Side Lighting Enhances Morphophysiology by Inducing More Branching and Flowering in Chrysanthemum Grown in Controlled Environment.

Light is one of the most important factors that influence plant growth and development. This study was conducted to examine how lighting direction affects plant morphophysiology by investigating plant growth parameters, leaf anatomy, epidermal cell elongation, stomatal properties, chloroplast arrangement, and physiological changes. In closed-type plant factory units, the rooted cuttings of two chrysanthemum (Chrysanthemum morifolium Ramat.) cultivars, 'Gaya Glory' and 'Pearl Egg', were subjected to a 10 h photoperiod with a 300 µmol∙m-2∙s-1 photosynthetic photon flux density (PPFD) provided by light-emitting diodes (LEDs) from three directions relative to the plant including the top, side, and bottom. Compared to the top or bottom lighting, the side lighting greatly enhanced the plant growth, improved the leaf internal structure and chloroplast arrangement, induced small stomata with a higher density, and promoted stomatal opening, which is associated with an increased stomatal conductance and photosynthetic efficiency. It is worth noting that the side lighting significantly enhanced the induction of branching and flowering for both cultivars., The plants grown with side lighting consistently exhibited the greatest physiological performance. We conclude that the lighting direction had a profound effect on the morphophysiological characteristics of chrysanthemum, and that side lighting dramatically promoted their growth and development, especially in their branching and flowering.

Induction of novel inflorescence traits in Chrysanthemum through 60Co gamma irradiation.

PURPOSE: The novelty in flower color or inflorescence form is recognized as a valuable trait in Chrysanthemum - a potential commercial flower crop with significant worth in global cut flower trade. This study was conducted to irradiate white and orange flowered cultivars of Chrysanthemum with an objective to identify and isolate desirable types representing novelty in flower color and inflorescence form from the irradiated populations. The terminal rooted cuttings of Chrysanthemum exposed to γ-irradiation at 10 or 15 Gy doses were found effective for inducing novel flower color variants in cultivars Thiching Queen and Purnima. The mutant progeny evolved with novel inflorescence traits of these cultivars will enrich the existing germplasm of Chrysanthemum for further utilization in breeding programs. MATERIALS AND METHODS: Two standard type Chrysanthemum cultivars, Thiching Queen and Purnima were exposed to varied doses of γ-rays (0, 5, 10, 15, and 20 Gy) using Cobalt 60 (60Co) as irradiation source for treating rooted cuttings. The irradiated mutant population was evaluated for likely variation in various vegetative and flowering characters compared to non-irradiated (control) plants. RESULTS: In Chrysanthemum cultivars Thiching Queen, seven and 'in Purnima', two flower color variants were isolated from the irradiated populations that were reportedly novel in color and desirable for commercial aspect. The leaf abnormalities were observed in mutant populations exhibiting variation in flower color, shape, and size of leaves. Certain floral abnormalities were also observed in inflorescence that reportedly progressed with increase in dosage of γ-rays irradiation. CONCLUSIONS: This study developed a gamma ray (60Co) induced mutagenesis protocol with potential application to develop novel and desirable mutants in Chrysanthemum.

Spectral quality of monochromatic LED affects photosynthetic acclimation to high-intensity sunlight of Chrysanthemum and Spathiphyllum.

Vertical farming using light-emitting diode offers potential for the early production phase (few weeks) of young ornamental plants. However, once transferred to the greenhouse, the photosynthetic acclimation of these young plants might depend on this initial light regime. To obtain insight about this acclimatization, Chrysanthemum (sun species) and Spathiphyllum (shade species) were preconditioned in growth chambers for 4 weeks under four light qualities: blue (B), red (R), red/blue (RB, 60% R) and white (W) at 100 µmol m-2 s-1 . Monochromatic light (R and B) limited leaf development of both species, which resulted in a lower leaf mass per area when compared to multispectral light (RB for Chrysanthemum, RB and W for Spathiphyllum). R-developed leaves had a lower photosynthetic efficiency in both species. After the light quality pretreatment, plants were transferred to the greenhouse with high-intensity natural light conditions. On the first day of transfer, R and B preconditioned leaves of both species had an inhibited photosynthesis. After 1 week in natural light condition, rapid light curve parameters of Chrysanthemum leaves that developed under B acclimated to sunlight had a similar level than RB-developed leaves unlike R-leaves. Spathiphyllum leaves showed a decrease in maximum electron transport rate and this was most pronounced for the R pretreatment. After 1 month, R-preconditioned Chrysanthemum had the lowest dry mass, while no effects on the dry weight of Spathiphyllum with respect to the pretreatments were observed. Light quality during preconditioning affected the leaf ability to acclimate to natural high light intensities in greenhouse environment.

Adventitious rooting of Chrysanthemum is stimulated by a low red:far-red ratio.

Adventitious rooting, a critical process in the vegetative propagation of many ornamentals, can be affected by both light intensity and light quality. We investigated the use of spectral light quality to improve adventitious rooting of Chrysanthemum morifolium cuttings by applying different combinations of blue, red and far-red light. Additionally, unrooted cuttings were treated before planting with two auxin transport inhibitors (TIBA and NPA) to study the effect of light quality on auxin biosynthesis and/or transport. Results showed that lowering the R:FR ratio (decreasing the phytochrome photostationary state, PSS) improved rooting significantly and decreased the inhibiting effect of the auxin transport inhibitor NPA. An extra decrease of PSS by adding blue light to a red + far-red spectrum further enhanced rooting. In contrast, adding blue light to solely red light decreased rooting, an effect which was more pronounced in combination with the auxin transport inhibitors TIBA and NPA. Our results show that phytochrome plays a role in adventitious root formation through the action of auxin, but that also blue light receptors interact in this process.

Regulation of flowering by green light depends on its photon flux density and involves cryptochromes.

Photoperiodic lighting can promote flowering of long-day plants (LDPs) and inhibit flowering of short-day plants (SDPs). Red (R) and far-red (FR) light regulate flowering through phytochromes, whereas blue light does so primarily through cryptochromes. In contrast, the role of green light in photoperiodic regulation of flowering has been inconsistent in previous studies. We grew four LDP species (two petunia cultivars, ageratum, snapdragon and Arabidopsis) and two SDP species (three chrysanthemum cultivars and marigold) in a greenhouse under truncated 9-h short days with or without 7-h day-extension lighting from green light (peak = 521 nm) at 0, 2, 13 or 25 µmol m-2  s-1 or R + white (W) + FR light at 2 µmol m-2  s-1 . Increasing the green photon flux density from 0 to 25 µmol m-2  s-1 accelerated flowering of all LDPs and delayed flowering of all SDPs. Petunia flowered similarly fast under R + W + FR light and moderate green light but was shorter and developed more branches under green light. To be as effective as R + W + FR light, saturation green photon flux densities were 2 µmol m-2  s-1 for LDP ageratum and SDP marigold and 13 µmol m-2  s-1 for LDP petunia. Snapdragon was the least sensitive to green light. In Arabidopsis, cryptochrome 2 mediated promotion of flowering under moderate green light, whereas both phytochrome B and cryptochrome 2 mediated that under R + W + FR light. We conclude that 7-h day-extension lighting from green light-emitting diodes can control flowering of photoperiodic ornamentals and that in Arabidopsis, cryptochrome 2 mediates promotion of flowering under green light.

Red to Far-Red Light Ratio Modulates Hormonal and Genetic Control of Axillary bud Outgrowth in Chrysanthemum (Dendranthema grandiflorum 'Jinba').

Single-flower cut Chrysanthemum (Dendranthema grandiflorum 'Jinba') holds a unique status in global floriculture industry. However, the extensive axillary bud outgrowth presents a major drawback. Shade is an environment cue that inhibits shoot branching. Present study was aimed at investigating the effect of ratio of red to far-red light (R:FR) in regulating the lateral bud outgrowth of Chrysanthemum and the detailed mechanism. Results showed that the fate of axillary buds at specific positions in stem exhibited difference in response to R:FR. Decreasing R:FR resulted in elevation of abscisic acid (ABA) accumulation in axillary buds. Expression of ABA, indole-3-acetic acid (IAA) and strigolactones (SL) -related metabolism and signal transduction genes was significantly changed in response to low R:FR. In addition, low R:FR caused the re-distribution of sucrose across the whole plant, driving more sucrose towards bottom buds. Our results indicate that low R:FR not always inhibits bud outgrowth, rather its influence depends on the bud position in the stem. ABA, SL and auxin pathways were involved in the process. Interestingly, sucrose also appears to be involved in the process which is necessary to pay attention in the further studies. The present study also lays the foundation for developing methods to regulate axillary bud outgrowth in Chrysanthemum.

Chrysanthemum morphology, photosynthetic efficiency and antioxidant capacity are differentially modified by light quality.

The effect of light quality on leaf morphology, photosynthetic efficiency and antioxidant capacity of leaves that fully developed under a specific spectrum was investigated in Chrysanthemum cv. Four light treatments were applied at 100µmolm-2s-1 and a photoperiod of 14h using light-emitting diodes, which were 100% red (R), 100% blue (B), 75% red with 25% blue (RB) and white (W), respectively. Intraspecific variation was investigated by studying the response of eight cultivars. Overall, red light significantly decreased the leaf area while the thinnest leaves were observed for W. Chlorophyll content and Chl a/b ratio was highest for W and lowest under R. B and RB resulted in the highest maximum quantum yield (Fv/Fm) and quantum efficiency (ΦPSII). A negative correlation between heat dissipation (NPQ) and ΦPSII was found. Blue light induced the highest hydrogen peroxide content, which is a proxy for total ROS generation, followed by W and RB while low contents were found under R. The antioxidative response was not always correlated with hydrogen content and differed depending on the light quality treatment. Blue light enhanced the proline levels, while carotenoids, total flavonoid and phenolic compounds were higher under W. Intraspecific variation in the responses were observed for most parameters with exception of leaf thickness; this intraspecific variation was most pronounced for total phenolic and flavonoid compounds.

Light quality and quantity regulate aerobic methane emissions from plants.

Studies have been mounting in support of the finding that plants release aerobic methane (CH4 ), and that these emissions are increased by both short-term and long-term environmental stress. It remains unknown whether or not they are affected by variation in light quantity and quality, whether emissions change over time, and whether they are influenced by physiological parameters. Light is the primary energy source of plants, and therefore an important regulator of plant growth and development. Both shade-intolerant sunflower and shade-tolerant chrysanthemum were investigated for the release of aerobic CH4 emissions, using either low or high light intensity, and varying light quality, including control, low or normal red:far-red ratio (R:FR), and low or high levels of blue, to discern the relationship between light and CH4 emissions. It was found that low levels of light act as an environmental stress, facilitating CH4 release from both species. R:FR and blue lights increased emissions under low light, but the results varied with species, providing evidence that both light quantity and quality regulate CH4 emissions. Emission rates of 6.79-41.13 ng g-1 DW h-1 and 18.53-180.25 ng g-1 DW h-1 were observed for sunflower and chrysanthemum, respectively. Moreover, emissions decreased with age as plants acclimated to environmental conditions. Since effects were similar in both species, there may be a common trend among a number of shade-tolerant and shade-intolerant species. Light quantity and quality are influenced by factors including cloud covering, so it is important to know how plants will be affected in the context of aerobic CH4 emissions.

Comparative analyses of light-induced anthocyanin accumulation and gene expression between the ray florets and leaves in chrysanthemum.

Light is one of the key environmental factors that affect anthocyanin biosynthesis. However, the underlying molecular mechanism remains unclear, and many problems regarding phenotypic change and corresponding gene regulation have not been solved. In the present study, comparative analyses of light-induced anthocyanin accumulation and gene expression between the ray florets and leaves were performed in Chrysanthemum × morifolium 'Purple Reagan'. After contrasting the variations in the flower color phenotype and relative pigment content, as well as expression patterns of structural and regulator genes responsible for anthocyanin biosynthesis and photoreceptor between different plant organs under light and dark conditions, we concluded that (1) both the capitulum and foliage are key organs responding to light for chrysanthemum coloration; (2) compared with flavones, shading makes a greater decrease on the anthocyanins accumulation; (3) most of the structural and regulatory genes in the light-induced anthocyanin pathway specifically express in the ray florets; and (4) CmCHS, CmF3H, CmF3'H, CmANS, CmDFR, Cm3GT, CmMYB5-1, CmMYB6, CmMYB7-1, CmbHLH24, CmCOP1 and CmHY5 are key genes for light-induced anthocyanin biosynthesis in chrysanthemum ray florets, while on the transcriptional level, the expressions of CmPHYA, CmPHYB, CmCRY1a, CmCRY1b and CmCRY2 are insignificantly changed. Moreover, the inferred comprehensive effect of multiple signals on the accumulation of anthocyanins and transmission channel of light signal that exist between the leaves and ray florets were further discussed. These results further our understanding of the relationship between the gene expression and light-induced anthocyanin biosynthesis, and lay foundations for the promotion of the molecular breeding of novel flower colors in chrysanthemums.

Physiological controls of chrysanthemum DgD27 gene expression in regulation of shoot branching.

KEY MESSAGE: DgD27 was cloned from D. grandiflorum for the first time and played an important role in shoot branching of chrysanthemum. Shoot branching plays an important role in determining plant architecture. D27 was previously proven to be involved in the strigolactone biosynthetic pathway in rice, Arabidopsis, and Medicago. To investigate the role of D27 in shoot branching of chrysanthemum, we isolated the D27 homolog DgD27. Functional analysis showed that DgD27 was a plastid-localized protein that restored the phenotype of Arabidopsis d27-1. Gene expression analysis revealed that DgD27 was expressed at the highest levels in stem, and was up-regulated by exogenous auxin. Decapitation could down-regulate DgD27 expression, but this effect could be restored by exogenous auxin. DgD27 expression was significantly down-regulated by dark treatment in axillary buds. In addition, DgD27 transcripts produced rapid responses in shoots and roots under conditions of phosphate absence, but only mild variation in responses in buds, stems, and roots with low nitrogen treatment. DgBRC1 transcripts also showed the same response in buds under low nitrogen conditions. Under phosphate deficiency, indole-3-acetic acid (IAA) levels increased, zeatin riboside levels decreased, and abscisic acid (ABA) levels increased in the shoot, while both IAA and ABA levels increased in the shoot under low nitrogen treatments. Gibberellin acid levels were unaffected by phosphate deficiency and low nitrogen treatments. Taken together, these results demonstrated the diverse roles of DgD27 in response to physiological controls in chrysanthemum shoot branching.

Morphogenesis, Flowering, and Gene Expression of Dendranthema grandiflorum in Response to Shift in Light Quality of Night Interruption.

The impact of shifts in the spectral quality of light on morphogenesis, flowering, and photoperiodic gene expression during exposure to light quality of night interruption (NI) was investigated in Dendranthema grandiflorum. The circadian rhythms of plants grown in a closed walk-in growth chamber were interrupted at night for a total of 4 h, using light-emitting diodes with an intensity of 10 µmol·m⁻²·s⁻¹ PPF. The light quality of the NI was shifted from one wavelength to another after the first 2 h. Light treatments consisting of all possible pairings of blue (B), red (R), far-red (Fr), and white (W) light were tested. Plants in the NI treatment groups exposed to Fr light grew larger than plants in other treatment groups. Of plants in NI treatment groups, those in the NI-WB treatment grew the least. In addition, the impact of shifts in the light quality of NI on leaf expansion was greater in treatment groups exposed to a combination of either B and R or R and W light, regardless of their order of supply. Flowering was observed in the NI-RB, NI-FrR, NI-BFr, NI-FrB, NI-WB, NI-FrW, NI-WFr, NI-WR, and SD (short-day) treatments, and was especially promoted in the NI-BFr and NI-FrB treatments. In a combined shift treatment of B and R or B and W light, the NI concluded with B light (NI-RB and NI-WB) treatment induced flowering. The transcriptional factors phyA, cry1 and FTL (FLOWERING LOCUS T) were positively affected, while phyB and AFT were negatively affected. In conclusion, morphogenesis, flowering, and transcriptional factors were all significantly affected either positively or negatively by shifts in the light quality of NI. The light quality of the first 2 h of NI affected neither morphogenesis nor flowering, while the light quality of the last 2 h of NI significantly affected both morphogenesis and flowering.

Biochemical traits and proteomic changes in postharvest flowers of medicinal chrysanthemum exposed to enhanced UV-B radiation.

The article studied UV-B effects on biochemical traits and proteomic changes in postharvest flowers of medicinal chrysanthemum. The experiment about UV-B effects on biochemical traits in flowers included six levels of UV-B treatments (0 (UV0), 50 (UV50), 200 (UV200), 400 (UV400), 600 (UV600) and 800 (UV800) µWcm(-2)). UV400, UV600 and UV800 treatments significantly increased the contents of hydrogen peroxide, malondialdehyde and UV-B absorbing compounds, and the activity of phenylalanine ammonia lyase enzyme over the control. The contents of chlorogenic acid and flavone in flowers were significantly increased by UV-B treatments (except for UV50 and UV800). Two-dimensional gel electrophoresis was utilized to analyze proteomic changes in flowers with or without UV-B radiation. Results indicated that 43 protein spots (>1.5-fold difference in volume) were detected, including 19 spots with a decreasing trend and 24 spots with an increasing trend, and 19 differentially expressed protein spots were successfully indentified by MALDI-TOF MS. The indentified proteins were classified based on functions, the most of which were involved in photosynthesis, respiration, protein biosynthesis and degradation and defence. An overall assessment using biochemical and differential proteomic data revealed that UV-B radiation could affect biochemical reaction and promote secondary metabolism processes in postharvest flowers.

CsTFL1, a constitutive local repressor of flowering, modulates floral initiation by antagonising florigen complex activity in chrysanthemum.

Chrysanthemums require repeated cycles of short-day (SD) photoperiod for successful anthesis, but their vegetative state is strictly maintained under long-day (LD) or night-break (NB) conditions. We have previously demonstrated that photoperiodic flowering of a wild diploid chrysanthemum (Chrysanthemum seticuspe f. boreale) is controlled by a pair of systemic floral regulators, florigen (CsFTL3) and anti-florigen (CsAFT), produced in the leaves. Here, we report the functional characterisation of a local floral regulator, CsTFL1, a chrysanthemum orthologue of TERMINAL FLOWER 1 gene in Arabidopsis. Constitutive expression of CsTFL1 in C. seticuspe (CsTFL1-ox) resulted in extremely late flowering under SD and prevented up-regulation of floral meristem identity genes in shoot tips and leaves. Bimolecular fluorescence complementation assay showed that both CsTFL1 and CsFTL3 interacted with CsFDL1, a bZIP transcription factor FD homologue, in the nucleus. The transient gene expression assay indicated that CsTFL1 suppresses flowering by directly antagonising the flower inductive activity of the CsFTL3-CsFDL1 complex. Our results suggest that strict maintenance of vegetative state under non-inductive photoperiod is achieved by the coordinated action of both the systemic floral inhibitor and local floral inhibitor CsTFL1, which is constitutively expressed in shoot tips.

Transcriptomic analyses reveal species-specific light-induced anthocyanin biosynthesis in chrysanthemum.

BACKGROUND: The flower colour of agricultural products is very important for their commercial value, which is mainly attributed to the accumulation of anthocyanins. Light is one of the key environmental factors that affect the anthocyanin biosynthesis. However, the deep molecular mechanism remains elusive, and many problems regarding the phenotypic change and the corresponding gene regulation are still unclear. In the present study, Chrysanthemum × morifolium 'Purple Reagan', a light-responding pigmentation cultivar, was selected to investigate the mechanism of light-induced anthocyanin biosynthesis using transcriptomic analyses. RESULTS: Only cyanidin derivatives were identified based on the analyses of the pigmentation in ray florets. Shading experiments revealed that the capitulum was the key organ and that its bud stage was the key phase responding to light. These results were used to design five libraries for transcriptomic analyses, including three capitulum developmental stages and two light conditions. RNA sequences were de novo assembled into 103,517 unigenes, of which 60,712 were annotated against four public protein databases. As many as 2,135 unigenes were differentially expressed between the light and dark libraries with 923 up-regulated and 1,212 down-regulated unigenes in response to shading. Next, interactive pathway analysis showed that the anthocyanin biosynthetic pathway was the only complete metabolic pathway both modulated in response to light and related to capitulum development. Following the shading treatment, nearly all structural genes involved in the anthocyanin biosynthetic pathway were down-regulated. Moreover, three CmMYB genes and one CmbHLH gene were identified as key transcription factors that might participate in the regulation of anthocyanin biosynthesis under light conditions based on clustering analysis and validation by RT-qPCR. Finally, a light-induced anthocyanin biosynthesis pathway in chrysanthemums was inferred. CONCLUSION: The pigmentation of the ray florets of chrysanthemum cultivar 'Purple Reagan' is dependent on light. During the light-induced pigmentation process, the expression of seven structural genes in the anthocyanin biosynthetic pathway (regulated by at least four transcription factors in response to light) are the main contributors to the pigmentation of chrysanthemums. This information will further our understanding of the molecular mechanisms governing light-induced anthocyanin biosynthesis in ornamental plants.

The changes in quality ingredients of Qi chrysanthemum flowers treated with elevated UV-B radiation at different growth stages.

The paper mainly reported the changes in quality ingredients of Qi chrysanthemum flowers treated with elevated UV-B radiation at different growth stages. The experiment included two levels of UV-B radiation (ambient UV-B, a 10% increase in ambient UV-B). Elevated UV-B radiation was carried out for 10-days during seedling, vigorous growth, bud and flower stages of Qi chrysanthemum, respectively. Elevated UV-B treatments applied during four development stages did not significantly affect flower yield, the rate of superoxide radical production and malondialdehyde concentration in flowers, while increased free amino acid concentration. The amino acid concentration induced by elevated UV-B radiation applied during bud stage was higher than that during the other stages. Elevated UV-B radiation applied during vigorous growth (except for flavone), bud and flower stages of chrysanthemum significantly increased hydrogen peroxide concentration, phenylalanine ammonia lyase enzyme activity, vitamin C, chlorogenic acid and flavone concentrations in flowers. These results suggested that active and nutritional ingredients in flowers of chrysanthemum could be increased by elevated UV-B radiation applied during the later growth stages of chrysanthemum. The paper supplied a simple and environmental-friendly method to improve quality of medicinal plants.

Using the quantum yields of photosystem II and the rate of net photosynthesis to monitor high irradiance and temperature stress in chrysanthemum (Dendranthema grandiflora).

Under a dynamic greenhouse climate control regime, temperature is adjusted to optimise plant physiological responses to prevailing irradiance levels; thus, both temperature and irradiance are used by the plant to maximise the rate of photosynthesis, assuming other factors are not limiting. The control regime may be optimised by monitoring plant responses, and may be promptly adjusted when plant performance is affected by extreme microclimatic conditions, such as high irradiance or temperature. To determine the stress indicators of plants based on their physiological responses, net photosynthesis (Pn) and four chlorophyll-a fluorescence parameters: maximum photochemical efficiency of PSII [Fv/Fm], electron transport rate [ETR], PSII operating efficiency [F'q/F'm], and non-photochemical quenching [NPQ] were assessed for potted chrysanthemum (Dendranthema grandiflora Tzvelev) 'Coral Charm' under different temperature (20, 24, 28, 32, 36 °C) and daily light integrals (DLI; 11, 20, 31, and 43 mol m(-2) created by a PAR of 171, 311, 485 and 667 µmol m(-2) s(-1) for 16 h). High irradiance (667 µmol m(-2) s(-1)) combined with high temperature (>32 °C) significantly (p < 0.05) decreased Fv/Fm. Under high irradiance, the maximum Pn and ETR were reached at 24 °C. Increased irradiance decreased the PSII operating efficiency and increased NPQ, while both high irradiance and temperature had a significant effect on the PSII operating efficiency at temperatures >28 °C. Under high irradiance and temperature, changes in the NPQ determined the PSII operating efficiency, with no major change in the fraction of open PSII centres (qL) (indicating a QA redox state). We conclude that 1) chrysanthemum plants cope with excess irradiance by non-radiative dissipation or a reversible stress response, with the effect on the Pn and quantum yield of PSII remaining low until the temperature reaches 28 °C and 2) the integration of online measurements to monitor photosynthesis and PSII operating efficiency may be used to optimise dynamic greenhouse control regimes by detecting plant stress caused by extreme microclimatic conditions.

Identification and characterization of the CONSTANS-like gene family in the short-day plant Chrysanthemum lavandulifolium.

The CONSTANS (CO) and CONSTANS-like (COL) genes play key roles in the photoperiodic flowering pathways, and studying their functions can elucidate the molecular mechanisms underlying flowering control in photoperiod-regulated plants. We identified eleven COL genes (ClCOL1-ClCOL11) in Chrysanthemum lavandulifolium and divided them into three groups that are conserved among the flowering plants based on phylogenetic analysis. Most of the ClCOL genes are primarily expressed in the leaf and shoot apices, except for ClCOL6-ClCOL9, which belong to Group II. The expression levels of ClCOL4-ClCOL5 and ClCOL7-ClCOL8 are up-regulated under inductive short-day (SD) conditions, whereas ClCOL6 is down-regulated under inductive SD conditions. The ClCOL genes exhibit four different diurnal rhythm expressions (Type I-Type IV). The Type I genes (ClCOL4-ClCOL5) are highly transcribed under light. The Type II genes (ClCOL1-ClCOL2, ClCOL10) display increased expression in darkness and are rapidly suppressed under light. Transcripts of ClCOL6-ClCOL9 and ClCOL11, belonging to Type III, are abundant in the late light period or at the beginning of the dark period. ClCOL3 belongs to Type IV, with high expression in the early light period and dark period. The peak expression levels of ClCOL4-ClCOL6 are decreased and postponed in the non-inductive night break (NB) and under long-day (LD) conditions, indicating that those genes may play an essential role in the flowering regulation of C. lavandulifolium. The overexpression of ClCOL5 promotes the flowering of Arabidopsis grown under LD conditions, suggesting that ClCOL5 may function as a flowering enhancer in C. lavandulifolium. This study will be useful not only for the study of the C. lavandulifolium photoperiod-dependent flowering process but also for the genetic manipulation of flowering time-related genes to change the flowering time in the chrysanthemum.

Spectral effects of supplementary lighting on the secondary metabolites in roses, chrysanthemums, and campanulas.

To investigate the effect of the light spectrum on photosynthesis, growth, and secondary metabolites Rosa hybrida 'Scarlet', Chrysanthemum morifolium 'Coral Charm', and Campanula portenschlagiana 'BluOne' were grown at 24/18°C day/night temperature under purpose-built LED arrays yielding approximately 200 µmol m(-2)s(-1) at plant height for 16 h per day. The four light treatments were (1) 40% Blue/60% Red, (2) 20% Blue/80% Red, (3) 100% Red, and (4) 100% White (Control). The plant height was smallest in 40% Blue/60% Red in roses and chrysanthemums, while the biomass was smallest in the white control in roses and in 100% Red in chrysanthemums. The total biomass was unaffected by the spectrum in campanulas, while the leaf area was smallest in the 40% Blue/60% Red treatment. In 100% Red curled leaves and other morphological abnormalities were observed. Increasing the blue to red ratio increased the stomatal conductance though net photosynthesis was unaffected, indicating excess stomatal conductance in some treatments. With higher blue light ratio all phenolic acids and flavonoids increased. In view of the roles of these secondary metabolites as antioxidants, anti-pathogens, and light protectants, we hypothesize that blue light may predispose plants to better cope with stress.

The effects of UV-B radiation intensity on biochemical parameters and active ingredients in flowers of Qi chrysanthemum and Huai chrysanthemum.

The article studied UV-B effects on biochemical parameters and active ingredients in flowers of Qi chrysanthemum and Huai chrysanthemum during the bud stage. The experiment included four UV-B radiation levels (CK, ambient UV-B; T1, T2 and T3 indicated a 5%, 10% and 15% increase in ambient UV-BBE, respectively) to determine the optimal UV-B radiation intensity in regulating active ingredients level in flowers of two chrysanthemum varieties. Flower dry weight of two cultivars was not affected by UV-B radiation under experimental conditions reported here. UV-B treatments significantly increased the rate of superoxide radical production, hydrogen peroxide (H2O2) (except for T1) and malondialdehyde concentration in flowers of Huai chrysanthemum and H2O2 concentration in flowers of Qi chrysanthemum. T2 and T3 treatments induced a significant increase in phenylalanine ammonia lyase enzyme (PAL) activity, anthocyanins, proline, ascorbic acid, chlorogenic acid and flavone content in flowers of two chrysanthemum varieties, and there were no significant differences in PAL activity, ascorbic acid, flavone and chlorogenic acid content between the two treatments. These results indicated that appropriate UV-B radiation intensity did not result in the decrease in flower yield, and could regulate PAL activity and increase active ingredients content in flowers of two chrysanthemum varieties.

Influences of four different light-emitting diode lights on flowering and polyphenol variations in the leaves of chrysanthemum (Chrysanthemum morifolium).

Light-emitting diodes (LEDs) are an efficient alternative to traditional lamps for plant growth. To investigate the influence of LEDs on flowering and polyphenol biosynthesis in the leaves of chrysanthemum, the plants were grown under supplemental blue, green, red, and white LEDs. Flower budding was formed even after a longer photoperiod than a critical day length of 13.5 h per day under blue light illumination. The weights of leaves and stems were highest under the white light illumination growth condition, whereas the weight of roots appeared to be independent of light quality. Among nine polyphenols characterized by high-performance liquid chromatography-tandem mass spectroscopy, three polyphenols were identified for the first time in chrysanthemum. A quantitation and principal component analysis biplot demonstrated that luteolin-7-O-glucoside (2), luteolin-7-O-glucuronide (3), and quercetagetin-trimethyl ether (8) were the highest polyphenols yielded under green light, and dicaffeoylquinic acid isomer (4), dicaffeoylquinic acid isomer (5), naringenin (7), and apigenin-7-O-glucuronide (6) were greatest under red light. Chlorogenic acid (1) and 1,2,6-trihydroxy-7,8-dimethoxy-3-methylanthraquinone (9) were produced in similar concentrations under both light types. The white and blue light appeared inefficient for polyphenol production. Taken together, our results suggest that the chrysanthemum flowering and polyphenol production are influenced by light quality composition.