Narrow-leaved plantain (Plantago lanceolata L.), a perennial forb, flowers for virtually the full length of the growing season in temperate latitudes and as a result it is exposed to widely variable intensities of shortwave (UV-B) radiation. In order to determine effects of spring and summer levels of UV-B exposure on growth and development, representatives of 42 maternal families were grown for 85 days at 3.2 and 6.4 kJ m−2 day−1 BE300, levels corresponding to early spring and mid-summer in central Illinois. Impacts on early vegetative stages were most pronounced; early vegetative growth was decreased by higher levels of UV-B and both leaf angle (a measure of erectness) and leaf hair density were increased. At harvest, vegetative growth was significantly affected by higher levels of UV-B as well; the mass of senescent leaves and crown tissue were both decreased. Although exposure to higher levels of UV-B decreased inflorescence number by nearly 15%, it did not significantly alter reproductive biomass. Significant variation attributable to maternal families was present in nearly all measurements and the range of variation among families was wider than among UV-B treatments. A marginally significant (p=0.07) maternal family by UV-B interaction was found for the number of inflorescences, suggesting that, within populations of this plant, some small amount of genetic variation exists to allow for differential reproductive performance under a regime simulating spring and summer differences in UV-B exposure. For the most part, however, in this cosmopolitan species the level of adaptation to natural levels of variation in UV-B radiation does not differ dramatically among maternal families.
summaryIn Nicotiana sylvestris Spegazzini and Comes (Solanaceae), we examined the relationships among wounding, endogenous leaf jasmonic acid (JA) pools, and whole‐plain (WP) nicotine accumulation over a range of wounding intensities and spatial distributions, in order to explore optimal defence (OD) theory predictions. We quantitatively wounded one or four leaves and then quantified: (1) JA in damaged and undamaged leaves 90 min after wounding; (2) WP nicotine concentration after 5 d (the times when JA and nicotine attain the largest wound‐induced concentrations). We find: (1) statistically significant, positive relationships on a leaf‐by‐leaf basis among the number of leaf punctures, endogenous leaf JA, and WP nicotine accumulation; (2) that young, undamaged leaves have a higher concentration of JA than do older, undamaged leaves, and produce a greater amount of JA per puncture than older leaves, but that all leaves have the same JA content (ng JA per leaf); and (3) that a damaged leaf produces less JA when other leaves in the canopy are wounded than when it is the onh wounded leaf in the canopy, but that when it is the only wounded leaf, the phylotactically adjacent, undamaged leaves do not increase their JA concentrations. The observation that younger leaves produce more JA per puncture than do older leaves is consistent with OD theory predictions. The observation that a small amount of damage localized to a single leaf is as effective as a larger amount of damage dispersed across the canopy in increasing leaf JA and WP nicotine accumulation shows the plant's ability to differentiate between dispersed and localized damage. Because the quantity of JA in a wounded leaf 90 min after wounding is a reliable indicator of the WP nicotine response to wounding, this trait provides insight into how plants integrate information about environmental insults and tailor their defence responses.
Jasmonic acid (JA) is thought to be part of a signal-transduction pathway which dramatically increases de-novo nicotine synthesis in the roots and increases whole-plant (WP) nicotine pools in response to the wounding of the leaves in Nicotiana sylvestrisSpegazzini and Comes (Solanaceae). We report the synthesis of a doubly labeled JA ([1, 2-13C]JA) and use it as an internal standard to quantify by gas chromatography-mass spectrometry the changes in root and shoot JA pools in plants subjected to differing amounts of standardized leaf wounding. Wounding increased JA pools 10-fold locally in damaged leaves within 90 min and systemically in the roots (3.5-fold) 180 min after wounding. If JA functions as an intermediary between stimulus and response, quantitative relationships among the stimulus, JA, and the response should exist. To examine these relationships, we varied the number of punctures in four leaves and quantified both the resulting JA in damaged leaves after 90 min and the resulting WP nicotine concentration after 5 d. We found statistically significant, positive relationships among number of leaf punctures, endogenous JA, and WP nicotine accumulation. We used two inhibitors of wound-induced nicotine production, methyl salicylate and indole-3-acetic acid, to manipulate the relationships between wound-induced changes in JA and WP nicotine accumulation. Since wounding and the response to wounding occur in widely separated tissues, we applied inhibitors to different plant parts to examine their effects on the local and systemic components of this response. In all experiments, inhibition of the wound-induced increase in leaf JA 90 min after wounding was associated with the inhibition of the nicotine response 5 d after wounding. We conclude that wound-induced increases in leaf JA are an important component of this long-distance signal-transduction pathway.
Both herbivory and mechanical damage result in increases in the concentration of the wound-signal molecule, jasmonic acid (JA), and the defense metabolite, nicotine, in native tobacco plants, Nicotiana sylvestris Speg. et Comes (Solanaceae). We found that higher concentrations of JA resulted from herbivory by Manduca sexta (L.) larvae than from the mechanical damage designed to mimic the herbivory. While both herbivory and mechanical damage increased JA concentrations in roots of wounded plants, herbivory did not induce either higher root JA or nicotine responses than mechanical damage. In a separate experiment in which mechanical damage was not designed to mimic herbivory, JA responses to herbivory were higher than those to mechanical damage, but the whole-plant (WP) nicotine responses were smaller. Furthermore, when regurgitants from M. sexta larvae were applied to standardized mechanical leaf wounds, leaf JA responses were dramatically amplified. However, neither the root JA response nor the WP nicotine response was comparably amplified by application of regurgitants. Our findings demonstrate that the response of N. sylvestris to herbivory is different from its response to mechanical damage; moreover, oral secretions from larvae may be partly responsible for the difference. During feeding, M. sexta larvae appear to modify the plant's normal defensive response to leaf wounding by reducing the systemic increase in root JA after leaf damage and the subsequent WP nicotine response.
1. Foraging patterns of the squash beetle, Epilachna borealis Fabricius (Coleoptera: Coccinellidae), in field cages are not consistent with the hypothesis that this beetle avoids leaves in which its feeding damage has caused the induction of cucurbitacins.2. E. borealis feeding does not induce accumulation of cucurbitacins inside or outside of feeding trenches.3. Cucurbitacins do not reduce the survival or growth rate of E. borealis larvae.4. High concentrations of cucurbitacins fail to deter E.borealis feeding; rather, cucurbitacins are feeding stimulants for squash beetles.5. Trenching behaviour in E.borealis is facultative and correlates with leaf turgor and the degree to which sticky phloem sap oozes from damaged leaves.6. Mucilaginous phloem sap in many curcurbit species inhibits feeding by E. borealis when it is applied to beetle mouthparts.
Projected decreases in stratospheric ozone may result in increases in shortwave ultraviolet (UVB) irradiation at the earth's surface. Furanocoumarins, phototoxic compounds found inCitrus jambhiri foliage, increase in concentration when these plants are grown under enhanced UVB. Survivorship schedules ofTrichoplusia ni (Lepidoptera: Noctuidae) caterpillars reared on plants in the presence and absence of enhanced UVB regimes differ significantly; larvae develop more slowly in early life when reared on plants exposed to increased UVB. This same developmental pattern is observed whenT. ni larvae are reared on artificial diets amended with ecologically appropriate amounts of furanocoumarins. Thus, anthropogenically derived changes in stratospheric ozone and concomitant changes in UV light quality at the earth's surface may influence ecological interactions between insects and their host plants by altering secondary metabolism and hence foliage quality for herbivores.
Two ecotypes of Arabidopsis thaliana, collected from Libya and Norway, were grown in the greenhouse under. UV-B doses of 0 and 10.5 kJ m[sup [minus]2] UV-B[sub BE]. The high UV-B dose simulated midsummer ambient conditions over Libya and a 40% reduction in stratospheric ozone over Norway. The Libyan ectotype, which originated from latitudes where solar UV-B is high, showed no UV-B induced damage to plant growth. However the Norwegian ecotype, which originated from latitudes where solar UV-B is low, showed a significant reduction in plant height, inflorescence weight, and rosette weight in response to enhanced UV-B. Although fruit and seed number for both ecotypes were unaffected by enhanced UV-B radiation the germination success of the seeds harvested from the irradiated Norwegian plants were significantly reduced. The two ecotypes also differed with respect to their accumulation of kaempferol, a putative UV-B protective filter. The Libyan ecotype increased kaempferol concentration by 38% over the 0 kJ treatment whereas the Norwegian ecotype increased by only 15%. These data suggest that, for these ecotypes, variation in UV-B sensitivity may be explained by the differential induction of UV-absorbing leaf pigments.
Extracts ofCitrus jambhiri foliage exposed to and shielded from UV-B radiation were assayed for phytochemical changes and phototoxicity against four fungal pathogens, two of which (Fusarium solani andF. oxysporum) are causative agents of root rots and two of which (Penicillium italicum andP. digitatum) are associated with fruit rots. Conidial pigment mutants of these four fungal species were assayed to determine whether pigments play a role in protecting fungi against plant photosensitizers. Exposure to 10.2 kJ/ day UV-B radiation for 95 days significantly reduced phototoxicity of leaf extracts to fungi. Although furanocoumarin levels were reduced by UV-B, analysis of covariance revealed that variation in phototoxicity of the extracts cannot be attributed entirely to variation in furanocoumarin content; thus, the possibility exists that nonfuranocoumarin phototoxic constituents, as yet unidentified, respond to UV-B exposure and contribute to overall phototoxic defense ofC. jambhiri against pathogens. Root rot fungi were substantially more sensitive to furanocoumarin phototoxicity than were fruit rot fungi, a pattern consistent with the amount of light exposure normally experienced by these fungi when associated with phototoxic plants. Although pigmented strains of all four species displayed greater resistance to phototoxicity of pure furanocoumarins, no strain differences were detected in assays of foliar extracts; this finding also suggests that nonfuranocoumarin constituents may be involved in the phototoxic defense ofC. jambhiri against pathogens.
Extracts of foliage exposed to and shielded from UV-B radiation were assayed for phytochemical changes and phototoxicity against four fungal pathogens, two of which ( and) are causative agents of root rots and two of which ( and) are associated with fruit rots. Conidial pigment mutants of these four fungal species were assayed to determine whether pigments play a role in protecting fungi against plant photosensitizers. Exposure to 10.2 kJ/ day UV-B radiation for 95 days significantly reduced phototoxicity of leaf extracts to fungi. Although furanocoumarin levels were reduced by UV-B, analysis of covariance revealed that variation in phototoxicity of the extracts cannot be attributed entirely to variation in furanocoumarin content; thus, the possibility exists that nonfuranocoumarin phototoxic constituents, as yet unidentified, respond to UV-B exposure and contribute to overall phototoxic defense of against pathogens. Root rot fungi were substantially more sensitive to furanocoumarin phototoxicity than were fruit rot fungi, a pattern consistent with the amount of light exposure normally experienced by these fungi when associated with phototoxic plants. Although pigmented strains of all four species displayed greater resistance to phototoxicity of pure furanocoumarins, no strain differences were detected in assays of foliar extracts; this finding also suggests that nonfuranocoumarin constituents may be involved in the phototoxic defense of against pathogens.
Seeds of P. lanceolata were collected from a local population and 4 replicates of 42 maternal families were grown for 90 days in the greenhouse with at two levels of supplemental UVB radiation (6 and 12 kJ day[sup [minus]1] BE[sub 300]). Higher UVB radiation increased leaf hair density and decreased plant size during early growth; family identity affected these also. Leaves excised from a subset of the plants were fed to ultimate instar larvae of J. coenia and assayed for iridoids. Increased UVB radiation did not alter the iridoid content of the leaves or the growth of the larvae. In a separate experiment, P. lanceolata growing under the two levels of UVB irradiation were infested with neonate larvae and larval growth was monitored. Larval growth was not markedly altered by enhanced UVB. These findings suggest that increased UVB is unlikely to alter the suitability of P. lanceolata as a host for J. coenia.
Rooted cuttings of C. jambhiri were grown under enhanced levels of UVB radiation for 95 days. Bacterial phototoxicity and furanocoumarin content were determined in extracts made from various tissues from the aboveground biomass. Young, newly expanded leaves contained significantly higher concentrations of furanocoumarins than older leaves and stems. Additionally, the proportional concentration of psoralen was higher in young leaves than in old leaves. While treatment with UVB did not result in a change in the overall level of furanocoumarins, it did cause an increase in the ratio of psoralen to bergapten. Bacterial phototoxicity paralleled the distribution of furanocoumarin content among tissue types; analysis of covariance suggested that the phototoxic properties of the extracts could be accounted for on the basis of furanocoumarin content alone.
Abstract. 1. Reproductive costs associated with flight capability were evaluated in the wing dimorphic planthopper, Prokelisia dolus Wilson, by comparing the life history of traits of winged (macropterous) and flightless (brachypterous) females under controlled laboratory conditions.2. Macropters with large thoraces and fully developed wings maintain a greater investment in flight apparatus than brachypters with small thoraces and reduced wings.3. Associated with greater flight capability in the macropter of P.dolus are shorter adult life, decreased total fecundity, and delayed age at first reproduction compared to brachypterous females.4. Under field conditions where mortality is high, the difference in realized fecundity between the two wing forms living on similar resources is further exaggerated with the brachypter having the greater advantage.5. When the life history traits of the wing forms of P. dolus are compared with traits for nine other species of planthoppers, two similarities emerge. First, the preoviposition period of the macropterous wing form is always longer than that for the brachypter resulting in a reproductive delay. Second, most studies show that macropters are less fecund than brachypters.6. There is no general tendency among planthopper species for macropterous adults to live fewer days or develop more slowly as nymphs compared to their flightless counterparts.7. The reproductive delay and reduced fecundity of the volent wing form of planthoppers supports the notion that flight capability is costly and that phenotypic trade‐offs between flight and reproduction exist.
The relationship between body length (mm) and average daily fecundity (no. eggs per female per day) of field-collected Prokelisia marginata was investigated in the greenhouse. There is significant positive relationship between body length and average daily fecundity, and the functional relationship is the same for both macropterous and brachypterous wing forms. Because female body size differs among the natural habitats of this species, the predicted fecundity varies as well. We suggest that habitat variation in body size-mediated fecundity in part forms the adaptive basis for migration in P. marginata.