Bud abortion in tulip bulbs studied by magnetic resonance imaging

After storage and subsequent planting of flower bulbs, the flower bud frequently appears to be aborted. This physiological aberration is probably caused by a change in the water status of the bulb and may be initiated during storage. The development of bud abortion in tulip bulbs was studied during long-term dry storage of the bulbs at 5 degrees C. The anatomy of individual tulip bulbs was followed non-invasively with T2-weighted NMR imaging, which allowed the monitoring of the growth of the shoot and daughter bulbs. Quantitative maps of T1 and T2 relaxation times of individual bulbs were used to assess regional changes in the water status of different tissues. Parallel to the NMR measurements, bulbs were planted to assess the ultimate flower quality. Moreover, water content, osmolality of tissue sap and ion leakage of excised shoot and scale tissues were determined to obtain information about the water status and viability of the bulbs. Significant decreases during long-term storage were found in T1 and T2 relaxation times in the shoot and particularly in the stamens. An increase in the osmolality of tissue sap and the decrease in relaxation times in the shoot below a certain threshold value attained after 24 weeks of storage, could be indicative for the emergence of bud abortion in tulips.     

Endogenous levels of abscisic Acid and decanoic Acid in dutch iris bulbs and the influence of abscisic Acid and decanoic Acid on iris meristems cultured in vitro

Abscisic acid (ABA) and decanoic acid inhibited shoot elongation and floral development of Dutch iris (Iris hollandica Hoog. cv Ideal) meristems cultured in vitro. No synergism with respect to inhibition of leaf growth between ABA and decanoic acid was observed. With monthly harvest dates, from July 10, 1981 to October 10, 1981, there was a progressive decrease in endogenous level of free ABA in `Ideal' iris bulbs. Bulbs subjected to a full set of the usual preplanting storage conditions flowered, on average, 46 days after planting versus 194 days after planting for bulbs planted directly after harvest. ABA levels at harvest were 4- to 5-fold those after the preplanting storage treatment. In general, ABA levels did not correlate well with the length of time from planting until flowering of iris bulbs. Endogenous decanoic acid levels did not follow any pattern with respect to harvest date or postharvest treatment. After the postharvest high temperature treatment, there was about a 3-fold increase in nonscale decanoic acid concentration. Decanoic acid levels, in nonscale tissue, remained high after each of the other postharvest treatments. It is concluded that there is no good evidence to support the contention that either ABA or decanoic acid is directly involved in iris bulb dormancy.     

Compositional changes in cell wall polysaccharides from chilled and non-chilled cucumber fruit

Cell wall carbohydrate composition and 1-aminocyclopropane-1-carboxylic acid (ACC) content have been determined in chilled (2.5°) and non-chilled (12.5°) cucumber fruit. The major compositional change that accompanied the increased capability for ACC synthesis during chilling was a diminished loss of galactose residues, relative to the loss which occurred at 12.5°. However, the loss of galactose residues increased markedly when fruit were transferred from 2.5° to 20°, and wall galactose levels eventually declined to similar levels in both chilled and non-chilled fruit. Rhamnose, arabinose, xylose, mannose and cellulose content of walls was similar in chilled and non-chilled fruit and did not change substantially upon transfer of fruit to 20°. Upon transfer of chilled fruit from 2.5° to 20°, an increase in the relative amount of galacturonic acid in cell walls occurred; this change did not occur in non-chilled fruit. Thus, chilling stress results in a rapid change in the neutral sugar and galacturonic acid composition of cell wall pectic polysaccharides upon warming.     

Measurement of endogenous ABA levels in chilled somatic embryos of carrot by immunoassay

Somatic embryos of carrot, Daucus carota L. Royal Chantenay, were chilled at 4°C for the last 3 days of development in order to harden torpedo stage embryos to increase embryo survival during desiccation. ABA levels in chilled and non-chilled embryos were measured using a polyclonal radioimmunoassay and a monoclonal enzyme-linked immunosorbent assay (ELISA). The monoclonal ELISA is the preferred technique due to superior sensitivity and specificity. ABA levels, measured by either technique, were similar in chilled and non-chilled embryos. The relative water content was lower in chilled embryos than in non-chilled embryos and chilling altered protein secretion of one cell line.List of abbreviations ABA abscisic acid - 2,4-D 2,4-dichlorophenoxyacetic acid - BHT 2,5-di-tert-butyl-4-methyl phenol - BSA bovine serum albumin - ELISA enzyme-linked immunosorbent assay.     

低温胁迫下二倍体杂交种高山松光系统Ⅱ功能稳定性研究

以人工气候室内的高山松(Pinus densata)及其亲本油松(P.tabulaeformis)和云南松(P.yunnanensis)实生苗为材料,通过测定低温胁迫和恢复后其幼苗的快速叶绿素荧光诱导动力学(JIP-test)参数,分析高山松及其亲本的光系统Ⅱ(PSⅡ)功能稳定性以及对低温胁迫的响应差异,以探讨高山松适应高海拔低温环境的光合生理生态机制。结果表明:经过33 d的低温(12℃/5℃,昼/夜)处理后,高山松PSⅡ功能参数单位截面积吸收的能量(ABS/CSo)、捕获的能量(TRo/CSo)、电子传递的能量(ETo/CSo)、耗散的能量(DIo/CSo)、有活性反应中心的数量(RC/CSo),以及原初光化学反应的最大量子产额(φPo)、吸收光能用于QA-以后的电子传递的能量比例(φEo)、捕获光能用于QA-以后的电子传递的能量比例(Ψo)、光能利用能力参数(PIabs)、2 ms时相对可变荧光(VJ)、荧光上升的初始斜率(Mo)、荧光曲线与Fm所围面积(Area)、介于F0和Fm之间的相对面积(Sm)和QA氧化还原次数(N)均与处理前无显著差异;油松和云南松的φPo、φEo、Ψo、PIabs、Area与对照差异显著,云南松的VJ、Mo、Sm、N、ETo/CSo、DIo/CSo、ABS/CSo也与对照存在显著差异;温度恢复之后(25℃/18℃,昼/夜),这些荧光参数的恢复程度表现为高山松>油松>云南松。可见,低温胁迫显著影响了油松和云南松的光能传递和光能利用能力以及受体库大小,且云南松遭受伤害的程度高于油松,而高山松的PSⅡ功能及其稳定性在遭受低温胁迫时明显高于亲本油松和云南松,且胁迫解除后恢复程度高,从而使得高山松能够在高海拔低温环境中累积代谢所需的有机物,保证其正常的生长和发育。     

Regulation of summer dormancy by water deficit and ABA in Poa bulbosa ecotypes

BACKGROUND AND AIMS: Survival of many herbaceous species in Mediterranean habitats during the dry, hot summer depends on the induction of summer dormancy by changes in environmental conditions during the transition between the winter (growth) season to the summer (resting) season, i.e. longer days, increasing temperature and drought. In Poa bulbosa, a perennial geophytic grass, summer dormancy is induced by long days, and the induction is enhanced by high temperature. Here the induction of summer dormancy in a Mediterranean perennial grass by water deficit under non-inductive photoperiodic conditions is reported for the first time. METHODS: Plants grown under 22/16 degrees C and non-inductive short-day (9 h, SD) were subjected to water deficit (WD), applied as cycles of reduced irrigation, or sprayed with ABA solutions. They were compared with plants in which dormancy was induced by transfer from SD to inductive long-day (16 h, LD). Responses of two contrasting ecotypes, from arid and mesic habitats were compared. Dormancy relaxation in bulbs from these ecotypes and treatments was studied by comparing sprouting capacity in a wet substrate at 10 degrees C of freshly harvested bulbs to that of dry-stored bulbs at 40 degrees C. Endogenous ABA in the bulbs was determined by monoclonal immunoassay analysis. KEY RESULTS: Dormancy was induced by WD and by ABA application in plants growing under non-inductive SD. Dormancy induction by WD was associated with increased levels of ABA. Bulbs were initially deeply dormant and their sprouting capacity was very low, as in plants in which dormancy was induced by LD. Dormancy was released after 2 months dry storage at 40 degrees C in all treatments. ABA levels were not affected by dormancy relaxation. CONCLUSIONS: Summer dormancy in P. bulbosa can be induced by two alternative and probably additive pathways: (1) photoperiodic induction by long-days, and (2) water deficit. Increased levels of endogenous ABA are involved in both pathways.     

Bulbil production in Narcissus: the effect of temperature and duration of storage on bulb unit development and subsequent propagation by twin-scaling

The growth of bulb components and carbohydrate content of Narcissus bulbs was followed during storage at 5–30 ^oC, and the productivity of twin-scale propagules cut from bulbs after these storage treatments was studied. Growth of daughter bulb unit components was greatest during storage at 9–15 ^oC, and greatly inhibited at 30 ^oC. Concentrations of soluble sugars were highest following 30 ^oC storage, and higher in intermediate bulb scales than in outer or inner scales. Bulbil production on twin-scales was greatest on propagules cut from the intermediate scales, and in bulbs previously stored at 15 or 30 ^oC.     

Exogenous GA<Subscript>3</Subscript> overcomes bud deterioration in tulip (<Emphasis Type="Italic">Tulipa gesneriana</Emphasis> L.) bulbs during dry storage by promoting endogenous IAA activity in the internodes

Endogenous free IAA was examined with an immunohistochemical method for its involvement in the reduction of bud deterioration after GA₃ was injected into the bulbs. We found that tulip bulbs stored at 20°C constantly developed severe bud deterioration, whereas the symptoms of deterioration was lighter in the bulbs with GA₃ injection and not observed in the bulbs with 4°C treatment. 73% success in overcoming bud deterioration was achieved in 20°C with GA₃ treatment after 8 weeks of bulb storage, and the success rate was 7% after 12 weeks of storage. IAA was detected in the parenchyma cells in the internodes of the shoot after the bulbs were stored at 4°C or at 20°C with GA₃ injection for 4 weeks, but little was detected in the bulbs stored at 20°C constantly. Moreover, a weak IAA signal was present in between the cells of the internodes irrespective of bulb treatment. After planting, the bulbs that had been treated differently exhibited different flowering ability. The bulbs stored at 4°C for 4, 8 and 12 weeks attained high flowering percentage, which was lower in the 20°C with GA₃ treatment and lowest in the 20°C treatment. It may be concluded that GA₃ injection decreases bud deterioration of tulip bulbs during dry storage at 20°C by promoting the endogenous IAA in the internodes.     

Changes in free polyamines in Tulipa gesneriana flower stalks during dry-storage and after planting of bulbs

Tulip bulbs cv. Apeldoorn are dry-stored at 5°C for 12 weeks to ensure sufficient elongation of the flower stalk, when subsequently planted at higher temperatures (17–20°C). To investigate whether free polyamines are involved in this process, flower stalk internodes were analyzed during dry-storage and after planting of the bulbs.During dry-storage for 12 weeks at 5°C (cooled) and 17°C (non-cooled), the free putrescine, spermidine and spermine amounts per flower stalk increased. The putrescine amount increased at 5°C significantly more than at 17°C, whereas the opposite was found for the spermine amount. These differences developed early during dry-storage and disappeared rapidly at subsequent higher temperatures.After planting, the lower- and uppermost flower stalk internodes of the pre-cooled bulbs elongated much faster than those of the non-cooled ones. In the pre-cooled bulbs, the free polyamine amounts per internode increased with time after planting, but the time course of these changes was different. In the non-cooled bulbs, the free polyamine amounts increased to a much lesser extent or remained more or less constant.It is argued that the observed changes in the free polyamine contents are probably not required for the cold-induced extension growth of tulips cv. Apeldoorn.Abbreviations PA polyamine - Put putrescine - Spd spermidine - Spm spermine     

MRI can reveal metabolic changes in lily bulbs in vivo during dormancy release

The factors influencing dormancy release in lily bulbs strongly affect commercialization success, but the mechanism of dormancy release is still unclear. Magnetic resonance imaging (MRI) can detect changes in morphology and water status in a living plant bulb and aid in investigating release factors. To evaluate whether MRI could be used to detect intra-bulb metabolic changes during the dormant period in Oriental Lilies (Lilium ??Sorbonne??), a series of MRI and sugar concentration measurements were performed weekly on bulbs stored for 11 weeks at 4°C. The image quality of intra-bulb structure obtained using T ₁-weighted imaging was superior to that obtained using T ₂-weighted imaging and had a higher signal-to-noise ratio (0.97±0.01). Magnetization transfer ratio values for the bud and basal plate declined during the first eight weeks of cold storage (P>0.05), and were well correlated with concentration of soluble sugar in the bud (R ²=0.95) and basal plate (R ²=0.93). Thus, MRI can serve as a valuable tool for observation and analysis of dynamic morphological and metabolic changes in vivo during dormancy release. This information is potentially useful as a guide in the improvement of horticultural product quality.     

Carbohydrate metabolism in Nerine sarniensis bulbs developing in liquid culture

Anatomical and physiological changes accompanying enhanced Nerine sarniensis cv. Salmon Supreme bulb growth in vitro were examined. Small bulbs, 2–3 mm in diameter, grown in vitro on a semi-solid medium were subcultured to liquid medium with elevated levels of sucrose (Suc) and inorganic phosphate. Bulbs' fresh and dry weights, carbohydrate contents and the activities of enzymes related to carbohydrate metabolism were determined at different stages of bulb development. Starch was the dominant storage carbohydrate in these bulbs, and the leaf bases parenchyma cells were the principal storage tissue. During the first month of bulb growth, only small changes in starch content were detected. However, an increase in starch level was observed at later stages of development. The activity of ADP-glucose pyrophosphorylase (EC 2.7.7.27), a key enzyme of starch synthesis, increased just before the increase in starch accumulation. Sucrose was the dominant soluble sugar in the bulbs, only traces of glucose and fructose were detected. The activity of alkaline invertase (INV, EC 3.2.1.26) was higher than that of acid INV during the growth period. Sucrose synthase (EC 2.4.1.13) exhibited the highest Suc-degrading activity during bulb growth. Suc was hydrolyzed in the medium by the cell wall bound acid INV during the growing period. The results are discussed in relation to enhanced nerine bulb growth and development in vitro.     

细叶百合的生物量和营养分配

 以栽培的2年生细叶百合（Lilium pumilum）为材料,于2000年的生长季从蕾期至种子成熟期进行6次取样,对其各器官生物量和氮、磷元素的配置进行了动态研究。结果表明,细叶百合虽然以种子繁殖为主,但在整个生长季用于生殖器官的生物量投资的比例并不很大,大量干物质分配到地下器官鳞茎中（平均为60.17%）;茎、叶的生物量分配比例仅次于鳞茎;雄蕊生物量分配比例明显高于雌蕊。在叶萌动及展叶初期植株全氮百分含量最高;从春季萌动至秋季果实成熟,叶中的氮呈逐渐递减的趋势;茎和生殖器官的全氮含量在蕾期最大;生殖器官与叶、鳞茎的全氮含量相关显著。磷在生殖器官的含量较高,这与磷在植物有性生殖过程中的重要作用相一致;生殖器官与茎的全磷含量相关显著。地下器官全氮、全磷随季节变化有增多的趋势;地上各器官全氮、全磷相关显著,随季节变化有明显减少的趋势。     

仙客来组织培养中再生器官类型及增殖稳定性比较

对‘胜利女神’(Cyclamen persicum cv. Victoria’)和‘大红’(C. persicum cv. ‘Dahong’)两个仙客来品种成苗叶片诱导的愈伤组织,在6-BA与NAA配合的12个分化培养基上继代培养,第一次继代主要产生大量的不定芽芽点,从第二次继代后会得到5种类型的再生器官,分别为不定芽芽点、多芽球茎、多芽新梢、单芽球茎和叶片状芽丛。通过对不同再生器官增殖稳定性的试验表明,‘大红’品种的多芽球茎和多芽新梢在继代培养中再生相同器官的稳定性比较强,由于多芽球茎的增殖率大于多芽新梢,在继代中选择多芽球茎为再生体系较理想;不定芽芽点在以后的继代中会分散产生各种再生器官,不宜作为增殖器官;单芽球茎和叶片状芽丛继代分散性强,为不正常再生器官。‘胜利女神’品种多芽球茎和多芽新梢继代的稳定性比‘大红’品种差,易于产生多种不正常器官。     

Cell water balance of white button mushrooms (Agaricus bisporus) during its post-harvest lifetime studied by quantitative magnetic resonance imaging

A combination of quantitative water density and T2 MRI and changes therein observed after infiltration with 'invisible' Gd-DTPA solution was used to study cell water balances, cell water potentials and cell integrity. This method was applied to reveal the evolution and mechanism of redistribution of water in harvested mushrooms. Even when mushrooms did not lose water during the storage period, a redistribution of water was observed from stipe to cap and gills. When the storage condition resulted in a net loss of water, the stipe lost more water than the cap. The water density in the gill increased, probably due to development of spores. Deterioration effects (i.e. leakage of cells, decrease in osmotic water potential) were found in the outer stipe. They were not found in the cap, even at prolonged storage at 293 K and R.H.=70%. The changes in osmotic potential were partly accounted for by changes in the mannitol concentration. Changes in membrane permeability were also indicated. Cells in the cap had a constant low membrane (water) permeability. They developed a decreasing osmotic potential (more negative), whereas the osmotic potential in the outer stipe increased, together with the permeability of cells.     

Neoseiulus paspalivorus,a predator from coconut,as a candidate for controlling dry bulb mites infesting stored tulip bulbs

The dry bulb mite, Aceria tulipae, is the most important pest of stored tulip bulbs in The Netherlands. This tiny, eriophyoid mite hides in the narrow space between scales in the interior of the bulb. To achieve biological control of this hidden pest, candidate predators small enough to move in between the bulb scales are required. Earlier experiments have shown this potential for the phytoseiid mite, Neoseiulus cucumeris, but only after the bulbs were exposed to ethylene, a plant hormone that causes a slight increase in the distance between tulip bulb scales, just sufficient to allow this predator to reach the interior part of the bulb. Applying ethylene, however, is not an option in practice because it causes malformation of tulip flowers. In fact, to prevent this cosmetic damage, bulb growers ventilate rooms where tulip bulbs are stored, thereby removing ethylene produced by the bulbs (e.g. in response to mite or fungus infestation). Recently, studies on the role of predatory mites in controlling another eriophyoid mite on coconuts led to the discovery of an exceptionally small phytoseiid mite, Neoseiulus paspalivorus. This predator is able to move under the perianth of coconuts where coconut mites feed on meristematic tissue of the fruit. This discovery prompted us to test N. paspalivorus for its ability to control A. tulipae on tulip bulbs under storage conditions (ventilated rooms with bulbs in open boxes; 23 °C; storage period June–October). Using destructive sampling we monitored predator and prey populations in two series of replicated experiments, one at a high initial level of dry bulb mite infestation, late in the storage period, and another at a low initial dry bulb mite infestation, halfway the storage period. The first and the second series involved treatment with N. paspalivorus and a control experiment, but the second series had an additional treatment in which the predator N. cucumeris was released. Taking the two series of experiments together we found that N. paspalivorus controlled the populations of dry bulb mites both on the outer scale of the bulbs as well as in the interior part of the bulbs, whereas N. cucumeris significantly reduced the population of dry bulb mites on the outer scale, but not in the interior part of the bulb. Moreover, N. paspalivorus was found predominantly inside the bulb, whereas N. cucumeris was only found on the outer scale, thereby confirming our hypothesis that the small size of N. paspalivorus facilitates access to the interior of the bulbs. We argue that N. paspalivorus is a promising candidate for the biological control of dry bulb mites on tulip bulbs under storage conditions in the Netherlands.     

Exogenous ethylene inhibits sprout growth in onion bulbs

Background and Aims

Exogenous ethylene has recently gained commercial interest as a sprouting inhibitor of onion bulbs. The role of ethylene in dormancy and sprouting of onions, however, is not known.

Methods

A cultivar (Allium cepa ‘Copra’) with a true period of dormancy was used. Dormant and sprouting states of onion bulbs were treated with supposedly saturating doses of ethylene or with the ethylene-action inhibitor 1-methylcyclopropene (1-MCP). Initial sprouting was determined during storage at 18 °C by monitoring leaf blade elongation in a specific size class of leaf sheaths. Changes in ATP content and sucrose synthase activity in the sprout leaves, indicators of the sprouting state, were determined. CO₂ and ethylene production of onion bulbs during storage were recorded.

Key results

Exogenous ethylene suppressed sprout growth of both dormant and already sprouting onion bulbs by inhibiting leaf blade elongation. In contrast to this growth-inhibiting effect, ethylene stimulated CO₂ production by the bulbs about 2-fold. The duration of dormancy was not significantly affected by exogenous ethylene. However, treatment of dormant bulbs with 1-MCP caused premature sprouting.

Conclusions

Exogenous ethylene proved to be a powerful inhibitor of sprout growth in onion bulbs. The dormancy breaking effect of 1-MCP indicates a regulatory role of endogenous ethylene in onion bulb dormancy.Key words: Bulb dormancy, Allium cepa, onion, sprout growth, ethylene, CO₂ production, respiration, 1-methylcyclopropene     

Differentiation in Lilium bulbscales grown in vitro. Effects of activated charcoal, physiological age of bulbs and sucrose concentration on differentiation and scale leaf formation in vitro

Organ differentiation and growth of tissue cultured bulbscales of Lilium auratum Lindl. was investigated. Benzyladenine stimulated bulb and bulbscale differentiation but inhibited root formation. Addition of activated charcoal to the medium negated the effect of BA on differentiation, while the growth of bulbs was markedly stimulated. Organ formation was also influenced by the physiological age of bulbs. From bulbscales of a 3-week-old bulb, a large number of bulbs and roots was formed but callus was not produced. A high sucrose concentration (90 g/1) reduced the subsequent sprouting after planting in soil. Scale leaf formation was also regulated. Sucrose (30 g/1) stimulated scale leaf formation but 90 g/1 were inhibitory.     

The effects of nitrogen, phosphorus, potassium and magnesium on the growth of tulips during the second season of treatment and on the chemical composition of the bulbs

In factorial nutrition treatments, for a second season, with three rates of nitrogen, phosphorus and potassium, and some additional treatments with magnesium, on two bulb sizes of Golden Harvest and Elmus tulip, the effect of nitrogen was again found to be dominant. Nitrogen deficiency decreased bulb yields, leaf area, stem length (with one exception) and flower size; it also retarded flowering: the majority of small nitrogen-deficient bulbs produced only single leaves. Deficiencies of phosphorus, potassium and magnesium had more marked effects than in the first season of treatment and generally reduced bulb yields and leaf area; in some instances stem length and flower size were also reduced. High phosphorus advanced the flowering of Elmus bulbs. The relationships between nutrition and mineral composition of the bulbs showed consistent trends determined by rates and duration of treatments, planting size and variety.     

Developmental changes and tissue distribution of lectin in Tulipa

A sensitive enzyme-immunoassay was developed to quantify the tulip lectin and used to follow its distribution during the life cycle of tulips cv. Attila.The tulip lectin is predominantly located in the bulbs. At planting time the absolute lectin concentration is approximately the same in all bulb scales. However, as the shoot grows and the plant turns on to flowering, the lectin concentration rapidly decreases, first in the inner bulb scales but later also in the outer bulb scale. Soon after flowering the lectin rapidly accumulates in the new daughter bulbs.Lectin levels in leaves, stems and flowers are very low. The lectin in these tissues is already present before the sprout emerges. During the first two weeks after planting, there is a small increase in lectin concentration, followed by a rapid decrease as the plant turns on to flowering. By flowering time all the lectin has disappeared from the aerial parts.Abbreviations DW dry weight - ELISA enzyme-linked immunosorbent assay - FW fresh weight - PBS phosphate-buffered saline - PBSN phosphate-buffered saline containing 0.02% sodium azide - PBST phosphate-buffered saline containing 0.02% sodium azide and 0.05% Tween 20 - TL tulip lectin - Tris 2-amino-2-(hydroxymethyl)-1,3-propanediol     

Relationships between Bulb Size, Apex Size and Flowering in Bulbous Iris cv. Ideal

Large, retarded bulbs of the Iris cv. Ideal flower readily after exposure to appropriate preplanting conditions, whereas smaller bulbs flower with decreased frequency. Flower initiation occurs when apices from large, retarded bulbs are cultured on either Murashige-Skoog medium supplemented with gibberellic acid, or on the same medium without gibberellic acid on which scales from large bulbs have been incubated. Since floral initiation seldom occurs in explants from small bulbs, it is likely that reduced flowering of small bulbs relates in part to characteristics of the apical meristem. Apical dome diameter is one characteristic of retarded (and freshly dug) bulbs that is positively correlated with bulb size. However, whereas prolonged storage at retarding temperature increases the frequency of flowering of smaller bulbs, there is not a concomitant increase in apical dome diameter. Moreover, the ratio of apical dome diameter to bulb size in freshly dug bulbs does not increase measurably with later digging date indicating that apical dome size is not correlated with bulb maturity.