Defoliation of the annual herb Abutilon theophrasti: mechanisms underlying reproductive compensation

A number of studies have shown that under some conditions plants may fully or partially compensate for leaf tissue loss; however, the mechanisms underlying compensatory responses are not well understood. Previous work demonstrated that the annual herb Abutilon theophrasti fully compensated for 75% defoliation, but only when grown in the absence of stem competition. We examined potential mechanisms of compensatory response and how they are influenced by resource limitation (i.e., competition for light). Full compensation for these annual plants was defined as equal final reproductive output in defoliated and control plants. In the current study we observed substantial compensation in defoliated plants growing at low density, despite losing 75% of leaf area prior to the onset of flowering. Plant responses associated with compensation included (1) increased reproductive efficiency, which may in turn may have resulted from increased canopy light penetration and transient increases in leaf-level photosynthetic rates; (2) greater allocation to reproduction (RA); (3) changes in biomass allocation from roots to shoots; (4) lower leaf longevity, and (5) increased percent fruit set. Although some of these responses were also observed in defoliated plants grown at high density, the inability of high-density plants to compensate appeared to result from competition for light; these plants delayed reproduction and continued to produce new leaves late in the growing season after low-density, defoliated plants had shifted allocation of resources to reproduction. Received: 20 June 1996 / Accepted: 12 February 1997     

Mechanisms and traits associated with compensation for defoliation in <Emphasis Type="Italic">Ruellia nudiflora</Emphasis>

A full understanding of the ecology and evolution of plant tolerance to damage requires the measurement of a diversity of traits (including multiple fitness-correlates) and underlying mechanisms. Here, we address the compensatory response to defoliation in the perennial herb Ruellia nudiflora, measure biomass allocation patterns and relate them to compensation, and address multiple mechanisms and traits that determine compensatory ability. We used maternal full-sib lines of R. nudiflora and conducted a defoliation experiment in which half the plants of each line were subjected to removal of 40% of leaf area (the other half remained undamaged). Fitness-correlated traits, physiological traits, and leaf longevity were measured during a 2-month period after defoliation. Using another set of plants, we conducted a second defoliation experiment and measured the concentration of non-structural carbohydrates to test for root-to-shoot carbon mobilization as a compensatory mechanism. R. nudiflora showed full compensation in terms of fruit output, and compensatory ability was positively correlated with investment in root biomass in the absence of damage. In addition, defoliated plants produced shorter-lived leaves and had a greater concentration of starch in roots, suggesting that reduced leaf longevity and accumulation of below-ground carbon reserves act as compensatory mechanisms. By measuring multiple fitness-correlates and induced traits, we provide a comprehensive evaluation of R. nudiflora compensatory responses to herbivory.     

Analytical studies on the responses of sunflower (Helianthus annuus) to various defoliation treatments

The effects of defoliation treatments on plant growth in sunflower (Helianthus annuus) were studied in the field. Four defoliation treatments, 0 (control), 37.4, 56.1 and 93.4% of the total leaf dry weight, were applied to plants that had small third leaves. Decreased leaf weight/whole plant weight (F/W) ratios in defoliated plants rapidly recovered to almost the same ratio as that observed in the control within 12 to 16 days after defoliation according to the degree of defoliation. The mechanism involved in the recovery of the F/W ratio in defoliated plants mainly consisted of three parameters: enhancement of (1) carbon distribution ratios in the leaves, (2) photosynthetic activity in the remaining leaves, and (3) retranslocation of carbon from the stem and/or roots to leaves. Inhibitive effects of defoliation on relative growth rate and net assimilation rate were seen at an early stage, but subsequently both rates became larger in defoliated plants than in controls. Defoliated plants tended to show rapid development and expansion of new leaves, and to show increased specific leaf area and protein synthesis in individual leaves. The sugar content of leaves in defoliated plants was higher than that in controls, while the content in both stem and roots was lower. These responses seem to be advantageous for development of the photosynthetic system. Heights of defoliated plants were clearly depressed according to the degree of defoliation, and this was attributed largely to differences in the elongation rates of the internodes resulting from defoliation.     

The effects of high temperature on isoprene synthesis in oak leaves

Isoprene emission from plants is highly temperature sensitive and is common in forest canopy species that experience rapid leaf temperature fluctuations. Isoprene emission declines with temperature above 35 °C but the temperature at which the decline begins varies between 35 and 44 °C. This variability is caused by the rate at which leaf temperature is increased during measurement with lower temperatures associated with longer measurement cycles. To investigate this we exposed leaves of red oak (Quercus rubra L.) to temperature regimes of 35–45 °C for periods of 20–60 min. Isoprene emission increased during the first 10 min of high temperature exposure and then decreased over the next 10 min until it reached steady state. This phenomenon was common at temperatures above 35 °C but was not noticeable at temperatures below that. The response was reversible within 30 min by lowering leaf temperature to 30 °C. Because there is no storage of isoprene inside the leaf, this behaviour indicates regulation of isoprene synthesis in the leaf. We demonstrated that the variability in isoprene decline results from regulation and explains the variability in the temperature response. This is consistent with our theory that isoprene protects leaves from damage caused by rapid temperature fluctuations.     

Pattern of defoliation and its effect on photosynthesis and growth of Goldenrod

1. Leaf area was removed from Solidago altissima in either a dispersed pattern (half of every leaf removed) or a concentrated pattern (every other leaf removed) and effects on leaf gas exchange, vegetative growth and flowering were examined relative to undefoliated controls. Gas exchange was measured for leaves remaining after defoliation and for regrowth leaves that developed post-damage (at 7, 16 and 26 days post-defoliation).
2. Area-based photosynthetic rates of leaves remaining after defoliation were not affected by either dispersed or concentrated damage, but damage of both types enhanced area-based photosynthesis of regrowth leaves at 16 days post-defoliation and to a lesser extent at 26 days post-defoliation.
3. Dispersed damage, but not concentrated damage, stimulated mass-based photosynthesis of undamaged leaves remaining after defoliation. Undamaged leaves remaining after defoliation and regrowth leaves on damaged plants had higher specific leaf area (leaf area/leaf mass) than comparable leaves on control plants. Mass-based photosynthesis was more strongly elevated by defoliation than area-based photosynthesis because of this increase in specific leaf area.
4. Plants with dispersed damage recovered more quickly from defoliation; they had higher relative growth rates in the first week post-defoliation than plants with concentrated damage. Both types of defoliation caused similar reductions in flower production.
5. These results add to accumulating evidence that dispersed damage is generally less detrimental to plants than concentrated damage and suggest that physiological changes in leaves may be part of the reason.     

Stress-induced changes in carbon sources for isoprene production in Populus deltoides

Isoprene is emitted from leaves of numerous plant species and has important implications for plant metabolism and atmospheric chemistry. The ability to use stored carbon (alternative carbon sources), as opposed to recently assimilated photosynthate, for isoprene production may be important as plants routinely experience photosynthetic depression in response to environmental stress. A CO₂‐labelling study was performed and stable isotopes of carbon were used to examine the role of alternative carbon sources in isoprene production in Populus deltoides during conditions of water stress and high leaf temperature. Isotopic fractionation during isoprene production was higher in heat‐ and water‐stressed leaves (?8.5 and ?9.3‰, respectively) than in unstressed controls (?2.5 to ?3.2‰). In unstressed plants, 84–88% of the carbon in isoprene was derived from recently assimilated photosynthate. A significant shift in the isoprene carbon composition from photosynthate to alternative carbon sources was observed only under severe photosynthetic limitation (stomatal conductance < 0.05 mol m^?2 s^?1). The contribution of photosynthate to isoprene production decreased to 77 and 61% in heat‐ and water‐stressed leaves, respectively. Across water‐ and heat‐stress experiments, allocation of photosynthate was negatively correlated to the ratio of isoprene emission to photosynthesis. In water‐stressed plants, the use of alternative carbon was also related to stomatal conductance. It has been proposed that isoprene emission may be regulated by substrate availability. Thus, understanding carbon partitioning to isoprene production from multiple sources is essential for building predictive models of isoprene emission.     

Plants utilize isoprene emission as a thermotolerance mechanism

Isoprene is a volatile compound emitted from leaves of many plant species in large quantities, which has an impact on atmospheric chemistry due to its massive global emission rate (5 x 10(14) carbon g year(-1)) and its high reactivity with the OH radical, resulting in an increase in the half-life of methane. Isoprene emission is strongly induced by the increase in isoprene synthase activity in plastids at high temperature in the day time, which is regulated at its gene expression level in leaves, while the physiological meaning of isoprene emission for plants has not been clearly demonstrated. In this study, we have functionally overexpressed Populus alba isoprene synthase in Arabidopsis to observe isoprene emission from transgenic plants. A striking difference was observed when both transgenic and wild-type plants were treated with heat at 60 degrees C for 2.5 h, i.e. transformants revealed clear heat tolerance compared with the wild type. High isoprene emission and a decrease in the leaf surface temperature were observed in transgenic plants under heat stress treatment. In contrast, neither strong light nor drought treatments showed an apparent difference. These data suggest that isoprene emission plays a crucial role in a heat protection mechanism in plants.     

Biomass allocation and leaf chemical defence in defoliated seedlings of Quercus serrata with respect to carbon-nitrogen balance

BACKGROUND AND AIMS: Both nutrient availability and defoliation affect the carbon-nutrient balance in plants, which in turn influences biomass allocation (e.g. shoot-to-root ratio) and leaf chemical composition (concentration of nitrogen and secondary compounds). In this study it is questioned whether defoliation alters biomass allocation and chemical defence in a similar fashion to the response to nutrient deficiency. METHODS: Current-year seedlings of Quercus serrata were grown with or without removal of all leaves at three levels of nutrient availability. KEY RESULTS: Plant nitrogen concentration (PNC), a measure of the carbon-nutrient balance in the plant, significantly decreased immediately after defoliation because leaves had higher nitrogen concentrations than stems and roots. However, PNC recovered to levels similar to or higher than that of control plants in 3 or 6 weeks after the defoliation. Nitrogen concentration of leaves produced after defoliation was significantly higher than leaf nitrogen concentration of control leaves. Leaf mass per plant mass (leaf mass ratio, LMR) was positively correlated with PNC but the relationship was significantly different between defoliated and control plants. When compared at the same PNC, defoliated plants had a lower LMR. However, the ratio of the leaf to root tissues that were newly produced after defoliation as a function of PNC did not differ between defoliated and control plants. Defoliated plants had a significantly lower concentration of total phenolics and condensed tannins. Across defoliated and control plants, the leaf tannin concentration was negatively correlated with the leaf nitrogen concentration, suggesting that the amount of carbon-based defensive compounds was controlled by the carbon-nutrient balance at the leaf level. CONCLUSIONS: Defoliation alters biomass allocation and chemical defence through the carbon-nutrient balance at the plant and at the leaf level, respectively.     

Roles of the fructans from leaf sheaths and from the elongating leaf bases in the regrowth following defoliation of Lolium perenne L

The study of carbohydrate metabolism in perennial ryegrass (Lolium perenne L. cv. Bravo) during the first 48 h of regrowth showed that fructans from elongating leaf bases were hydrolysed first whereas fructans in mature leaf sheaths were degraded only after a lag of 1.5 h. In elongating leaf bases, the decline in fructan content occurred not only in the differentiation zone (30–60 mm from the leaf base), but also in the growth zone. Unlike other soluble carbohydrates, the net deposition rate of fructose remained positive and even rose during the first day following defoliation. The activity of fructan exohydrolase (FEH; EC 3.2.1.80) was maximal in the differentiation zone before defoliation and increased in all segments, but peaked in the growth zone after defoliation. These data strongly indicate that fructans stored in the leaf growth zone were hydrolysed and recycled in that zone to sustain the refoliation immediately after defoliation. Despite the depletion of carbohydrates, leaves of defoliated plants elongated at a significantly higher rate than those of undefoliated plants, during the first 10 h of regrowth. This can be partly attributed to the transient increase in water and nitrate deposition rate. The results are discussed in relation to defoliation tolerance. Received: 16 June 2000 / Accepted: 17 October 2000     

Isoprene synthesis in plants: lessons from a transgenic tobacco model

Isoprene is a highly reactive gas, and is emitted in such large quantities from the biosphere that it substantially affects the oxidizing potential of the atmosphere. Relatively little is known about the control of isoprene emission at the molecular level. Using transgenic tobacco lines harbouring a poplar isoprene synthase gene, we examined control of isoprene emission. Isoprene synthase required chloroplastic localization for catalytic activity, and isoprene was produced via the methyl erythritol (MEP) pathway from recently assimilated carbon. Emission patterns in transgenic tobacco plants were remarkably similar to naturally emitting plants under a wide variety of conditions. Emissions correlated with photosynthetic rates in developing and mature leaves, and with the amount of isoprene synthase protein in mature leaves. Isoprene synthase protein levels did not change under short-term increase in heat/light, despite an increase in emissions under these conditions. A robust circadian pattern could be observed in emissions from long-day plants. The data support the idea that substrate supply and changes in enzyme kinetics (rather than changes in isoprene synthase levels or post-translational regulation of activity) are the primary controls on isoprene emission in mature transgenic tobacco leaves.     

Isoprene emission from aspen leaves : influence of environment and relation to photosynthesis and photorespiration

Isoprene emission rates from quaking aspen (Populus tremuloides Michx.) leaves were measured simultaneously with photosynthesis rate, stomatal conductance, and intercellular CO₂ partial pressure. Isoprene emission required the presence of CO₂ or O₂, but not both. The light response of isoprene emission rate paralleled that of photosynthesis. Isoprene emission was inhibited by decreasing ambient O₂ from 21% to 2%, only when there was oxygen insensitive photosynthesis. Mannose (10 millimolar) fed through cut stems resulted in strong inhibition of isoprene emission rate and is interpreted as evidence that isoprene biosynthesis requires either the export of triose phosphates from the chloroplast, or the continued synthesis of ATP. Light response experiments suggest that photosynthetically generated reductant or ATP is required for isoprene biosynthesis. Isoprene biosynthesis and emission are not directly linked to glycolate production through photorespiration, contrary to previous reports. Isoprene emission rate was inhibited by above-ambient CO₂ partial pressures (640 microbar outside and 425 microbar inside the leaf). The inhibition was not due to stomatal closure. This was established by varying ambient humidity at normal and elevated CO₂ partial pressures to measure isoprene emission rates over a range of stomatal conductances. Isoprene emission rates were inhibited at elevated CO₂ despite no change in stomatal conductance. Addition of abscisic acid to the transpiration stream dramatically inhibited stomatal conductance and photosynthesis rate, with a slight increase in isoprene emission rate. Thus, isoprene emission is independent of stomatal conductance, and may occur through the cuticle. Temperature had an influence on isoprene emission rate, with the Q₁₀ being 1.8 to 2.4 between 35 and 45°C. At these high temperatures the amount of carbon lost through isoprene emission was between 2.5 and 8% of that assimilated through photosynthesis. This represents a significant carbon cost that should be taken into account in determining midsummer carbon budgets for plants that are isoprene emitters.     

Effect of water availability on tolerance of leaf damage in tall morning glory,Ipomoea purpurea

Resource availability may limit plant tolerance of herbivory. To predict the effect of differential resource availability on plant tolerance, the limiting resource model (LRM) considers which resource limits plant fitness and which resource is mostly affected by herbivore damage. We tested the effect of experimental drought on tolerance of leaf damage in Ipomoea purpurea, which is naturally exposed to both leaf damage and summer drought. To seek mechanistic explanations, we also measured several morphological, allocation and gas exchange traits. In this case, LRM predicts that tolerance would be the same in both water treatments. Plants were assigned to a combination of two water treatments (control and low water) and two damage treatments (50% defoliation and undamaged). Plants showed tolerance of leaf damage, i.e., a similar number of fruits were produced by damaged and undamaged plants, only in control water. Whereas experimental drought affected all plant traits, leaf damage caused plants to show a greater leaf trichome density and reduced shoot biomass, but only in low water. It is suggested that the reduced fitness (number of fruits) of damaged plants in low water was mediated by the differential reduction of shoot biomass, because the number of fruits per shoot biomass was similar in damaged and undamaged plants. Alternative but less likely explanations include the opposing direction of functional responses to drought and defoliation, and resource costs of the damage-induced leaf trichome density. Our results somewhat challenge the LRM predictions, but further research including field experiments is needed to validate some of the preliminary conclusions drawn.     

The regulation of isoprene emission responses to rapid leaf temperature fluctuations

Isoprene emission from leaves is temperature dependent and may protect leaves from damage at high temperatures. We measured the temperature of white oak ( Quercus alba L.) leaves at the top of the canopy. The largest short-term changes in leaf temperature were associated with changes in solar radiation. During these episodes, leaf temperature changed with a 1 min time constant, a measure of the rate of temperature change. We imposed rapid temperature fluctuations on leaves to study the effect of temperature change rate on isoprene emission. Leaf temperature changed with a 16 s time constant; isoprene responded more slowly with a 37 s time constant. This time constant was slow enough to cause a lag in isoprene emission when leaf temperature fluctuated rapidly but isoprene emission changed quickly enough to follow the large temperature changes observed in the oak canopy. This is consistent with the theory that isoprene functions to protect leaves from short periods of high temperature. Time constant analysis also revealed that there are two processes that cause isoprene emission to increase with leaf temperature. The fastest process likely reflects the influence of temperature on reaction kinetics, while the slower process may reflect the activation of an enzyme.     

Delayed induced changes in the biochemical composition of host plant leaves during an insect outbreak

In birch, Betula pubescens, herbivore-induced delayed induced resistance (DIR) of defoliated trees may cause a strong reduction in the potential fecundity of a geometrid folivore Epirrita autumnata. In this study, we examined the biochemical basis of DIR in birch leaves during a natural outbreak of E. autumnata. A set of experimental trees was defoliated at four sites by wild larvae in the peak year of the outbreak, whereas control trees were protected from defoliation by spraying with an insecticide. The biochemical composition of leaves was analysed in the following year and, although the DIR response was weak during this outbreak, causing less than a 20% reduction in the potential fecundity of E. autumnata, some consistent relationships between defoliation, biochemistry and pupal mass of E. autumnata suggested a general biochemical basis for the defoliation-induced responses in birch leaves. Total concentrations of nitrogen, sugars and acetone-insoluble residue (e.g. cell wall polysaccharides, cell-wall-bound phenolics, protein, starch, lignin and hemicellulose) were consistently lower, and total concentrations of phenolics, especially of gallotannins and soluble proanthocyanidins, were higher in the leaves of trees defoliated in the previous year than in those protected from defoliation. The capacity of tannins to precipitate proteins correlated with contents of gallotannins, and was highest in defoliated trees. The pupal mass of E. autumnata showed a strong, positive correlation with concentrations of nitrogen and sugars, and a negative correlation with the acetone-insoluble residue and gallotannins in foliage. Correlations with other measured biochemical traits were weak. The correlation coefficients between biochemical traits and pupal mass consistently had similar signs for both defoliated and insecticide–sprayed trees, suggesting that variation in leaf quality due to defoliation in the previous year was based on similar biochemical traits as variation for other reasons. We suggest that DIR is associated with reduced growth activity of leaves, and may be seen as a delay in the biochemical maturation of leaves in defoliated trees. This explains the high concentration of gallotannins in defoliated trees, a characteristic feature of young leaves. However, the lower content of nitrogen and the higher content of soluble proanthocyanidins in defoliated trees are traits usually characterising mature, not young, leaves, indicating defoliation-induced changes in chemistry in addition to modified leaf age. Our results emphasise the importance of understanding the natural changes in chemistry during leaf maturation when interpreting defoliation-induced changes in leaf biochemistry. Received: 26 January 1998 / Accepted: 10 April 1998     

Interactive effects of elevated CO2 and soil fertility on isoprene emissions from Quercus robur

The effects of global change on the emission rates of isoprene from plants are not clear. A factor that can influence the response of isoprene emission to elevated CO₂ concentrations is the availability of nutrients. Isoprene emission rate under standard conditions (leaf temperature: 30°C, photosynthetically active radiation (PAR): 1000 μmol photons m^?2 s^?1), photosynthesis, photosynthetic capacity, and leaf nitrogen (N) content were measured in Quercus robur grown in well‐ventilated greenhouses at ambient and elevated CO₂ (ambient plus 300 ppm) and two different soil fertilities. The results show that elevated CO₂ enhanced photosynthesis but leaf respiration rates were not affected by either the CO₂ or nutrient treatments. Isoprene emission rates and photosynthetic capacity were found to decrease with elevated CO₂, but an increase in nutrient availability had the converse effect. Leaf N content was significantly greater with increased nutrient availability, but unaffected by CO₂. Isoprene emission rates measured under these conditions were strongly correlated with photosynthetic capacity across the range of different treatments. This suggests that the effects of CO₂ and nutrient levels on allocation of carbon to isoprene production and emission under near‐saturating light largely depend on the effects on photosynthetic electron transport capacity.     

Effect of defoliation on concentrations of allelochemicals and soluble sugars in leaves of weeping birch (<Emphasis Type="Italic">Betula pendula</Emphasis> roth)

Effect of strong (75%) and complete (100%) artificial defoliation of weeping birch Betula pendula Roth on the dynamics of soluble sugars and phenols—flavonols, catechins, and tannins in leaves of damaged plants was investigated. Within the first 15 days after strong defoliation of birch, no changes were found in leaf contents of flavonol, catechin, and tannin. The concentration of sugars first increased but, on the 10th day after defoliation, it returned to the normal level. One year after strong defoliation, the lead concentrations of catechins and tannins in damaged trees increased, while the concentrations of flavonols and sugars did not differ from that in leaves of control trees. In two years after strong damage, the increased concentration of tannins was retained, while catechins and sugars remained at the control level. One year after complete (100%) artificial defoliation, the leaf concentrations of flavonols and sugars in damaged plants did not differ from that in control plants, while the leaf concentrations of catechins and tannins exceeded those in control plants. Two years after complete damage, the leaves contained an increased amount of tannins, whereas the amounts of catechins, flavonols, and sugars did not differ from the control levels.     

Isoprene emission by plants is affected by transmissible wound signals

Isoprene (2-methyl 1,3-butadiene) is emitted from many plants, but the signals regulating isoprene emission are unknown. Mounting leaves in a gas exchange chamber or taking small leaf punches for biochemical analysis was found to reduce the rate of isoprene emission (Loreto & Sharkey 1993). This phenomenon was investigated by putting terminal leaflets of velvet bean (Mucuna deeringeniana L.) and kudzu [Pueraria lobaia (Willd) Ohwi.] into a gas exchange chamber and monitoring isoprene emission and photosynthesis. Lateral leaflets or remote leaves were then wounded or mechanically stimulated. The rate of isoprene emission was reduced after 1 min by up to 75% by burning a lateral leaflet with a match. Even a 7 ms^?1 (25km h^?1) wind imposed on a lateral leaflet reduced isoprene emission from the terminal leaflet by 18%. Photosynthesis rates were either unaffected by these treatments or reduced more slowly than isoprene emission rates, indicating that the effect of isoprene emission rates was not a consequence of changes in photosynthetic activity. Isoprene emission from a terminal leaflet was reduced by burning leaves above and below the monitored leaflet when on the same stem. The effect was much reduced if the burned leaf (all three leaflets) was on a different stem from the monitored leaflet. Reduction of the rate of isoprene emission was observed even when the burned leaf was 52 cm distant from the measured leaflet. Increasing the distance between the stressed leaf and the monitored leaf caused the effect to be slower and smaller. It is speculated that a signal is generated by wounding which propagates through the plant at 1.3 mm s^?1. This velocity was consistent throughout the measurements and is similar to the rate of propagation of electrical signals such as action potentials and variation potentials. The effect of the environmental stress, and particularly the wind effect, can be frequent in nature and should be considered when estimating local and regional emission of isoprene for modelling atmospheric chemistry. If leaf samples used for isoprene determination are exposed to the type of stress we investigated, isoprene emission inventories based on leaf level measurements will be underestimated.     

Delayed Onset of Isoprene Emission in Developing Velvet Bean (Mucuna sp.) Leaves

Isoprene (2-methyl-1,3-butadiene) is one of the major volatile hydrocarbons emitted by plants, but its biosynthetic pathway and role in plant metabolism are unknown. Mucuna sp. (velvet bean) is an isoprene emitter, and leaf isoprene emission rate increased as much as 125-fold as leaves developed, and declined in older leaves. Net CO₂ assimilation and stomatal conductance, under different growth and environmental conditions, increased 3 to 5 days prior to an increase in isoprene emission rate, indicating that photosynthetic competence develops before significant isoprene emission occurs.     

Defoliation of alders (Alnus glutinosa) affects herbivory by leaf beetles on undamaged neighbours

The effects of defoliation of alder (Alnus glutinosa) on subsequent herbivory by alder leaf beetle (Agelastica alni) were studied in ten alder stands in northern Germany. At each site, one tree was manually defoliated (c. 20% of total foliage) to simulate herbivory. Subsequent damage by A. alni was assessed on ten alders at each site on six different dates from May to September 1994. After defoliation, herbivory by A. alni increased with distance from the defoliated tree. Laboratory experiments supported the field results. Not only leaf damage in the field, but also the extent of leaf consumption in laboratory feeding-preference tests and the number of eggs oviposited per leaf in another laboratory test were positively correlated with distance from the defoliated tree. Resistance was therefore induced not only in defoliated alders, but also in their undamaged neighbours. Consequently, defoliation of alders may trigger interplant resistance transfer, and therefore reduce herbivory in whole alder stands.     

Fractionation of Carbon Isotopes during Biogenesis of Atmospheric Isoprene

The stable carbon isotope composition of isoprene emitted from leaves of red oak (Quercus rubra L.) was measured. Isoprene was depleted in ¹³C relative to carbon recently fixed by photosynthesis. The difference in isotope composition between recently fixed carbon and emitted isoprene was independent of the isotopic composition of the source CO₂. β-Carotene, an isoprenoid plant constituent, was depleted in ¹³C relative to whole leaf carbon to the same degree as isoprene, but fatty acids were more depleted. Isoprene emitted from leaves fed abscisic acid was much less depleted in ¹³C than was isoprene emitted from unstressed leaves. We conclude that isoprene is made from an isoprenoid precursor that is derived from acetyl-CoA made from recent photosynthate. The carbon isotope composition of isoprene in the atmosphere is likely to be slightly more negative (less ¹³C) than C₃ plant material but when plants are stressed the isotopic composition could vary.