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Photosynthetic Responses of Swiss Chard, Kale, and Spinach Cultivars to Irradiance and Carbon Dioxide Concentration

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Photosynthetic Responses of Swiss Chard, Kale, and Spinach Cultivars to Irradiance and Carbon Dioxide Concentration HORTSCIENCE 52(5):706–712. 2017. doi: 10.21273/HORTSCI11799-17 of reduced or increased irradiance or CO2 on yield. We also desired to determine Photosynthetic Responses of Swiss whether leafy green varieties differed in the Pn responses to irradiance and CO2 and whether some varieties were more suited to Chard, Kale, and Spinach Cultivars supplemental, or reduced, irradiance and CO2 levels. to Irradiance and Carbon Dioxide We acknowledge that translating instan- taneous Pn measurements on a per-unit-area basis to whole-plant photosynthesis has lim- Concentration itations (see Discussion). Yet, an initial com- John Erwin1,3 and Esther Gesick2 parative study exploring the variation in the Department of Horticultural Science, University of Minnesota, 305 Alderman instantaneous Pn on a per-unit-area basis is valuable in that it provides some insight into Hall, 1970 Folwell Avenue, St. Paul, MN 55108 the degree of variation among species and vari- eties. These data also provide some guidance on Additional index words. Brassica, Beta, Spinacea, light saturation point, CO2 saturation point, respiration irradiance and CO2 levels that maximize the Pn on the uppermost leaves, especially when plants Abstract. The impact of irradiance (0–1200 mmol·mL2·sL1) and carbon dioxide concen- are young and interior leaf shading is limited. tration (CO2; 50–1200 ppm) on kale (Brassica oleracea and B. napus pabularia; three cultivars), Swiss chard (chard, Beta vulgaris; four cultivars), and spinach (Spinacea Materials and Methods oleracea; three cultivars) photosynthetic rate (Pn; per area basis) was determined to facilitate maximizing yield in controlled environment production. Spinach, chard, and Chard, [‘Rhubarb’, ‘Fordhook Giant’, L2 L1 kale maximum Pn were 23.8, 20.3, and 18.2 mmol CO2·m ·s fixed, respectively, across ‘Bright Yellow’, and ‘Bright Lights’ (red)], varieties (400 ppm CO2). Spinach and kale had the highest and lowest light compensation kale [‘Toscano’, ‘Winterbor’, and ‘Red points [LCPs (73 and 13 mmol·mL2·sL1, respectively)] across varieties. The light Russian’ (B. napus pabularia)], and spinach saturation points (LSPs) for chard and kale were similar at 884–978 mmol·mL2·sL1, (‘Melody’, ‘Harmony’, and ‘Bloomsdale but for spinach, the LSP was higher at 1238 mmol·mL2·sL1. Dark respiration was lowest LS’) seeds were sown in 10.5-cm-diameter L2 L1 on kale and highest on spinach (L0.83 and L5.00 mmol CO2·m ·s , respectively). The plastic pots (5 seeds/pot) in premoistened spinach CO2 compensation point (CCP) was lower (56 ppm) than the chard or kale CCP LC-8 soilless growing media (Sun Gro Horti- (64–65 ppm). Among varieties, ‘Red Russian’ kale Pn saturated at the lowest CO2 culture, Bellevue, WA) and placed in a green- concentration (858 ppm), and ‘Bright Lights’ chard saturated at the highest (1266 ppm; house (24 ± 2 °C day and 16 ± 2 °C night L2 L1 300 mmol·m ·s ). Spinach Pn was more responsive to increasing irradiance than to temperatures; St. Paul, MN). Kale and chard CO2. Kale Pn was more responsive to increasing CO2 than to irradiance, and chard Pn seeds were obtained from Johnny’s Selected was equally responsive to increasing CO2 or irradiance. Implications and limitations of Seeds (Winslow, ME), and spinach seeds this work when ‘‘upscaling’’ to whole-plant responses are discussed. were obtained from W. Atlee Burpee & Co. (Warminster, PA). Seeds germinated in 4–7 d. Greenhouse daylight (0800–1400 HR) was Leafy green vegetable options are in- photosynthesis increases (Bjorkman,€ 1981; supplemented with 75 mmol·m–2·s–1 high- creasing as communities become more eth- Chagvardieff et al., 1994; Dorais, 2003). The pressure sodium lighting when daylight (at nically or racially diverse or both, as the primary inputs into the photosynthetic pro- plant level) was below 200 mmol·m–2·s–1. health and nutritional benefits of greens cess are light (irradiance), CO2, and water After 7 d, the three most uniform (similar consumption are reported (Bertoia et al., (Bjorkman,€ 1981). Therefore, maximizing size) seedlings were left to grow, whereas the 2015; Hu and Rimm, 2015), and as interest photosynthesis in leafy greens to maximize others were removed. As kale and chard flower after they unfold a specific leaf num- in year-round locally produced foods in- yield would require that irradiance, CO2, creases (Feldmann and Hamm, 2015). Three or water not be limited (Fu et al., 2017; ber, after a cool temperature exposure, or increasingly popular leafy vegetables are kale Gaudreau et al., 1994; Gent, 2016). both, daylength was extended with 75 m –2 –1 (Brassica oleracea and B. napus pabularia), In northern climates, year-round leafy mol·m ·s from 1600–2200 HR as flower- spinach (Spinacea oleracea), and Swiss chard green production requires protected cultiva- ing was not a concern [16 h photoperiod; (chard, Beta vulgaris). tion during the late fall, winter, and early mean daily light integral (DLI) = 12.4 · –2· –1 Kale, spinach, and chard leaves are har- spring when temperatures drop below mol m d ] to simulate a typical production vested and sold on a fresh-weight basis. The freezing. Irradiance and CO in protected environment to maximize yield (J. Erwin, 2 personal observation). In contrast, as long ability of plants to increase fresh weight, cultivation often vary, intentionally and days can promote flowering on spinach early or mass, is associated with photosynthesis unintentionally, depending on covering type, in development, spinach seedlings were where plant mass generally increases as plant spacing, degree of ventilation, whether grown under short days (8 h photoperiod; air is circulated, and whether supplemental opaque cloth pulled over plants from 1400– lighting or CO2 are supplied (Kretchen and · –2· –1 Received for publication 20 Jan. 2017. Accepted 0800 HR daily; mean DLI = 10.7 mol m d ) Howlett, 1970). Little work has been con- to inhibit flowering. After 30 d, plants were for publication 5 Apr. 2017. ducted on the effects of irradiance and CO The authors acknowledge and appreciate the 2 transplanted into 7.6-l plastic pots. Through- financial support of the Minnesota Agriculture on the Pn of leafy greens other than lettuce out, plants were watered as needed with Experiment Station, USDA-ARS Floriculture and (Lactuca sativa; Dorais, 2003; Fu et al., 2017; irrigation water containing 250 ppm N from Nursery Research Initiative (FNRI), National In- Gaudreau et al., 1994) and recent work by 15N–0P–15K fertilizer (Peter’s Dark Weather stitute of Food and Agriculture (NIFA), and mem- Gent (2016) on spinach. An understanding of Feed; The Scotts Co., Marysville, OH). All bers of the University of Minnesota through the how irradiance and CO2 impact the Pn of such plants were watered at the same time to ensure Floriculture Research Alliance including Altman greens would facilitate maximizing the Pn to similar media nutritional status (confirmed Plants, Inc., Rocket Farms, Inc., Smith Gardens, Inc., maximize yield in protected cultivation. The and Green Circle Growers, Inc. with soil tests) among species and varieties. 1Professor. objective of the research presented here was to Photosynthetic rate determination. After 2Research Fellow. determine the Pn responses of spinach, kale, kale plants unfolded seven true leaves (>45° 3Corresponding author. E-mail: erwin001@umn. and chard to irradiance and CO2 and to inform angle from the stem), chard plants unfolded edu. producers of the advantages or disadvantages four leaves, and spinach plants unfolded eight 706 HORTSCIENCE VOL. 52(5) MAY 2017 –2 –1 leaves (4 weeks across species), the impact (irradiance when estimated Pn =0mmol·m ·s Results of irradiance and CO2 on instantaneous Pn on CO2 fixed), and ‘‘k’’ was a constant that a per-unit-leaf-area basis was determined on represented the ratio of the quantum yield Photosynthetic responses to irradiance. the second leaf below the uppermost fully (q) to the Pn at the LCP (Marino et al., 2010). Pmax (identified by providing saturating irradi- expanded unfolded leaf on five plants of each Rd (dark respiration) was calculated as ance levels at 400 ppm CO2) differed among species and variety. The Pn was measured the estimated Pn (I) when irradiance was species, varieties within a species, and across using a LI-COR LI6400XT portable photo- 0 mmol·m–2·s–1. all varieties (Table 2). Spinach had the highest m –2 –1 synthesis meter (LI-COR, Inc., Lincoln, NE) Pmax at 23.8 mol CO2·m ·s fixed, whereas ÀkCðÞÀC 2 ðÞ À 0 m –2 –1 using a cuvette (6 cm ) with a built-in vari- Pn C = Pmax 1 e Eq. [2] for kale it was 20.3 mol CO2·m ·s fixed and m · –2· –1 able LED light source. The Pn at 0, 100, 200, for chard it was 18.2 mol CO2 m s fixed 400, 600, 800, 1000, and 1200 mmol·m–2·s–1 Eq. [2] shows Pn responses [(Pn (C)] to (across varieties; Fig. 1; Table 2). Among kale irradiance was determined. The Pn at 50, 200, CO2. Pn responses to increasing CO2 were varieties, ‘Red Russian’ and ‘Toscano’ had m · –2· –1 400, 600, 800, 1000, and 1200 ppm CO2 was asymptotic; ‘‘Pmax’’ in Eq. [2] estimates the ahigherPmax (21.0–22.3 mol CO2 m s ) m · –2· –1 also determined. The Pn was recorded 5 min asymptote. ‘‘C0’’ is the estimated CCP (CO2 than ‘Winterbor’ (17.4 mol CO2 m s ; m · –2· –1 after a change in irradiance or CO2 after the when Pn =0 mol m s CO2 fixed), and Table 2). Among spinach varieties, ‘Melody’ m · –2· –1 Pn had stabilized. Throughout, cuvette tem- ‘‘k’’ is a constant. The CO2 concentration at had a higher Pmax (26.6 mol CO2 m s ) perature was maintained at 24 °C, and the 95% of Pmax approximated the CO2 satura- than ‘Harmony’ or ‘Bloomsdale LS’ (22.3– –1 m –2 –1 atmospheric flow rate was 400 mL·min .
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