Enhanced UV-B radiation,artificial wounding and leaf chemical defensive potential in Phlomis fruticosa L.

The combined effects of additional UV-B radiation and artificial wounding on leaf phenolics were studied in a short term field experiment with the drought semi-deciduous Mediterranean shrub Phlomis fruticosa L. The seedlings were grown under ambient or ambient plus supplemental UV-B radiation (biologically equivalent to a 15% ozone depletion over Patras, 38.3° N, 29.1° E) for 7 months before wounding. Unexpectedly, supplemental UV-B radiation decreased leaf phenolics. Subsequently, wounding was effected by removing leaf discs from some of the plants, while the rest remained intact and served as controls. Wounding significantly increased phenolics of the wounded leaves and the increase was more pronounced under supplemental UV-B radiation. In addition, wounding had a significant positive effect on the phenolics of the opposite, intact leaf, but only under additional UV-B radiation. We conclude that UV-B radiation, wounding and their combination may affect the chemical defensive potential of Phlomis fruticosa. In addition, increased levels of phenolics after herbivore attack under field conditions may afford extra protection against enhanced UV-B radiation levels.     

Variation in floral morphology within a population of Aster hispidus var. tubulosus (Asteraceae,Astereae)

The family Asteraceae has a particular inflorescence, the capitulum, consisting of ray florets and disc florets. The ray florets function as petals that attract pollinators. Marked variation in the ray floret morphology is known in a natural population of Aster hispidus var. tubulosus (Asteraceae). We analyzed the variation and found two distinct types in the ray florets, the long tubular ray floret and the ligulate ray floret. In this species, therefore, the variation in floral morphology among capitula, each of which is the basic pollination unit, is caused by the variation in the composition of the two ray floret types among capitula. We evaluated the sources of the observed variation in the floral morphology among capitula within a population using a hierarchical analysis that separated within‐individual (i.e. among capitula within each individual) and between‐individual components of the variation. We found that the main source of the variation lay at the between‐individual level, not at the between‐capitulum level nested within individuals. This finding will provide the basic knowledge that enables future study exploring whether the between‐individual variation in floral morphology caused by the compositional variation of the ray floret types leads to differential pollination success of individual plants in species of Asteraceae.     

Reduction of ambient UV-B radiation does not affect growth but may change the flowering pattern of Rosmarinus officinalis L.

The effects of sub-ambient levels of UV-B radiation on the shrub Rosmarinus officinalis L. were investigated in a field filtration experiment in which the ambient UV-B was manipulated by a combination of UV-B transmitting and UV-B absorbing filters. As a result, the plants were receiving near-ambient or drastically reduced UV-B radiation doses. Drastic reduction of UV-B radiation had no effect on mean, total and maximum stem length, number of stems per plant, dry mass of leaves, stems and roots and leaf nitrogen and phenolic contents. However, flowering was more pronounced under reduced UV-B radiation during the winter period which coincides with ascending ambient UV-B radiation. In contrast, during autumn and early winter, a period which coincides with descending ambient UV-B radiation, flowering was unaffected by reduced UV-B radiation. We can conclude that natural UV-B radiation does not affect growth of Rosmarinus officinalis, but its reduction could influence the flowering pattern of the species.     

Enhanced UV-B radiation,flower attributes and pollinator behaviour in Cistus creticus: a Mediterranean field study

The aim of this investigation was to examine the reasons for the higher pollination success in Cistus creticus under enhanced UV-B radiation (Stephanou & Manetas 1998). Thus, a selected array of floral attributes as well as the frequency and duration of insect visits were studied in the field under ambient or ambient plus supplemental UV-B radiation, simulating a 15% ozone depletion over Patras (38.3° N, 29.1° E). Video-recording revealed two categories of visitors, i.e. true pollinators (bees) and nectar thieves. The frequency of visits to both control and UV-B treated plants was the same and independent of whether the UV-B tubes were on or off during video recording. UV-B radiation had no effect on gross floral morphology (petal surface area, number of pollen grains, stamens and ovules, optical properties of petals and stamens), yet nectary size was almost doubled. In addition, the duration of insect visits was significantly longer on UV-B treated plants, provided that the UV-B tubes were off during monitoring. The differences were abolished during the part of the day that the tubes were on, indicating that the insects were annoyed by supplemental UV-B radiation. These results are consistent with the nectaries producing larger quantities of nectar, which caused the insects to stay longer on flowers of UV-B treated plants and improved pollination success.     

Effects of exogenous hormones on floret development and grain setin wheat

At specific stages during floret development, solutions of IAA,GA₃, zeatin and ABA were injected into the leaf sheath around theyoung spike of wheat (Triticum aestivum L.) to study theregulating effects of exogenous hormones on floret development. Zeatin promotedfloret development and significantly increased the number of fertile florets aswell as grain set, especially at the stage of anther-lobe formation. Zeatinalsoincreased the sugar concentrations in spikes at anthesis. In contrast, IAA,GA₃ and ABA inhibited floret development, with different patternsforeach of the hormones. IAA inhibited the development of the whole spike and allflorets in the spikelets such that grain loss occurred in all positions in thespikelets. GA₃ increased the number of fertile florets per spike,butdecreased grain set of the third floret in each spikelet, especially whenapplied at terminal spikelet formation. ABA inhibited floret development, anddecreased the number of fertile florets and grain set at almost all developmentstages, except at anther-lobe formation. The inhibitory effect of ABA wasmainlyon the first and third florets in each spikelet.     

Enhanced UV-B radiation increases the reproductive effort in the Mediterranean shrub Cistus creticus under field conditions

Seedlings of the Mediterranean shrub Cistus creticus L. were grown in the field under ambient or ambient plus supplemental UV-B radiation (simulating a 15% ozone depletion over Patras, 38.3°W, 29.1°E) for 20 months. During this period, measurements of photosynthetic capacity, photochemical efficiency of PS II, chlorophylls and carotenoids were performed once per season. Supplemental UV-B radiation had no significant effect on these parameters nor on the total, above ground biomass accumulation, plant height and leaf specific mass measured at plant harvest. It was observed, however, that UV-B supplementation increased the number of seeds per fruit as well as mean individual seed mass. As a result, seed number and total seed mass per plant were considerably increased. Germination rates of produced seeds were not affected. We may conclude that C. creticus is a UV-B resistant plant whose competitive ability may be improved by enhanced UV-B radiation through an increase in its reproductive effort and a higher contribution to the seed bank.     

Decreased Ultraviolet-B Radiation Alters the Vertical Biomass Distribution in Cocksfoot

A greenhouse experiment was conducted to investigate whether small differences in UV-B irradiance would lead to changes in competition between cocksfoot (Dactylis glomerata L. cv. Athos) and white clover (Trifolium repens L. cv. Mervi). Plants were grown in greenhouses covered with different thicknesses of UV-transmittant plexi (3 and 5 mm) resulting in 82 % and 88 % of ambient UV-B radiation. Aboveground biomass was harvested at 4-week intervals and the vertical distribution of biomass, leaf thickness and specific leaf area were determined. Tillering, stubble and root biomass and crop height were also measured. There was only one significant effect: at 88 % of ambient UV-B radiation a larger fraction of the biomass was present in the lower layers and a smaller fraction was present in the upper layers.     

Different responses to UV-B enhanced solar radiation in radish and carrot

Radish (Raphanus sativus L.) and carrot (Daucus carota L.), plants with underground storage organs grown in the field, were exposed to either ambient (UVA) or 20 % UV-B (UVE) enhanced solar radiation till their root yield stage. In radish, UVE produced a significant increase in shoot and root fresh mass (FM), increase in the contents of chlorophyll, carotenoids, flavonoids, and total proteins per unit FM, Hill reaction rate, and root yield. In contrast, carrot responded negatively to UVE showing a loss in the above parameters.     

Effects of enhanced UV-B radiation on the hormonal content of vegetative and reproductive tissues of two tomato cultivars and their relationships with reproductive characteristics

The effects of enhanced UV-B radiation on hormone changes in vegetative and reproductive tissues of tomato (Lycopersicon esculentum Mill.) and their relationships with reproductive characteristics were studied. Two cultivars, TongHui (TH) and XiaGuang (XG), were grown in the field for one growing season under ambient (Control), ambient plus 2.54 kJ m^–2 d^–1 (T1) or ambient plus 4.25 kJ m^–2 d^–1 (T2) of supplemental ultraviolet-B (280–320 nm). The number of open flowers increased significantly in the TH cultivar under T2 while it declined in the XG cultivar under T1. Although pollen germination from both cultivars was inhibited by UV-B treatment, fruit number was enhanced in the TH cultivar at both UV-B doses and in the XG cultivar at the low dose. On the other hand, seed size (dry weight) was reduced in the XG cultivar by both UV-B doses and in the TH cultivar at the low UV-B dose. The final germination rates of seeds from control and UV-B treated plants of both cultivars showed no significant differences (p > 0.05), while germination was delayed in the TH cultivar at both doses of UV-B and in the XG cultivar only for T2. To determine the mechanism of UV-B's effects on developmental processes, hormone concentrations in leaves, pistils and seeds were analyzed using ELISA on partially purified extracts. The results suggested that enhanced UV-B radiation induced hormone changes in both vegetative and reproductive tissues. The alteration of flower number may be associated with the changes of ZR in leaves under enhanced UV-B radiation and the delayed germination may due to the changes in seed ABA and GAs.     

Effects of the deficiens allele on ear development and yield in barley

Lines partially-isogenic for the deficiens (V^tV^t and two-rowed (VV) barley ear types were developed in the genetic background of cv. Kym, for comparisons of ear development and yield components. The suppression of lateral floret growth by the V^t allele during pre-anthesis ear development occurred at a relatively late stage and only occasionally was median floret survival enhanced in the deficiens line. The final size of lateral florets in both deficiens and normal ear types appeared to be largely governed by assimilate supply. At anthesis, the lateral florets comprised about 10% of the dry matter of normal ears, whereas in the deficiens ears the median florets were correspondingly heavier. After anthesis, grain growth in the deficiens line was slower than in the normal line, although final grain weight was not affected. In another experiment the V^t allele occasionally increased the number of grains per ear, but consistently decreased the weight per grain. In drilled plot trials, the mean yield of the deficiens line was 8% lower than the normal. Excision of lateral florets at anthesis showed that photosynthesis in these organs did not contribute to the yield advantage of the normal ear type. It is suggested that the inferior performance of the deficiens type was attributable to the greater physical constriction of endosperm expansion imposed by the development of thicker, more rigid, palea and lemma tissues.     

Variation of Apigenin Quantity in Diploid and Tetraploid Chamomilla recutita (L.) Rauschert

Abstract: Apigenin, a pharmacologically important flavonoid of the chamomile plant, was analyzed at two ploidy levels during a three-year period. This flavonoid accumulates in the ligulate florets of the anthodium. Higher percentages of apigenin were found in the ligulate florets of a diploid cultivar, in comparison with tetraploid plants. However, when the total apigenin (mg of compound) in the anthodium was evaluated, tetraploid individuals accumulated significantly more flavonoid. Moreover, in contrast to morphological quantitative characteristics of the anthodium, which varied significantly in different years, apigenin percentage in the ligulate florets was constant and not influenced by environmental conditions. Apigenin content was also found to change during inflorescence ontogeny. It represents the highest percentage of dry mass in young developing florets and anthodia of both cultivars. The total apigenin content of the anthodium, however, increases during flowering, although at later stages apigenin forms only a minor part of ligulate floret and anthodium dry mass.     

Effects of elevated ultraviolet radiation on Quercus robur and its insect and ectomycorrhizal associates

Saplings of pedunculate oak (Quercus robur L.) were exposed at an outdoor facility to modulated levels of elevated UV-B radiation (280–315 nm) under treatment arrays of cellulose diacetate-filtered fluorescent lamps which also produced UV-A radiation (315–400 nm). Saplings were also exposed to UV-A radiation alone under control arrays of polyester-filtered lamps and to ambient levels of solar radiation under arrays of unenergized lamps. The UV-B treatment corresponded to a 30% elevation above the ambient level of erythemally weighted UV-B radiation. Sapling growth and the occurrence of associated organisms were examined over two years. In both years, leaves of saplings exposed to UV-B treatment were thicker and smaller in area relative to leaves exposed to ambient and control levels of radiation. UV-B treatment also retarded bud burst at one sampling in the first year of the study. Some responses were recorded which were common to both treatment and control arrays, implying that UV-A radiation, or some other factor associated with energized lamps, was responsible for the observed effects. Saplings under treatment and control arrays were taller in the first year of the study, suffered greater herbivory from chewing insects, and had lower root dry weights and greater insertion heights of secondary branches than saplings exposed to ambient levels of radiation. Exposure of saplings to elevated UV-A radiation alone under control arrays increased estimated leaf volumes in the second year of the study and reduced the number of secondary branches and the total number of branches per sapling after two years, relative to both treatment and ambient arrays. There were no effects of elevated ultraviolet radiation on shoot or total plant weight, root/shoot ratios, stem diameter, the numbers or insertion heights of primary or tertiary branches, total leaf number, timing of leaf fall or frequency of ectomycorrhizas. Our study suggests that any increases in UV-B radiation as a result of stratospheric ozone depletion will influence the growth of Q. robur primarily through effects on leaf morphology.     

Effects of UV-B radiation and additional irrigation on the Mediterranean evergreen sclerophyll Ceratonia siliqua L. under field conditions

Seedlings of Ceratonia siliqua L. were grown for 1 year in the field under ambient or ambient plus supplemental UV-B radiation (corresponding to 15% ozone depletion over Patras) and received two levels of additional irrigation during the summer dry period. The experiment was started during February 1998 and two major samplings were performed, the first at the end of the dry period (September 1998) and the second at the end of the experiment (January 1999). Plants receiving additional irrigation showed significantly higher leaf number, plant height and chlorophyll content at the end of the summer, but these differences were abolished at the final harvest. Plants growing under enhanced UV-B radiation had significantly fewer leaves and less nitrogen content at the end of the dry period, but these effects were also abolished at the final harvest, during which significant UV-B induced increases in stem dry mass were observed. None of the other measured parameters (mean leaf area, leaf dry mass, leaf thickness, UV-B absorbing compounds, phenolics, tannins and photochemical efficiency of PSII) were affected by either treatment. Combined UV-B / water effects were not significant. We may conclude that although some minor responses to enhanced UV-B radiation were evident, C. siliqua is resistant against UV-B radiation damage at the level applied.     

Influence of UV-light on carthamin accumulation and floret elongation in a cultivar of saffron thistle (Carthamus tinctorius L.)

Pre-matured florets of Benibana, a cultivar of saffron thistle (Carthamus tinctorius L.) was irradiated with UV-B (280–320 nm) or UV-C (254 nm) light for 48 h at 23±1 °C and the influence of UV-light on carthamin accumulation and floret elongation was investigated. UV-C light enhances carthamin accumulation most prominently, showing a specific value of 52.3 nmol carthamin·dm⁻³·h⁻¹·25μm⁻² (13.9 times of control), while it restricts floret elongation by a light-suppression manner (net elongation: 0.058 mm·h⁻¹, one ninth of control). UV-B light is also promotive for the red colour appearance (25.0 nmol carthamin·dm⁻³·h⁻¹·25 μm⁻², 6.7 times of control) with suppressing floret elongation (net elongation: 0.17 mm·h⁻¹, one third of control). Heterogeneous productivity of carthamin was seen in floret tissues after continuous treating under UV-C light. Carthamin accumulation, heterogeneous carthamin productivity and decrease of floret elongation restraint under UV-lights are discussed.     

Combined effects of enhanced UV-B radiation and additional nutrients on growth of two Mediterranean plant species

Seedlings of two Mediterranean plants, the slow-growing, evergreen sclerophyll Ceratonia siliqua L. and the fast growing drought semi-deciduous Phlomis fruticosa L., were grown for one year in the field at ambient or ambient plus supplemental UV-B radiation (equivalent to a 15% ozone depletion) and two levels of applied fertilizers (NPK). The effects on growth, morphological, anatomical and physiological parameters were measured at final plant harvest. Additional nutrients increased leaf nitrogen, improved growth and reduced the root/shoot ratio in both plants, yet these effects were more pronounced in the fast growing P. fruticosa. A nutrient-induced increase in chlorophyll content was also observed in this plant. The growth responses to UV-B radiation were different for the two species. Growth in C. siliqua was not affected by UV-B radiation at both nutrient levels and the same was true for P. fruticosa at low nutrients. However, at the high nutrient level, supplemental UV-B radiation improved growth in P. fruticosa, indicating a strong interaction between the treatments. Photosystem II (PSII) photochemical efficiency, methanol-extractable UV-B absorbing capacity, total phenolics and tannins were not affected by either treatment in both plants. It is concluded that nutrient levels can strongly modify the UV-B radiation effects on growth of P. fruticosa. We presume that this may be correlated to the fast growing habit of this species.     

Self fertilization and seed set in Trifolium repens L. by in situ and in vitro pollination

Summary The frequency of seed formation has been determined from self-pollination in situ (by hand) and in vitro for Trifolium repens. Selfing in situ was measured over a period of 3 years in which plants were held either at 35 °C for 24 h post-pollination (1984 and 1985) or held at ambient temperatures (1986). Mean yield of self-seed per 100 florets was 2.8 in 1984, 5.2 in 1985 and 2.2 in 1986. This was based on over 15,000 selfings per year with seven varieties and a total of 166 genotypes. In general, seed set following self-pollination was low; 53% of all genotypes set less than one seed per 100 florets selfed. Selfing of 340 excised florets in vitro with six genotypes gave a mean of 30.6 seeds per 100 florets. Temperature treatments (post-pollination) had no significant effect on seed yield in vitro. Treatment of florets in vitro for 24 h post-pollination with 0.1% CO₂ increased the yield of self seed with three genotypes but had no effect on a fourth genotype.     

A comparison of Hordeum bulbosum-mediated haploid production efficiency in barley using in vitro floret and tiller culture

Summary A high efficiency of Hordeum bulbosum-mediated haploid production in barley has been achieved using a floret culture technique in which florets pollinated with Hordeum bulbosum are cultured on modified N₆ media containing 0.5 mg/l kinetin and 1.2 mg/l2,4-D. Cultures were maintained at 25 °C with a 16 h photoperiod for 9 days before embryo rescue. In a comparison of haploid production efficiency using five F₁ hybrids from winter x winter and winter x spring barley crosses, 41.6 haploid plants/100 florets pollinated were produced using floret culture. Using detached tiller culture, 13.5 haploid plants/100 florets pollinated were produced. Higher efficiencies achieved with floret culture are attributed to the formation of larger, differentiated embryos. Such embryos lead to higher frequencies of plant regeneration. The F₁ from a winter x winter cross was inferior in haploid production compared to F₁s from winter x spring crosses. No genotype x technique interaction was observed.Oregon Agricultural Experiment Station Technical Paper No. 8653     

Respiration and heat production by the inflorescence of Philodendron selloum Koch

During a 2-d sequence of anthesis, the spadices of the thermogenic arum lily, Philodendron selloum, regulated maximum temperature within a small range (37–44°C) by reversible thermal inhibition of respiratory heat production. This response protects the inflorescence and the attracted insects from thermal damage. Heat production by whole spadices, measured by O₂ respirometry, equalled heat loss, measured by gradient layer calorimetry, which confirmed the heat equivalence of O₂ consumption (20.4 J ml^-1). This also indicated that there was no net phosphorylation during thermogenesis, heat production being the primary function of high rates of respiration. The sterile male florets consumed about 30 ml g^-1 h^-1 and the average 124-g spadix produced about 7 W to maintain a 30°C difference between spadix and ambient temperature. Most of the energy for thermogenesis is present in the florets before anthesis. Despite high respiratory rates, thermogenesis is an energetically inexpensive component of the reproductive process.     

The direct effects of UV-B radiation on Betula pubescens litter decomposing at four European field sites

A co-ordinated series of field experiments were conducted to consider the effects of elevated UV-B radiation applied directly to decomposing plant litter. Betula pubescens was decomposed under ambient and elevated UV-B (simulating a 15% ozone depletion) using outdoor irradiation facilities at Adventdalen, Norway (78° N), Abisko, Sweden (68° N), Amsterdam, The Netherlands (52° N,) and Patras, Greece (38° N). There was no significant effect of treatment on mass loss for samples collected after 2, 12 and 14 months decomposition at Amsterdam, or after 4 months decomposition at Adventdalen. Significant reductions in the mass loss of litter decomposing under elevated UV-B compared to ambient were found at the other 2 sites. The only effect of treatment on litter chemistry during decomposition was a significant reduction in the N concentration of material at Abisko and a significant increase in C:N at Patras for litter decomposing under elevated UV-B. Significant differences were found in the structure of the fungal community decomposing litter in Sweden, the only site to be tested. These data, and the few published studies of the response of decomposition to UV-B incident on litter suggest that, in the ecosystems and climates that have been studied, such direct effects are typically confined to the initial stages of decomposition, and are rather small in magnitude.     

Influence of Temperature on the Effects of Artificially Enhanced UV-B Radiation on Aquatic Bryophytes Under Laboratory Conditions

We examined, under laboratory conditions, the influence of temperature (2 °C vs. 10 °C) on the physiological responses of two aquatic bryophytes from a mountain stream to artificially enhanced UV-B radiation for 82 d. These organisms may be exposed naturally to relatively low temperatures and high levels of UV-B radiation, and this combination is believed to increase the adverse effects of UV-B radiation. In the moss Fontinalis antipyretica, UV-B-treated samples showed severe physiological damages, including significant decreases in chlorophyll (Chl) and carotenoid (Car) contents, Chl a/b and Chl/phaeopigment ratios, Chl a fluorescence parameters F_v/F_m and _PS2, electron transport rate (ETR_max), and growth. In the liverwort Jungermannia cordifolia, UV-B radiation hardly caused any physiological change except for growth reduction. Thus, this liverwort seemed to be more tolerant to UV-B radiation than the moss under the specific experimental conditions used, maybe partly due to the accumulation of UV-B absorbing compounds. The influence of temperature on the effects of UV-B radiation depended on the species: the higher the UV-B tolerance, the lower the influence of temperature. Also, different physiological variables showed varied responses to this influence. Particularly, the lower temperature used in our study enhanced the adverse effects of UV-B radiation on important physiological variables such as F_v/F_m, growth, and Chl/phaeopigment ratios in the UV-B-sensitive F. antipyretica, but not in the more UV-B-tolerant J. cordifolia. Thus, the adverse effects of cold and UV-B radiation were apparently additive in the moss, but this additiveness was lacking in the liverwort. The Principal Components Analyses (PCA) conducted for both species with the physiological data obtained after 36 and 82 d of culture confirmed the above results. Under natural conditions, the relatively high water temperatures in summer might facilitate the acclimation of aquatic bryophytes from mountain streams to high levels of UV-B radiation. This may be relevant to predict the consequences of concomitant global warming and increasing UV-B radiation.