Physiological integration in the clonal perennial herb Trifolium repens L.

Summary Translocation of ¹⁴C-labelled carbohydrates between the parent stolon and branches, and among branches, of Trifolium repens plants was investigated in two glasshouse experiments to determine patterns of physiological organisation in this clonal species. Differential defoliation treatments were applied to the parent stolon and/or branches to test the sensitivity of translocation to the short-term carbon needs of defoliated sinks. Strong reciprocal exchange of carbohydrate between the parent stolon and branches was observed, with 18 41% of the ¹⁴C exported from leaves on the parent stolon moving to branches, while branches simulta-neously exported 25% (for old source branches) to 54% (for young source branches) of the ¹⁴C they assimilated to the parent plant, including translocation to other branches. Branch-to-branch translocation occurred both acropetally and basipetally. Parent-to-branch, branch-to-parent and branch-to-branch carbon fluxes all increased in response to defoliation of the sink, at the expense of carbon supply to stolon tissue or roots of the source module. Reduced export to stolon tissue of the parent axis played a major role in facilitating C reallocation from leaves on the parent stolon to defoliated branches. The observed patterns of C allocation and translocation could be adequately explained by accepted source-sink theory, and are consistent with a high degree of intra-plant physiological integration in resource supply and utilisation. This information provides mechanistic explanations for aspects of the growth dynamics and ecological interactions of T. repens in the patchy environment of a grazed pasture.     

Effect of Carbohydrate Demand on the Remobilization of Starch in Stolons and Roots of White Clover (Trifolium repens L.) after Defoliation

White clover plants were grown from stolon tips in growth cabinetsand then defoliated. Thereafter, changes in the contents ofnon-structural carbohydrates such as starch, sucrose, glucose,fructose, maltose, and pinitol in stolons and roots were monitored.Initial contents of carbohydrate reserves, photosynthetic supplyof new carbohydrates and carbohydrate demand after defoliationwere varied by growing the plants at various CO₂ partial pressures,by varying the extent of defoliation and by removing eitherroots or stolon tips at the time of defoliation. Remobilization of carbohydrate reserves in stolons increasedproportionally to their initial contents and was greater whenplants had been severely defoliated, suggesting that carbohydrateswere remobilized according to availability and demand. Starchwas the predominant reserve carbohydrate. Starch degradationwas associated with decreased contents of sucrose, glucose andfructose in young stolon parts and roots but not in old stolonparts suggesting that starch degradation was not strictly controlledby the contents of these sugars. A decrease in the demand forcarbohydrates by removal of roots did not decrease starch degradationbut increased the contents of sucrose, glucose, and fructose.Removal of stolon tips decreased starch degradation and contentsof sucrose, glucose, and fructose. The results suggest thatstarch degradation was controlled by a factor other than sucrose,glucose, and fructose which was exported from stolon tips, e.g.gibberellin. Key words: White clover, storage carbohydrates, remobilization, regrowth     

The Growth and Carbon Allocation Patterns of White Clover (Trifolium repens L.) Plants of Contrasting Branching Structure

The growth, morphology and carbon allocation patterns of F1progeny white clover (Trifolium repens L.) plants selected foreither low (‘LBF’) or high (‘HBF’) frequencyof stolon branching were compared in two controlled-environmentexperiments. Selections from within both a small-leaved (‘GrasslandsTahora’) and a large-leaved (‘Grasslands Kopu’)clover cultivar were compared, and plants were grown under arelatively lenient defoliation treatment (expt 1) or under threelevels of defoliation seventy (expt 2). Carbon allocation patternswere measured by ¹⁴CO₂ pulse-chase labelling using fully unfoldedleaves on the main (parent) stolon. LBF and HBF displayed consistent differences in the selectedcharacter though, within cultivars, the difference between selectionswas most pronounced for Kopu. The selections developed fundamentallydifferent branching structures resulting from differences inbranching frequency, with total branch weight per plant averaging122 mg for LBF and 399 mg for HBF (mean of both experiments).More C moved from parent stolon leaves to branches in HBF thanin LBF (mean 22.6% vs. 15.1% respectively of the ¹⁴C exportedfrom source leaves). More C also moved to stolon tissue in HBF,but, counterbalancing this and the difference in allocationto branches, less moved to developing leaves and roots on theparent stolon itself compared to LBF. However, the total weightof developing leaves and roots per parent stolon was generallygreater in HBF than in LBF, probably reflecting greater C importby these sinks from the higher number of branches present perplant in the former selection. HBF plants were consistentlylarger at harvest than LBF plants. There were no defoliationtreatment x selection interactions in C allocation patternsin expt 2. The implications of the results for plant performancein grazed pastures are discussed. Branching, carbon translocation, defoliation, growth, morphology, Trifolium repens, white clover     

The Effect of Defoliation on the Nitrogen Economy of White Clover: Regrowth and the Remobilization of Plant Organic Nitrogen

Single plants of white clover (Trifolium repens) were establishedfrom stolon cuttings rooted in acid-washed silver sand. Allplants were inoculated with Rhizobium trifolii, and receivednutrient solution containing 0·5 mg ¹⁵N as either ammoniumor nitrate weekly for 12 weeks (i.e. 6 mg ¹⁵N in total). Plantswere then leniently defoliated or left intact, and the labelledN supply was replaced with unlabelled N. Lenient defoliationremoved fully expanded leaves only, leaving immature leaveswhich accounted for 50–55% of the total; growing pointnumbers were not reduced. Nodules, leaves and growing pointswere counted over the following 21 d period, and d. wts, N contents,and ¹⁵N enrichments of individual plant organs were determined. Defoliated plants had fewer nodules, but numbers of growingpoints were unaffected by defoliation. The rates of both leafemergence and expansion were accelerated in defoliated plants;in consequence the number of young leaves remained less thanin intact plants until day 21. Total dry matter (DM) and N accumulationwere less in defoliated plants, and a greater proportion oftotal plant DM was invested in roots. About 97 % of plant totalN was derived from fixed atmospheric N, but there was incompletemixing of fixed and mineral N within the plant. Relatively moremineral N was incorporated into roots, whereas there was relativelymore fixed N in nodules. There was isotopic evidence that Nwas remobilized from root and stolon tissue for leaf regrowthafter defoliation; approximately 2 % of plant N turned overdaily in the 7-d period after defoliation, and this contributedabout 50% of the N increment in leaf tissue. White clover, Trifolium repens L. cv. SI84, lenient defoliation, N economy, regrowth, N remobilization     

Carbon and Nitrogen Yield, and N2 Fixation in White Clover Plants Receiving Simulated Continuous Defoliation in Controlled Environments

Single plants of white clover grown in controlled environments,and dependent for nitrogen on N, fixation, were defoliated at1 or 2 d intervals to 3, 2 and 1 expanded leaves per stolon(Expt 1), and to 1,0.5 (1 leaf on every alternate stolon) and0 expanded leaves per stolon (Expt 2), for 43–50 days Plants adapted to severe defoliation by developing much smallerleaves with a slightly reduced specific leaf area, more stolons,a smaller proportion of weight in leaf, root and nodules anda greater proportion of weight in stolons. The daily yield (materialremoved by defoliation) of d. wt and nitrogen generally decreasedwith severity of defoliation, as did the residual plant weight.However, the ‘efficiency’ of yield (daily yield/residualweight x 100) of dry matter and nitrogen was greater in themost severely defoliated treatments, attaining a maximum of5–6 % All plants adapted to the imposed defoliation regimes, howeversevere, with the result that even plants maintained withoutany fully expanded leaves invested a similar fraction of theirmetabolic resources in shoot and root as less severely defoliatedplants, and continued to grow and fix N₂, albeit at a very reducedrate of 1–2 mg Nd^–11. The energetic cost of N₂ fixation(acetylene reduction) remained constant in all treatments at31 mole CO₂ mole C₂H₄^–1, but there was some evidence thatrate of N₂ fixation per unit of nodule weight declined in themost harshly defoliated treatment. Trifolium repens, white clover, continous defolation, growth, N₂ fixation     

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.     

Regrowth of spring canola (Brassica napus) after defoliation

Aims

Regrowth of dual-purpose canola after grazing is important for commercial success and the aim of this research was to investigate the effects of defoliation on the development, growth, photosynthesis and allocation of carbohydrates.

Methods

We conducted two pot experiments in which defoliation was conducted at multiple intensities with scissors. Experiment 1 determined changes in flowering date due to defoliation while Experiment 2 investigated the effects of defoliation on growth, photosynthesis and allocation of carbohydrates in canola.

Results

Time to the appearance of the first flower was delayed by up to 9 days after the removal of all leaves at the start of stem elongation (GS30), and up to 19 days if the elongating bud was also removed. Stem growth rate decreased by 56–86 % due to defoliation and tap roots did not increase in mass when plants were completely defoliated. Leaf area continued to expand at the same rate as in un-defoliated plants. The new leaf area established per gram of regrowth biomass over 20 days was 158 cm².g^-1 for the complete defoliation treatments compared with 27 cm².g^?1 for the half-defoliated treatment and 13 cm².g^?1 for the un-defoliated treatment. Despite a reduction in total biomass of up to 60 %, the proportion of dry matter partitioned to the leaves was 18 % for all treatments within 20 days after defoliation. Total non-structural carbohydrate levels were reduced rapidly in the stem by day two (predominately sucrose) and the tap root by day four (predominately starch) after defoliation and did not recover to match un-defoliated plant levels within 20 days. Residual leaves on defoliated plants maintained photosynthetic rate compared with the same leaf cohorts on un-defoliated plants in which photosynthetic rate decreased to 39 % by day 12.

Conclusions

The rapid recovery of leaf area in defoliated canola was facilitated by the sustained high photosynthetic rate in remaining leaves, rapid mobilisation of stored sugars (stem) and starch (root), and a cessation of root and stem growth.     

Effects of defoliation and atmospheric CO2 depletion on nitrate acquisition,and exudation of organic compounds by roots of Festuca rubra

The aim of this study was to investigate the physiological basis of increased root exudation from Festuca rubra, in response to defoliation. The hypothesis, that assimilate supply to roots is a key determinant of the response of root exudation to defoliation was tested by imposing CO₂-deplete (< 50 mol mol^–1) atmospheres to F. rubra. This was done as a non-destructive means of preventing supply of new assimilate to roots of intact and defoliated plants. F. rubra was grown in axenic sand systems, with defoliation and CO₂-depletion treatments applied to plants at 14 and 35 days after planting. Root exudation and NO₃ ^– uptake were quantified throughout, and post-treatment uptake and allocation of N were determined from the distribution of ¹⁵N label, supplied as ¹⁵NO₃ ^–. Defoliation of F. rubra resulted in significantly (P <0.01) increased root exudation, CO₂-depletion did not result in increased exudation from plants of either age. When treatments were applied to F. rubra after 14 days, defoliation and CO₂-depletion each reduced NO₃ ^– uptake significantly (P <0.05). However, in older plants, uptake of NO₃ ^– was less sensitive to defoliation and CO₂-depletion. The results indicate that increased root exudation following defoliation is not related directly to reduced assimilate supply to roots. This was evident from the lack of effect of CO₂-depletion on root exudation, and the absence of correlation between root-C efflux and the rate of NO₃ ^– uptake. The physiological basis of increased exudation following defoliation remains uncertain, but may be dependent on physical damage, either directly or as a consequence of systemic responses to wounding.     

The Influence of Leaf Position and Defoliation on the Assimilation and Translocation of Carbon in White Clover (Trifolium repens L.) 1. Carbon Distribution Patterns

The assimilation of carbon (C) by, and distribution of ¹⁴C from,leaves at each end of an unbroken sequence of ten mature leaveson the main stolon of clonal plants of white clover (Trifoliumrepens L.) were measured to identify intra-plant factors determiningthe direction of C movement from leaves. Leaves at two intermediatepositions were also measured. Localized movement of ¹⁴C to sinks at the same node as, or atthe one to two nodes immediately behind, the fed leaf accountedfor 40–50% of the total ¹⁴C exported by all measured leaves.A further 50–60% of exported ¹⁴C was therefore availablefor more-distant sinks, and the direction of translocation ofthis C was determined by the relative total strength or demand(number x size x rate of activity or growth) of sinks forwardof, or behind, the leaf in question. Thus 85% of the ¹⁴C exportedfrom the youngest measured leaf moved toward the base of thestolon, while about 60% of the ¹⁴C exported from the oldestleaf moved acropetally. Defoliating plants to leave just one mature leaf on the mainstolon (at any one of the same four positions studied in undefoliatedplants), and no leaves on branches, resulted in: (1) increasednet photosynthetic rate in all residual leaves: (2) increased%export of fixed C from one of the four leaves; (3) increasedexport to the main stolon apex from all except the eldest leaf;(4) increased export to branches from three of the four leaves;and (5) decreased export to stolon tissue and roots from allleaves, within 3 d of defoliation. These responses would seemto ensure the fastest possible replacement of lost leaf areaand, thus, restoration of homeostatic growth. The observed patternsof C assimilation and distribution in both undefoliated anddefoliated white clover plants are consistent with the generalrules of source-sink theory; the distance between sources andcompeting sinks, and relative sink strength, emerge as the mostimportant intra-plant factors governing C movement. These resultsemphasize the need to consider plant morphology, and the modularnature of plant growth, when interpreting patterns of resourceallocation in clonal plants, or plant responses to stressessuch as partial defoliation. Trifolium repens L, white clover, photosynthesis, assimilate translocation, defoliation     

Impact of defoliation intensities on plant biomass,nutrient uptake and arbuscular mycorrhizal symbiosis in Lotus tenuis growing in a saline‐sodic soil

The impact of different defoliation intensities on the ability of Lotus tenuis plants to regrowth, mobilise nutrients and to associate with native AM fungi and Rhizobium in a saline‐sodic soil was investigated. After 70 days, plants were subjected to 0, 25, 50, 75 and 100% defoliation and shoot regrowth was assessed at the end of subsequent 35 days. Compared to non‐defoliated plants, low or moderate defoliation up to 75% did not affect shoot regrowth. However, 100% treatment affected shoot regrowth and the clipped plants were not able to compensate the growth attained by non‐defoliated plants. Root growth was more affected by defoliation than shoot growth. P and N concentrations in shoots and roots increased with increasing defoliation while Na⁺ concentration in shoots of non‐defoliated and moderately defoliated plants was similar. Non‐defoliated and moderately defoliated plants prevented increases of Na⁺ concentration in shoots through both reducing Na⁺ uptake and Na⁺ transport to shoots by accumulating Na⁺ in roots. At high defoliation, the salinity tolerance mechanism is altered and Na⁺ concentration in shoots was higher than in roots. Reduction in the photosynthetic capacity induced by defoliation neither changed the root length colonised by AM fungi nor arbuscular colonisation but decreased the vesicular colonisation. Spore density did not change, but hyphal density and Rhizobium nodules increased with defoliation. The strategy of the AM symbiont consists in investing most of the C resources to preferentially retain arbuscular colonisation as well as inoculum density in the soil.     

Refoliation of deciduous canopy trees following severe insect defoliation: comparison of <Emphasis Type="Italic">Fagus crenata</Emphasis> and <Emphasis Type="Italic">Quercus crispula</Emphasis>

Deciduous trees can survive severe defoliation by herbivores and often refoliate in the same season. Refoliation following severe defoliation represents compensatory regrowth to recover foliage biomass. Although the relationship between defoliation intensity and degree of refoliation at the individual level has been quantified following artificial defoliation for saplings and small trees, no study has examined the relationship for canopy trees and interspecific differences in this relationship. In this study, defoliation by gypsy moths in an outbreak year and subsequent refoliation were visually surveyed for canopy trees of Fagus crenata (n?=?80) and Quercus crispula (n?=?113) in central Japan. Defoliation and refoliation estimates were scored in 10% classes as the ratio to foliage present before defoliation. The degree of refoliation and the proportion of refoliated trees were high in severely defoliated trees. For 60 and 100% defoliated trees, respective refoliations were 2 and 66% for F. crenata, and 37 and 88% for Q. crispula. All of the 90 and 100% defoliated trees refoliated. These results indicate that severely defoliated trees show an increased need for refoliation to maintain metabolism. Beta regression analysis showed that Q. crispula possessed higher refoliation capability than F. crenata. This is likely associated with the relatively large storage reserves and recurrent growth flush pattern of oak species, which are strong characteristics of oaks and adaptive for response to herbivory and catastrophic disturbances. Interspecific differences in refoliation capability may exert differential effects on forest ecosystem processes, such as influencing the growth of understory species.     

Retranslocation of carbon reserves from the woody storage tissues into the fruit as a response to defoliation stress during the ripening period in Vitis vinifera L.

A technique for reliable labeling of the carbon reserves of the trunk and roots without labeling the current year's growth of grapevines was developed in order to study retranslocation of carbon from the perennial storage tissues into the fruit in response to defoliation stress during the ripening period. A special training system with two shoots was used: the lower one (feeding shoot) was cut back and defoliated to one single leaf (¹⁴CO₂-feeding leaf) while the other (main shoot) was topped to 12 leaves. The potted plants were placed in a water bath at 30 °C to increase root temperature and therefore their sink activity. Additionally, a cold barrier (2–4 °C) was installed at the base of the main shoot to inhibit acropetal ¹⁴C translocation. Using this method, we were able to direct labeled assimilates to trunk and roots in preference to the current year's growth. On vines with root and shoot at ambient temperature, 44% of the ¹⁴C activity was found in the main shoot 16 h after feeding whereas only 2% was found in the temperature-treated vines. At the onset of fruit ripening, and at three-week intervals thereafter until harvest, potted grapevines were fed with ¹⁴CO₂ using the temperature treatment described above. Sixteen hours after feeding, half of the vines of each group were defoliated by removing all except the two uppermost main leaves. Three weeks after each treatment, vines were destructively harvested and the dry weight and ¹⁴C incorporation determined for all plant parts. Under non-stressing conditions, there was no retranslocation of carbon reserves to support fruit maturation. Vines responded to defoliation stress by altering the natural translocation pattern and directing carbon stored in the lower parts to the fruit. In the three weeks following veraison (the inception of ripening in the grape berry), 12% of the labeled carbon reserves was translocated to the fruit of defoliated plants compared to 1.6% found in the clusters of control vines. Retranslocation from trunk and roots was highest during the middle of the ripening period, when 32% of the labeled carbon was found in the fruit compared to 0.7% in control plants. Defoliation during this period also caused major changes in dry-matter partitioning: the fruit represented 31% of total plant biomass compared to 21% measured in the control vines. Root growth was reduced by defoliation at veraison and during the ripening period. Defoliation three weeks before harvest did not affect dry matter or ¹⁴C partitioning.     

Foliar CO2photoassimilation and chloroplast linear electron transport rates in nitrogen-sufficient and nitrogen-limited soybean plants

Leaflets of soybean plants which are moderately inorganic nitrogen (N)-limited exhibit either no difference in the rate of net photosynthesis or as much as a 15–23% lower net photosynthesis rate per unit area than leaflets of N-sufficient plants [Robinson JM (1996) Photosynth Res 50: 133–148; Robinson JM (1997a) Int J Plant Sci 158: 32–43]. However, mature leaflets of N-limited soybean plants have a higher CO₂photoassimilation rate per unit chlorophyll than leaflets of N-sufficient soybean plants at both moderate light intensity (500 µmol m^-2s^-1) and saturating light intensity (1200 µmol m^-2s^-1) [Robinson JM (1996) Photosynth Res 50: 133–148]. This study was undertaken to determine whether chloroplast thylakoids isolated from the leaflets of nitrogen-limited soybean plants displayed similar or higher linear electron transport rates (H₂O ferredoxin NADP) per unit chlorophyll than thylakoids isolated from leaflets of N-sufficient plants. Chlorophyll concentration in reaction mixtures containing chloroplast thylakoids prepared from leaflets of N-limited plants was manipulated so that it was similar to the chlorophyll concentration in reaction mixtures of thylakoids prepared from leaflets of N-sufficient plants. Measurements of ferredoxin dependent, NADP dependent, O₂photo-evolution in thylakoid isolates were carried out in saturating light (1500 µmol m^-2s^-1) and with (an uncoupler) in the chloroplast reaction mixtures. Chloroplast thylakoids isolated from N-limited soybean plant leaflets routinely had a 1.5 to 1.7 times higher rate of uncoupled, whole chain electron transport per unit chlorophyll in saturating light than did chloroplast thylakoids isolated from leaflets of N-sufficient plants. The results suggest that the photosystems and photosynthetic electron transport chain components are more active per unit Chl in leaflet chloroplast thylakoids of N-limited soybean plants than in thylakoids of N-sufficient plants.     

Rapid N transport to pods and seeds in N-deficient soybean plants

Non-nodulated soybean (Glycine max (L.) Merr.) plants were cultivated hydroponically under N-sufficient (5 mM NaNO(3)) or N-deficient (0.5 mM NaNO(3)) conditions. (13)N- or (15)N- labelled nitrate was fed to the cut end of the stems, and the accumulation of nitrate-derived N in the pods, nodes and stems was compared. Real-time images of (13)N distribution in stems, petioles and pods were obtained using a Positron Emitting Tracer Imaging System for a period of 40 min. The results indicated that the radioactivity in the pods of N-deficient plants was about 10 times higher than that of N-sufficient plants, although radioactivity in the stems and nodes of N-deficient versus N-sufficient plants was not different. A similar result was obtained by supplying (15)NO(3) to cut soybean shoots for 1 h. The fact that the N translocation into the pods from NO(3) fed to the stem base was much faster in N-deficient plants may be due to the strong sink activity of the pods in N-deficient plants. Alternatively, the redistribution of N from the leaves to the pods via the phloem may be accelerated in N-deficient plants. The temporal accumulation of (13)NO(3) in nodes was suggested in both N-sufficient and N-deficient plants. In one (13)NO(3) pulse-chase experiment, radioactivity in the stem declined rapidly after transferring the shoot from the (13)NO(3) solution to non-labelled NO(3); in contrast, the radioactivity in the node declined minimally during the same time period.     

Ecophysiology of Zigadenus nuttallii,a toxic spring ephemeral in a warm season grassland: effect of defoliation and fire

Summary Zigadenus nuttallii, a highly toxic spring ephemeral in tallgrass prairie, was studied in 1985 to ascertain: 1) several ecophysiological characteristics of the species, 2) seasonal patterns of biomass accumulation, and 3) its response to defoliation and fire. The maximum photosynthetic rate of Z. nuttallii measured in unburned prairie was 13.2 moles CO₂ m^-2 s^-1 which occurred at 24–28° C and an incident quantum flux of 0.8–1.0 mmoles m^-2 s^-1. Maximum stomatal conductance measured was 5.4 mm s^-1. Early in the season, belowground storage organs (bulbs) decreased in mass and supplied much of the energy for growth of leaves, even though CO₂ uptake was possible. Buld mass did not increase until about 6 weeks after shoot emergence implying that, at this time, leaves had become a source rather than a sink for carbohydrates. The result of a single, severe defoliation event was a decrease in biomass of bulbs, leaves and reproductive structures in Z. nuttallii. Intrinsic compensatory mechanisms were not detected. In contrast, fire, which also defoliated plants, did not result in any biomass decrease at the end of the season. Improved post-fire microclimate and increased nutrient supply (extrinsic factors) may have contributed to higher photosynthetic rates and led to biomass compensation in burned prairie. These data support arguments that intrinsic compensatory mechanisms have evolved in response to chronic herbivory.     

The Effect of Lenient Defoliation on the Nitrogen Economy of White Clover: The Contribution of Mineral Nitrogen to Plant Nitrogen Accumulation During Regrowth

Single plants of white clover (Trifolium repens L.) were grownfrom stolon cuttings rooted in sand. All plants were inoculatedwith Rhizobium trifolii, and for 14 weeks received nutrientsolution containing 0.5 mg N each week, as either ammonium ornitrate. Plants were then leniently defoliated or were leftintact and a ¹⁵N-labelled N source was applied at intervalsof 4 d to replace the unlabelled N. Lement defoliation removedfully expanded leaves only; the remaining immature leaves accountedfor 39–44% of the total. At harvests over the following21 d, leaf numbers were counted and dry matter (DM), N contentsand ¹⁵N enrichments of individual plant organs were determined. Rates of leaf emergence and expansion were accelerated in defoliatedplants; numbers of young leaves were similar in defoliated andintact plants. Total DM and N content were less in defoliatedthan intact plants and were not affected by form of N supplied.DM of young leaves, growing points and stolons and N contentof young leaves were, however, greater when ammonium ratherthan nitrate N was supplied. Rates of increase in the contentof plant total N were 8.2 ± 1.36 mg N d^-1 and 10.2±1.82 mg N d^-1 in defoliated and intact plants respectively.The increases were predominantly due to N₂ fixation, since recoveryof ¹⁵N showed that less than 1% of the increment in plant totalN was assimilated mineral N. Nevertheless, the contributionof mineral N to plant total N was 50% more in defoliated thanin intact plants; higher amounts of mineral N were found particularlyin young leaves and growing points. Partitioning of mineralN to nodulated roots increased over time and was greater whenammonium rather than nitrate N was present. White clover, Trifolium repens L. cv. S184, lenient defoliation, N accumulation, N₂ fixation     

Allocation of carbon to shoots,roots, soil and rhizosphere respiration by barrel medic (Medicago truncatula) before and after defoliation

The allocation of carbon to shoots, roots, soil and rhizosphere respiration in barrel medic (Medicago truncatulaGaertn.) before and after defoliation was determined by growing plants in pots in a labelled atmosphere in a growth cabinet. Plants were grown in a ¹⁴CO₂-labelled atmosphere for 30 days, defoliated and then grown in a ¹³CO₂-labelled atmosphere for 19 days. Allocation of ¹⁴C-labelled C to shoots, roots, soil and rhizosphere respiration was determined before defoliation and the allocation of ¹⁴C and ¹³C was determined for the period after defoliation. Before defoliation, 38.4% of assimilated C was allocated below ground, whereas after defoliation it was 19.9%. Over the entire length of the experiment, the proportion of net assimilated carbon allocated below ground was 30.3%. Of this, 46% was found in the roots, 22% in the soil and 32% was recovered as rhizosphere respiration. There was no net translocation of assimilate from roots to new shoot tissue after defoliation, indicating that all new shoot growth arose from above-ground stores and newly assimilated carbon. The rate of rhizosphere respiration decreased immediately after defoliation, but after 8 days, was at comparable levels to those before defoliation. It was not until 14 days after defoliation that the amount of respiration from newly assimilated C (¹³C) exceeded that of C assimilated before defoliation (¹⁴C). This revised version was published online in June 2006 with corrections to the Cover Date.     

Changes in the Activity of Antioxidant Enzymes in Wheat Leaves and Roots as a Function of Nitrogen Source and Supply

The activity of enzymes participating in the systems of antioxidant protection was assayed in the second leaf and roots of 21-day-old wheat seedlings (Triticum aestivum L.) grown in a medium with nitrate (NO^– ₃ treatment), ammonium (NH⁺ ₄ treatment), or without nitrogen added (N-deficiency treatment). The activities of superoxide dismutase (SOD), peroxidase, ascorbate peroxidase, glutathione reductase, and catalase in the leaves and roots of the NH⁺ ₄ plants was significantly higher than in the plants grown in the nitrate medium. The activity of SOD decreased and ascorbate peroxidase markedly increased in leaves, whereas the activity of ascorbate peroxidase increased in the roots of N-deficient plants, as compared to the plants grown in nitrate and ammonium. Low-temperature incubation (5°, 12 h) differentially affected the antioxidant activity of the studied plants. Whereas leaf enzyme activities did not change in the NH⁺ ₄ plants, the activities of SOD, peroxidase, ascorbate peroxidase, and catalase markedly increased in the NO^– ₃ plants. In leaves of the N-deficient plant, the activity of SOD decreased; however, the activity of other enzymes increased. In response to temperature decrease, catalase activity increased in the roots of NO^– ₃ and NH⁺ ₄-plants, whereas in the N-deficient plants, the activity of peroxidase increased. Thus, in wheat, both nitrogen form and nitrogen deficiency changed the time-course of antioxidant enzyme activities in response to low temperature.     

Clonal plants in a spatially heterogeneous environment: effects of integration on Serengeti grassland response to defoliation and urine-hits from grazing mammals

It has been hypothesized that clonal integration between ramets inenvironments with spatially variable rates of herbivory and nutrientavailabilities leads to increased growth and fitness in the genet. An increasein genet growth could potentially influence ecosystem processes such as primaryproductivity and nutrient cycling. I tested the idea that clonal integrationwould lead to greater aboveground productivity, compensatory response todefoliation, and N re-distribution in a factorial experiment in fieldplots in the Serengeti Ecosystem, Tanzania. Each plot had either all stolonssevered or left intact, had repeated defoliation (to simulate grazing bythe African buffalo) or none, and was either located next to a plot thatreceived urea (to simulate a urine-hit) or next to a plot thatremained untreated. Plots that received stolon severing treatments had32% less peak biomass than did connected control plots, and this suggeststhat plants grew better when ramets remained connected. However, compensatoryresponse by plots to repeated defoliation was inconsistent with the hypothesizedbenefits of ramet integration at the ecosystem level: productivityresponse to defoliation was similar between plots with connected vs. severedramets (i.e. no stolon severing × defoliation interaction wasfound). When averaged across other treatments, defoliated plots hadincreased productivity compared to nondefoliated plots during the growingseason. Thus, ramet connection and defoliation did increase productivity whenthey were considered alone, but productivity response to defoliation wasunaffected by ramet connections. Urea additions, which led to a 78%increase in productivity in adjacent urine-hits, had noconsistent effect on productivity but did increase leaf percent N in adjoiningstudy plots. Thus, in the Serengeti, urine hits probably have very localizedeffects on productivity during the initial growing season.     

Induced tree response to herbivory influences the foraging preference and performance of a conifer‐defoliating sawfly

Herbivory often alters the growth and development of woody plants and can thereby render hosts less susceptible to subsequent herbivores. We carried out field surveys and experiments to investigate how previous herbivory influences adult egg lay, larval feeding preference, and associated survival of the yellowheaded spruce sawfly, Pikonema alaskensis (Rohwer) (Hymenoptera: Tenthredinidae), within crowns of black spruce, Picea mariana (Mill.) Britton et al. (Pinaceae). Pikonema alaskensis females laid nearly twice as many eggs, and late‐instar larvae consumed nearly twice as much foliar biomass, on undefoliated vs. defoliated branches. The major factor driving the lower incidence of egg lay on defoliated branches was a significant reduction in the availability of preferred size classes of shoots. In general, adult females preferred medium‐sized shoots, whereas late instars preferred large shoots; previous defoliation generally decreased the availability of medium and large shoots. In a field bioassay, late‐instar survival generally declined with increasing defoliation, which again corresponded to significant reductions in shoot length on defoliated branches. The tendency of P. alaskensis to avoid shoots on severely defoliated branches is likely to increase the time lag between herbivory and the feedback effects of induction and could thus contribute to fluctuations in sawfly population density during outbreak. Our results differ from numerous past studies of closely related diprionid sawflies in conifers, most of which have reported weak or positive effects of previous herbivory on host susceptibility; we attribute the observed responses of spruce to the unusual sawfly feeding preference of P. alaskensis for developing conifer foliage.