Foliar chemical concentrations in red alder stands of various ages

Summary Six stands of red alder (Alnus rubra Bong.) were sampled to assess the relationship between stand age and foliar chemical concentrations. The stands ranged in age from 9 to 45 years, and were located near Olympia, Washington, on sites of similar soil, aspect, and slope. Foliage was collected in July, and concentrations of most essential nutrients and aluminum were determined. Foliar concentrations of these elements varied among the different stands. Highest concentrations of N, P, Ca, S, Fe, and Al were found in trees of the youngest stand. Concentrations of N, P, S, and Fe were negatively correlated with stand age. The strongest correlation was obtained with P (r=–0.96,P<0.01), followed by N (r=–0.85,P=0.03). Results suggest that declining growth and deterioration of red alder stands with age may be related, at least in part, to limiting supplies of essential nutrients, especially P, N, and S.     

Nodulation potential of soils from red alder stands covering a wide age range

Red alder (Alnus rubra Bong.) stands in the Pacific Northwest are the common first stage in succession following disturbance. These stands are highly productive and contribute a large amount of N to the soils as a result of their N₂-fixing symbiosis with Frankia. As these alder stands age, the soils not only increase in total N, but concentrations of NO^– ₃ increase and pH decreases as a result of nitrification. The objective of this study was to determine how the nodulation capacity of Frankia varies as red alder stands age and if differences in nodulation capacity are related to changes in soil properties. Nodulation capacity was determined by a red alder seedling bioassay for soils from red alder stands in the Oregon coast range covering a wide range of ages. Six chronosequences were sampled, each containing a young, an intermediate, and an older alder stand. Soil total N, total C, NO^– ₃, NH⁺ ₄, and pH were measured on the same soil samples. These factors as well as alder stand characteristics were compared with nodulation capacity in an attempt to identify soil characteristics typical in developing alder stands that most strongly affect nodulation capacity. Soil pH and NO^– ₃ concentration were highly correlated with nodulation capacity and with each other. Cluster analysis of the sites using these two variables identified two groups with distinctly different nodulation capacities. The cluster with the higher nodulation capacity was lower in NO^– ₃ and higher in pH than the other cluster, which included the majority of sites. There was substantial overlap in the age ranges for the two clusters and there was no significant correlation between age and nodulation capacity. Thus nodulation capacity appears to be most closely related to soil properties than to stand age.     

Nutrient relations in coppiced black cottonwood and red alder

Two-year-old coppice of black cottonwood and red alder, grown in pure culture and in mixture, were compared using terminal twigs and leafless shoots harvested in the winter. Terminal twigs were taken with buds intact; they were about 15 cm long. Leafless shoot samples included all above-ground components. In pure culture, dry weights of the leafless shoots per plant were similar for the two species. In mixture with alder, however, weight of the cottonwood plants was enhanced and that of alder was reduced, but neither response was statistically significant. Nutrient concentration, content per plant, and utilization varied by the plant tissues analyzed, cultural treatment (purevs. mixed), and species. In general, nutrient concentrations were higher in the terminal twigs than in the leafless shoots of both species. Cultural treatment did not significantly affect nutrient concentration in cottonwood twigs or in the leafless shoots of either species. Concentrations of N and Fe were significantly higher and those of Mn were lower in twigs of mixed alder than in twigs of pure alder. Twigs of cottonwood were significantly higher than those of alder in concentration of P and Zn, and lower in N, Mn, and Cu. Compared with alder, cottonwood leafless shoots were significantly higher in concentration of Ca, but lower in N, S, Cu, and Mn. With few exceptions, nutrient content was highest in the shoots of the large plants of mixed cottonwood, intermediate in medium-sized pure cottonwood and pure alder, and lowest in the small mixed alder. Cottonwood was significantly more efficient than alder in use of N, S, and Cu, and less efficient in use of Ca. Some of the differences between cultural treatments and species may be associated directly or indirectly with the N₂-fixing ability of red alder. Mixed culture of the two species appears promising because of the increased growth of cottonwood. Planted separately in pure culture, the choice between cottonwood and alder may be determined, in part, by the nutritional status of the soil where plantations are established.     

Inoculation and production of container-grown red alder seedlings

Summary The inoculation ofAlnus rubra (red alder) withFrankia sp. can lead to a highly efficient symbiosis. Several factors contribute to the successful establishment of nitrogenfixing nodules: (1) quantity and quality ofFrankia inoculant; (2) time and method of inoculation; (3) nutritional status of the host plant.Frankia isolates were screened for their ability to nodulate and promote plant growth of container-grown red alder. Inoculations were performed on seedlings and seeds. Apparent differences in symbiotic performance could be seen when seeds or seedlings were inoculated. Plants inoculated at planting performed significantly better than those inoculated four weeks later in terms of shoot height, nodule number and shoot dry weight. If inoculation was delayed further, reduction in shoot height, nodule number and shoot dry weight resulted. The effect of fertilizer was also investigated with regard to providing optimal plant growth after inoculation. Plants receiving 1/5 Hoagland's solution minus nitrogen showed maximal plant growth with abundant nodulation. Plants receiving 1/5 Hoagland's solution with nitrogen showed excellent plant growth with significantly reduced nodulation.     

Nitrogen cycling in dense plantings of hybrid poplar and black alder

Summary Nitrogen cycling was studied during the third growing season in pure and mixed plantings (33×33 cm spacing) of hybrid poplar and black alder in southeastern Canada. After 3 years, hybrid poplar growth and N content of living tissues in a plot and of individual hybrid poplar plants increased with the proportion of black alder in a planting. No differences were detected among N contents of individual alder plants regardless of plot treatment. Black alder allocated a larger portion of its N to roots than hybrid poplar. Symbiotic nitrogen fixation was estimated to account for 80% of the nitrogen in aboveground alder tissues in the pure treatment using natural¹⁵N dilution. N return in leaf litter was estimated to be 70kg ha^–1 in the pure alder treatment and decreased to a minimum of 20 kg ha^–1 in the pure hybrid poplar plots. No difference was detected among treatments for throughfall N content. Nitrogen concentration in roots and leaf litterfall of black alder was higher than hybrid poplar. Significant soil N accretion occurred in mixed plantings containing two alders to one poplar and pure black alder plantings. Nitrogen availability (NO₃–N) increased with the amount of black alder in a plot. Results suggest that the early increase in nitrogen accumulation of hybrid poplar in mixed treatments can be attributed to an increase of total soil N availability resulting from the input of large amounts of N from easily mineralizable alder tissue.     

Genetic diversity and phylogenetic relationships in alder,Alnus firma, revealed by AFLP

Molecular markers for alder,Alnus firma Sieb. et Zucc, have not been studied extensively. Here, we used amplified fragment length polymorphism (AFLP) to investigate genetic relationships among 15 natural populations. EcoRI-ACG + Msel-CTG combinations revealed the highest polymorphism (62.2%). A total of 171 DNA fragments were identified. On average, 58.1% of the AFLP markers that were generated using four primer pairs were polymorphic. Diversity was insignificant among the populations. The combination of a wind-pollinated, outcrossing breeding system along with large population sizes, and the ability to regenerate by stump sprouting may explain the high level of genetic diversity within this species. The majority (98%) of the genetic variance resided within populations. The average number of individuals that were exchanged between populations per generation was very high (N _em = 12.3). Gene dispersal in alder is apparently by seed dispersalvia water and human activity as well as through pollen. Five individuals per population were claded in the same cluster.     

Nitrogen in shoot litter,root litter and root exudates from nitrogen-fixingAlnus incana

Summary A pot experiment withAlnus incana (L.) Moench growing in sand was set up to compare the amounts of nitrogen released from plants shoot litter with that released below ground as root litter and/or root exudation. No nitrogen fixation by free-living microorganisms was found in the sand and the increased nitrogen content of the plant + soil system was therefore due to nitrogen fixation byFrankia in the alder root-nodules. Most of the nitrogen released from the plants was in the nitrogen-rich leaf and other shoot litter. Only small amounts of nitrogen were found in the drainage water from the pots and were recorded as increased nitrogen content of the sand.     

Distribution of spore-positive and spore-negative nodules in stands ofAlnus glutinosa andAlnus incana in Finland

Summary The distribution of spore positive (Sp⁺) and spore negative (Sp⁻) nodules on the two native alder species (A. incana andA. glutinosa) in Finland was investigated. Nodules were collected throughout the country from different ecosystems (forests, swamps, lake- sea- and riversides, old pastures and fields as well as from alder plantations). OnA. incana Sp⁺ nodules predominated, whereas onA. glutinosa the vast majority of the nodules were of the Sp⁻ type. Sp⁺ nodules onA. glutinosa were found only at sites where the two alder species grew close together. This distribution pattern indicates an association of nodule type with alder species, the reasons for which are discussed. Indications of saprophytic growth in the Sp⁻ strain were also found.     

N2 fixation by red alder (Alnus rubra) and scotch broom (Cytisus scoparius) planted under precommerically thinned Douglas-fir (Pseudotsuga menziesii)

Summary From acetylene reduction assays over a 10-month period starting in April 1979, nodule activities averaged 18.78 (se 4.67) moles C₂H₄ g nodule dw^–1 h^–1 forAlnus rubra and 59.95 (se 12.14) moles C₂H₄ g nodule dw^–1 h^–1 forCytisus scorparius. Plant rates were 1.91 (se. 47) moles C₂H₄ plant^–1 h^–1 forA. rubra and 0.55 (se. 17) moles C₂H₄ plant^–1 h^–1 forC. Scoparius. Plant activity and total leaf N were strongly correlated with the dw of other plant parts, but nodule activity and percent leaf N were not. Plant and nodule activities were not associated with temperature, moisture stress, precipitation events or percent light for either species over the growing season nor for 54A. rubra sampled in mid-season 1979 on one replication. After 5 to 6 growing seasons, 14A. rubra on the same site ranged from 30 to 332 cm in height and showed strong correlation between nodule dw, leaf dw, plant size and total leaf N. Results from this study and others indicate logistic equations may be modified to predict the effect of adding a N₂ fixing plant to a population of non N₂ fixing trees.     

The improvement and utilization in forestry of nitrogen fixation by actinorhizal plants with special reference toAlnus in Scotland

Summary Alnus species are used widely in Britain for land reclamation, forestry and other purposes. Rapid juvenile growth of the AmericanAlnus rubra makes it an attractive species for planting on N-deficient soils, particularly those of low organic content. In small plot trials, this species is nodulated by indigenous soil frankiae as effectively asAlnus glutinosa. Over a three year period both species return similar amounts of N to the ecosystem, estimated at up to 10–12 kg N ha^–1. Several strains ofFrankia have been isolated from local (Lennox Forest)A. rubra nodules. These differ morphologically and in their growth on different culture media, both from each other and fromA. glutinosa nodule isolates. AllAlnus isolates, however, have a total cellular fatty acid composition qualitatively similar to some other Group B frankiae. Glasshouse tests in N free culture suggest thatA. rubra nodules formed after inoculation of seedlings with American spore (–) isolates are three times more effective in N fixation than those inoculated with LennoxA. rubra spore (+) nodule homogenates. By contrast, the early growth of seedlings inoculated with spore (–)Frankia strains suggests at best a 35% improvement in N fixing activity over seedlings inoculated with LennoxA. rubra nodule isolates. Nevertheless, this improvement in activity, together with the better performance of seedlings inoculated with isolates compared with those treated with crushed nodule preparations, suggest that it would be worthwhile commercially to inoculate nursery stock with a spore (–)Frankia strain.     

Nitrogen fixation and respiration by root nodules ofAlnus rubra Bong.: Effects of temperature and oxygen concentration

Summary Using a root nodule cuvette and a continuous flow gas exchange system, we simultaneously measured the rates of carbon dioxide evolution, oxygen uptake and acetylene reduction by nodules ofAlnus rubra. This system allowed us to measure the respiration rates of single nodules and to determine the effects of oxygen concentration and temperature on the energy cost of nitrogen fixation. Energy cost was virtually unchanged (2.8–3.5 moles of carbon dioxide or oxygen per mole of ethylene) from 16 to 26°C (pO₂=20 kPa) while respiration and nitrogenase activity were highly temperature dependent. At temperatures below 16°C, nitrogenase activity decreased more than did respiration and as a result, energy cost rose sharply. Acetylene reduction ceased below 8°C. Inhibition of nitrogenase activity at low temperatures was rapidly reversed upon return to higher temperatures. At high temperatures (above 30°C) nitrogenase activity declined irreversibly, while respiration and energy cost increased.Energy cost was nearly unchanged at oxygen partial pressures of 5 to 20 kPa (temperature of 20°C). Respiration and nitrogenase activity were strongly correlated with oxygen tension. Below 5 kPa, acetylene reduction and oxygen uptake decreased sharply while production of carbon dioxide increased, indicating fermentation. Fermentation alone was unable to support nitrogenase activity. Acetylene reduction was independent of oxygen concentration from 15 to 30 kPa. Nitrogenase activity decreased and energy cost rose above 30 kPa until nearly complete inactivation of nitrogenase at 70–80 kPa. Activity declined gradually, such that acetylene reduction at a constant oxygen concentration was stable, but showed further inactivation when oxygen concentration was once again increased. Alder nodules appear to consist of a large number of compartments that differ in the degree to which nitrogenase is protected from excess oxygen.Supported by United States Department of Agriculture Grant 78-59-2252-0-1-005-1     

Estimation of carbon storage, carbon inputs, and soil CO2 efflux of alder plantations on granite soil in central Korea: comparison with Japanese larch plantation

Alder is a typical species used for forest rehabilitation after disturbances because of its N₂-fixing activities through microbes. To investigate forest dynamics of the carbon budget, we determined the aboveground and soil carbon content, carbon input by litterfall to belowground, and soil CO₂ efflux over 2 years in 38-year-old alder plantations in central Korea. The estimated aboveground carbon storage and increment were 47.39 Mg C ha⁻¹ and 2.17 Mg C ha⁻¹ year⁻¹. Carbon storage in the organic layer and in mineral soil in the topsoil to 30 cm depth were, respectively, 3.21 and 66.85 Mg C ha⁻¹. Annual carbon input by leaves and total litter in the study stand were, respectively, 1.78 and 2.68 Mg C ha⁻¹ year⁻¹. The aboveground carbon increment at this stand was similar to the annual carbon inputs by total litterfall. The diurnal pattern of soil CO₂ efflux was significantly different in May, August, and October, typically varying approximately twofold throughout the course of a day. In the seasonally observed pattern, soil CO₂ efflux varied strongly with soil temperature; increasing trends were evident during the early growing season, with sustained high rates from mid May through late October. Soil CO₂ efflux was related exponentially to soil temperature (R ² = 0.85, P < 0.0001), but not to soil water content. The Q ₁₀ value for this plantation was 3.8, and annual soil respiration was estimated at 10.2 Mg C ha⁻¹ year⁻¹. An erratum to this article can be found at     

Egg distribution and mortality in the alder aphid, Pterocallis alni

Alder aphid oviparae produced 14 eggs at constant temperature but in the field the average was 6. Favoured oviposition sites were bud axils and bark crevices. Summer pruning of host trees reduced the availability but not the choice of site. Oviparae reared on leaf regrowth of summer pruned alder were smaller and contained fewer eggs than those on mature leaves. Winter pruning resulted in the loss of 41% buds and 25% eggs. Total winter egg mortality was 65%, being greatest in early winter due to insect predation. Spring egg hatch was synchronised with bud burst and both were affected by temperature.

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Distribution des oeufs et mortalité Pterocallis alni

Résumé Les ovipares de P. alni ont pondu 14 oeufs à température constante mais dans la nature la moyenne n'était que de 6. La taille estivale a réduit les disponibilités, mais n'a pas modifié le choix des lieux de ponte. Les ovipares élevés sur feuilles de repousses estivales d'aulnes taillént plus petits et contenaient moins d'oeufs que ceux élevés sur feuilles adultes. La taille hivernale a provoqué la perte de 41% des bourgeons et 25% des oeufs. La mortalité hivernale totale des oeufs a été de 65%, la partie la plus importante, due à la prédation par les insectes, ayant eu lieu au début de l'hiver. L'éclosion des oeufs au printemps était synchronisée avec l'éclatement des bourgeons, tous les deux étant conditionnés par la température.

     

Litter production,nutrient recycling and litter accumulation inPinus caribaea Morelet var.hondurensis stands in the northern Guniea savanna of Nigeria

Summary The seasonal pattern and quantity of litterfall were studied during a two-year period in two unthinned stands ofPinus caribaea Morelet var. hondurensis Barr. and Golf. in Nigeria. Although pine needles were cast continuously throughout the year, the peak period of litterfall occurred in the dry months of March and April. Mean values of annual litterfall were 3068 and 3665 kg/ha in the two stands aged 7–9 and 9–11 years respectively. Nutrient returns in litterfall in the stands had mean values of 15.0, 0.6, 17.3, 18.2 and 6.3 kg/ha of N, P, K, Ca and Mg respectively. Comparatively low amounts of N and P returned in litterfall were attributed to soil deficiencies of the two elements.Measurements of ground litter showed considerable dry matter accumulation (11378 kg/ha) in the litter layers. Estimates of litter decomposition rate and recycling time showed that it would take 3 to 4 years for the organic matter in annual litterfall to decompose completely as contrasted to about 2 to 5 months often reported under mixed nautral savanna vegetation in the same climatic environment. Similar estimates of nutrient recycling time also showed that between 2 to 4 years were required to mineralize nutrient elements in the annual litterfall; the relative mobilities of the elements were in the order K>Mg>P>NCa.     

Gall mite (Eriophyes laevis) infestation and leaf removal affect growth of leaf area in black alder (Alnus glutinosa) short shoots

Summary We measured the effects ofEriophyes laevis mite galls on the relative growth of short shoot leaf area ofAlnus glutinosa. A portion of leaves was artificially removed from a set of short shoots with both high and low gall density to cause local stress conditions. Nontreated high and low gall density short shoots were used as controls. The results show that the relative growth of leaf area measured for short shoots is negatively affected by high gall density. Artificial leaf removal, on the other hand, had positive effects on leaf area growth. Interestingly, the growth of leaf area did not differ for high gall density short shoots with leaf removal and noninfested short shoots with no leaf removal. This result may be caused by the combined, opposite effects of leaf removal and gall infestation.     

Nitrate reductase and glutamine synthetase activities in relation to growth and nitrogen assimilation in red oak and red ash seedlings: effects of N-forms,N concentration and light intensity

The effects of growing seedlings of red oak (Quercus rubra) and red ash (Fraxinus pennsylvanica) with Hoagland solutions containing five N-regimes, differing in the N-forms (NH₄, NO₃) and concentrations (High and Low), in relation to light intensity were investigated by the utilization of enzymatic markers of the N assimilation pathway, nitrate reductase (NR) and glutamine synthetase (GS). Red oak and red ash showed different patterns of N-assimilation. Red oak seedlings assimilated NO₃ in low amounts in their roots and leaves, whereas red ash seedlings assimilated high amounts of NO₃, mostly in the leaves. A significant amount of constitutive NR activity was found in red oak seedlings supplied with NH₄ N-regime. This could be characteristic of a species adapted to soils that are poor in nitrogen. Root GS activity was lower in red oak seedlings than in red ash seedlings, indicating that the rate of NH₄ assimilation differed in these two hardwood species. Low irradiance reduced growth of both hardwood species, but greatly affected the specific leaf area of red ash and reduced NO₃ assimilation (when data are expressed per leaf area). Both species reacted similarly to N-regimes in terms of relative growth rate.     

Distribution of roots and root nodules and biomass allocation in young intensively managed grey alder stands on a peat bog

Development of below-ground biomass and biomass allocation were studied in two different stands of young grey alder stands growing on a peat bog. Both stands were given the same fertilization and irrigation treatment. The roots were investigated from 1) open plastic tubes enclosing the complete root systems in 1982, and 2) root cores 1984–86. Coarse roots (diameter>1 mm) were mainly found close to the trunk of the trees while fine roots (≤1 mm) were more evenly distributed in the stands. Root nodules were intermediate in distribution. The root systems were shallow, with more than 90% of the biomass in the uppermost 9–10 cm of the soil, probably because of low oxygen availability in the peat soil. The biomass allocation to the above-ground parts increased during the study period.     

Growth and biomass turnover ofHydrocharis dubia L. cultured under different nutrient conditions

Growth of a floating-leaved plant,Hydrocharis dubia L., was examined under varying nutrient conditions between 0.3 and 30 mgN l⁻¹ total inorganic nitrogen.H. dubia plants cultured under the most nutrient-rich condition showed the highest maximum ramet density (736 m⁻²), the highest maximum biomass (80.4 g dry weight m⁻²), and the highest total net production (185 g dry weight m⁻² in 82 days). Plants under nutrient-poor conditions had a relatively large proportion of root biomass and a small proportion of leaves with a long life span. Compared with other floating-leaved and terrestrial plants, the maximum biomass ofH. dubia was relatively small. This, and the rapid biomass turnover, was related to the short life span of leaves (13.2–18.7 days) and large biomass distribution to leaves.     

Fine root growth and nutrient release in decomposing leaf litter in three contrasting vegetation types in central Amazonia

We tested the hypothesis that the growth of fine roots upward into the leaf litter, forming a ‘surface root mat’, found widely in Amazonian forests, is of adaptive value for plants of poor soils because it makes possible more rapid uptake of limiting nutrients. We assessed the effect of invasion by fine roots on the rates of loss of dry mass and nutrient content of leaf litter over 1 year in three plots in each of three contrasting forest types in central Amazonia: the stunted facies of heath forest known as campina (SHF), the taller facies of heath forest known as campinarana (THF), both on spodosols, and the surrounding lowland evergreen rain forest (LERF) on ultisol. Pairs of bags filled with freshly fallen leaves from the trees of Clitoria fairchildiana (Fabaceae) were placed on the litter layer; in order to prevent roots entering the control bag in each pair that bag was lifted from the forest floor and turned over each week, while the treatment bag was left undisturbed. From each plot, four pairs of litter bags were retrieved after 30, 60, 120, 180, 270 and 360 d, and all roots that had grown into the litterbags were carefully removed. The leaves and roots from each bag were oven-dried for nutrient analysis. In no forest type was there a significant difference in the rate of loss of dry matter from the litter between the bags with and without roots. The amounts of roots invading the litter bags increased significantly in the sequence SHF < THF < LERF. In no forest did the presence of roots have a significant effect on the rate of disappearance of N or P from the leaf material, or on the rate of accumulation of Fe and Al. In the SHF there was no significant effect of invasion by roots on the rates of disappearance of Ca, Mg, Mn or Zn, but in the THF, the rates of disappearance of these four elements between 270 and 360 d were significantly greater in the presence of roots. In the LERF the results were similar, but the effects of roots started earlier—the rates of disappearance of Ca and Mg were significantly enhanced between 120 and 360 d. The results from the SHF may be interpreted to suggest that the growth of fine roots (and their fungal associates) upward into leaf litter is unlikely to be explained wholly by their role in the efficient recovery of mineral nutrients.     

The roles of temperature and food quality in affecting the performance of the alder aphid, Pterocallis alni

Six successive summer generations of the alder aphid (Pterocallis alni (DeGeer) (Homoptera: Callaphididae)) were reared simultaneously in the field and under controlled temperature conditions in the laboratory. The growth and reproduction of each generation were recorded. The available food quality for the aphids was measured by weekly analysis of foliar soluble nitrogen concentrations. Although there was a significant change in leaf soluble nitrogen during the season, the decline was only 23% and this did not appear to have an adverse effect on the performance of this aphid. Instead, the major environmental factor affecting the aphid is temperature. This is evidenced by the facts that when aphids were reared at constant temperatures, there were no differences in generation performance, even though food quality varied seasonally. In addition, all generations of the aphid posess the same number of ovarioles, indicating that there is no pre-programmed anticipation of a seasonal deterioration in food quality in this aphid species.

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Résumé P. alni DeGeer (Homopt. Callaphididae) vit sur Alnus glutinosa Gaertner pendant toute l'année. L'aulne étant un arbre fixant l'azote, cette étude a été entreprise pour examiner la croissance et la reproduction de ce puceron sur cet arbre afin d'expliquer les changements de la dynamique de sa population. 6 générations successives estivales ont été élevées dans la nature et en température constante au laboratoire. La qualité de l'aliment a été mesurée chaque semaine par une microkjeldahl analyse des concentrations foliaires en azote soluble. La qualité de l'aliment varie saisonnière, mais ne diminue que de 23%; elle semble n'avoir que peu d'effets sur le puceron, puisque la fécondité dans la nature n'est réduite que pendant une génération. Quand la température est maintenue constante, il n'y a pas de différences entre les performances des différentes générations de pucerons. Ainsi, les changements saisonniers dans la croissance et la reproduction des pucerons sont provoqués par la température et non par la qualité de l'aliment.