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A necrotic disorder occurs on upper leaves of many oriental hybrid lily (Lilium L.) cultivars, including the most-widely-grown `Star Gazer'. We term this disorder “upper leaf necrosis” (ULN) and hypothesize that it is a calcium (Ca) deficiency. We demonstrated that Ca concentration in necrosed tissues was nearly six-fold below that of normal leaves (0.10% vs. 0.57% dry weight), and that Ca concentration was negatively associated with percentage necrosed leaf area. It was concluded that ULN is a Ca deficiency disorder. When the symptoms were slight, early ULN symptoms appeared as tiny depressed spots on the lower surface of the leaf, or as water-soaked areas when the disorder was severe. Most commonly, ULN began on the leaf margin. The injured areas turned brown, leading to leaf curling, distortion, or tip death. ULN occurred on leaves associated with flower buds and leaves immediately below the flower buds. For the plants grown from 16-18 cm circumference bulbs, the five leaves directly below the flower buds and larger leaves associated with the 1st and the 2nd flower buds were most susceptible. In general, flower buds were not affected by ULN, and continued to develop and flower normally, even though they were associated with subtending, highly distorted leaves. Eighty-five percent of plants began to exhibit ULN symptoms 30-40 days after planting (i.e., 24-34 days after shoot emergence). This was the stage when the 6th or 7th leaf under the bottom flower bud was just unfolded. Light was not the main factor that initiated ULN, however, ULN severity was greatly increased by light reduction, as leaf transpiration was reduced.

Upper leaf necrosis (ULN) on Lilium `Star Gazer' has been recently demonstrated to be a calcium (Ca) deficiency disorder. In the current studies, we confirmed this by using a Ca-free nutrient regime to reproduce ULN symptoms. The ability of a bulbous storage organ to supply calcium to a growing shoot is poorly understood. Therefore, we conducted experiments to determine Ca partitioning during early growth stages, and under suboptimal Ca levels to determine how the bulb affects the symptomatology. The results indicated that ULN is originally caused by an insufficient Ca supply from the bulb. In the most susceptible period, bulb dry matter decreased dramatically and Ca concentrations in immature folded leaves dropped to very low levels. Consequently, necrosis began to appear on the upper, young leaves. The bulb was able to supply Ca to other organs, but only to a limited extent since Ca concentration in bulbs was low (0.04% w/w). To confirm this result, we cultivated lilies with low-Ca or Ca-free nutrient solution and obtained bulbs with extremely low internal Ca concentrations. Upon forcing these low-Ca bulbs, we found, as expected, prominent necrosis symptoms on the lower and middle leaves. Data suggested the lower and middle leaves relied more on Ca supplied from the bulb, while upper leaves and flowers relied more on Ca uptake from the roots. Different organs have different Ca requirements, and tissue sensitivity to Ca deficiency varies according to the growth stage.

Upper leaf necrosis (ULN) on Lilium `Star Gazer' has been shown to be a calcium (Ca) deficiency disorder. Initial symptoms of ULN tend to appear on leaf margins. Before flower buds are visible, young expanding leaves are congested and overlap each other on the margin. In the current study, we examined the relationship between leaf enclosure, transpiration, and upper leaf necrosis. We demonstrated that low transpiration rate and enclosure of young leaves played an important role in the occurrence of ULN. Young expanding leaves are low transpiration organs. The younger the leaf, the lower the transpiration rate and Ca concentration. Leaf enclosure further reduced transpiration of these young leaves and promoted ULN. Upper leaf necrosis was suppressed by manually unfolding the leaves using a technique we refer to as artificial leaf unfolding (ALU). ALU minimized leaf congestion, exposing leaves that were previously enclosed. We demonstrated that the effect of ALU was not the consequence of thigmomorphogenesis, as ULN was not reduced by mechanical perturbation in lieu of ALU. With ALU, transpiration of upper leaves was significantly increased and Ca concentration of the first leaf immediately below the flower buds was increased from 0.05% to 0.20%. We concluded that leaf enclosure promoted ULN occurrence, and ALU suppressed ULN primarily by increasing transpiration. The use of overhead fans to increase airflow over the tops of the plants significantly reduced both ULN incidence and severity.

Plants of Phalaenopsis orchid are known for their great resilience and ability to flower under less than ideal conditions, including long periods without fertilization. Significant nutrient storage is thought to account for this characteristic; however, the use of stored nutrients in Phalaenopsis has not been fully studied. We used ¹⁵N-labeled Johnson’s solution to trace the use of stored nitrogen (N) and recently absorbed fertilizer N in Phalaenopsis given various fertilizer levels during forcing. By separately labeling fertilizer N applied to Phalaenopsis Sogo Yukidian ‘V3’ plants 6 weeks before and 6 weeks into forcing, we found in the inflorescence that the ratio of N derived from fertilizer applied 6 weeks before forcing to the N derived from fertilizer applied 6 weeks into forcing was 43% to 57%. With 90% reduction in fertilizer concentration during the reproductive stage, the ratio increased to 89% to 11%, indicating that stored N becomes a significant N source for inflorescence development when fertility becomes limited. Reducing fertilizer level during the reproductive stage from full-strength Johnson’s solution down to zero decreased the dry weight of newly grown leaves, reduced the number of flowers from 10.8 to 8.9, and slightly increased the time required between initiation of forcing and anthesis. However, the overall effect of reduced fertilization on the growth and flowering of Phalaenopsis Sogo Yukidian ‘V3’ plants in this study was slight, because under little or no fertilization, more stored N was mobilized and this was sufficient to meet most of the N demand for inflorescence development.

Pseudobulbs are carbohydrate storage organs in Oncidesa. A current pseudobulb forms on a developing vegetative shoot in each growth cycle and it becomes a back pseudobulb when the next vegetative shoot emerges. Both current and back pseudobulbs store carbohydrates, but their functions might differ because the inflorescence emerges from the new (current) shoot after the shoot has developed to a certain stage. This study investigated carbohydrate storage and use in current and back pseudobulbs. We analyzed carbohydrates in the current pseudobulb at five stages during inflorescence development. Glucose and fructose were the highest in the current pseudobulb in the first two stages, when the inflorescence was 10 to 35 cm tall. Then, both glucose and fructose decreased in the following stages to support inflorescence development, but starch increased at that time. In addition, we used Oncidesa with one or two new vegetative shoots to study the use of carbohydrates in pseudobulbs during growth cycles. In both plants with one or two shoots, glucose and fructose accumulated when current pseudobulbs formed, but plants with two new shoots had smaller current pseudobulbs and lower monosaccharide concentrations. Plants with two shoots also consumed more starch in all back pseudobulbs, whereas in the plants with one new shoot, starch only decreased significantly in the first back pseudobulb, which was closer to the new shoot. In addition, if an inflorescence did not develop in the previous growth cycle, new shoots used the monosaccharides that remained in the youngest back pseudobulb for growth; at the same time, starch accumulated in all back pseudobulbs. The current pseudobulb was the actively growing part. Its main carbohydrates were monosaccharides, which accounted for 25% of dry weight and Oncidesa used these carbohydrates mainly for inflorescence growth. After monosaccharides in the pseudobulb were used, the pseudobulb began to store starch. Back pseudobulbs, in which >50% of dry weight was starch, were the primary storage organs that supported new vegetative shoot growth and partly supported later inflorescence development that emerged from the new (current) shoot.

Oncidesa Gower Ramsey ‘Honey Angel’ is a cut flower crop of high economic value worldwide. The regulation of flowering is important for cut flower production scheduling. However, its flowering transition mechanism is still unclear. Oncidesa usually flowers at the end of the growth cycle for each pseudobulb; this timing is probably related to carbohydrate accumulation. During this study, we investigated the carbohydrates in the pseudobulbs from juvenile plants to adult plants and compared the carbohydrates in flowering and nonflowering adult plants. The current pseudobulb and back pseudobulbs of the plants at 0, 0.5, 1.0, 1.5, and 2.0 years after having been moved out of the tissue culture flask were collected. The first pseudobulb formed at 0.5 years, and plants had fulfilled four growth cycles and flowered at 2.0 years. Each successive current shoot grew larger and the back shoot number progressively increased after each new growth cycle. The concentration of total soluble sugars in the current shoot increased from 5.5% of dry weight at 0.5 years to 20.2% of dry weight at 1.5 years. Conversely, the starch concentration decreased in the current pseudobulb as the plants matured. The starch concentration in the back pseudobulbs did not change when the plant grew a new shoot. The starch concentrations in the back pseudobulbs ranged from 33.2% to 57.5% of dry weight, but the combined content of starch in all of the back pseudobulbs increased significantly from 168 mg at 0.5 years to 4608 mg at 2.0 years because of the increasing number of back shoots. The starch in the first back pseudobulb of the nonflowering adult plants accounted for 18.0% of dry weight, which was lower than that of the flowering plants (48.3%). There was no significant difference in total soluble sugars in the current pseudobulb of the nonflowering and flowering plants. Overall, we revealed that the increase in the back shoot number increased the total amount of reserve carbohydrates as the plant reached reproductive maturity. A low starch level was observed in nonflowering adult plants. In both cases, flowering plants had higher starch storage in the back pseudobulbs, suggesting that carbohydrates might regulate the flowering of Oncidesa Gower Ramsey ‘Honey Angel’.

The vase life of Eustoma cut flowers can be extended by adding sugars to the vase solution, but the exact role of sugars and how they are translocated in tissues are not clear. Thus, we observed the preserving effect of different sugars in vase solutions on Eustoma and compared sugar concentrations in vase solutions and in the flowers as well as stems and leaves of cut flowers in a solution containing 200 mg·L⁻¹ 8-hydroxyquinoline sulfate (8-HQS) with and without 20 g·L⁻¹ sucrose during different flowering stages. Inclusion of glucose, fructose, or sucrose in the vase solution extended the vase life of cut flowers with no significant differences among sugar types. During flower opening, the concentration of added sucrose in the vase solution dropped, and the fresh weight (FW), glucose concentration, and sucrose concentration of flowers in sucrose solutions increased, whereas flowers in solutions without sucrose had lower FW and glucose concentrations. During flower senescence, sugar concentration in the vase solution did not change much, but the FW and sucrose concentrations in all flowers declined, although the FW of sucrose-treated flowers fell more slowly. For stems and leaves in the sucrose solution, sugar concentrations increased during the first 7 days with only glucose slightly declining during senescence, whereas the FW was maintained during the entire vase life. In contrast, FWs of those in the solution without sucrose gradually declined. In conclusion, sucrose in the vase solution promoted flower opening and maintained the water balance of Eustoma cut flowers. Glucose and fructose also extended the vase life, likely in similar ways.

Phalaenopsis (Phalaenopsis spp.) is the most important indoor potted plant worldwide. Tissue analysis is very important for managing fertilization practices but the effects of sampling position and plant maturity must be considered. However, there has been little research on the distribution of tissue carbon (C) and nitrogen (N) among leaves and changes of tissue C and N composition during various developmental stages in phalaenopsis. In this study, we thus determined the effects of leaf age, plant maturity, and cultivars on C and N partitioning in phalaenopsis. Overall, C concentration was more uniform and was less affected by the abovementioned factors investigated, whereas N concentration significantly decreased as leaves aged or as plants matured. In P. Sogo Yukidian ‘V3’, new expanding leaf had the highest N concentration of 2.72% of dry weight (DW) and seventh mature leaf had the lowest value of 1.48% DW. Results also indicate that N was not evenly distributed within a leaf, whereas N concentration gradually decreased from the leaf tip to the leaf base. The middle section of the second mature leaf is an appropriate tissue for sampling to obtain the representative N and C concentrations in phalaenopsis. As for the changes in C and N composition through five developmental stages, two cultivars were compared, including the large, white-flowered P. Sogo Yukidian ‘V3’ and the small, purple-flowered P. Sogo Lotte ‘F2510’. As the large-flowered ‘V3’ grew from deflasked plantlet to fully matured plant (18 months after deflasking) in a 10.5-cm pot, whole-plant N concentration decreased from 4.63% DW to 1.67% DW and C/N thus increased from 9.1 to 26.1. Despite the large difference in plant size, the small-flowered ‘F2510’ had a similar trend and values during vegetative growth stages. However, the two cultivars had different trends during reproductive stages. Tissue N concentration and C/N did not further change as mature large-flowered ‘V3’ plants were forced to flower. By contrast, tissue N concentration in small-flowered ‘F2510’ further decreased and C/N thus further increased, which was due to its small stored N pool. Major N sink organ shifted from roots to inflorescences during reproductive growth and the stored N in roots as well as in leaves was then used for flower development.

In the commercial production of phalaenopsis orchids, the cultivation time after deflasking is used to describe the plant age and maturity. Carbon-to-nitrogen (C/N) ratio is often used as an indicator of plant growth and flowering potential. High C/N ratios are considered to promote reproductive growth, and low C/N ratios are associated with the early vegetative growth or even inhibiting flowering. This study investigated how plant age and maturity affected flowering ability and flower quality of phalaenopsis and their relationship to C/N ratio. The plant materials of various ages were the purple, small-flowered Phalaenopsis Sogo Lotte ‘F2510’ and white, large-flowered P. Sogo Yukidian ‘V3’, which were 2 to 7 months and 10 to 20 months after deflasking, respectively. Plants were placed under 25/20 °C for 4 months to force flowering and investigate the flowering-related parameters. The leaf C/N ratio of both varieties increased in general with the increase of plant age. The spiking (flower-stalk emergence) rate of P. Sogo Lotte ‘F2510’ 2 months after deflasking was only 42%, which indicates that these plants were not completely out of their juvenile phase, whereas that of those 3 to 7 months after deflasking was 100%, indicating that plants had acquired full flowering ability. No linear correlation was found between the C/N ratio and days to spiking, to first visible bud, to first flower open, and to 90% flower opening in the white, large-flowered P. Sogo Yukidian ‘V3’. However, there was a positive correlation between the C/N ratio and inflorescence length, flower-stalk diameter, first flower diameter, and flower count. Thus, the C/N ratio is feasible to be used as an indicator for assessing the flowering quality in phalaenopsis.

Growers realize the importance of nitrogen (N) on the vegetative growth of phalaenopsis orchids (hybrids of Phalaenopsis sp.), but often overlook its influence on reproductive growth. Low N may result in slow plant growth, pale-green leaves, abscission of lower leaves, and few flowers in phalaenopsis. Increasing N concentration up to 200 mg·L⁻¹ promotes leaf growth and increases flower count. High N concentration promotes lateral branching on the flowering stalk, thereby greatly increasing the total flower count and elevating the commercial value. It is important that N be continually applied during the forcing period for best flowering performance, particularly for those that had undergone international shipping. For the vegetative phalaenopsis plants that are induced to flower without being shipped internationally, the N that is already in the plant before spiking provides 43% and the N being absorbed by roots after cooling provides 57% of the total N in the inflorescence at time of visible bud. When insufficiently fertilized or no fertilization is applied during the forcing period, more of the existing N in a plant is mobilized for inflorescence development. Phalaenopsis roots can take up all three forms of N [i.e., nitrate (NO₃-N), ammonium (NH₄-N), and urea] directly. In two studies, phalaenopsis plants were supplied with the same amount of total N but with varying NO₃-N from 100%, 75%, 50%, 25%, to 0% (a common N concentration was achieved by the substitution of the respective balance with NH₄-N). Plants were smaller when receiving 75% or 100% NH₄-N with a tendency of decreasing top leaf width and whole-plant leaf spread as NO₃-N decreased from 100% to 0%. Spiking was delayed and spiking rate decreased when plants were grown in sphagnum moss, but not a bark mix, and received more than 50% of the N in NH₄-N. As the ratio between NO₃-N and NH₄-N increased, flowers became increasingly larger. The negative effects of low ratios of NO₃-N to NH₄-N were more severe in the second flowering cycle. When supplied with 50% or more NH₄-N, the absorption of cations by phalaenopsis roots declined, with reduced concentrations of calcium and magnesium in plants, while symptoms of ammonium toxicity appeared, including growth retardation, chlorotic leaves, and necrotic roots. In conclusion, adequate N and its continual supply during both vegetative and reproductive stages are recommended for the best growth and flowering of phalaenopsis. Since phalaenopsis plants prefer N in the NO₃-N form, it is suggested that growers choose and apply a fertilizer with nitrate as the major N source.