Effects of bison grazing on Andropogon gerardii and Panicum virgatum in burned and unburned tallgrass paririe

Summary Responses to clipping and bison grazing in different environmental contexts were examined in two perennial grass species, Andropogon gerardii and Panicum virgatum, on the Konza Prairie in northeastern Kansas. Grazed tillers had lower relative growth rates (RGR) than clipped tillers following defoliation but this difference was transient and final biomass was not affected by mode of defoliation. Grazed tillers of both species had higher RGR throughout the season than ungrazed tillers, resulting in exact compensation for tissue lost to defoliation. However, A. gerardii tillers which had been grazed repeatedly the previous year (1988) had reduced relative growth rates, tiller biomass and tiller survival in 1989. This suggests that the short-term increase in aboveground relative growth rates after defoliation had a cost to future plant growth and tiller survival.In general, the two species had similar responses to defoliation but their responses were altered differentially by fire. The increase in RGR following defoliation of A. gerardii was relatively greater on unburned than burned prairie, and was influenced by topographic position. P. virgatum responses to defoliation were similar in burned and unburned prairie. Thus grazing, fire, and topographical position all interact to influence tiller growth dynamics and these two species respond differently to the fire and grazing interaction. In addition, fire may interact with grazing pattern to influence a plants' grazing history and thus its long-term performance.     

Consequence of ramet defoliation on plant clonal propagation and biomass allocation: Example of five rhizomatous grassland species

We investigated the effects of ramet defoliation frequency on clonal propagation and the patterns of biomass production and allocation on five rhizomatous species (Carex divisa Hude., Eleocharis palustris L., Juncus articulatus L., Juncus gerardii Lois. and Elytrigia repens L.). Plants were grown during an 18-week experiment in greenhouse conditions. The above ground parts of ramets were clipped following three treatments: frequent (every 2 weeks), moderate (every 4 weeks) and unclipped (control). The growth of C. divisa, J. articulatus and E. repens was strongly affected by defoliation whereas E. palustris and J. gerardii maintained a similar performance when defoliated. The latter were able to compensate for the biomass loss even after six consecutive clippings. Defoliation frequency had a significant effect on total biomass production for C. divisa, J. articulatus and E. repens while J. gerardii and E. palustris maintained total biomass production. Most of the studied species showed a decrease in clonal traits when defoliated. Clipped plants displayed fewer and shorter rhizomes. Defoliation had a strong influence in biomass production with a decrease in rhizome mass in all clipped species. A greater allocation to aerial parts and a lower to rhizomes were also detected. Moderate defoliation entailed intermediate response in 1/3 of detected significant effects of defoliation on plant traits. Finally, in the experimental conditions, E. palustris and J. gerardii were the most tolerant species to defoliation, while J. articulatus was intermediate and C. divisa and E. repens had the lowest tolerance.     

Clonal integration and effects of simulated herbivory in old-field perennials

Summary We compared the growth, phenology and leaf demography of partly defoliated, connected shoots with that of partly defoliated, severed shoots in four old-field perennials (Solidago canadensis, S. altissima, S. gigantea, Aster lanceolatus) with differing genet architectures (rhizome systems), in a common garden and in the field. Our main hypothesis was that defoliation would have fewer negative effects on shoot performance if shoots were connected than if their rhizomes were severed. Since degree of clonal integration is related to differences in genet architecture, our second hypothesis was that the effects of defoliation would be less pronounced in more integrated than in less integrated clones. Removing about 50% of the total leaf area from shoots had different effects depending on plant species, shoot density, and in particular whether rhizome connections between shoots were left intact or severed. In agreement with our prediction, experimentally isolated shoots in the field or in high density clumps in the garden suffered the most from defoliation, while shoots with intact connections or in low density clumps suffered the least. Our second prediction was neither confirmed nor falsified in the present study. Solidago altissima showed overcompensation in response to simulated herbivory in the common garden, i.e. defoliated shoots grew faster and were larger at harvest than their non-defoliated neighbours.     

Phenological changes in rate of respiration and annual carbon balance in a perennial herbaceous plant, <Emphasis Type="Italic">Primula sieboldii</Emphasis>

Primula sieboldii E. Morren is a clonal herbaceous species with a short foliar period from spring to early summer. We have studied the temperature-dependence of the rate of respiration at the whole-ramet level throughout the phenological stages of P. sieboldii to reveal its photosynthate-utilization strategy. P. sieboldii ramets were grown in a chamber enabling simulation of seasonal changes in temperature. Rates of respiration were measured at three phenological stages—the foliar period, the before-chilling defoliated (BCD) period, and the after-chilling defoliated (ACD) period. In the foliar period the rate of respiration, on a biomass basis at 20°C (R ₂₀), of the above-ground plant parts was 2.5 times that of the below-ground parts. The R ₂₀ of the below-ground parts in the foliar period was 6.5 times that in the BCD period and 1.6 times that in the ACD period. Estimation of the ramet carbon balance under these growth conditions showed that ramets respired 87% of total photosynthate production during the experimental period (8.5 months). Respiratory consumption in the foliar period accounted for 70% of the yearly total, whereas 24 and 6% were consumed in the BCD and ACD periods, respectively. An extremely low rate of respiration during the long defoliated period led to a positive net annual carbon balance for P. sieboldii ramets.     

Effects of moisture availability on clonal growth in bamboo Pleioblastus maculata

The effects of moisture availability on clonal growth and biomass investment in the bamboo Pleioblastus maculata were investigated over a four-year period by transplanting Pleioblastus maculata clones into soils with different levels of moisture availability in the field. The results showed that: (1) The higher the moisture availability, the greater the total biomass of P. maculata clones. Although fewer culms are produced at the higher moisture levels, mean tiller biomass is greater. (2) Under different levels of moisture availability, obvious differences in the total rhizome length (p < 0.01), spacer length (p < 0.05) and the sizes of bamboo culms (height, p < 0.01; diameter, p < 0.01) were observed. Thus, the higher the moisture availability, the shorter the rhizomes and the larger ramets. (3) In microhabitats with low moisture availability, bamboo allocated more biomass to underground organs, which promotes elongation of rhizomes and increases root production, thereby helping to capture underground resources essential to growth. In microhabitats of high moisture availability, the biomass is primarily allocated to the aboveground growth of ramets. (4) We suggest that soil moisture availability effects the foraging strategies of bamboo, that bamboo plants growing with low moisture availability produce longer rhizomes (that is, more, although shorter, spacers) with more biomass allocation than plants in high moisture and have a better ability to forage to increase the probability of locating adequate moisture patches. Also, longer length distance between shoots (that is, longer spacers) in high soil moisture than in low is adapted to avoid intense competition from faster growing aboveground growth in high moisture patches.     

Mineral allocation to reproduction in Sorhum bicolor and Sorghum halepense in relation to parental nutrient supply

Summary This experiment investigated the effect of parental nutrient shortage on the allocation of five nutrients to seeds and rhizomes in Sorghum halepense, a perennial, noxious weed, and to seeds in Sorghum bicolor, an annual, cultivated species. Plants from both species were grown from seeds and supplied with fertilizer at three concentrations. The allocation of biomass and nutrients (N, P, K, Ca and Mg) to reproductive and vegetative parts was determined. Relative biomass allocation to reproduction (either sexual or vegetative) remained constant in S. halepense in spite of large differences in total plant weight. In S. bicolor, however, biomass allocation to sexual reproductive structures decreased significantly with decreasing nutrient supply. Individual seed weight was not modified by parental nutrient supply in S. halepense, but it increased with decreasing nutrient availability in S. bicolor. Important differences in mineral allocation to seeds were found between the two species. While S. bicolor seeds were largely buffered from the differences in parental nutrient status, concentration of nutrients in S. halepense seeds decreased significantly with decreasing supply for all the nutrients analyzed except Ca. However, mineral nutrient concentration in S. halepense rhizomes remained remarkably constant despite differences in the external supply, evincing the priority given to vegetative reproduction at the expense of sexual reproduction. Overall, the pattern of nutrient allocation in S. bicolor seeds under different nutrient supply resembled the pattern observed in S. halepense rhizomes, but it had little resemblance to the pattern of nutrient allocation in S. halepense seeds. The results are discussed in terms of differences and similarities in the reproductive strategy of these two species.     

Effects of the loss of clonal integration on four sedges that differ in ramet aggregation

Although clonal growth is a dominant mode of plant growth in wetlands, the importance of clonal integration, resource sharing among ramets, to individual ramet generations (mother and daughter) and entire clones of coexisting species has not been well investigated. This study evaluated the significance of clonal integration in four sedge species of varying ramet aggregations, from clump-forming species (Clumpers –Carex sterilis, Eleocharis rostellata), with tightly aggregated ramets (rhizomes<1cm), to runner species (Runners –Schoenoplectus acutus, Cladium mariscoides), with loosely aggregated ramets. We manipulated clonal integration by either severing connections between target mother and daughter ramets or leaving connections intact, and then planted them in an intact neighborhood of a fen in Michigan, USA. We measured growth parameters of original and newly produced ramets over two growing seasons and conducted a final biomass harvest, to address four hypotheses. First, we expected integrated clones to accumulate more biomass than severed clones. However, final clone-level biomass and ramet production were the same for both treatments in all species although severing initially stimulated ramet production by Schoenoplectus and produced a more compact ramet aggregation in Cladium. Second, we hypothesized that mother ramets would experience a cost of integration, through reduced ramet or biomass production, while daughters would experience a benefit, through increased resource availability from mothers. Mother ramets of Cladium suffered a cost from integration, while Schoenoplectus mothers suffered a slight cost and Carex daughters saw a slight benefit. Finally, we hypothesized that integration would be more active in runner species than in clumper species. Indeed, we documented more active integration in runners than clumpers, but none of the study species were dependent upon integration for growth or survival once daughter ramets were established with their own roots and shoots. This study demonstrates that integration between established ramets may not be the most important advantage to clonal growth in this wetland field site. The loss of integration elicited varied responses among coexisting species in their natural habitat, somewhat but not completely related to their growth form, suggesting that a combination of plant life history traits contributes to the dependence upon clonal integration among established ramets of clonal species.     

Growth and allocation of the arctic sedges Eriohorum angustifolium and E. vaginatum: effects of variable soil oxygen and nutrient availability

In arctic tundra soil, oxygen depletion associated with soil flooding may control plant growth either directly through anoxia or indirectly through effects on nutrient availability. This study was designed to evaluate whether plant growth and physiology of two arctic sedge species are more strongly controlled by the direct or indirect effects of decreased soil aeration. Eriophorum angustifolium and E. vaginatum, which originate from flooded and well-drained habitats, respectively, were grown in an in situ transplant garden at two levels of soil oxygen, nitrogen, and phosphorus availability over two growing seasons. In both species, N addition had a stronger effect on growth and biomass allocation than P addition or soil oxygen depletion. Net photosynthesis and carbohydrate concentrations were relatively insensitive to changes in these factors. Biomass reallocated from shoots to below-ground parts in response to limited N supply was equally divided between roots (nutrient acquisition) and perennating rhizomes (storage tissue formation) in E. angustifolium. E. Vaginatum only increased its allocation to rhizomes. In the flood-tolerant E. angustifolium, growth was improved by soil anoxia and biomass allocation among plant parts was not significantly affected. Contrary to our initial hypothesis, whole-plant growth in E. vaginatum improved in flooded soils; however, it only did so when N availability was high. Under low N availability growth in flooded soils was reduced by 20% compared to growth in the aerobic environment. Reduced biomass allocation to rhizomes and thus to storage potential under anaerobic conditions may reduce long-term survival of E. vaginatum in flooded habitats.     

Woody plant invasion of grasslands: establishment of honey mesquite (Prosopis glandulosa var.glandulosa) on sites differing in herbaceous biomass and grazing history

Summary Emergence and survival of honey mesquite (Prosopis glandulosa var.glandulosa Torr.) seedlings was quantified on sites with contrasting grazing histories: long-term continuous grazing (LTG) and long-term protection (LTP) from grazing by cattle. On each site, different levels of heroaceous defoliation were imposed at monthly intervals (no defoliation=ND, moderate=MD and heavy=HD). The two weeks following seed dissemination appeared to be the most critical toProsopis establishment on LTP-ND plots. Openings in the herbaceous layer created by moderate defoliation of grasses on the LTP site increased germination and/or survival 7-to 8-fold during this period. However, increasing the degree of defoliation from moderate to heavy did not stimulate additional emergence on either the LTP or LTG site. Emergence from scarified seed placed in cattle dung (17 to 30%) was lower than that of bare seed placements in various microhabitats (43–60%). However, deposition of scarifiedProsopis seed in dung in conjunction with graminoid defoliation may be the most likely combination of events when livestock are present. Emergence from seeds transported into grasslands by other fauna likely would be low, unless seeds were deposited in areas where grasses had been defoliated.Prosopis survival was comparably high in dung and bare seed placements after one growing season. survival of seedlings present two weeks after seed dissemination ranged from 74 to 97% at the end of the second growing season. Seedling survival and shoot development (biomass, leaf area and height) were similar on LTP and LTG sites, regardless of the level of herbaceous defoliation or seed placement. In addition, the magnitude and patterns of net photosynthesis, stomatal conductance and xylem water potential were comparable among one-year-old seedtings on ND, MD and HD plots, even though differences in herbaceous species composition and above- and below-ground biomass between these treatments were substantial. Such data suggest competition for soil resources between grasses andProsopis may be minimal early in the life cycle ofProsopis. High rates ofProsopis emergence and establishment on LTP-MD plots are counter to the widespread assumption that long-term and/or heavy grazing is requisite forProsopis encroachment into grasslands. Results are discussed with regard to factors contributing to the recent, widespread invasion of this woody legume into grasslands of southwestern North America.Abbreviations LTG long-term grazed - LTP long-term protected from grazing - ND non-defoliated - MD moderate defoliation - HD heavy defoliation     

Defoliation and below-ground herbivory in the grass Muhlenbergiaquadridentata : Effects on plant performance and on the root-feeder Phyllophaga sp. (Coleoptera, Melolonthidae)

In this study we evaluated (1) the combined effects of simulated defoliation and below-ground herbivory (BGH) on the biomass and nitrogen content of tillers and roots of the bunchgrass Muhlenbergia quadridentata and (2) the effect of defoliation on the survival of third-instar root-feeder larvae of Phyllophaga sp. The experiment was performed in a pine forest area at an altitude of 3200 m above sea level. The grass and the root-feeder species were native and dominant in the understory and in the macroarthropod root-feeder communities, respectively. Plants were established in pots in the field and were subjected to the following treatments in a factorial design: simulated defoliation (three levels) and BGH (with or without root-feeder larvae) with ten replicates per treatment. Plants were defoliated three times at 2-month intervals. The interaction between defoliation and root herbivory was significant for all components of plant biomass. In every case, light defoliation with BGH decreased live above-ground, root and total plant biomass, and the number of live tillers by more than 50% with respect to the same defoliation level without root-feeders. Plants apparently did not compensate for the carbon drain by root-feeders when a high proportion of older leaves were not removed by defoliation. Plants under heavy defoliation were not affected by the presence of root-feeders and showed a greater live/dead above-ground biomass ratio than lightly defoliated and control plants. Defoliation and BGH did not change tiller and root N concentrations but root herbivores did decrease live-tiller N content in lightly defoliated plants. Root-feeders but not defoliation decreased the root/shoot ratio by 40% and the live/dead above-ground biomass ratio by 45% through increased tiller mortality. Survivorship and final biomass of Phyllophaga sp. larvae were not affected by defoliation treatments during the 6-month study period. Received: 17 May 1996 / Accepted: 1 November 1996     

松嫩平原野古草种群构件结构动态

野古草是根茎型无性系禾草,在松嫩平原草甸经常形成单优种群落。采用单位面积挖掘取样、分株按营养繁殖世代划分龄级、根茎按实际生活年限划分龄级的方法,对松嫩平原单优群落和混生群落的野古草种群构件结构进行了调查与分析。结果表明,在生长季初期两群落野古草种群均以春性分株和根茎芽占优势,且分株及芽构件结构相对稳定,芽库的输出率单优群落为80.4%,混生群落为62.5%;整个生长季分株由2—3个龄级组成,1a分株数量是2a的2.9—10.2倍,其生物量各月份所占比例平均为93%,随着龄级的增加依次明显减少,呈增长型年龄结构;根茎由3—4个龄级组成,根茎累积长度及生物量均以2a占绝对优势,为稳定型年龄结构;分株生产力1a明显高于2a,对种群贡献最大;根茎贮藏力除个别月份以3a、4a最高外,两群落大部分以2a最高,在生长季后期,1a根茎物质积累的速率最快。     

Responses to simulated herbivory and water stress in two tropical C4 grasses

Summary The African grass Hyparrhenia rufa has established itself successfully in South American savannas (Llanos) and displaced dominant native grasses such as Trachypogon plumosus from the wetter and more fertile habitats. Several ecophysiological traits have been related to the higher competitive capacity of H. rufa. To further analyze the behavior of both species, their growth, biomass allocation, physiological and architectural responses to defoliation and water stress were compared under controlled conditions. Although total, aerial and underground biomass decreased under defoliation in both grasses, increases in clipped-leaf biomass and area compensated for defoliation in H. rufa but not in T. plumosus. This difference was due mainly to a higher proportion of assimilates being directed to leaf and tiller production and a higher leaf growth rate in the African grass as compared to T. plumosus, which showed incrased senescence under frequent defoliation. In both species, water stress ameliorated the effects of defoliation. The ability to compensate for defoliated biomass in H. rufa is possibly related to its long coevolution with large herbivores in its original African habitat and is apparently one of the causes of its success in Neotropical savannas.     

The effect of resource integration on fitness related traits in a clonal dune perennial,Hydrocotyle bonariensis

Summary The costs and benefits of resource integration in Hydrocotyle bonariensis were examined by comparing severed and intact clones grown across multiple resource gradients. Basipetal movement of water, nitrogen and photosynthates was demonstrated to occur between two rhizome branch systems interconnecting hundreds of ramets within a clone. Hydrocotyle clones of this size and larger have been shown to span highly patchy and unpredictable resource conditions in coastal dune environments. The extensive movement of water and nitrogen to portions of a clone deficient in these resources, resulted in a significant net benefit to the clone in terms of fitness-related traits: total biomass, ramet proliferation and seed production. The translocation of photosynthates across light gradients allowed for sexual reproduction and clonal expansion in the shade although this incurred a net cost to the clone in terms of ramet and seed production.     

Production and nitrogen responses of the African dwarf shrub Indigofera spinosa to defoliation and water limitation

Summary The dwarf shrub Indigofera spinosa Forsk. (Papilionacea), a native forage species of arid Northwest Kenya, was propogated from seed, grown in a controlled environment, and subjected to three treatments of defoliation and watering frequencies in a factorial experimental design. Biomass production and nitrogen accumulation in tissue components were measured to determine defoliation responses in a water-limited environment. We hypothesized that plants would maintain biomass and nitrogen flows despite removal of aboveground meristems and tissues by defoliation. Principal experimental results included a slight reduction (11%; P=0.08) of total biomass production by clipping ca. 1/3 or 2/3 of new leaves and stems and all apical meristems every month. Total aboveground production was not affected by clipping, while final root biomass was reduced 17% by the 2/3 clipping. The least water stressed plants were affected most negatively by defoliation, and the unclipped plants responded more negatively to greater water limitation. Plants achieved partial biomass compensation through alterations in shoot activity and continued allocation of photosynthate to roots. A smaller fraction of leaf production was directed to litter in clipped plants although clipping only removed the youngest tissues, suggesting that clipping increased leaf longevity. In turn, each leaf probably contributed a greater total quantity of photosynthate. Photosynthetic rates were also likely to have been increased by clipping water-stressed plants. In contrast to biomass, plants overcompensated for nitrogen lost to defoliation. Total nitrogen uptake by individual plants was stimulated by defoliation, as there was more total nitrogen in leaves and stems. Increased nitrogen uptake was achieved by clipping stimulation of total uptake per unit of root rather than of total root mass.     

Do rhizome severing and shoot defoliation affect clonal growth of Leymus chinensis at ramet population level?

Leymus chinensis (Trin.) Tzvel. is a perennial species of Gramineae, usually subject to defoliation from grazing and mowing. We examined whether shoot defoliation and rhizome severing affected rhizome and ramet growth, and vegetative bud outgrowth of L. chinensis ramet populations. We also tested the hypothesis that clonal growth of the ramets subject to defoliation would benefit from clonal integration between interconnected ramets besides from possible compensatory growth. To 48 experimental plots, we applied six treatments resulting from interactions between two rhizome connection states (unsevered/severed) and three defoliation regimes (non-defoliated, mildly-defoliated and heavily-defoliated). Defoliation affected rhizome growth and bud outgrowth, but had little effect on shoot growth. Mild and heavy defoliation exerted similar effects on rhizome growth. Only heavy defoliation significantly reduced bud outgrowth while mild defoliation did not. The fact that shoot growth did not change after defoliation and that the bud numbers remained unchanged after mild defoliation suggest that the compensatory response enable the species to tolerate grazing to some extent. Neither rhizome severing nor the interaction of rhizome severing and defoliation had effect on any tested variables. Lack of the effect of rhizome severing falsified the first half of our hypothesis, that is, clonal integration was unimportant in our experiment. The probable reasons were suspected to be the short duration of the experiment and/or the buffer effect of carbohydrate reserves in rhizomes for shoot growth and bud production in time of defoliation.     

Trade-offs among growth, clonal, and sexual reproduction in an invasive plant Spartina alterniflora responding to inundation and clonal integration

The purpose of this article was to study the trade-offs among vegetative growth, clonal, and sexual reproduction in an aquatic invasive weed Spartina alterniflora that experienced different inundation depths and clonal integration. Here, the rhizome connections between mother and daughter ramets were either severed or left intact. Subsequently, these clones were flooded with water levels of 0, 9, and 18 cm above the soil surface. Severing rhizomes decreased growth and clonal reproduction of daughter ramets, and increased those of mother ramets grown in shallow and deep water. The daughter ramets disconnected from mother ramets did not flower, while sexual reproduction of mother ramets was not affected by severing. Clonal integration only benefited the total rhizome length, rhizome biomass, and number of rhizomes of the whole clones in non-inundation conditions. Furthermore, growth and clonal reproduction of mother, daughter ramets, and the whole clone decreased with inundation depth, whereas sexual reproduction of mother ramets and the whole clones increased. We concluded that the trade-offs among growth, clonal, and sexual reproduction of S. alterniflora would be affected by inundation depth, but not by clonal integration.     

Contribution of flexible allocation priorities to herbivory tolerance in C4 perennial grasses: an evaluation with 13C labeling

The ability of plants to rapidly replace photosynthetic tissues following defoliation represents a resistance strategy referred to as herbivory tolerance. Rapid reprioritization of carbon allocation to regrowing shoots at the expense of roots following defoliation is a widely documented tolerance mechanism. An experiment was conducted in a controlled environment to test the hypothesis that herbivory-sensitive perennial grasses display less flexibility in reprioritizing carbon allocation in response to defoliation than do grasses possessing greater herbivory tolerance. An equivalent proportion of shoot biomass (60% dry weight) was removed from two C₄ perennial grasses recognized as herbivory-sensitive, Andropogon gerardii and Schizachyrium scoparium, and two C₄ perennial grasses recognized as herbivory-tolerant, Aristida purpurea and Bouteloua rigidiseta. Both defoliated and undefoliated plants were exposed to ¹³CO₂ for 30 min, five plants per species were harvested at 6, 72 and 168 h following labeling, and biomass was analyzed by isotope ratio mass spectrometry. The tallgrass, A. geraiddii, exhibited inflexible allocation priorities while the shortgrass, B. rigidiseta, exhibited flexible allocation priorities in response to defoliation which corresponded with their initial designations as herbivory-sensitive and herbivory-tolerant species, respectively. A. gerardii had the greatest percentage and concentration of ¹³C within roots and lowest percentage of ¹³C within regrowth of the four species evaluated. In contrast, B. rigidiseta had a greater percentage of ¹³C within regrowth than did A. gerardii, the greatest percentage of ¹³C within new leaves of defoliated plants, and the lowest concentration of ¹³C within roots follwing defoliation. Although both midgrasses, S. scoparium and A. purpurea, demonstrated flexible allocation priorities in response to defoliation, they were counter to those stated in the initial hypothesis. The concentration of ¹³C within new leaves of S. scoparium increased in response to a single defoliation while the percentage and concentration of ¹³C within roots was reduced. A. purpurea was the only species in which the percentate of ¹³C within new leaves decreased while the percentage of ¹³C within roots increased following defoliation. The most plausible alternative hypothesis to explain the inconsistency between the demonstrated responsiveness of allocation priorities to defoliation and the recognized herbivory resistance of S. scoparium and A. purpurea is that the relative ability of these species to avoid herbivory may make an equal or greater contribution to their overall herbivory resistance than does herbivory tolerance. Selective herbivory may contribute to S. scoparium's designation as a herbivorysensitive species even though it possesses flexible allocation priorities in response to defoliation. Alternatively, the recognized herbivory resistance of A. purpurea may be a consequence of infrequent and/or lenient herbivory associated with the expression of avoidance mechanisms, rather than the expression of tolerance mechanisms. A greater understanding of the relative contribution of tolerance and avoidance strategies of herbivory resistance are required to accurately interpret how herbivory influences plant function, competitive interactions, and species abundance in grazed communities.     

Effects of manipulation of plant carbon nutrient balance on tall goldenrod resistance to a gallmaking herbivore

Summary Ramets from six Solidago altissima clones of known resistance to the stem gallmaker Eurosta solidaginis were grown with and without supplemental nutrients. In a greenhouse experiment, mated female Eurosta were allowed to oviposit in ramets that were subsequently grown through flowering and then harvested to determine biomass allocation. Supplemental nutrients increased plant biomass but did not affect resistance to this gallmaking herbivore. This result does not conform to the plant carbon/nutrient balance hypothesis which predicts that enhanced mineral nutrition will indirectly cause a reduction in carbon-based defensive chemistry. Our results indicate a strong genetic basis to ball gallmaker resistance since modification of host phenotype did not influence susceptibility. We suggest that evaluating the degree of genotypic or environmental control of plant resistance will be especially helpful in clarifying the patterns of defensive chemical responses.     

Comparative responses of the Savanna grasses Cenchrus ciliaris and Themeda triandra to defoliation

Summary Two perennial tussock grasses of savannas were compared in a glasshouse study to determine why they differed in their ability to withstand frequent, heavy grazing; Cenchrus ciliaris is tolerant and Themeda triandra is intolerant of heavy grazing. Frequent defoliation at weekly intervals for six weeks reduced shoot biomass production over a subsequent 42 day regrowth period compared with previously undefoliated plants (infrequent) in T. triandra, but not in C. ciliaris. Leaf area of T. triandra expanded rapidly following defoliation but high initial relative growth rates of shoots were not sustained after 14 days of regrowth because of reducing light utilising efficiency of leaves. Frequently defoliated plants were slower in rate of leaf area expansion and this was associated with reduced photosynthetic capacity of newly formed leaves, lower allocation of photosynthate to leaves but not lower tiller numbers. T. triandra appears well adapted to a regime where defoliation is sufficiently infrequent to allow carbon to be fixed to replace that used in initial leaf area expansion. In contrast, C. ciliaris is better adapted to frequent defoliation than is T. triandra, because horizontally orientated nodal tillers are produced below the defoliation level. This morphological adaptation resulted in a 10-fold higher leaf area remaining after defoliation compared with similarly defoliated T. triandra, which together with the maintenance of moderate levels of light utilising efficiency, contributed to the higher leaf area and shoot weight throughout the regrowth period.     

The effect of previous defoliation of pole-stage lodgepole pine on plant chemistry,and on the growth and survival of pine beauty moth (Panolis flammea) larvae

Summary A study of the effects of defoliation by insects on the chemistry of lodgepole pine (Pinus contorta), and on the performance of Panolis flammea (Lepidoptera; Noctuidae) larvae, was carried out in a forest in northwest Scotland I year after a severe outbreak of P. flammea had caused extensive defoliation. Larval weight and survival were not significantly different on trees that had experienced different levels of defoliation in 1986. The nitrogen and tannin content of current and previous years' pine needles was not significantly affected by defoliation (although both were slightly greater in the foliage of defoliated trees). Phosphorus content of young pine foliage was lower (but not significantly lower except on one occasion) on heavily defoliated trees. On all sampling occasions, however, the nitrogen: phosphorus ratio was significantly higher on heavily defoliated trees. There were large differences in monoterpene composition of the previous year's shoots associated with defoliation intensity, but these differences had largely disappeared in the new growth. The results are discussed in relation to other studies on the effects of insect damage on plant chemistry and insect performance and in relation to the abundance of P. flammea in Scotland.