Nitrogen fertilization affects interactions between the components of an insect–fungus–plant tripartite system

1. The chrysomelid beetle, Gastrophysa viridula and the rust fungus, Uromyces rumicis both occur on leaves of Rumex obtusifolius growing in a wide range of soil nutrient conditions. We investigated the effect of fertilizing plants with eight nitrate and four ammonium concentrations on the components of this tripartite interaction in a controlled environment.
2. Leaf weight, area and total plant weight increased as both nitrate and ammonium concentrations increased up to 15mmol l^–1. Between 15 and 50mmol l^–1 added nitrate, leaf and total plant weight decreased. Total plant weight was unaffected by increasing ammonium fertilization from 15 to 25mmol l^–1.
3. The density and percentage of U. rumicis pustules sporulating 8 days after infection decreased with increasing nitrate but were unaffected by increasing the concentration of ammonium fertilization.
4. Leaf area consumed and number of eggs laid by G. viridula decreased as the concentration of nitrate fertilization increased. Increasing the concentration of ammonium decreased leaf area eaten.
5. First instar mortality of G. viridula was increased and gregariousness and adult fecundity was decreased additively by the combination of U. rumicis infection and decreasing the nitrate concentration fed to plants from 10 to 1mmol l^–1.     

The effect of nitrogen fertilization and rust fungus infection, singly and combined, on the leaf chemical composition of Rumex obtusifolius

1. The chrysomelid beetle Gastrophysa viridula occurs on Rumex obtusifolius growing in a range of nutrient conditions and also on plants infected with the foliar fungus Uromyces rumicis . In a controlled environment, we investigated the effect of fertilizing plants with eight nitrate and four ammonium concentrations, with or without infection, on leaf nutritional quality.
2. Increasing nitrate fertilization increased leaf oxalate, total nitrogen and nitrate concentrations and water content, and decreased total non-structural carbohydrate (NSC) concentrations. Increasing ammonium fertilization increased leaf nitrogen concentration and water content, decreased nitrate and NSC concentrations, and had no effect on oxalate concentrations.
3. Infection produced a mainly additive effect to fertilization, increasing NSC and oxalate, and decreasing nitrate and nitrogen concentration in whole plants fed nitrate, and increasing nitrate and NSC in whole plants fed ammonium.
4. Young leaves on infected plants remained uninfected and had greater nitrogen and NSC concentrations, and lower oxalate and nitrate concentrations, than infected leaves on the same plant.
5. These results are discussed in relation to changes in C:N and NSC:organic nitrogen ratios, the effect of nitrate and oxalate, and the known feeding and oviposition preferences of the beetle. The results suggest that there is an optimum nitrogen fertilization level for G. viridula development.     

Induced resistance of Gossypium australe against its most abundant folivore,Bucculatrix gossypii

Induced resistance of cultivated Gossypium to its exotic, agricultural pests is well studied but little is known about whether native cottons respond to damage by endemic herbivore populations. This study examined induced responses of Gossypium australe to its most abundant folivore, Bucculatrix gossypii. Prior damage did not affect the number of new mines initiated. Survival of miners on damaged, young leaves and cotyledons was reduced compared with survival on young leaves and cotyledons of undamaged plants. However, the induced resistance was not systemic; survival of miners on older, undamaged leaves of damaged seedlings was not different from survival on older leaves of undamaged controls. This localized induced resistance did not produce an overdispersed distribution of either mines or successful mines. On the contrary, the distributions tended towards clumped, although they were not statistically distinguishable from random. Although a localized induced response affected miner survival, no effects on behaviour were observed.     

Variation in damage to cotton affecting larval feeding preference of Spodoptera littoralis

Feeding experiments with larvae of Spodoptera littoralis were performed with leaves from cotton plants subjected to damage and from undamaged plants. In the experiments, four different time intervals (1, 3, 7, and 14 days) after damage induction and two different levels (high and low) of herbivore damage were tested. Seven days after damage induction larvae fed less on the young top leaves from damaged plants for both levels of damage. At the high damage level, the larvae fed less on leaves from the damaged plants after just three days, and this effect still remained 14 days after damage infliction. When mature leaves from the middle of the plant were compared, no difference between treatments was observed.Two plant sizes were tested, small plants with 4–5 true leaves and large plants with 8–10 true leaves. In small plants the induced changes affecting larval feeding were found mainly in the youngest leaf at the top of the plant, while in large plants the induced effects were found in both the youngest and the second youngest leaves.In plants subjected to artificial damage, larvae fed less on top leaves of the damaged plants when compared to leaves from undamaged plants. When leaves from plants that had been artificially damaged were directly compared with leaves from plants damaged by herbivores, larvae fed more on the youngest leaves from artificially damaged plants when the plants were large. In small plants no significant difference was found when comparing artificial and herbivore damage.     

Within-plant distribution of induced resistance in apple seedlings: rapid acropetal and delayed basipetal responses

Induction of plant resistance by herbivory is a complex process, which follows a temporal dynamic and varies spatially at the within-plant scale. This study aimed at improving the understanding of the induction process in terms of time scale and within-plant allocation, using apple tree seedlings (Malus × domestica) as plant model. Feeding preferences of a leaf-chewing insect (Spodoptera littoralis) for previously damaged and undamaged plants were assessed for six different time intervals with respect to the herbivore damage treatment and for three leaf positions. In addition, main secondary defense compounds were quantified and linked to herbivore feeding preferences. Significant herbivore preference for undamaged plants (induced resistance) was first observed 3 days after herbivore damage in the most apical leaf. Responses were delayed in the other leaf positions, and induced resistance decreased within 10 days after herbivore damage simultaneously in all tested leaf positions. Chemical analysis revealed higher concentrations of the flavonoid phloridzin in damaged plants as compared to undamaged plants. This indicates that herbivore preference for undamaged apple plants may be linked to phloridzin, which is the main secondary metabolite of apple leaves. The observed time course and distribution of resistance responses within plants contribute to the understanding of induction processes and patterns, and support the optimal defense theory stating young tissue to be prioritized. Moreover, induced resistance responses occurred also basipetally in leaves below the damage site, which suggests that signaling pathways involved in resistance responses are not unidirectional.     

Defoliation effects on isoprene emission from Populus deltoides

Isoprene emission from plants is one of the principal ways in which plant processes alter atmospheric chemistry. Despite the importance of this process, few long-term controls over basal emission rates have been identified. Stress-induced changes in carbon allocation within the entire plant, such as those produced by defoliation, have not been examined as potential mechanisms that may control isoprene production and emission. Eastern cottonwood (Populus deltoides) saplings were partially defoliated and physiological and growth responses were measured from undamaged and damaged leaves 7 days following damage. Defoliation reduced isoprene emission from undamaged and damaged leaves on partially defoliated plants. Photosynthetic rates and leaf carbon and nitrogen pools were unaffected by damage. Photosynthetic rate and isoprene emission were highly correlated in undamaged leaves on undamaged plants and damaged leaves on partially defoliated plants. There was no correlation between photosynthetic rate and isoprene emission in undamaged leaves on partially defoliated plants. Isoprene emission was also highly correlated with the number of source leaves on the apical shoot in damage treatments. Increased carbon export from source leaves in response to defoliation may have depleted the amount of carbon available for isoprene synthesis, decreasing isoprene emission. These results suggest that while isoprene emission is controlled at the leaf level in undamaged plants, emission from leaves on damaged plants is controlled by whole-branch allocation patterns. Received: 12 May 1998 / Accepted: 9 November 1998     

Autumnal changes in tissue nitrogen of autumn olive,black alder and eastern cottonwood

Two experiments were conducted to determine patterns of N change in tissues of autumn olive (Elaeagnus umbellata Thunb.) and black alder (Alnus glutinosa [L.] Gaertn.) during autumn in central Illinois, U.S.A. In the first study leaf nitrogen concentrations of autumn olive decreased 40% at an infertile minespoil site and 39% at a fertile prairie site throughout autumn whereas nitrogen concentrations in respective bark samples increased by 39% and 37%. Salt-extractable protein concentrations increased in bark and decreased in leaves over the sampling period. Free amino acid concentrations of autumn olive leaves decreased over the course of the experiment from peak concentrations in August. Asparagine, glutamic acid and proline were major constituents of the free amino acid pools in leaves. Total phosphorus concentrations of autumn olive leaves declined by 40–46% during autumn while bark concentrations of P did not significantly change.In the second experiment non-nodulated seedlings of alder receiving a low level of N-fertilization did not exhibit net resorption of leaf N during autumn whereas foliar N concentration of contrasting nonactinorhizal cottonwood plants (Populus deltoides Bartr. ex. Marsh) under the same fertilization regime decreased by 27% after the first frost. A gradual but significant decrease of 38% in foliar N concentration of nodulated alder seedlings grown under a low N-fertilization regime was associated with the cessation of nitrogenase activity during autumn in nodules. Compared with the low N fertilization regime, the higher level of N-fertilization resulted in smaller autumnal decreases of foliar N concentration in nodulated alder (17%) and in cottonwood (20%); but there was no decrease in foliar N concentration in non-nodulated alder. The higher level of N-fertilization promoted a greater accumulation of N in the roots than in the bark of both tree species after the first frost.Our results suggest that black alder lackingFrankia symbionts does not exhibit net leaf N resorption and that autumnal decreases in leaf N ofFrankia-nodulated black alder result primarily from declining foliar N import relative to export due to low temperature inhibition of N₂ fixation. In contrast, autumn olive exhibited greater and more precipitous autumnal declines in foliar N concentration than those of alder, and the pattern of N decline was unaffected by site fertility.     

Consequences of the Elimination of Old Leaves upon Spring Phenological Events and the New Leaves Nutrient Concentration in a Wintergreen Woody Species in the Southern Hemisphere

Previous studies analyzed the importance of old leaves conservancy for wintergreen species plant growth only after early spring old leaves elimination. However, carbon and nutrient resources for growth could have already been translocated from old leaves to shoots during autumn. In this work, the effect of old leaves absence on the leaf mass per area (LMA, g m⁻²) and nutrient concentration of new spring leaves, shoot growth, and flowering was studied in Aristotelia chilensis, an Andean Patagonic woody wintergreen species of Argentina. Plants were studied after autumn defoliation (AD) or late winter defoliation (WD) and results were compared to those of undamaged control plants (CO). The new leaves LMA and mineral nutrient (N, P, K, and Mg) concentration values did not decrease in AD or WD compared to CO plants. Conversely, CO plants showed higher flowering intensity and shoot lengthening compared to AD or WD plants. There were not remarkable differences regarding the defoliation time, though non-flowering shoots grew in a lesser degree than the flowering shoots in WD plants. It was concluded that A. chilensis old leaves cohort is an important source to shoot growth and flowering but their absence does not affect the new leaves structure or nutritional status from early spring in either AD or in WD plants. New leaves formation probably is guaranteed by resources (carbon and nutrients) previously stored in stems or even in the buds containing the preformed leaves since March, by the end of summer. Provided the availability of complete resources for the new leaf flush independently of the old leaves A. chilensis would restore the carbon balance as soon as possible to resume the growth of heterotrophic tissues at normal rates. Endogenous response to counterbalance the old leaves absence on non-flowering shoots was more effective when there was greater lag time between defoliation and shoot growth resume. Flowering and non-flowering shoots compete for the available resources when A. chilensis have not yet expanded leaves and shoots supporting reproductive structures were stronger sinks compared to non-flowering shoots in WD plants.     

Microfungi associated with Abies needles and Betula leaf litter in a subalpine coniferous forest

We investigated microfungal assemblages on leaf litter within a subalpine forest in central Japan and their variation with season, litter depth, and litter species. Microfungal assemblages were compared for Abies needles and Betula leaf litter collected from litter and fermentation layers of the forest floor during the growing season in spring, summer, and autumn. A total of 35 and 42 species were isolated from Abies needles and Betula leaf litter, respectively. The observed variation in microfungal assemblages was primarily attributable to seasonal differences. The frequencies of Trichoderma viride, Volutella ciliata, Mucor sp., and Umbelopsis ramanniana increased in summer, leading to a high degree of similarity of microfungal assemblages in different litter depths and litter species. Microfungal assemblages on Abies needles in spring and autumn and those on Betula leaves in spring were characterized by Trichoderma viride, V. ciliata, Thysanophora penicillioides, Trichoderma polysporum, and (or) Mortierella alpina. Microfungal assemblages on Betula leaves in autumn were characterized by the absence of these species and the occurrence of Cladosporium cladosporioides. The results were discussed with an emphasis on the role of microfungi in decomposition processes and the impact on fungi of predicted future increases in global temperature.     

Simulated herbivory advances autumn phenology in Acer rubrum

To determine the degree to which herbivory contributes to phenotypic variation in autumn phenology for deciduous trees, red maple (Acer rubrum) branches were subjected to low and high levels of simulated herbivory and surveyed at the end of the season to assess abscission and degree of autumn coloration. Overall, branches with simulated herbivory abscised ～7 % more leaves at each autumn survey date than did control branches within trees. While branches subjected to high levels of damage showed advanced phenology, abscission rates did not differ from those of undamaged branches within trees because heavy damage induced earlier leaf loss on adjacent branch nodes in this treatment. Damaged branches had greater proportions of leaf area colored than undamaged branches within trees, having twice the amount of leaf area colored at the onset of autumn and having ～16 % greater leaf area colored in late October when nearly all leaves were colored. When senescence was scored as the percent of all leaves abscised and/or colored, branches in both treatments reached peak senescence earlier than did control branches within trees: dates of 50 % senescence occurred 2.5 days earlier for low herbivory branches and 9.7 days earlier for branches with high levels of simulated damage. These advanced rates are of the same time length as reported delays in autumn senescence and advances in spring onset due to climate warming. Thus, results suggest that should insect damage increase as a consequence of climate change, it may offset a lengthening of leaf life spans in some tree species.     

Response of cowpea (Vigna unguiculata) to inoculation with VA-mycorrhizal fungi and to rock phosphate fertilization in some unsterilized Nigerian soils

Summary The effects on cowpea of inoculation with vesicular-arbuscular (VA) mycorrhizal fungi and rock phosphate (RP) fertilization were studied in pots using Alagba and Araromi series soils and in the field on Alagba, Apomu and Egbeda series soils. Inoculation of the plants with VA-mycorrhizal fungi caused very rapid infection of the roots. A higher per cent mycorrhizal infection was maintained during subsequent plant growth in the field. RP application reduced the degree of infection without affecting plant growth in the field and in pot experiments. Nodulation, nitrogen fixation and utilization of RP were increased by inoculation with mycorrhizal fungi in the pot experiments but not in the field experiments. In the pot experiments, inoculated plants supplied with RP flowered earlier, and took up more phosphorus than either inoculated plants without RP or uninoculated plants. The largest response to inoculation in terms of shoot dry matter, nodule yield and nitrogen content of shoots was obtained in Alagba soil under both pot and field conditions.IITA Journal Series Paper No. 136.     

Seasonal dynamics in a yeast population on the Oxalis acetosella L. leaves

Analysis of an epiphytic yeast population on the leaves of the evergreen common wood sorrel Oxalis acetosella L. throughout a year showed that the density and the species composition of this population underwent regular seasonal changes. There were almost no yeasts on the young spring leaves. However, the yeast population on the mature leaves tended to increase in the autumn, reaching a maximum after the formation of continuous snow cover. Then the yeast population on the leaves tended to decrease, reaching a minimum in the spring. The species diversity of the yeasts was maximum in the autumn. The population of the epiphytic yeast species Cystofilobasidium capitatum, Rhodotorula fujisanensis, Leucosporium scottii, and Cryptococcus flavus peaked in the autumn. On the other hand, the population of the widespread epiphytic species Cryptococcus laurentii on the wood sorrel leaves peaked in January. The relative abundance of the red-pigmented phytobionts Rhodotorula glutinis and Sporobolomyces roseus virtually did not change throughout the year. The relative abundance of the euribiotic species Cryptococcus albidus showed irregular monthly variations. The data obtained show that the epiphytic microbial population of various plants can be comprehensively studied only by analyzing this population throughout the vegetative period of the plants.     

Colonisation of pitcher plant leaves at several spatial scales

Abstract.  1. The effect of meso-scale (zone within bog and local plant density) and fine-scale (leaf length and resource availability) factors on the colonisation of pitcher plant leaves by arthropods was examined in an eastern Canadian bog.
2. In spring, the abundances of three arthropods, the mosquito Wyeomyia smithii , the midge Metriocnemus knabi , and the mite Sarraceniopus gibsoni , were determined for plots with low, moderate, and high densities of pitcher plants. All overwintering inhabitants were then removed from the plots. Newly opening leaves were colonised from outside the plots, and arthropod abundances were assessed again in autumn.
3. Pitcher plant fauna varied in their response to the meso-scale factors. In autumn (soon after colonisation), midges were more abundant in areas with high densities of pitcher plants. The relationship between mosquito abundance and plant density, and the variation in abundance among zones within the bog in the spring, were probably due to overwintering mortality.
4. All taxa responded to the fine-scale factors, leaf length, and capture rate, in the autumn, but the strength of the responses frequently depended on a meso-scale factor (plant density), in which responses were usually strongest where plants were sparse. Thus, the interaction between meso- and fine-scale processes needs to be considered when interpreting patterns of species abundance within arthropod assemblages in pitcher plant leaves.     

The Effect of Changes in Organic Nitrogen Concentration of Rumex obtusifolius L. on the Population Dynamics of Aphis rumicis L. (Homoptera: Aphididae)

ABSTRACT The study of plant-herbivore dynamics was carried out using Aphis rumicis L. The insect is oligophagous to the genus Rumex. It utilizes its food plant by means of phloem sucking. Here, in this study, organic nitrogen as a form of a pellet fertilizer was used as a limiting factor. The major aim of the study is to find out the tripartite relationship among soil nitrogen concentration, the plant, and the insect. Observations were made from both the natural habitat site and the manipulated field site. The result shows that Aphis rumicis responded positively to increased N concentration of leaves (p< 0.005). The application of organic nitrogen did not alter the pattern of the population dynamics of Aphis rumicis over the study period. However, it may determine the time for the optimal exploitation of resources by the insect. It is also suggested that the growth efficiency of the insect relates to the C: N ratio in the phloem.     

铲草和施肥对降香黄檀与印度檀香混交林土壤氮素矿化淋溶的影响

珍贵树种降香黄檀与印度檀香混交种植是当前华南地区人工林发展的一种重要模式.本研究设置对照(不做处理)、铲草、施肥、铲草+施肥4个处理,研究抚育措施对林地土壤净矿化速率、净硝化速率、净铵化速率和氮素淋溶速率的影响.结果表明:4个处理0~10 cm土层在春、秋季有最大净氮矿化速率,分别为18.92、18.13 mg·kg^-1·month^-1;在春、秋季有最大硝化速率,分别为20.35、18.85 mg·kg^-1·month^-1;夏、冬季有最大铵化速率,分别为0.22、0.26 mg·kg^-1·month^-1;秋季的氮素淋溶最严重,为15.98 mg·kg^-1·month^-1,全年最大淋溶为86.69 mg·kg^-1.铲草、施肥、铲草+施肥都在一定程度上抑制了土壤氮的净矿化和硝化速率,铲草、施肥、铲草+施肥处理年氮矿化量分别下降26.2%、16.1%、6.3%,年氮硝化量分别下降17.1%、16.6%、1.4%,同时也抑制了土壤铵态氮的累积.铲草、施肥、铲草+施肥处理全年氮素淋溶量依次减少25.2%、8.6%、6.1%.相对于铲草、施肥、铲草+施肥抚育措施,季节因素对土壤氮素矿化和淋溶过程的影响更显著.铲草、施肥、铲草+施肥措施在一定程度上抑制了土壤氮素硝化和铵化过程,减少了土壤氮素的矿化和淋溶损失量,有利于土壤肥力的保存和氮素的累积.     

Influence of Canopy Height and Fertilization Levels on the Resistance of Lycopersicon Hirsutum to Aculops Lycopersici (Acari: Eriophyidae)

The objective of this work was to study the effect of NK fertilization levels and canopy height on the resistance of Lycopersicon hirsutum and Lycopersicon esculentum to Aculops lycopersici (Acari: Eriophydae). The effects of NK fertilization levels and canopy height in the leaf size and density of trichomes and their effects on tridecan-2-one (2-TD) and undecan-2-one (2-UD) limiting the attack of A. lycopersici on tomato plants were assessed. Different NK fertilization levels had no effect on the resistance of L. hirsutum to A. lycopersici. No significant differences were found in attack rates of this mite on leaves of the top and median parts of L. hirsutum canopy. The type and density of trichomes were the main determining factor of A. lycopersici attack on tomato plants. High trichome densities and type VI glandular trichomes which produce tridecan-2-one are important resistance factors on tomato plants. L. hirsutum showed a high resistance level to A. lycopersici due to high densities of type VI glandular trichomes and consequently higher levels of tridecan-2-one in its leaves.     

Evidence of induced plant defences in a pondweed

SUMMARY. 1. Palatability changes, induced by herbivore damage, have received much attention in terrestrial plant-insect research but negligible investigation in freshwater systems.
2. Potamogeton coloratus (Potamogetonaceae) and the folivorous caddis larva Triaenodes bicolor (Insecta: Trichoptera) were used to duplicate bioassay techniques common in terrestrial plant-insect research.
3. Leaves of P. coloratus were artificially damaged. Grazing on damaged leaves, and undamaged leaves from damaged plants, was compared to that on control leaves from undamaged plants up to 10 days after initial damage.
4. The palatability of damaged and undamaged leaves from damaged plants declined compared to control leaves. The differences were significant at 5 days after damage, but had recovered to approximately equal palatability after 10 days.     

Accumulation and Loss of Nitrogen in White Clover (Trifolium repens L.) Plant Organs as Affected by Defoliation Regime on Two Sites in Norway

In order to improve the basis for utilising nitrogen (N) fixed by white clover (Trifolium repens L.) in northern agriculture, we studied how defoliation stress affected the N contents of major plant organs in late autumn, N losses during the winter and N accumulation in the following spring. Plants were established from stolon cuttings and transplanted to pots that were dug into the field at Apelsvoll Research Centre (60°42′ N, 10°51′ E) and at Holt Research Centre (69°40′ N, 18°56′ E) in spring 2001 and 2002. During the first growing season, the plants were totally stripped of leaves down to the stolon basis, cut at 4 cm height or left undisturbed. The plants were sampled destructively in late autumn, early spring the second year and after 6 weeks of new spring growth. The plant material was sorted into leaves, stolons and roots. Defoliation regime did not influence the total amount of leaf N harvested during and at the end of the first growing season. However, for intensively defoliated plants, the repeated leaf removal and subsequent regrowth occurred at the expense of stolon and root development and resulted in a 61–85% reduction in the total plant N present in late autumn and a 21–59% reduction in total accumulation of plant N (plant N present in autumn + previously harvested leaf N). During the winter, the net N loss from leaf tissue (N not recovered in living nor dead leaves in the spring) ranged from 57% to 74% of the N present in living leaves in the autumn, while N stored in stolons and roots was much better conserved. However, the winter loss of stolon N from severely defoliated plants (19%) was significantly larger than from leniently defoliated (12%) and non-defoliated plants (6%). Moreover, the fraction of stolon N determined as dead in the spring was 63% for severely defoliated as compared to 14% for non-defoliated plants. Accumulation in absolute terms of new leaf N during the spring was highly correlated to total plant N in early spring (R² = 0.86), but the growth rates relative to plant N present in early spring were not and, consequently, were similar for all treatments. The amount of inorganic N in the soil after snowmelt and the N uptake in plant root simulator probes (PRS^TM) during the spring were small, suggesting that microbial immobilisation, leaching and gas emissions may have been important pathways for N lost from plant tissue.     

Effect of Potassium Chloride vs. Potassium Sulphate Fertilization at Different Soil Moisture on Take-All of Wheat

A pot experiment was conducted with two moisture levels to study the influence of KC1vs. K₂SO₄ fertilization on the growth of healthy and take-all infected spring wheat.Transpiration and diffusive resistance were measured in the flag leaves. The flag leaves of healthy plants had the highest rate of transpiration.Transpiration declined with higher incidence of Gaeumannomyces grminis var. trutici (Ggt).With low soil moisture and also with sulphate fertilization transpiration tended to be lower.The difference between sulphate and chloride fertilization was relatively greater in more severely affected than in healthy plants, in particular with lower soil moisture. The flag leaves of plants were maintained in a green state for a longer period of time by chloride fertilization. This indicates a higher and prolonged metabolic activity which is reflected in the grains yields. Yield depressions as a consequence of Ggt infection were smaller with KC1 than with K₂SO₄ fertilization and more marked under water stress than with optimal water supply.