Factors affecting calculation of nutritional induces for foliage-fed insects: an experimental approach

To determine how nutritional indices for insects fed leaves are affected by the experimental conditions and the physiology of the plant material, we used larvae of the buckmoth, Hemileuca lucina Hy. Ed. (Saturniidae) and their hostplant Spiraea latifolia Ait. Bork (Rosaceae). Under experimental conditions identical to those used to determine larval nutritional indices, we found that the age of leaves (new versus mature) significantly affected their metabolism and water loss, but simulated herbivory did not directly affect leaf metabolism. Over a 6-day test, nitrogen concentration showed an initial increase followed by a gradual decline, and was higher in new leaves compared to mature leaves. New leaves increased in protein concentration and then gradually returned to the initial level, whereas mature leaves changed little over the 6-day test. These changes in percent nitrogen and protein may largely reflect the disproportional changes in non-nitrogenous materials. Solitary and grouped larvae had similar growth rates on new leaves, but they differed on mature leaves. Deliberate manipulation of larvae during the course of an experiment significantly reduced relative growth rates by increasing duration of the stadium rather than by decreasing biomass gained. Two methods of estimating larval gut contents at mid-stadium were compared: weight of frass produced and weight of digestive tract and contents. After the end of the 4-day test period used to determine nutritional indices, the digestive tracts with food accounted for 10.8% of the larval dry weight. Larval frass produced in 24 h after the end of the test period comprised 9.3% of the larval dry weight. Correction factors for plant metabolism changed nutritional indices by 1 to 8%, while those for larval gut contents altered indices by 2 to 15%.     

Variation in phenology and nutritional quality between host plants and its effect on larval performance in a specialist butterfly,Zerynthia rumina

The main host plants of the butterfly Zerynthia rumina L. (Lepidoptera: Papilionidae) in southern Spain occur in different habitats and in general do not grow sympatrically. Therefore, each single local butterfly population uses the particular host available within its range. Aristolochia longa L. is a tuberous perennial herb available only in the spring, while A. baetica L. is an evergreen perennial vine with indeterminate growth. However, because of the toughness of older leaves, newly hatched larvae feed only on new leaves of A. baetica, and most of these leaves are produced well before the larvae hatch. In laboratory experiments, caterpillars feeding on either new or mature A. longa leaves grew faster and converted food into biomass more efficiently than those feeding on new A. baetica leaves. These differences are related to variation in nutritional quality among the host plants. Estimates of butterfly abundance were lower in sites where Z. rumina uses A. baetica, compared with those where the host is A. longa. The potential differential effect of these two food plants on the densities of local butterfly populations relying on them is discussed here.     

The timing of bud burst and its effect on tree growth

A phenology model for estimating the timings of bud burst – one of the most influential phenological phases for the simulation of tree growth – is presented in this study. The model calculates the timings of the leafing of beech (Fagus sylvatica L.) and oak (Quercus robur L.) and the May shoot of Norway spruce (Picea abies L.) and Scots pine (Pinus sylvestris L.) on the basis of the daily maximum temperature. The data for parameterisation and validation of the model have been taken from 40 climate and 120 phenological stations in southern Germany with time series for temperature and bud burst of up to 30 years. The validation of the phenology module by means of an independent data set showed correlation coefficients for comparisons between observed and simulated values of 54% (beech), 55% (oak), 59% (spruce) and 56% (pine) with mean absolute errors varying from 4.4 days (spruce) to 5.0 days (pine). These results correspond well with the results of other – often more complex – phenology models. After the phenology module had been implemented in the tree-growth model BALANCE, the growth of a mixed forest stand with the former static and the new dynamic timings for the bud burst was simulated. The results of the two simulation runs showed that phenology has to be taken into account when simulating forest growth, particularly in mixed stands.     

The effect of flutter on the temperature of poplar leaves and its implications for carbon gain

The leaf temperatures of two poplar species (Populus tremuloides Michx. and P. fremontii Wats.) were characterized by attaching thermocouples to leaves that were either constrained to a fixed position or allowed to flutter naturally. There were no observed temperature differences between fluttering and constrained leaves in the lower canopy, but fluttering leaves at the top of the canopy were as much as 2–4°C cooler than constrained leaves. An increase in heat transfer, a decrease in light interception or both could account for these observed differences in the temperature of fluttering versus constrained leaves. Fluttering can increase the boundary-layer conductance to convective heat exchange by as much as 50 and 20% for laminar and turbulent flow, respectively. The benefit that these leaf temperature differences may provide to the carbon economy of a poplar canopy was dependent on the ambient temperature. Populus fremontii, which is frequently exposed to daytime temperatures exceeding 35°C during summer months in the central valley of California, USA, could show an increase in carbon gain as a result of lower upper canopy leaf temperatures. For aspen, the benefit would be much smaller and often negative because of much lower air temperatures. Lower leaf temperatures may also increase the water use efficiency of poplars. However, the maintenance of lower leaf temperatures may not be the primary adaptive significance of leaf flutter.     

The effect of neem allelochemicals on nutritional physiology of larval Spodoptera litura

Neem allelochemicals azadirachtin, salannin, nimbinene and nimbin were administered to different larval instars of the tobacco armyworm, Spodoptera litura orally in artificial diet, topically or via injection. Nutritional analyses revealed strong antifeedant and growth regulatory effects of azadirachtin which were independent of each other. While salannin and nimbinene induced concentration dependent feeding deterrence only; nimbin was inactive to the 1000 ppm level against this insect species. One of the causes of the reduced growth rate of azadirachtin treated insects was due to an increase in the costs associated with growth. This was relative to a drastic reduction in the activity of gut trypsin. Salannin and nimbinene, however, did not interfere with the trypsin activity of the gut. These results and those from nutritional studies suggest that salannin and nimbinene have no toxicity mediated effects on S. litura larvae and antifeedant activity is a result of the effects on deterrent and other chemoreceptors. The fact that azadirachtin directly or indirectly inhibits the secretion of trypsin by the enzyme-secreting cells of the gut is discussed.     

Control of plant growth by nitrogen: differences between cereals and broadleaf species

Abstract. In four dicotyledonous species low levels of N strongly inhibited leaf expansion during the day but had little or no effect at night. In contrast, daytime and night-time expansion were equally affected in four cereal species. The results support the general concept that in dicotyledons, N controls leaf expansion through its effects on hydraulic conductivity. In such N-limited plants, water deficits generated by transpiration may inhibit daytime cell expansion. In cereals, cell expansion and transpiration occur in separate zones of the leaf and are apparently unrelated.
Growth analysis showed that low levels of N inhibited leaf area growth more strongly in dicotyledons than in cereals, but had similar effects on net assimilation rates of plants in the two groups. As a result, dry matter production was more efficient in cereals than in dicotyledons when N was limiting.     

Effect of dietary nitrogen on gypsy moth larval nutritional indices, development and lipid content

Nutritional indices, development rates, percent dry weights and total lipids were determined in gypsy moth larvae (Lymantria dispar L.) reared on a high wheat germ (HWG) diet or diets prepared from lyophilized, ball-milled oak or pine foliage as the only source of dietary nitrogen (N). With regard to both total and proteinaceous N content, HWG diet>oak diet>pine diet. All nutritional indices measured were significantly lower in second instars fed pine diet vs. oak diet. Protein supplementation of pine diet with either casein or ovalbumin to bring total N up to the level present in oak diet resulted in small increased in approximate digestibility (AD) and effciency of conversion of ingested food (ECI), but relative growth rate (RGR) remained unaffected. The low RGR of larvae fed pine diet (unsupplemented or protein supplemented), as compared to those fed HWG or oak diet, was accompanied by significantly lower larval percent dry weight and percent total lipid. In contrast, RGR, larval percent dry weight and total lipid values were comparable in second instars fed HWG or oak diet. Insects reared from the first through the final instar on oak diet exhibited lower pupal weights compared to those reared on HWG. Casein addition to oak diet generally resulted in even more extended larval development times and further reduced pupal weights, but wheat germ addition to oak diet did not alter development rates and caused an increase in pupal weights.     

The influence of ancymidol on morphology,anatomy, and chlorophyll levels in developing and mature Helianthus annuus leaves

Ancymidol foliar spray at 132 mg·liter^–1 a.i. modified leaf anatomy of developing Helianthus annuus L. Mammoth Russian leaves, but not of mature leaves. Ancymidol was effective in retarding plant growth when applied at the young seedling or at a more mature stage of growth. Ancymidol increased leaf weight per unit area and chlorophyll content on an area and unit weight basis, regardless of stage of leaf development. Total chlorophyll per leaf was also increased in mature leaves. Thus, darker green foliage due to increased chlorophyll content and modified leaf anatomy responses were determined to be independent effects.Texas Agricultural Experiment Station paper no. 22927.     

Influence of host plant, foliar phenology and larval dietary history on Lymantria dispar larval nutritional indices

Quantitative nutritional indices were measured in 3rd instar gypsy moths, Lymantria dispar L. Analyses of the main and interactive effects of host plant consumed (oak or pine), foliar phenology (early or late season) and larval dietary history (whether fed artificial diet or foliage as second instars) indicate that larval performance was significantly affected by all three variables tested. Relative Growth Rate (RGR) was higher on oak than pine early in the season, but the reverse occurred one month later, owing largely to the slower Relative Consumption Rate (RCR) of late season oak feeders. Efficiency of Conversion of Digested food (ECD) was depressed in larvae fed pine foliage for two instars compared with those fed oak. The interaction between larval dietary history and foliar phenology produced significant effects on all indices measured. Foliar analyses showed that, as the season progressed, oak had a higher percent dry weight than did pine. On a dry weight basis, pine foliage had significantly less total and protein nitrogen than did oak collected at the same time.

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Résumé Les indices nutritionnels larvaires ont été évalués sur des chenilles de L. dispar L. du troisième stade. L'analyse des effets et des interactions de la nature de la plante consommée (Quercus microcarpa) ou Pinus strobus, de la phénologie foliaire (en début de saison, c'est-à-dire seconde quinzaine de mai, ou en fin de saison, c'est-à-dire à la mi-juin), de l'alimentation antérieure (régime artificiel de germes de blé ou feuillage pendant le second stade) montre que les performances larvaires sont fortement modifiées par ces 3 variables. Le taux de croissance relative (RGR) est plus élevé sur chêne que sur pin en début de saison, mais s'inverse un mois plus tard avec une forte diminution sur chêne. L'efficacité de conversion de l'aliment (ECD) est diminuée sur pin pendant les second et troisième stades. L'interaction entre l'alimentation antérieure et le phénologie foliaire a des effets significatifs sur la digestibilité approchée (AD), sur l'efficacité de la conversion de l'aliment ingéré (ECI), et sur ECD, RCR et RGR.Les analyses foliaires ont montré que, lorsque la saison avance, le poids sec du chêne devient supérieur à celui du pin. En considérant le poids sec, le feuillage de pin contient moins d'aliments et de protéines que le chêne récolté au même moment. La teneur en azote tend à diminuer chez les deux espèces, de 17% pour le pin et 26% pour le chêne.Nous pensons que certains de ces facteurs déterminent la valeur de ces plantes pour L. dispar. Les chenilles âgées peuvent compenser plus ou moins la faible assimilation sur pin par des RCR plus élevés. Le même comportement ne semble pas possible sur chêne, une plus grande quantité de chêne avancé en saison que de feuilles jeunes ne pouvant être consommée pour compenser la diminution de la digestibilité.

     

Variation in the temperature preference and growth rate of individual fish reconciles differences between two growth models

1. A growth model, originally developed for brown trout (Salmo trutta), has now been fitted to data for Atlantic salmon (S. salar) and stone‐loach (Barbatula barbatula) from English populations, and Arctic charr (Salvelinus alpinus) from Sweden. The model relates growth rate to temperature for a fish of standard size and the functional relationship has a triangular shape with a sharp peak at the optimal temperature for growth and zero growth at the base of the triangle. It was unsuitable for growth data for Norwegian salmon, and a curvilinear Ratkowsky model provided a better fit, though the experimental protocol was different in the Norwegian and English experiments. 2. The Norwegian salmon were kept in groups in each tank, had to compete for food, and had to be divided into slow, moderate and fast growers before the Ratkowsky model could be fitted. Each English salmon was kept in its own tank and fed individually. For replicate experiments, fish of similar size were selected. Variation among fish kept under similar conditions was therefore small, and the triangular model was essentially for individual fish, not groups of fish. 3. The present simulation study tests the hypothesis that individual differences in the growth response could account for the curvilinear growth‐temperature relationship for the Norwegian salmon. The triangular model was used to generate the growth response to temperature for a group of salmon, each fish having a slightly different temperature preference and growth rate. The result was a curvilinear response, well approximated by the Ratkowsky model (adjusted R² = 0.96). When the variability in individual temperature preference was increased, the Ratkowsky model was an even better fit (adjusted R² = 0.98). Therefore, the apparent discrepancy between the two models was reconciled by allowing for individual differences in temperature preference and growth rate within groups of fish.     

Effect of temperature and leaf age on growth versus moulting time of a generalist caterpillar fed plantain (Plantago lanceolata)

Abstract.

• 1 The simultaneous effects of daytime temperature (20°C versus 30°C) and leaf age (new versus intermediate-aged) on a generalist insect herbivore were examined. Fourth-instar Spilosoma congrua caterpillars were tested on plantain (Plantago lanceolata), one of this lepidopteran species’host plants, for which the major defensive chemicals, iridoid glycosides (aucubin and catalpol), could be quantified.
• 2 Cool temperature depressed amount of food eaten, amount of frass, and consumption and growth rates, and increased the proportion of time spent in the non-feeding period (from head-capsule slippage to ecdysis).
• 3 Average iridoid glycoside concentration was 2.8% dry weight (d.w.) in new leaves and 1.6% d.w. in intermediate-aged leaves. When fed new leaves, the caterpillars had a higher efficiency of conversion of ingested food to biomass and a higher growth rate than those fed intermediate-aged leaves. Furthermore, the proportion of time spent in the non-feeding period was prolonged by a diet of intermediate-aged leaves.
• 4 A second experiment showed that the percentage dry weight of aucubin, catalpol and total iridoid glycosides increased over 24 h in incubated, excised leaves, which meant that the caterpillars in the first experiment experienced somewhat higher iridoid glycoside concentrations than the levels in fresh leaves.
• 5 Overall, these results indicate that this generalist should prefer new plantain leaves over older leaves even though new leaves contain higher concentrations of iridoid glycosides.

     

The effect of carbon dioxide concentration on respiration of growing and mature soybean leaves

Soybean plants were grown continuously at 350 and 700cm³m^?3 CO₂ at constant temperature. Respiration rates of third trifoliolate leaves were measured at the growth CO₂ concentration for the whole dark period from 5d before through to 5d after full area expansion. The short-term response of respiration rate to the measurement CO₂ concentration was also determined at each age. Respiration rates per unit of dry mass declined with age and were significantly less at a given age or RGR in leaves grown and measured at the elevated CO₂. The difference in respiration rate was largest in mature leaves and resulted from the different measurement CO₂ concentrations. The respiratory costs of the tissue synthesis, estimated from the elemental composition of the tissue, did not differ substantially between CO₂ treatments. The response of respiration rate to carbon dioxide concentration was not strongly affected by the form of nitrogen supplied. Maintenance respiration calculated by subtracting growth respiration from total respiration was negative in rapidly growing leaves for both CO₂ treatments. This indicates that CO₂ efflux in the dark does not accurately reflect the average 24 h rate of energy expenditure on growth and maintenance for soybean leaves.     

The effect of temperature differences between porometer head and leaf surface on stomatal conductance measurements

It is generally known that instantaneous values of leaf conductance as measured with a dynamic porometer need to be corrected for the temperature difference, ΔT, between the porometer cup and the sampled leaf. Leaf conductances, obtained with a Delta-T AP4 dynamic porometer, with and without correction for ΔT are compared for a bush species (Guiera senegalensis) and two forb species (Jacquemontia tamnifolia and Mitracarpus scaber). With temperature differences predominantly varying within the ±2.5 °C recommended by the manufacturer, it appears that the differences between uncorrected and corrected conductances are very large, up to 100% on average, especially for the two forbs. Furthermore, it is shown that, using the Mitracarpus data, a relatively small error of ±0.5 °C in ΔT can cause a difference of 25–50% in the final conductance value, in particular for the high conductance range. An error of ±0.5 °C may easily occur: the accuracy of ΔT as measured by the thermistors in the porometer is 0.2 °C and the temperature variation within a leaf can be much larger. This result will have implications for upscaling of leaf conductances to canopy values or may explain why upscaled values appear not to correspond with down-scaled values, obtained from eddy correlation measurements and an inverted canopy transpiration model.     

Impacts of population growth and economic development on the nitrogen cycle in Asia

Asia is the major consumer of fertilizer nitrogen and energy in the world, and consequently shares a considerable proportion of the world creation of reactive nitrogen (Nr). However, if estimated on per capita basis, Asia is characterized by a lower arable land area, fertilizer nitrogen consumption, energy consumption, and gross domestic product, as well as lower daily protein intake. To meet the increasing needs for food and energy for the growing population combined with the improvement of living standards, Nr will inevitably increase. The present study estimates the creation of Nr and the emissions of various N compounds into environment in Asia currently and in 2030. In comparison with the world averages, the lower fertilizer nitrogen and energy use efficiencies, and the lower use of animal wastes for agriculture imply that there is potential for moderating the increase in Nr and its impacts on the environment. Strategies for moderating the increase are discussed.     

Effects of leaf position and nitrogen supply on the expansion of leaves of field grown sunflower (Helianthus annuus L.)

Leaf growth responses to N supply and leaf position were studied using widely-spaced sunflower plants growing under field conditions. Both N supply (range 0.25 to 11.25 g added N per plant) and leaf position significantly (p=0.001) affected maximum leaf area (LA_max) of target leaves through variations in leaf expansion rate (LER); effects on duration of expansion were small. Specific leaf nitrogen (SLN, g N m^-2) fell quite rapidly during the initial leaf expansion phase (LA < 35% LA_max) but leveled off during the final 65% increase of leaf area. This pattern held across leaf positions and N supply levels. Leaf nitrogen accumulation after 35% LA_max continued up to achievement of LA_max; reductions in the higher SLN characteristic of the initial phase were insufficient to cover the nitrogen requirements for expansion during the final phase. LER in the quasi-linear expansion phase (35 to 100% of LA_max) was strongly associated with SLN above a threshold that varied with leaf position (mean 1.79±0.225 g N m^-2). This contrasts with the response of photosynthesis at high irradiance to SLN, which has previously been shown to have a threshold of 0.3 g N m^-2; in the present work saturation of photosynthetic rate was evident when SLN reached 1.97 g N m^-2. Thus, once the area of a leaf exceeds 35% of LA_max, expansion proceeds provided SLN values are close to the levels required for maximum photosynthesis. However, growth of leaves during the initial expansion phase ensures a minimum production of leaf area even at low N supply levels.     

Impacts of population growth and economic development on the nitrogen cycle in Asia

Asia is the major consumer of fertilizer nitrogen and energy in the world, and consequently shares a considerable proportion of the world creation of reactive nitrogen (Nr). However, if estimated on per capita basis, Asia is characterized by a lower arable land area, fertilizer nitrogen consumption, energy consumption, and gross domestic product, as well as lower daily protein intake. To meet the increasing needs for food and energy for the growing population combined with the improvement of living standards, Nr will inevitably increase. The present study estimates the creation of Nr and the emissions of various N compounds into environment in Asia currently and in 2030. In comparison with the world averages, the lower fertilizer nitrogen and energy use efficiencies, and the lower use of animal wastes for agriculture imply that there is potential for moderating the increase in Nr and its impacts on the environment. Strategies for moderating the increase are discussed.