260 μmol·m-2·s-1 (Carter et al. 1997;Cho et al. 2019;He and Bazzaz 2003;Reekie and Bazzaz 1991). Excessive sugars delay the onset of flowering and control the flowering mechanism (Springer and Ward 2007). The flowering was delayed by 8 to 10 days in the plants on media containing 5% sucrose compared with those on media containing 1% sucrose in Arabidopsis (Ohto et al. 2001). In this study, Phalaenopsis had a higher PN under elevated CO2, which might have influenced the flowering signals of plants and delayed flowering time. Excessive sugars may delay flowering time through 20 weeks of CO2 treatments (Cho et al. 2019), but the flowering time results through 16 weeks of CO2 treatments differed from previously reported. The photosynthetic rate, total carbohydrate, and foliar starch content of pigeon pea increased under elevated CO2 of 550 μmol·mol-1 and delayed flowering by 8 to 9 days but had no adverse effect on final yield (Sreeharsha et al. 2015). The number of flowers was 8.7, 7.3, 2.2, and 3.3 more under elevated CO2 than those under ambient CO2 with SOP1 to SOP4, respectively (data not shown), which indicated that Phalaenopsis Queen Beer ‘Mantefon’ also had no adverse effects on final production.

    Plants mobilized the excessive soluble sugars to potential sinks of roots at reproductive stages. The enhancement of plant root biomass by elevated CO2 was related to higher leaf photosynthetic capacity (Figs. 3 and 4). Besides representing a greater sink strength, the higher root biomass under elevated CO2 in Phalaenopsis could lead to greater nutrient uptake, enhancing PN. Elevated CO2 increases the number of fine and lateral roots and subapical root hair development (Li et al. 2012). The changes in root morphology may enlarge the root biomass and thus aid the nutrient and water uptake. K uptake from the soil under elevated CO2 may be reduced due to decreased t r or ameliorate the effects of drought stress by reducing the stomatal resistance (Asif et al. 2018). S is mainly bounded to carbon to form organic molecules along with N and P in plants (Li et al. 2020). While K decreased with S in shoots, the changes in K and S contents in Chinese cabbage were correlated with linear regression (Reich et al. 2016). This can explain why elevated CO2 with SOP2 and SOP3 triggered a significant increase in plant root biomass and PN. The increased WUE in elevated CO2 with SOP3 is attributed to the higher net photosynthesis compared with ambient CO2, underscoring its potential for higher productivity of Phalaenopsis.

    We detected a lower a* value in elevated CO2 than in ambient CO2. The a* of strawberry under the elevated CO2 treatment decreased by 20% compared to those under the ambient air during the storage due to lower anthocyanin concentration and down-regulated phenylpropanoid and flavonoid biosynthesis under CO2 stress (Li et al. 2019). Also, elevated CO2 reduced a* of petal color in the leaves of pigmented lettuce ‘Oat leaf’ to a greater extent in saline conditions than in non-salinity conditions (Pérez-López et al. 2013); this scenario would explain the lowest a* of Phalaenopsis Queen Beer ‘Mantefon’ under elevated CO2 with SOP4. However, the anthocyanins in strawberry fruit accumulated dramatically after removing CO2 stress, suggesting that elevated CO2 didn’t have a residual effect on anthocyanins synthesis if CO2 elevation was removed (Li et al. 2019). Since long-term exposure to elevated CO2 concentrations results in reduced flower numbers and increased floral bud abortion, Phalaenopsis subjected to elevated CO2 should display before flower spike induction (Kim et al. 2017). Therefore, Phalaenopsis Queen Beer ‘Mantefon’ flowers are sensitive to elevated CO2, but the redness of the petal color itself could be restored after appropriate control of CO2 supply term.

    In conclusion, elevated CO2 levels stimulated Phalaenopsis root biomass production with a nutrient range from SOP2 to SOP3 but did not affect flower spike development in this study. As a consequence of higher PN and WUE, there were increases in root biomass production which acted as a sink organ. Plants grown under elevated CO2 conditions required greater amounts of SOP2 to SOP3, consisting of 300 to 500 mg·L-1 K, than those under ambient CO2. Consequently, nutrients ranging from SOP2 to SOP3 should be supplied for Phalaenopsis Queen Beer ‘Mantefon’ cultivation to improve the photosynthesis during the reproductive growth stage under elevated CO2 condition of 720 μmol·mol-1.


    This study was carried out with the grants (NRF− 2018R1A2B6007834) from National Research Foundation of Korea and research funds of Seoul Women’s University, Korea (2022-0126).



    CO2 concentrations in a commercial greenhouse for Phalaenopsis Queen Beer ‘Mantefon’ with ambient and elevated CO2 over 20 weeks of 2 weeks intervals from June 8 to November 24, 2018 (150 days). Error bars represent standard deviation of the mean (n = 10).


    Changes in first (A) and second flower spike length (B), and the number of lateral branches (C) and floral buds (D) of Phalaenopsis Queen Beer ‘Mantefon’ grown under ≈ 400 (Ambient CO2) and ≈ 720 μmol·mol-1 (Elevated CO2) as affected by SOP supply. Each data point and error bars represent the mean and standard error of 11 plants. The different letters represent statistically significant differences as determined by the results of Duncan’s multiple range test (p < 0.05). F and P (NS = not significant, *p < 0.05, **p < 0.01, and ***p < 0.001) values of the two-way ANOVAs at 20 weeks of treatments are presented, C: CO2 treatments, S: SOP treatments, and C × S: CO2 × SOP treatments interaction. The lateral branch and floral bud first appeared at 12 weeks after treatments.


    The shoot (A), root (B), and spike biomass (C) partitioning of Phalaenopsis Queen Beer ‘Mantefon’ grown under under ≈ 400 (Ambient CO2) and ≈ 720 μmol·mol-1 (Elevated CO2) as affected by SOP supply at 20 weeks after treatments. Each data point and error bars represent the mean and standard error of three plants. The different letters represent statistically significant differences as determined by the results of Duncan’s multiple range test (p < 0.05). F and P (NS = not significant, *p < 0.05, **p < 0.01, and ***p < 0.001) values of the two-way ANOVAs are presented, C: CO2 treatments, S: SOP treatments, C × S: CO2 × SOP treatments interaction.


    The net CO2 uptake (PN), stomatal conductance (g s), transpiration rate (t r), and water-use efficiency (WUE) of Phalaenopsis Queen Beer ‘Mantefon’ grown under under ≈ 400 (Ambient CO2) and ≈ 720 μmol·mol-1 (Elevated CO2) as affected by SOP supply at 20 weeks after treatments. Each data point and error bars represent the mean and standard error of three plants. The different letters represent statistically significant differences as determined by the results of Duncan’s multiple range test (p < 0.05). F and p (NS = non-significant, *p < 0.05, and ***p < 0.001) values of the two-way ANOVAs are presented, C: CO2 treatments, S: SOP treatments, and C × S: CO2 × SOP treatments interaction.


    Electrical conductivity (EC), pH, the concentrations of nitrate (NO3), ammonium (NH4), and calcium (Ca), phosphate (P), potassium (K), and sulfur (S) supplied by potassium sulfate (SOP).

    Effects of CO2 levels with potassium sulfate (SOP) supply on the number of flower spikes and flowers and external petal color of Phalaenopsis Queen Beer ‘Mantefon’ at 20 weeks after treatments.


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    2. Journal Abbreviation : 'Flower Res. J.'
      Frequency : Quarterly
      Doi Prefix : 10.11623/frj.
      ISSN : 1225-5009 (Print) / 2287-772X (Online)
      Year of Launching : 1991
      Publisher : The Korean Society for Floricultural Science
      Indexed/Tracked/Covered By :

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    4. Template DOWNLOAD

      국문 영문 품종 리뷰
    5. 논문유사도검사

    6. KSFS

      Korean Society for
      Floricultural Science

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      Flower Research Journal

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