Acclimatisation in the green spruce aphid, Elatobium abietinum

Thin layer chromatography separation of 80% ethanol extracts of adult Elatobium abietinum revealed the presence of the polyhydric alcohol mannitol in aphids overwintering outdoors but not in aphids kept permanently indoors at 15°C. After 3 days at 15°C no traces of mannitol were left in overwintering aphids. Mean freezing temperatures of outdoor, unfed instar I nymphs were about 4°C lower than those of unfed instar I nymphs produced at 15°C. Mean freezing temperatures of overwintering adults were considerably higher than those of unfed instar I nymphs and showed no changes associated with time at 15°C following transference indoors. Similarly, mean freezing temperatures of Sitka spruce needles transferred to 15°C did not change. It was concluded that, although freezing was mainly avoided by supercooling, the presence of mannitol lowered the true freezing temperature of aphid haemolymph and, consequently, the actual freezing temperatures of nymphs produced under cold conditions. However, the considerable increase in freezing point temperatures caused by imbibition of plant sap masked these acclimatisation changes in feeding aphids.     

Flight regulation in the green spruce aphid (Elatobium abietinum)

In Britain the flight of alate parthenogenetic viviparae of Elatobium abietinum was found to occur only in early summer. Data for this study were extracted from suction trap catches and weather records obtained from meteorological stations adjacent to the traps. The duration of the flight period increased with latitude. At 51° N 95 % of the trap catch was taken in 24 days and at 55° N in 32 days. A flight threshold temperature of 12–13°C was deduced from trap catches taken within the forest crop. Displacements in median flight days at various sites in Britain could be related to values of accumulated day-degrees but not to differences in photoperiod. The median flight day can be predicted empirically by accumulating day-degree values over 8°C from 1 January up to a total of 115 day-degrees. The implications of seasonal flight activity on crop reinvasion is discussed.     

Supercooling and the low-temperature survival of the green spruce aphid Elatobium abietinum

Laboratory investigations into the low-temperature tolerance of the green spruce aphid, Elatobium abietinum, revealed that the insect was killed by freezing. Aphids and host Sitka spruce needles showed similar seasonal changes in supercooling ability. A noticeable increase in this ability occurred between June and October. Aphids were more susceptible to low temperatures when attached to the plant. It is suggested that mortality resulted from ice which formed in the sap of the host needles and spread into the feeding aphids via their mouthparts. Neither the chlorotic banding of needles, caused by aphid feeding, nor needle length affected needle supercooling. Increased duration of exposure increased the probability of freezing of supercooled needles at low temperatures. A small percentage of first-instar nymphs supercooled to much lower temperatures than the remainder of the population. These were newly born nymphs whose high supercooling ability markedly decreased when they began to feed.     

Damage and performance of the green spruce aphid, Elatobium abietinum on twenty spruce species

Field and laboratory observations of the relationships between the performance of Elatobium abietinum (Walker), Homoptera, Aphididae, and various species of spruce were undertaken from January 1980 to April 1981. The study included sampling for aphids in established field plots of spruce during May and June respectively before and after the migration period in spring. The aphid's performance (weight and mean relative growth rate) at different seasons on pot grown plants of selected spruce species was monitored, covering, in all, 20 species of spruce.Aphid performance was greatest on the North American spruces, especially Picea sitchensis (Bong) Carr and P. mexicana Martinez; the Asian spruces were the least favoured, especially P. glehnii (Schmidt) Mast. Between these two geographical groups the Eurasian spruce species (sensu Wright, 1955) demonstrated an intermediate aphid performance.

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Résumé Les observations dans la nature et au laboratoire sur les performances d'E. abietinum Walker et différentes espèces d'épicéas ont été effectuées de janvier 1980 à avril 1981. Les observations dans la nature comprenaient des prélèvements de pousses d'épicéa pour dénombrer les pucerons avant (mai) et après (juin) la période de migration du printemps. Les performances des pucerons (poids et taux moyen de croissance relative) ont été examinées à différentes saisons sur des plantes en pot sur un total de 20 espèches sélectionnées d'épicéas.Les performances du puceron ont été supérieures sur les espèces d'épicéa de l'Amérique du nord Picea sitchensis et P. mexicana; les espèces asiatiques étaient les moins favorables, particulièrement P. glehnii. Entre ces deux groups géographiques, les espèces eurasiennes (sensu Wright, 1955) ont permis des performances intermédiares chez les pucerons.

     

Summer survival of the green spruce aphid,Elatobium abietinum,in North East Scotland

Summary In North East Scotland small numbers of all developmental stages of the green spruce aphid survived on Sitka spruce needles during the summer months despite the nutritional inadequacy of these needles for aphid survival following population collapse in early summer. The surviving adults lost weight and fat reserves in response to time and contained low numbers of embryos. No metabolic acclimatization of the respiration rate occurred in response to exposure to different temperatures. Aphid respiration rates during summer were significantly higher than those in winter. Therefore, it was concluded that no summer aestivation occurred. Summer survival was dependent on the ability of a few aphids of all instars to survive on the nutritionally inadequate host, these aphids possibly possessing higher than normal fat reserves or feeding on marginally nutritionally superior trees or shoots.     

Effects of temperature on overwintering populations of the green spruce aphid Elatobium abietinum

Recorded minimum temperatures of – 7 ^oC or lower noticeably reduced overwintering populations of the green spruce aphid, Elatobium abietinum, in north-east Scotland. It is suggested that, at these temperatures, ice formation in the needles of the host Sitka spruce, caused attached aphids to freeze. Aphid mortality also occurred when maximum temperatures did not rise above + 6 ^oC for prolonged periods, possibly as a result of starvation following an extended chill coma. A diagrammatic representation of the main factors affecting anholocyclic populations of E. abietinum during the winter is presented to emphasize the governing role played by temperature. The balance between mortality and recruitment determines the size of the population at the end of the winter, and this in turn determines the subsequent summer infestation. It should be possible, therefore, to predict aphid outbreaks either from winter temperatures or from the number of aphids present at the end of the winter. Temperature records obtained from integrating thermometers indicated that the inside of the lower crown tended to be the warmest part of the tree during the winter, resulting in greater aphid survival in the lower branches.     

Factors determining the distribution of the green spruce aphid, Elatobium abietinum, on young and mature needles of spruce

1. The green spruce aphid Elatobium abietinum , was less active and more likely to feed when on mature rather than young needles of Sitka spruce.
2. Compared with survival on mature needles, survival of aphids on young needles was significantly reduced, especially on newly flushed needles, but became less so as the needles matured. The survival of aphids kept close to, but not in contact with, young needles was similarly reduced.
3. On young needles stripped of epicuticular wax, aphids were less active and fed and survived for longer than aphids on untreated young needles. In contrast, the supply of additional amino acids to shoots bearing young needles did not significantly improve the survival of aphids on these needles.
4. In a choice test, aphids were able to discriminate between epicuticular wax from young and mature needles. The reciprocal transfer of wax between young and mature needles reversed the aphids' marked preference for mature needles.
5. These results indicate that the aphid is deterred from settling and feeding on young needles by a volatile substance present in the epicuticular wax.     

Genetic differentiation of the green spruce aphid (Elatobium abietinum Walker), a recent invader to Iceland

1 The RAPD method (Random Amplified Polymorphic DNA) was used to investigate genetic diversity of the green spruce aphid, Elatobium abietinum Walker, a pest introduced recently to Iceland. 2 This aphid in Iceland comprised two polymorphic populations, one in the east and the other in the west of the country. The genetic variation between sites within a population was continuous and appeared to be in good agreement with geographical distances. 3 In the eastern population the variation was greater between sites than within sites, whereas in the western population the pattern of variation appeared to be the opposite. This overall greater genetic variation in the eastern population could be due to its having been colonized earlier than the western one. 4 The study also demonstrated a close relationship between the green spruce aphid in Iceland and aphids from Denmark, which agrees with their assumed origin. The differences in introduction time, adaptation and competitiveness between the two Icelandic populations are discussed.     

Systematic differences in the population density of green spruce aphid, Elatobium abietinum in a provenance trial of Sitka spruce, Picea sitchensis

Quantitative bioassay techniques were used to measure the susceptibility of Heliothis armigera to three nuclear polyhedrosis viruses (NPVs): H. armigera singly-enveloped NPV (HaSNPV), H. zea SNPV (HzSNPV) and H. armigera multiply-enveloped NPV (HaMNPV). Viruses were identified by EcoRI restriction endonuclease analysis. Electrophoretic profiles of DNA fragments revealed that the HaSNPV isolate was a previously undescribed genotypic variant. Bioassays with neonate and 6-day-old larvae measured small but significant differences in virulence between the three viruses. HzSNPV was the most virulent for neonate larvae with a median lethal dose (LD₅₀) of five polyhedra. HaMNPV was least virulent for 6-day-old larvae, with a LD₅₀ of 1400 polyhedra compared with 640–670 polyhedra for HaSNPV and HzSNPV. In addition, the median lethal time (LT₅₀) for infection with HaMNPV in neonate larvae was approximately 1·7 days longer than for the other viruses. Although they varied in virulence, each of the three viruses was sufficiently virulent to have considerable potential as a microbial control agent of H. armigera.     

Consistency of resistance to attack by the green spruce aphid (Elatobium abietinum Walker) in different ontogenetic stages of Sitka spruce

Abstract 1 The susceptibility of different genotypes of 29‐year‐old Sitka spruce to damage by the green spruce aphid, Elatobium abietinum, was investigated in a progeny trial where aphid damage on individual trees had previously been assessed twice in an earlier stage of ontogenetic development. The progeny trial comprised 14 open‐pollinated families originating from a clonal seed orchard that had been established using mature spruce trees selected for aphid resistance. 2 Previous investigations had demonstrated that resistance was inherited by the offspring, and that differences in resistance between progenies of the individual orchard clones were highly significant. 3 Susceptibility to aphid attack was recorded as the percentage loss of previous year's needles. Differences in susceptibility recorded between the juvenile trees were found to persist after the trees had developed into the closed‐canopy, sexually reproducing stage. Needle loss of the families was significantly less than that of the reference population of Sitka spruce. 4 Hybrids between Sitka spruce and white spruce were defoliated more heavily than pure Sitka spruce, and the difference increased with age. 5 Family heritability of resistance was estimated as 0.60 compared to 0.73 when the trees were assessed in the juvenile stage. The genetic correlation based on family means between damage in the juvenile and sexually reproducing stand was high (0.83), indicating a high consistency of resistance to the aphid over years and ontogenetic stages. 6 A skewed distribution of defoliation indicated that major genes are involved in the expression of resistance, and that the genetics behind resistance has a nonadditive component.     

Interactions between green spruce aphid (Elatobium abietinum (Walker)) and Norway and Sitka spruce under high and low nutrient conditions

1 Green spruce aphid (Elatobium abietinum) is a serious pest of spruce (Picea spp.) in north‐west Europe, causing defoliation of one‐year‐old and older needles. 2 Relationships between population development of E. abietinum, needle loss and tree growth were compared for five pure genotypes of Sitka spruce and mixed‐genotype material of Sitka and Norway spruce, grown under high and low nutrient conditions. 3 Despite wide differences in flushing date between spruce genotypes, E. abietinum populations peaked on the same date on each genotype and on the mixed‐genotype material, irrespective of nutrient supply. 4 Larger aphid populations developed on trees grown under high nutrient conditions than under low nutrients. However, more needles were lost per aphid in the low nutrient treatment and overall defoliation rates in the two nutrient treatments were similar. 5 Total aphid numbers differed significantly between Sitka spruce genotypes within nutrient treatments, but not in relation to bud‐burst or needle terpene content. Reductions in height growth caused by infestation were greater (15–44%), and related to mean aphid densities and defoliation, in the low nutrient treatment, but were smaller (11–27%) and not related to aphid density and defoliation in the high nutrient treatment. 6 Development of E. abietinum populations was similar on Norway and Sitka spruce, but Norway spruce lost fewer needles. However, the effects of infestation on tree growth were more closely related to aphid density and were similar for Norway and Sitka spruce. 7 Infestation caused a decrease in total root dry weight of Norway and Sitka spruce in proportion to the reductions observed in above‐ground growth.     

The effect of needle age on the acceptability of Sitka spruce needles to the aphid,Elatobium abietinum (Walker)

Summary A comparison of the feeding behaviour of E. abietinum on current year needles and previous year needles of P. sitchensis revealed that during the summer months the aphids preferentially settled on previous year needles, this response not being evident in winter. Aphids on current year needles in summer took a much longer period of time to commence sap uptake than in winter, intake ceasing following a very short feeding period. On previous year needles sap uptake in December commenced after a shorter feeding period than in June. Analysis of total and soluble nitrogen levels in Sitka spruce needles showed that current year needles had initially higher levels during shoot elongation in May and early June, but that previous year needles had higher levels for most of the remainder of the year. Quantitative analyses of amino acids revealed that in current year needles the levels were generally lower than in previous year needles. Less marked proportional differences were observed between previous year needles in May and in July/August when the needles were unsuitable. Addition of amino acids in solution into cut current year shoots resulted in increased longevity on shoots containing introduced iso-leucine, histidine and methionine and revealed a general imbalance of the amino acids.     

The effects of nutrition and density on the production of alate Elatobium abietinum on Sitka spruce

Summary Observations over a period of 10 years showed that, in Northeastern Scotland, alatae of E. abietinum regularly appeared in mid-May, the timing being unrelated to aphid density. The peak number of alatae produced was, however, correlated with aphid density. Following an initially high level the proportion of alatae dropped to virtually nil by mid-June, whilst over the same period the aphid population density increased. Amino acid levels in spruce needles were considerably higher during the period of alate formation than they were at the termination of alate production. It is suggested that a high amino acid level was the main factor controlling the formation of alatae and that population density affected the proportions of these alatae only when nutritional levels were favourable for alate formation.     

Effects of Elatobium abietinum on nutrient fluxes in Sitka spruce canopies receiving elevated nitrogen and sulphur deposition

1 In a field acid mist simulation experiment, Sitka spruce (Picea sitchensis) was sprayed with different pollutant treatments: N, NH₄NO₃; S, Na₂SO₄; NS Acid, NH₄NO₃ + H₂SO₄ and control, no spray. Treatment effects on the abundance of the green spruce aphid Elatobium abietinum and honeydew production were assessed. In addition, needles were sampled for phyllosphere micro‐organisms. In a manipulative experiment, shoots were established and maintained as with or without E. abietinum infestation in order to determine the effects of infestation on needle loss and throughfall nutrient fluxes. 2 Aphid numbers were highest during the end of May and early June, with almost twice as many needles infested in the NS Acid treatment compared with the other treatments. Honeydew production was not affected by the treatments. 3 On infested shoots, increasing numbers of yellowing and dead needles were recorded above the throughfall collectors as the season progressed. The numbers of dead needles falling into the collectors were significantly higher beneath infested shoots. There were strong positive correlations between aphid numbers above the throughfall collectors, the number of yellowing and dead needles on the shoots and the number of needles in the funnels of the throughfall collectors. Litter production was more affected by aphid number than by pollutant treatment. 4 Bacteria, yeasts and filamentous fungi were more prolific on infested needles and treatment effects on colony forming units (CFUs) were most pronounced in the NS Acid treatment. 5 Fluxes of inorganic nitrogen beneath infested shoots were generally lower than beneath uninfested shoots. This effect was more pronounced in those treatments that supplied N i.e. N, NS Acid. The combination of aphid infestation and N‐addition exerted the strongest influence on nutrient fluxes. The fluxes of potassium and of organic carbon (DOC) were higher beneath infested shoots in all treatments, through most of the survey period.     

The growth and decline of a population of the spruce aphid Elatobium abietinum during a three year study,and the changing pattern of fecundity,recruitment and alary polymorphism in a Northern Ireland Forest

The population size and structure of the green spruce aphid was followed throughout the spring — summer cycle on the same group of trees in a low-elevation coastal Sitka spruce forest for three consecutive years. The relationship between the pattern of change and the phenology of bud burst, which heralds a marked change in needle sap quality, suggests that yearly differences in the winter temperature regime may affect the duration of the population growth phase and hence the peak numbers attained in late spring.An index of population growth rate was sufficiently sensitive to aphid fecundity during the population cycles of two years to suggest that the changing rate of fertility was the decisive process governing changes in population size. The commitment of aphids to alate development was greater than that recorded elsewhere in Britain but did little to effect population decline since the contribution of alatae to larviposition was substantial while seasonally pulsed.     

Differences in the probing behaviour of Elatobium abietinum feeding on Sitka and Norway spruces

The probing behaviour of Elatobium abietinum (Walk.) on the needles of Norway spruce and Sitka spruce was directly related to the stomatal frequency accompanied by heavy deposition of saliva, and multiple branching and divergence of the salivary sheath, all of which were associated with the impenetrability of the endodermis. On Norway spruce these features were very greatly reduced where there was a higher rate of probing coupled with a higher rate of penetration into the phloem, which was not accompanied by early uptake of phloem sap when compared with aphids on Sitka spruce. These differences in probing behaviour are related to damage symptoms exhibited by the host trees.     

Influence of the green spruce aphid on defoliation and radial stem growth of Sitka spruce

Premature abscission of older Sitka spruce foliage on trees within forests in Northern Ireland was compared with green spruce aphid population levels 2 to 3 months earlier. Each aphid has a lower relative impact on needle loss at higher aphid population density; nevertheless, more aphids on a tree results in greater rates of needle loss. Evidence from horizontal stem increment sequences suggested that aphid feeding inhibited incremental growth, and that this inhibition might be delayed for up to two field seasons. However, the effect of aphids on stem growth in most years and on most trees seems relatively small compared with the potential of other tree-or year-specific processes.     

Northern Atlantic Oscillation effects on the temporal and spatial dynamics of green spruce aphid populations in the UK

1. The role of climate variability in determining the spatial and temporal patterns of numerical fluctuations is a central problem in ecology. The influence of the North Atlantic Oscillation (NAO) index on the population dynamics and spatial synchrony of the green spruce aphid Elatobium abietinum across the UK was shown. 2. Fifteen overlapping time series within the UK were analysed; we used nonparametric models for determining the feedback nonlinear structure and the climatic effects. The spatial synchrony of these populations and the relationship between synchrony and NAO was estimated. 3. From the 15 time series across the UK, 11 showed positive and significant NAO effects. In most of the cases the NAO effects were nonlinear showing strong negative effects of low values. The NAO variation improve the explained variance of the first-order feedback models in 14.5%; ranging from 0% to 48%. All data showed strong-nonlinear (concave) feedback structure. In most of the localities the explained variance by the first-order feedback was about 50-60%. 4. The spatial synchrony of the per capita growth rates and residuals is high across long distances for those populations affected by NAO. The correlation function predicts a spatial scale of synchrony of about 350-400 km for NAO influenced populations. 5. We think that simple population theoretical models describing the link between NAO fluctuations and green spruce aphid dynamics may be fundamental for predicting and simulating the consequences of different climatic scenarios of the future.     

Non-linear feedback processes and a latitudinal gradient in the climatic effects determine green spruce aphid outbreaks in the UK

The role of climatic fluctuations in determining the dynamics of insect populations has been a classical problem in population ecology. Here, we use long-term annual data on green spruce aphid populations at nine localities in the UK for determining the importance of endogenous processes, local weather and large-scale climatic factors. We rely on diagnostic and modelling tools from population dynamic theory to analyse these long-term data and to determine the role of the North Atlantic Oscillation (NAO) and local weather as exogenous factors influencing aphid dynamics. Our modelling suggests that the key elements determining population fluctuations in green spruce aphid populations in the UK are the strong non-linear feedback structure, the high potential for population growth and the effects of winter and spring weather. The results indicate that the main effect of the NAO on green spruce aphid populations is operating through the effect of winter temperatures on the maximum per capita growth rate (R_m). In particular, we can predict quite accurately the occurrence of an outbreak by using a simple logistic model with weather as a perturbation effect. However, model predictions using different climatic variables showed a clear geographical signature. The NAO and winter temperature were best for predicting observed dynamics toward the southern localities, while spring temperature was a much better predictor of aphid dynamics at northern localities. Although aphid species are characterized by complex life-cycles, we emphasize the value of simple and general population dynamic models in predicting their dynamics.