Suboptimal thermoregulation in male adders (Vipera berus) after hibernation imposed by spermiogenesis

In ectotherms, the main behavioural option for thermoregulation is the adjustment of daily and seasonal activity to the thermal quality of the environment. While active, ectotherms thermoregulate by shuttling in between thermally differing microhabitat patches. Here, we focused on the question of whether other behavioural or physiological processes could force ectotherms to maintain activity during thermally unfavourable periods, when accurate thermoregulation is impossible. Using laboratory experiments and field data we compared the thermoregulation of male adders ( Vipera berus ) between two periods in spring when (1) only males and (2) also females and juveniles had terminated their winter hibernation. We found that males thermoregulated actively both in the lab and in the field. Accurate thermoregulation was only possible during the second period because of the low thermal quality of the environment. Male adders maintained a lower mean body temperature in the field than in the laboratory within both periods, and in addition their body temperature during the first period was on average 4 °C lower than during the second period. The thermal qualities of the natural basking sites showed a similar pattern. We discuss the results in the context of a potential trade-off between spermiogenesis and thermoregulation, where the benefits of early spermiogenesis coupled with inaccurate thermoregulation are higher than the associated costs. The results support the contention that the earlier spring emergence of the male compared with female adders is explainable by natural selection favouring early initiation of spermiogenesis, and hence sex differences in phenology.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 19–27.     

Growth and reproduction of aphids and levels of free amino acids in Scots pine and Norway spruce in an open-air fumigation with ozone

To estimate the susceptibility of conifer seedlings to aphids under future tropospheric ozone levels, Scots pine (Pinus sylvestris L.) and Norway spruce (Picea abies (L.) Karst.) seedlings were exposed to ambient and elevated ozone levels in an open-air exposure system. Growth and reproduction of the aphids Schizolachnus pineti and Cinara pinea on Scots pine and Cinara pilicornis on Norway spruce were monitored. Levels of free amino acids in foliage and young shoots were used as indicators of host plant quality. In elevated treatment plots the ozone doses were between 1.2 and 1.7 times the dose in ambient plots in 1990–93. Half of the seedling material in 1992–93 was subjected to nitrogen fertilization treatment to evaluate the effects of increased N deposition. In 1990, population density of S. pineti on pine did not differ between ambient and elevated ozone treatments during growing season, but remained higher in the elevated ozone plot than in the ambient plot at the end of the growing season. This was associated with elevated levels of glutamic acid in foliage. In August 1992, the numbers of S. pineti were consistent between the two ambient ozone plots, but deviated highly between the two ozone-fumigated plots. Glycine concentration in pine foliage was elevated by ozone, but free amino acid concentrations were not related to aphid performance. In 1993, ozone and nitrogen did not significantly affect the relative growth rate (RGR) of S. pineti or C. pinea nymphs on Scots pine, but glutamic acid concentration in foliage was increased by nitrogen fertilization. On Norway spruce, fecundity of C. pilicornis females was higher in elevated ozone treatment, but RGR of nymphs was not affected in 1992. In 1993, RGR of C. pilicornis nymphs was increased by nitrogen fertilization in June, but not affected by ozone. Nitrogen fertilization increased the levels of total free amino acids, aspartic acid, glutamic acid and proline in elongating shoots of Norway spruce, and ozone reduced the concentrations of valine and γ-butyric acid. Our results suggest that availability of nitrogen from soil has a stronger impact on the concentrations of free amino acids in conifer seedlings than ozone. Some episodes of high ozone concentration may increase free amino acids in foliage. Aphid response to ozone was extremely variable, in agreement with previous laboratory experiments. The expected 20–70% increase in ambient concentrations of tropospheric ozone may in some occasions enhance aphid performance on Scots pine and Norway spruce seedlings, but in most cases the ozone effect on the susceptibility of conifer seedlings to sucking insect pests is not important.     

Carbon dioxide-induced changes in beech foliage cause female beech weevil larvae to feed in a compensatory manner

The phenology of Fagus sylvatica was unaffected by exposure to an atmosphere of elevated CO₂ (600 μL L^-1) after two years of fumigation. Non-significant changes in nitrogen and phenolic content of the leaves decreased the nutritional status of beech for female larvae in elevated CO₂ such that they responded by eating in a compensatory manner; males were unaffected. Rates of development, mortality and adult biomass of Rhynchaenus fagi were no different from those in ambient CO₂ concentrations (355 μL L ^-1). It is possible that, with the changes in leaf chemistry affecting the females, fecundity will be altered, with important consequences for populations of beech weevil.     

Severe photobiont injuries of lichens are strongly associated with air pollution

The photobiont ultrastructure of the epiphytic lichens Bryoria fuscescens and Bryoria fremontii was studied along the pollution gradient from two Cu-Ni smelters in Nikel and Monchegorsk in northern Finland and north-western Russia. The relationship between ultrastructural characteristics of B. fuscescens and environmental factors (i.e. climate, atmospheric SO₂ and bark element concentrations) was studied by using a principal component analysis (PCA) aiming to assess the air quality in a northern environment. Based on PCA, increased plasmolysis and mitochondrial changes in the Trebouxia photobiont were significantly correlated with elevated pollutant concentrations. Degenerated cells, showing altered chloroplasts and electron-translucent pyrenoglobuli, occurred in lichens growing 35–50 km from the Monchegorsk smelter. Cell wall and cytoplasmic lipid volumes, and size of pyrenoglobuli, positively correlated with the distance from the Monchegorsk smelter. Vacuoles and electron-opaque vacuolar deposits were significantly increased at the Finnish site in the vicinity of a pulp mill. Swelling of mitochondrial cristae and thylakoids showed little correlation with environmental factors, but indicated of initial stage of injuries and were observed at several slightly polluted sites in northern Finland and north-western Russia. The results suggest that the severe photobiont injuries of lichens are strongly associated with poor air quality.     

Do elevated atmospheric CO2 and O3 affect food quality and performance of folivorous insects on silver birch?

The individual and combined effects of elevated CO₂ and O₃ on the foliar chemistry of silver birch (Betula pendula Roth) and on the performance of five potential birch‐defoliating insect herbivore species (two geometrid moths, one lymantrid moth and two weevils) were examined. Elevated CO₂ decreased the water concentration in both short‐ and long‐shoot leaves, but the effect of CO₂ on the concentration of nitrogen and individual phenolic compounds was mediated by O₃ treatment, tree genotype and leaf type. Elevated O₃ increased the total carbon concentration only in short‐shoot leaves. Bioassays showed that elevated CO₂ increased the food consumption rate of juvenile Epirrita autumnata and Rheumaptera hastata larvae fed with short‐ and long‐shoot leaves in spring and mid‐summer, respectively, but had no effect on the growth of larvae. The contribution of leaf quality variables to the observed CO₂ effects indicate that insect compensatory consumption may be related to leaf age. Elevated CO₂ increased the food preference of only two tested species: Phyllobius argentatus (CO₂ alone) and R. hastata (CO₂ combined with O₃). The observed stimulus was dependent on tree genotype and the measured leaf quality variables explained only a portion of the stimulus. Elevated O₃ decreased the growth of flush‐feeding young E. autumnata larvae, irrespective of CO₂ concentration, apparently via reductions in general food quality. Therefore, the increasing tropospheric O₃ concentration could pose a health risk for juvenile early‐season birch folivores in future. In conclusion, the effects of elevated O₃ were found to be detrimental to the performance of early‐season insect herbivores in birch whereas elevated CO₂ had only minor effects on insect performance despite changes in food quality related foliar chemistry.     

Interactions of elevated carbon dioxide and temperature with aphid feeding on transgenic oilseed rape: Are Bacillus thuringiensis (Bt) plants more susceptible to nontarget herbivores in future climate?

Climate change factors such as elevated carbon dioxide (CO₂) and temperature typically affect carbon (C) and nitrogen (N) dynamics of crop plants and the performance of insect herbivores. Insect‐resistant transgenic plants invest some nutrients to the production of specific toxic proteins [i.e. endotoxins from Bacillus thuringiensis (Bt)], which could alter the C–N balance of these plants, especially under changed abiotic conditions. Aphids are nonsusceptible to Lepidoptera‐targeted Bt Cry1Ac toxin and they typically show response to abiotic conditions, and here we sought to discover whether they might perform differently on compositionally changed Bt oilseed rape. Bt oilseed rape had increased N content in the leaves coupled with reduced total C compared with its nontransgenic counterpart, but in general the C : N responses of both plant types to elevated CO₂ and temperature were similar. Elevated CO₂ decreased N content and increased C : N ratio of both plant types. Elevated temperature increased C and N contents, total chlorophyll and carotenoid concentrations under ambient CO₂, but decreased these under elevated CO₂. In addition, soluble sugars were increased and starch decreased by elevated temperature under ambient but not under elevated CO₂, whereas photosynthesis was decreased in plants grown under elevated temperature in both CO₂ levels. Myzus persicae, a generalist aphid species, responded directly to elevated temperature with reduced developmental time and decreased adult and progeny weights, whereas the development of the Brassica specialist Brevicoryne brassicae was less affected. Feeding by M. persicae resulted in an increase in the N content of oilseed rape leaves under ambient CO₂, indicating the potential of herbivore feeding itself to cause allocation changes. The aphids performed equally well on both plant types despite the differences between C–N ratios of Bt and non‐Bt oilseed rape, revealing the absence of plant composition‐related effects on these pests under elevated CO₂, elevated temperature or combined elevated CO₂ and temperature conditions.     

Effects of elevated carbon dioxide and ozone on aphid oviposition preference and birch bud exudate phenolics

The effect of atmospheric change on birch aphid ( Euceraphis betulae Koch) oviposition preference was examined and plant characteristics that are possibly responsible for the observed effects were investigated. It was hypothesized that the increasing concentrations of CO₂ and O₃ affect singly or in combination the oviposition of birch aphids via changes in host plant characteristics. Two genotypes of field-growing silver birch ( Betula pendula Roth) trees (clones 4 and 80), which were exposed to doubled ambient concentration of CO₂ and O₃, singly and in combination, in a 3-year open-top chamber experiment, were used in an aphid oviposition preference test. It was found that elevated CO₂, irrespective of ozone concentration, increased the number of aphid eggs laid on clone 4, but not in clone 80. Several flavonoid aglycones were identified from the exudate coating of birch buds. Although elevated CO₂ and O₃ affected these phenolic compounds in clone 4, the effects did not correlate with the observed changes in aphid oviposition. It is suggested that neither bud length, which was not affected by the treatments, nor surface exudate phenolics mediate birch aphid oviposition preference.     

Emission of herbivore-induced volatile terpenoids from two hybrid aspen (Populus tremula×tremuloides) clones under ambient and elevated ozone concentrations in the field

Tropospheric ozone levels are continuously rising due to human activities in the 21st century. Although the phytotoxic impact of ozone on plants has been well documented, the effect of ozone on plant emissions has received little attention. We have conducted a field-based investigation utilizing two clones of hybrid aspen ( Populus tremula L. × P . tremuloides Michx.) in a free-air ozone concentration enrichment (FACE) facility. The effects of chronic exposure to moderately increased concentrations of ozone on insect-induced terpene emissions by these trees were investigated. We used two herbivore species, Phyllobius piri , and Epirrita autumnata , both of which can reach outbreak levels on deciduous trees in Northern Europe. Our results indicated only very small changes in emissions due to increased ozone levels, but showed induction of some terpenes, particularly the monoterpene trans- β -ocimene and the homoterpene ( E )-4,8-dimethyl-1,3,7-nonatriene, in response to insect feeding. Here, we consider the positive aspects of conducting this type of study in the field and consider the possible influences of other field-based environmental factors.